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Songs of Experience
Music and the Brain
Songs of Experience
Music and the Brain
Organizers: Isabelle Peretz (University of Montréal), Robert Zatorre (McGill University), Virginia Penhune (Concordia University), Giuliano Avanzini (Istituto Neurologico "C. Besta"), and Luisa Lopez (Università di Roma "Tor Vergata")Promoted by the Pierfranco and Luisa Mariani FoundationReported by Kathleen McGowan | Posted September 30, 2008 Overview
Music has special access to the human brain. From our first weeks of life, we have a strong sense of rhythm and an acute sensitivity to melody. As adults, music is integral to both our most basic and some of our most sophisticated cognitive processes.
At certain times, clinicians have also seen hints that music has the power to cure. Melody and rhythm sometimes activate neurological abilities that have been lost to disease or damage. Such observations suggest that music may have broader applications in therapy. In addition, observing how music molds the brain's response to injury may answer more basic questions about neuroplasticity, the ability of the nervous system to reshape and reorganize itself in response to environmental changes.
This four-day conference, titled Neurosciences and Music III—Disorders and Plasticity, was held at McGill University on June 25–28, 2008, bringing together neurophysiologists, brain imaging researchers, rehabilitation specialists, musicologists, music educators, musicians, and psychologists, among others to share their work.
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Introduction
Music has special access to the human brain. From our first weeks of life, we have a strong sense of rhythm and an acute sensitivity to melody. As adults, music is integral to both our most basic and some of our most sophisticated cognitive processes. Listening to music changes everything from the way we talk to the way we move. It also has an extraordinary ability to evoke and modulate emotions, a deep structural relationship to language, and a profound hold on memory. For scientists of the brain and of human behavior, music offers a unique window on how we comprehend and respond to the world.
Music may be a new way to engage the brain's innate ability to heal itself.
At certain times, clinicians have also seen hints that music has the power to cure. Melody and rhythm sometimes activate neurological abilities that have been lost to disease or damage. People who can no longer speak may still be able to sing. Patients who have lost neurological control of their muscles may move fluidly to a beat. These isolated observations suggest that music may have broader applications in therapy. Music may turn out to be a new way to engage the brain's innate ability to heal itself. In addition, observing how music molds the brain's response to injury may answer more basic questions about neuroplasticity, the ability of the nervous system to reshape and reorganize itself in response to environmental changes.
Music's capacity to reshape the brain was the main focus of a conference titled The Neurosciences and Music III: Disorders and Plasticity, promoted by the Pierfranco and Luisa Mariani Foundation in partnership with Brams (International Laboratory for Brain, Music, and Sound Research), McGill University, the Montreal Neurological Institute and Hospital, and the Université de Montréal, with cooperation from the New York Academy of Sciences. This four-day conference, held at McGill in Montreal, Canada on June 25–28, 2008, brought together neurophysiologists, brain imaging researchers, rehabilitation specialists, musicologists, music educators, musicians, and psychologists, among others. It will also be the focus of an upcoming volume of the Annals of the New York Academy of Sciences.
Their investigations into our ability to perceive and produce music—the basic hardware of music processing—are revealing the relationship between human musical abilities and faculties such as speech, emotion, memory, and motor control. "In music, we think in sound," said Michael H. Thaut of Colorado State University. "Music is probably a biological language of the brain, and may be a precursor to higher cognitive function."
Advances in techniques such as functional magnetic resonance imaging (fMRI), which visualizes activity within the brain with great spatial precision, diffusion tensor imaging (DTI), which depicts the strength of connections between different brain areas, and electroencephalograms (EEG) and magnetoencephalography (MEG), which track brain responses with near real-time accuracy, allow new visualization of these circuits and responses. Investigating the responses of infants, professionally trained musicians, and people with music-specific deficits such as tone-deafness provides a wealth of information about the substrates and mechanisms of musical processing. But why we are such an intensely musical species—why we feel almost universally compelled to play, listen, sing, and dance—is still an open question.
Highlights from the meeting:
- Imaging studies are revealing the regions in the brain and the neural circuitry involved in human perceptions and responses to pitch, rhythm, and timbre in music.
- Studies of individuals with absolute pitch or deficits such as tone deafness or an inability to perceive music can offer unique insights into the genetics and physiology of music processing.
- The brain has a remarkable ability to remember and recognize musical phrases. Researchers are investigating how this occurs.
- The ability to perceive and respond to music appears to be present from birth. The musical quality of infant-directed speech can make it easier to acquire language.
- Studies have linked musical training to improvements in language, mathematical ability, and spatial reasoning. The question of whether musical training causes these improvements has not been proved definitively, but researchers are looking for activity in the brain that might provide some clues.
- Perhaps paradoxically, people with autism are often very engaged with music; researchers are seeking an understanding of why.
- Using music in rehabilitation programs for victims of stroke and traumatic brain injury has shown a number of positive outcomes.
- Music may have played an important role in the evolution of language and communication in our hominin ancestors, and persisted as a method of emotional communication and a means to facilitate social bonding.
Continue reading for a more detailed report on the conference, as well as slides and audio from several of the speakers' presentations.
The Circuitry of Music
Hearing and recognizing music come naturally to most of us, but are the result of complex cognition, requiring sensitive attention to pitch variations, rhythm processing, and the ability to distinguish timbre. Neuroscientists who study music and the brain are now mapping the basic neurocircuitry of these responses—which parts of the brain respond, in what order, and through what relationships.
Sensory and motor regions of the brain have traditionally been considered as two separate systems, but evidence from the science of music suggests this distinction may be overstated. Humans, unlike most other animals, have a powerful tendency to perceive meter and synchronize, or "entrain," with it, moving our bodies in rhythm to a regular beat. We often feel unconsciously compelled, for example, to tap our toes or bob our heads in time with music. New findings suggest this has a basis in our neurology. Music automatically activates regions of the brain involved in initiating the movements necessary for singing and dancing. This coupling between auditory and motor regions is strengthened in trained musicians, suggesting that these interconnections are flexible. Joyce Chen of McGill University has done experiments using fMRI to identify the activity of various brain regions as it processes rhythms of varying complexity. Her research suggests the superior temporal gyrus, and ventral premotor and dorsal premotor cortex are all engaged by action-related sounds.
Premotor cortex responds to auditory stimuli. The superior temporal gyrus, and ventral premotor and dorsal premotor cortex all show engagement with action-related sounds.
Mapping the circuitry of music is also leading to a more sophisticated understanding of intention and action-planning. We typically imagine that when we perform an action, higher cortical "planning" regions call the shots, and direct the activity of motor regions that carry out instructions. This schematic is being replaced by a new appreciation of the role of motor and premotor regions in higher-level processing.
Researchers have shown, for example, that when we listen silently to music, the neurons that control the larynx become more active. Others have documented that when piano players passively listen to piano music, the premotor regions of their fingers are activated. Imaging studies have identified a distributed network of brain regions, including Broca's areas, the insula, and the ventrolateral premotor cortex, involved in this audiomotor translation. Research by conference co-organizer Isabelle Peretz (University of Montreal) and colleagues shows that the supplementary motor area (SMA) is also activated when subjects listen silently to familiar melodies—they are, in a sense, mentally singing along with the tune. "These regions are really important in premotor planning," said Peretz.
Similarly, cognitive interpretations of the sounds we hear influence our basic sensory perception. Our brains respond differently when we hear the downbeat on the first rather than the second of two tones, even if the stimulus is objectively identical. "Not only does sound enable us to move rhythmically, but the way we hear the beat causes us to hear differently," said John Iversen of the Neurosciences Institute.
Perception of a stable periodicity in music is constructed in the mind.
When we listen to music, we perceive a steady beat that is intrinsic to the music itself, but that stable periodicity is a percept—something we construct in our minds. It is not clear how we create this sense of rhythm and where our timing abilities arise from. Functional MRIs show activity in regions including the premotor cortex, basal ganglia, cerebellum, prefrontal cortex, and putamen. Edward Large of Florida Atlantic University proposed that the human capacity to maintain rhythms in the absence of external input may be due to "neural oscillators." He posited that these could be an emergent property of groups of neurons that create periodic processes in the brain through alternating bursts of activity. Such a mechanism could underlie our ability to perceive rhythms in very ambiguous stimuli, such as highly syncopated music.
Other findings suggest that language and music processing share many neuronal networks and functions, and may have similar organizational structures in the brain. Along these lines, Luciano Fadiga of the University of Ferrara proposed a new description of Broca's area, the left hemispheric frontal-temporal region identified more than a century ago as essential to speech. This region is not limited to speech, but responds to both music and complex sentences in humans, and its homolog in monkeys is activated while executing or perceiving precision grips. Fadiga suggested it might function at the level of representation of actions or "potential motor acts." Humans with damage to Broca's area have nonfluent aphasias, but they also have difficulty ordering photographs that show a clear chain of events, he pointed out. "The premotor nature of Broca's region cannot be neglected," he said.
Absolute Pitch and Amusia
To understand cognitive function, neuroscientists often look to the exceptions: people with exceptional abilities or deficits. Tone-deafness, a specific deficit of melodic processing, may not present a major challenge to leading a normal life, but it's useful in the lab, said Aniruddh Patel of the Neurosciences Institute in San Diego: "It's exciting for cognitive neuroscience in general, because it gives us a system to trace the path between genes and complex cognition." Similarly, those rare people who possess absolute pitch—the ability to identify a tone in isolation—can help researchers identify the neural underpinnings of pitch discrimination, and the relative contributions of genetics and training.
Untrained singers believe they cannot carry a tune, although pitch and time analyses reveal that most are actually adequate.
Congenital amusia, a term coined by Isabelle Peretz, is a developmental deficit in melodic discrimination and pitch processing that probably affects between 2% and 4% of the population. Patel, exploring how this core deficit affects speech comprehension (making it difficult, for example, to distinguish between questions and statements in English), probes how amusics discriminate melodic contours, changes in the pace and direction of pitch during a sentence. He suggested that humans' unusual sensitivity to relative pitch in comparison with other animals could indicate that a natural range of genetic variations arose and was maintained in the population over the course of evolution. The most extreme variations give rise to this specific deficit. Amusia, most commonly described in terms of the failure to perceive or reproduce pitch change, he argued, may also have several subtypes, such as rhythmic deficits.
The science of bad singing is generating many provocative findings about the mechanisms that underlie musical production. Untrained singers generally believe they cannot carry a tune, although pitch and time analyses reveal that most are actually adequate singers. For those few who truly cannot sing, the cause of failure is not obvious: Is it a lapse of vocal control, or an inability to match perception with production? Simone Dalla Bella of the Warsaw University of Finance and Management, in describing the phenotypes of bad singers, has found a variety of selective errors, but in general, poor singers are more likely to sing out of tune than out of time.
Most people can reproduce the pitch and time intervals of a tune with reasonable accuracy, but bad singers who err by more than two standard deviations when reproducing pitch tend also to make mistakes in timing.
In related research, Steven Brown of Simon Fraser University investigates singing as an example of imitative or mirroring behavior. He posits that the basic problem of poor-pitch singing is a sensorimotor mistranslation: a failure to convert auditory information into appropriate motor signaling. "Vocal pitch imitation is perhaps the consummate mirror behavior," he said. "We can measure the accuracy of pitch imitation much better than things like imitation of facial expression, hand gestures, and the like." The larynx and the region of motor cortex that controls it could be key sites for this sensorimotor integration.
Cochlear implants, as a prosthetic for both congenitally deaf children and adults who lose their hearing, transmit a relatively crude signal from the outer ear to the auditory nerve that is interpreted by the brain as sound. The implants work very well for speech perception, in part because we are so overtrained in recognizing speech, said Robert Shannon of the House Ear Institute, but not as well for music recognition and interpretation of the emotional content of speech. Experiments with implant users show that linguistic and musical processing call upon some of the same mechanisms, an insight that can could help improve both music-listening and speech-perception. "We need to systematically understand the training regimes that would enable the shared resources to be capitalized upon," said Nina Kraus of Northwestern University. Interestingly, though, children with early cochlear implants are much more engaged in musical activities and actually enjoy both listening and making music. "Their performance in singing may be poor by our standards," said Sandra Trehub of University of Toronto at Mississauga, "but surely they can appreciate music."
The other extreme of human musical performance is absolute pitch. Much research has suggested that this remarkable ability is strongly determined by genetics, but training is also key: two-thirds of adults with absolute pitch began studying music before age 6. Mathias Oechslin of the University of Zurich has found that as many as 80% of young children undergoing Suzuki training show a capacity for tone discrimination and distinctive neural responses that presage the capacity to develop absolute pitch. The finding is controversial, but points the way toward our brains' powerful ability to be fine-tuned by environmental stimuli.
Music, Memory & Learning
As anyone who can belt out the lyrics of a Barry Manilow hit will tell you, music seems to have a unique place in memory. For this reason, musical behavior provides a good way to study learning and neural change.
Even if it seems obvious that "earworms" stick in your head, however, this basic phenomenon has yet to be proven, said Matthew Schulkind of Amherst College. "If these claims could be verified empirically, it would suggest there's something special about musical memory," he said. "It would suggest we can retain information for a very long time without rehearsal," which would challenge our findings and data about how memory works.
In general, memory research has focused on visual and verbal stimuli, and memory for music is poorly understood. Some research indicates that music can have a potentiating effect on memory in dementia patients, perhaps by increasing attention and arousal. And there's no doubt we are very good at recognizing familiar tunes: findings presented by Emmanuel Bigand of the Université de Bourgogne show that minute, fragmentary passages of music of as little as 50 milliseconds each can reliably generate a feeling of recognition in trained musicians, although they may not be able to identify the melody.
Humans' skills in music recognition begin developing very early. Young babies exposed to music rapidly learn to distinguish consonant from dissonant tones, variations in timbre, and other subtle musical attributes. An fMRI study of newborns just a few days old exposed to dissonant phrases and excerpts from Scarlatti and Schubert indicates that specialized systems dedicated to musical stimuli are present even at birth, suggesting a neural predisposition for musical processing. "The newborn brain appears to be able to extract regularities from musical stimuli," said Maria Cristina Saccuman of the University Vita-Salute San Raffaele, Milan.
By the age of six months, babies are highly attuned to "infant-directed speech," speaking with exaggerated prosody. "Why do they like it so much?" queried Jenny Saffran of the University of Wisconsin, Madison. Its musical quality seems to make language acquisition easier, her research suggests, as infants learn more quickly from sung speech than from spoken speech. Richer information may actually speed learning; infants also learn melodies with lyrics more quickly than those that don't have words. "We may underestimate how infants learn," said Saffran. "The additional complexity of natural speech and music may be particularly beneficial."
Infants attended more closely to a novel list of numbers when it was sung (right) rather than spoken (left)—and this richer information may facilitate learning.
A variety of evidence suggests that musical training induces lasting changes in the brain. The auditory cortex delivers stronger and earlier responses. The regions of sensory cortex in violinists that represent the most important fingers are enlarged. Conversely, piano training reduces activation in the cerebellum in pianists. Correlational studies have also linked musical training to improvements in language, mathematical ability, and spatial reasoning.
Debate persists as to whether training actually boosts these higher cortical functions, or whether a pre-existing and perhaps genetic predisposition accounts for both musical and other skills, but the specificity of the effects of musical training suggest that experience is essential, said Laurel Trainor of McMaster University and the Rotman Research Institute. Her research has linked music training to increased activity in the gamma-band that has been connected to attention, anticipation, and feature binding of stimuli. "It looks like this response might tell us about how the auditory cortex changes through musical practice in ways that affect other areas of brain," she said.
Multisensory information integration might be a key. Volunteers who learned melodies by playing them performed better and, with MEG imaging, showed distinct electrophysiological changes associated with learning, compared to volunteers who learned the melodies by listening only, showed the University of Münster's Christo Pantev. Motor input potentiated plastic change in the auditory cortex.
Northwestern University's Patrick Wong is studying how both formal musical training and informal exposure to different types of music relate to cognitive and neural function. His experiments suggest, for example, that although it is not essential, formal musical training can make it easier for individuals to distinguish tones in foreign languages. MRI studies also suggest that extensive training in a specific instrument can produce a cortical network of expertise specifically attuned to that instrument. Such effects may be similar in untrained, everyday music listeners for whom cultural experience can frame their responses to unfamiliar music of other cultures. A study comparing Westerners and emigrant Indians to people living in rural India who have had little exposure to Western music suggests that cultural circumstances may shape some people to be "bimusical": The brains of these subjects respond similarly to musical stimuli from both cultures.
Wong argued that such studies will eventually have educational and clinical implications, as researchers gain a more complete understanding of how music can contribute to learning. He also provided some early evidence that it may be important to account for differences in auditory behaviors between cultures.
The Way You Make Me Feel: Music and Emotion
Music has a direct line to the emotions, and can even evoke contradictory emotions simultaneously. "It can induce such sad feelings, but also induce joy," said Elvira Brattico of the University of Helsinki. She explores the brain processes that lie behind emotional and aesthetic responses to music, performing brain scans on volunteers while they listen to both beloved and loathed music. "Favorite music has powerful effects on the deep structures of brain," she said.
Temple Grandin, an author with autism, has said that she doesn't love music but merely finds it pretty. But many people with autism spectrum disorders, in which emotional processing and socialization are impaired, are very interested in and engaged with music, and many with even severe disabilities develop impressive musical skills. "The idea that [people on the autism spectrum] are interested in music and have emotional deficits can be seen as a paradox," said Daniel Levitin of McGill University, one that he is exploring in his research. When presented with short excerpts that most people would have no trouble describing as happy, sad, frightening, or peaceful, people with autism and related disorders are less skilled at making these associations. They are also unable to distinguish between different levels of expressivity in music.
Pamela Heaton, by contrast, found generally good performance, although a high level of variability, in her studies of high-functioning autistics asked to associate musical excerpts with emotions. In general, people with autism might be responding to nonemotional aspects of music, as Levitin suggested, or they may find in music an alternative to interpersonal emotional experiences. Autistic people "find interpersonal relations extremely stressful," said Heaton, and this might be why they are drawn to song: "Music is a living, emotional stimulus that we don't have to interact with."
Can Music Heal the Brain?
Music sometimes has apparently miraculous effects on patients with nervous system damage. For decades, the clinical literature has documented examples of aphasia patients who, suffering from a lesion in the left frontal or superior temporal lobe, can sing fluently but have almost entirely lost the ability to speak. Other isolated reports have described people with advanced senility who have forgotten even their own family, but may be able to remember tunes and lyrics. The fact that these abilities can be readily engaged by music suggests that these functions are not entirely lost, and could potentially be restored. Understanding how music naturally brings these capacities back online in the brain could lead to new therapeutic strategies for neurodegeneration and brain damage.
Music simultaneously activates sensory and motor systems in ways that may facilitate plasticity.
Beyond these anecdotal reports, there are other reasons to believe that music could be a uniquely powerful tool in rehabilitation. Learning and listening to music simultaneously activates sensory and motor systems in ways that can facilitate the plasticity necessary to regain lost functions. Increased attention and arousal can boost recovery in a more generalized way. Music helps people re-engage socially, and, because it is a lot more fun than most rehab programs, can both improve mood and make rehabilitation more appealing.
The effects of music on aphasia suggest that language might be supported by duplicate networks in the brain. After significant left hemispheric lesions, patients can begin to recover some function through two distinct routes, said Gottfried Schlaug of Beth Israel Deaconess Medical Center and Harvard Medical School. They may learn to recruit what is left of the damaged left hemisphere. Alternatively, the right hemisphere may take over some of the specialized language ability that was limited to the left side of the brain, and music may aid in this redevelopment and retraining process.
Schlaug used a program, Melodic Intonation Therapy, in which patients begin by humming phrases while tapping the hand along to a steady beat; humming is replaced by singing and then, over eight weeks, gradually by speaking with exaggerated musical prosody. "The idea is that melodic intonation leads to more right hemispheric activation," said Schlaug. Such was the hypothesis when the technique was first described; now it can be proven. Schlaug showed several impressive videos of stroke patients making progress with this technique. DTI scans indicate growth in the arctuate fasciculus, which connects temporal and premotor regions. The results are now being tested in a randomized clinical trial.
Here, diffusion tensor imaging reflects changes in the arctuate fasciculus of patients with left hemispheric damage, after training with Melodic Intonation Therapy has lead to improvements in speech.
Stroke or neurodegenerative diseases might also be effectively treated with music. Music could act "prosthetically" for Parkinson's disease patients, suggested Jessica Grahn, providing sensory stimulation that makes it easier to engage motor mechanisms. In stroke patients, Eckart Altenmüller of the University for Music and Theater, Hannover, found that three weeks of training on the piano led to strong improvements in upper limb fine motor control. Increases in event-related desynchronization in EEG directly before movement onset provided additional evidence that this technique was changing neuronal responses. Its success is "probably based on the optimal shaping of task demands," said Altenmüller. "Music is also great because it's self-rewarding, can line up very closely with actual abilities, and provides immediate auditory feedback"—all characteristics that can assist with rehabilitation.
Rhythmic stimulus provided by a metronome as part of a training program also helped stroke and traumatic brain injury patients regain the ability to walk, inducing improvements in velocity, stride length, and cadence superior to those following conventional training methods. "There's no magic to why these people suddenly get better when they hear music," said Michael Thaut of Colorado State University. The damage has made it difficult to control positional changes at the proper velocity, and rhythmic stimulation provides a more explicit timing cue.
Musical Man
The accumulating evidence that music processing connects into so many fundamental networks and functions in the brain raises a basic question: Why? Why would something that seems so incidental to survival be so deeply rooted in the brain, and so intimately intertwined with essential functions such as speech, emotional processing, learning, and even walking? "There are not many other activities to which people are so compulsively drawn, and those that exist usually have obvious survival value," said keynote speaker Steven Mithen, an anthropologist at the School of Human and Environmental Sciences at the University of Reading.
Mithen believes music is part of our evolutionary legacy. Our hominin ancestors, he proposed, communicated via variations in rhythm and pitch. As they became increasingly dependent on their social network to hunt and survive, understanding one another's emotions and mental states would have become more and more important. Musicality, said Mithen, would have both served as a communication method and a social glue. By the time of the Neanderthals, hominins may have developed a musical proto-language, using body and voice together—a system Mithen calls "Hmmm": "It would have been singing and dancing, their musicality critical to their social lives."
Some 200,000 to 70,000 years ago, that ancestral form diverged into two forms, what we now call speech and language. Speech became the primary method of transmitting information, whereas music remained a method of emotional communication and a means to facilitate social bonding. "Music was essential to the survival of our stone-age ancestors, and we have inherited the compulsion to engage in music," he said.
Mithen's persuasive argument is that music tells the story of where we came from. But the diverse research presented at this conference also suggests that music may point the way toward what we could be. Studying how music works in the brain might permit us to develop the treatments and therapies that could heal the sick—and to finally answer the question of who we truly are.
Open Questions
Why does musical training improve linguistic, mathematical, and spatial reasoning performance?
Does musical training improve attention and memory, and if so, by what cortical mechanisms?
What brain areas are activated during rhythm perceptions?
What neural computations give rise to rhythm perception?
What is a "beat," cognitively speaking? Does it have to do with expectancy, or attention?
Is bad singing a failure of vocal control, or an inability to match perception with production?
What are the neural mechanisms of out-of-tune singing?
Is a neural predisposition for processing music present at birth?
Is memory for music different than memory for other types of information?
Can music potentiate memory?
Why is infant-directed speech so compelling to babies, and does it promote language acquisition?
How and where is multisensory information processed in the brain?
Is absolute pitch a rare, genetically endowed trait, or the reproducible product of early training?
Are there cultural differences in how we process music, based on exposure?
Can we localize the musical lexicon in brain?
Why are very young babies so good at distinguishing music, when their auditory cortex is immature?
What is the purpose of "mirror" mechanisms in the brain?
Why are people with autism deficient in emotional processing yet intensely interested in and engaged with music?
How can people with cochlear implants perceive speech readily, even though they get only crude auditory information?
Why do humans love music?
Keynote Presentation
The Music Instinct: The Evolutionary History of Human Musical and Linguistic Ability
Speaker: Steven Mithen
School of Human and Environmental Sciences, University of Reading, UK
Why music? The enjoyment of music is as widespread among humans living today as the enjoyment of food, the use of language and any other trait that is recognised as universal. Music appears to be part of our biological constitution rather than a mere consequence of cultural history.
And yet music has been largely neglected by palaeoanthropologists. This is perhaps not surprising as the past is silent—how can we know what type of musical capacities were possessed by our hominin ancestors? However difficult, it is nevertheless essential to address the evolution of human musical ability; unless we do so, we are neglecting a fundamental part of the human condition, because we are a musical species, and music provides one of the key mechanisms for cultural transmission.
New research in the study of hominin fossils, archaeological remains as well as in the fields of neuroscience, developmental psychology and musicology are allowing to attain new insights into the evolutionary history of musicality. These suggest that communication by variations in pitch and rhythm most likely preceded that of speech, and may indeed have provided the foundation from which spoken language evolved.
This lecture reviews these developments and suggests how they can be pieced together in order to begin to gain some understanding of the musicality of human ancestors and how our current musical and linguistic abilities evolved.
Rhythms in the Brain: Basic Science and Clinical Perspectives
Coordinators: Joel S. Snyder, Virginia Penhune, and Edward W. Large
Introduction
Rhythm, along with pitch, is one of the fundamental dimensions of musical experience. Like pitch, rhythm encompasses a number of psychological phenomena (e.g., accents, beat, meter, and grouping) that are related to, but are not the same as, the physical dimensions of music (e.g., intensity variation and sequences of durations). Unlike pitch, which likely arises from place and temporal coding mechanisms in the peripheral and central auditory systems, it is less clear how or where rhythm is coded in terms of the anatomy or physiology of the auditory system. Furthermore, given that human timing abilities exist in other sensory modalities as well as in motor production, it is also unclear whether the fundamental mechanisms of musical rhythm processing occur strictly in auditory brain areas. Until recently, most of what we understood about rhythm perception and production came from studies of human behavior and from computational modeling. Lately, the tools of cognitive neuroscience have been applied to rhythm processing, which has provided new insights into where and how rhythm is coded in the human brain.
This symposium consists of speakers who have been leading this recent effort to understand the neural underpinnings of rhythmic behavior. Joyce Chen presents functional magnetic resonance imaging (fMRI) data showing what cortical brain areas are active and interact during perception and production of metrical patterns, particularly highlighting the importance of the premotor cortex. Jessica Grahn presents behavioral data from Parkinson's disease patients and fMRI data from healthy individuals, showing converging evidence for the importance of both cortical and subcortical brain areas in rhythm processing. Edward Large presents findings from studies using fMRI and electroencephalography (EEG) that reveal basic mechanisms of rhythmic processing and how these mechanisms inform theories of rhythmic expectancy and attention. Finally, John Iversen presents magnetoencephalography (MEG) data showing the effect of metrical interpretation on neural correlates of rhythm perception.
The Role of the Premotor Cortex in Sensorimotor Transformations for Music Production
Speaker: Joyce L. Chen*
BRAMS, McGill University, Montreal
The interplay between sounds and movements is not only important for music performance, but also for the acquisition of speech. Furthermore, these interactions between the auditory and motor systems may form the neural basis that underlies the use of music as a therapeutic tool in the facilitation of movements.
We performed a series of functional magnetic resonance imaging studies to examine the perception of, and synchronization to, musical rhythms, with the aim of elucidating the neural substrates mediating auditory-motor interactions. In studies 1 and 2, subjects tapped along with rhythms that parametrically varied in metric saliency and complexity, respectively. Study 3 probed the response of the motor system during actionperception coupling and decoupling: subjects listened passively, listened with anticipation to commit a motor act, and tapped along with musical rhythms. Three main results were found. First, the findings suggest that the pre-supplementary motor area, supplementary motor area and cerebellum may be engaged in the sequencing and timing of rhythmic actions. Second, we found that musicians additionally engage the prefrontal cortex, which may be related to their superior ability to deconstruct and organize musical rhythm in terms of working memory processes. Third and most important, our results reveal that the posterior superior temporal gyrus and the premotor cortex may be critical nodes for mediating the transformation of auditory information into motor actions. Specifically, the findings indicate a dissociation of function within ventral and dorsal components of the premotor system. Listening to and performing music may entail activation of motor programs associated with producing the music, enabled via auditory links with the ventral premotor cortex, whereas the dorsal premotor cortex may extract higher-order information from an auditory rhythm, such as its metrical structure, in order to implement temporally organized movements. We propose that this auditory-premotor circuit may be at the core of what links music, movement and language together, and may be a possible rationale for why auditory cues can be so effective in improving movements in those with motor disorders.
*Coauthors: Virginia B. Penhune, BRAMS, Concordia University, Montreal; and Robert J. Zatorre, BRAMS, McGill University, Montreal Neurological Institute.
The Role of the Basal Ganglia in Rhythm Processing: Evidence from Neuropsychology and Neuroimaging
Speaker: Jessica A. Grahn
Medical Research Council, Cognition and Brain Sciences Unit, Cambridge, UK
Perception of auditory rhythms activates brain areas typically associated with movement (premotor cortex / PMC, supplementary motor area / SMA, cerebellum, and basal ganglia), in addition to auditory areas (Schubotz, Friederici et al., 2000). The basal ganglia are preferentially activated during perception of rhythms with a regular beat (Grahn & Brett, 2007), but their necessity for beat-based rhythm processing has not been proven.
Patients with Parkinson's disease (PD) and controls were tested on a rhythm discrimination task to determine if basal ganglia dysfunction results in an impairment of processing rhythms that have a beat. In the control group, a significant advantage was observed for discrimination of rhythms with a beat compared to those without a beat. This advantage was significantly reduced in the PD group. Discrimination of rhythms without a beat, however, was not significantly different between the two groups, resulting in a group by rhythm-type interaction. This suggests that the basal ganglia are part of a system involved in detecting or generating a beat, and that this system is compromised in patients with Parkinson's disease.
An additional study was conducted to examine how the basal ganglia interact with other neural areas during beat perception, and if musical experience-dependent plasticity altered these interactions. fMRI was used to measure activation in musicians and non-musicians during perception of rhythms with and without a beat. The beat occurred in one of two ways: either explicitly emphasized with volume accents, or internally generated in accordance with duration accents in the rhythm (Povel & Okkerman, 1981; Povel & Essens, 1985; Patel, Iversen et al. 2005; Grahn & Brett, 2007).
Functional connectivity analyses showed increased coupling (for both musicians and non-musicians) between the basal ganglia and bilateral SMA, PMC, and superior temporal gyri (STG) during the beat conditions compared to the non-beat conditions. However, comparison within the two beat conditions revealed that the coupling of cortical motor (SMA and left PMC) and auditory (STG) areas was greater during duration-accented than volume accented conditions. More coupling between these cortical areas was observed for those with more musical experience, perhaps indicating experience-dependent plasticity in the cortical dynamics of the rhythm processing network.
References
Grahn, J. A. & Brett, M. (2007) Rhythm perception in motor areas of
the brain. Journal of Cognitive Neuroscience 19: 893-906.
Patel, A. D., Iversen, J. R. et al. (2005) The influence of metricality and modality on synchronization with a beat. Experimental Brain Research 163: 226-238.
Povel, D.-J. & Okkerman, H. (1981) Accents in equitone sequences. Perception & Psychophysics 30: 565-572.
Povel, D. J. & Essens, P. J. (1985) Perception of temporal patterns. Music Perception 2: 411-440.
Schubotz, R. I., Friederici, A. D. et al. (2000) Time perception and motor timing: a common cortical and subcortical basis revealed by fMRI. NeuroImage 11: 1-12.
Neural Correlates of Rhythmic Expectancy and Dynamic Attending
Speaker: Edward W. Large*
Florida Atlantic University, Boca Raton
The sounds that humans use for communication may be thought of as complex, temporally structured sequences of approximately discrete events, such as musical notes and speech syllables. At sequence time scales (i.e. hundreds of milliseconds), temporal patterning contributes to the perception of rhythmic attributes, including pulse and meter. Nonlinear oscillation is ubiquitous in neural systems, and over the past several years neural oscillation has been proposed as a fundamental mechanism of rhythm perception and attention.
We show how universal properties of nonlinear oscillation predict psychological properties of pulse and meter. We review recent evidence linking pulse and meter with dynamic allocation of attention to complex event sequences, and we discuss physiological evidence linking rhythm perception to neural dynamics.
*Coauthor: Joel S. Snyder, Department of Psychology, University of Nevada, Las Vegas.
The Effect of Metrical Interpretation on Brain Responses to Complex Rhythms
Speaker: John R. Iversen
The Neurosciences Institute, San Diego
The perceptual experience of a simple rhythm depends upon its metrical interpretation. Prior work has shown that brain activity reflects a listener's metrical interpretation of simple ambiguous rhythms: neural response to a note coinciding with the listener's internal sense of the beat is enhanced, specifically in the beta frequency range (20–30Hz). Additionally, the increase in beta due to the imagining of a beat resembles the increase in beta due to a physical accent, suggesting that the increase in beta may also reflect subjective accentuation. We will discuss recent work studying brain responses to more elaborate rhythms that vary in their perceived syncopation depending on the mental placement of the downbeat (which will be experimentally manipulated). We ask if the increase in transient neural activity in the beta range is a marker of the beat in these more complex rhythms, and consider if it can indicate internal beat placement even in absence of a physical input, as in syncopated patterns in which some beats are never marked by physical tones.
Normal and Impaired Singing
Coordinators: Simone Dalla Bella and Steven Brown
Introduction
A majority of people think that they sing poorly, even though many of them do not. Anecdotal evidence abounds regarding "tone deafness" or the tendency to sing out-of-tune. But what are the mechanisms that lead people to sing poorly? Research has focused on an analysis of both production and perception abilities, with an aim at elucidating the relationship between the two. Among production tasks, studies have examined both the singing of familiar songs from long-term memory and the imitation of novel melodic sequences from short-term memory. The goal of this symposium is to present recent evidence on poor singing in healthy individuals. Systematic research on poor singing in the general population is scant. However, recent studies have revealed that poor singing is unlikely to be a monolithic deficit, but is instead characterized by a diversity of singing "phenotypes", some of which involve impaired perceptual abilities (e.g., congenital amusia) and some of which do not. The speakers in this symposium describe the forms of poor singing thus far characterized, and attempt to explain the underlying cognitive and neural mechanisms. Implications for singing development are discussed as well.
In particular, Simone Dalla Bella describes his studies about performance on musical tasks, in a population of non-musicians, and the issue of poor singing with or without perceptual impairments. Steve Brown then carries the subject further: whether poor singing is related to a mismatch between correctly perceived tones and intended phonatory targets, thus putting the area of the larynx at the core of poor performance in this population.
Singing Proficiency in the Majority: Pitch and Time Accuracy Revealed by Acoustical Analysis
Speaker: Simone Dalla Bella
Department of Cognitive Psychology, University of Finance and Management, Warsaw
Acoustical methods can provide a reliable and objective estimate of singing proficiency in the general population, in terms of pitch and temporal accuracy (e.g., Dalla Bella, Giguère & Peretz, 2007). When asked to sing a well-known song (e.g., "Happy Birthday") non-musicians are as proficient as professional singers, at least when performing at a slow tempo. However, some of them exhibit poor singing, mostly limited to the pitch domain, and sometimes in the absence of perceptual deficits (see Pfordresher & Brown, 2007). More recently, singing proficiency has been thoroughly examined with tasks extending beyond singing familiar melodies. Forty participants without musical training were asked to repeat single pitches (Task 1), intervals (Task 2), and short melodies (Task 3). In addition, participants had to sing three well-known melodies (e.g. "Jingle Bells") at a natural tempo (Task 4) and at a fixed, slow tempo (Task 5). Additional tasks were performed in order to assess participants temporal processing (i.e., tapping tasks) and perception (i.e., Montreal Battery of the Evaluation of Amusia, MBEA). Each performance was analyzed with an acoustically based method, yielding objective measures of pitch and time accuracy. Results revealed that the majority could sing in tune and in time, thus confirming previous findings. Poor singers were mostly impaired in the pitch domain. Still, various patterns of poor singing emerged from the analyses of pitch and time accuracy (e.g., poor pitch singing with or without perceptual deficits). The results of previous and more recent studies on singing proficiency in non-musicians using acoustical methods are reviewed. Particular attention is paid to cases exhibiting poor singing, in order to elucidate the nature of their deficits. The relationship between perception and performance mechanisms in vocal performance is discussed.
Brain Areas Involved in Poor-pitch Singing
Speaker: Steven Brown
Department of Psychology, Simon Fraser University, Burnaby, BC
Cognitive studies carried out in collaboration with Peter Pfordresher (University at Buffalo) have shown that out-of-tune singing cannot be attributed to either perceptual or motor deficits. This has been shown by poor performance of non-musician subjects on imitative vocalization tasks in the absence of deficits in pitch discrimination. Hence, we favour a model in which poor singing in healthy individuals is due to a sensorimotor deficit, namely mistranslation between perceived pitches and intended phonatory targets. We are currently carrying out a functional magnetic resonance imaging (fMRI) study of vocal imitation, comparing the brain activity of accurate singers and poorpitch (tone-deaf) singers. Preliminary analysis of the imaging data shows that poor-pitch singers have normal levels of activation in auditory areas but deficient or anomalous activation in motor areas, including the larynx motor cortex, posterior cerebellum, and supplementary motor area.
The normal levels of auditory activation are consistent with our cognitive findings of unimpaired pitch discrimination. At the motor level, the greatest difference between good and bad singers is found in the larynx motor cortex. Given that several studies of auditory perception have shown activation in this region, the larynx area might turn out to be a key site for audiovocal integration, including vocal imitation. Hence, poor-pitch singing during imitative tasks might be due to some kind of disconnection or misconnection between auditory areas in the temporal lobe and phonatory areas in the larynx motor cortex.
Music Training and Induced Cortical Plasticity
Coordinator: Christo Pantev
Introduction
During the last years, music has increasingly been used as a tool for the investigation of human cognition and its underlying brain mechanisms. Music relates to many brain functions like perception, action, cognition, emotion, learning and memory and therefore music is an ideal tool to investigate how the human brain is working and how different brain functions interact. Novel findings have been obtained in the field of induced cortical plasticity by musical training.
The positive effects which music in its various forms has in the healthy human brain are not only important in the frame work of basic neuroscience, but they also strongly affect the practices in neurorehabilitation. Laurel Trainor reports on research on the effects of exposure to music in young age. Without formal training, children learn the differences between consonance and dissonance and learn what pitches are typical of their culture. The authors then used MEG measurements to explore the possibilities of specific learning processes both in the music and cognitive domains, when children are exposed to formal training. Christo Pantev in his presentation addresses the issue of plastic changes in the brain induces by performance vs listening training, as measured by Mismatch Negativity (MMN). The following lecture by Jäncke uses MMN to test whether musical training can enhance absolute pitch abilities and, in a second study, whether absolute pitch possessors have differences in white matter, as revealed by DTI. Mari Tervaniemi's lecture reviews the evidence available about the various neurocognitive profiles that different musicians display, given that musicians are usually referred to as homogeneous population.
Finally, Patrick Wong explores the impact of short- and long-term asymmetric linguistic and musical experiences on how our nervous system responds to complex sounds.
Domain-specific and Domain-general Effects of Musical Experience in Young Children
Speaker: Laurel J. Trainor*
Rotman Research Institute, Toronto, and McMaster University, Hamilton, ON
Musical experience in the form of lessons and daily practice has been shown to have specific effects on musical processing in older children and adults, as well as general effects on all cognitive domains tested by IQ measures. However, very few studies have directly examined the effects of musical lessons on various aspects of processing in preschool children.
We have demonstrated that when young children are exposed to a musical environment, as they are in everyday life, but not engaged in any formal musical training, they acquire sensitivity to various aspects of musical pitch structure in a logical order over a period of years.
Specifically, infants are initially sensitive to consonance and dissonance, young children readily learn what notes belong in the scales of the music of their culture, but children do not demonstrate sensitivity to implied harmony until about 6 years of age. The question of most interest here is whether sensitivity to harmonic structure can be learned at a younger age through specific training. We compared children between 3 and 5 years of age who were engaged in music lessons with those who were not (the children were matched in socioeconomic status and engagement in extracurricular activities).
While both groups showed thorough knowledge about what notes belong in a key, as would be expected from our previous data, those with musical backgrounds showed superior knowledge of harmonic structure.
Thus, we have shown that it is possible to develop sensitivity to harmony at an early age, and that musical training likely affects this process. We are continuing to follow these children to see whether they diverge further with additional musical training. The effects of musical training on other cognitive domains are more surprising, and the mechanism for such transfer is less clear. We are investigating the hypothesis that engaging in music lessons promotes more exercise in executive functioning and attentional processing in young children. Using magnetoencephalography (MEG), we measured MEG responses to musical tones in young children between 4 and 6 years of age four times over the course of their first year of training, as well as in a group of age-matched controls. We found that the negative response around 250 ms after stimulus onset, which is associated with sound categorization and attentional processing in children, changed more rapidly over the course of the year in the music students compared to in the controls. 15 This was accompanied by their superior improvement in working memory performance. These findings are consistent with the idea that musical training exerts its effects on other cognitive domains by training executive functions.
*Coauthors: Kathleen Corrigall, McMaster University, Hamilton, ON; Takako Fujioka, Rotman Research Institute, Toronto, and McMaster University, Hamilton, ON.
Interaction Between Brain Plasticity During Performance Training and Auditory Processing
Speaker: Christo Pantev
Institute for Biomagnetism and Biosignal Analysis, University of Münster
Besides processing of highly specific functions, musical training effects on brain plasticity were observed in simple functional representations in the sensorimotor and auditory cortices of musicians. However, it is yet unclear how sensorimotor practice and auditory practice interact with each other and how they induce cortical plasticity in each area during training. In a previous study we found that simple training in listening discrimination resulted in noticeable cortical plastic changes in auditory areas, as indicated by emerging of magnetoencephalographic (MEG) Mismatch Negativity response (MMNm). However, it is empirically true that intense active experiences enable us through musical training to acquire general schemes in auditory cognitive processing.
Here we address the question of how the effects of learning, in different brain areas, interact with each other, and how this influences the brain's plastic reorganization. Especially, the transformation of sensorimotor skills into musical processing activities has never been thoroughly investigated. Performance training in music needs strong integration of auditory, sensory and motor skills. This interaction greatly helps to acquire general schemes of musical structure such as melodic interval information. Therefore, we tested by MEG how the practice of motor skills contributes to enhance auditory memory traces when compared to mere training in listening as a control condition. Two groups, each of non-musicians, participated in pre- and post-training MEG recordings after a training session in either pure listening or performance. Performance trainings were carried out using a keyboard with correctness monitoring during the sessions. Training of listening was held using an adaptive 2AFC discrimination procedure with feedback to discriminate errors in the melody. Both trainings were continued until the subjects achieved above 90% correctness for each training task. We used recordings of the performance group as stimuli for the listening group. This kept the auditory material for both groups identical and allowed a discrimination task (correct vs. incorrect performance) that is as similar as possible for the listening group. The training results were estimated by means of MMNm and they reflected the brain plasticity changes in both groups. The outcome showed clear significant differences between the groups, indicating substantially larger reorganizational effects in the group of the performers as compared to the group obtaining only auditory training by listening.
Absolute Pitch
Speaker: Lutz Jäncke
Department of Neuropsychology, University of Zurich
Absolute pitch is an extraordinary ability, which rarely occurs. Several studies have focused so far on the neural and possible genetic underpinnings of this ability. Currently, most authorities would suppose that absolute pitch is strongly genetically determined. However, several claims have been made arguing that some kind of specific experience might also determine or at least foster the establishment of absolute pitch.
In addition, there is also growing interest in whether absolute pitch is somehow related to other cognition (e.g., language). In this talk I present the findings of two studies. One study focuses on the question of whether absolute pitch might be stimulated by musical training. The second study is an anatomical study in which we compared the fiber tracts of absolutepitch musicians with the fiber tracts measures of relative-pitch musicians and non-musicians. In the first experiment we examined children enrolled in Suzuki schools and measured the Mismatch Negativity in response to piano, violin, and sine tone. We found a remarkable change of the MMN amplitude during the course of music training with enhanced MMN amplitudes to the trained tones. In addition, we identified that the performance in labeling particular tones increased with performance, significantly correlating with MMN amplitude. In some students, performance was approximately similar to the performance found in young absolute-pitch musicians. In the second study we focused on the neural underpinnings of absolute pitch by measuring the fiber tracts using the DTI method. We found that absolutepitch musicians showed stronger temporo-parietal fiber tract connections indicated by increased FA values in the arcuate fiber tract. In addition, this stronger fiber tract was only found on the left hemisphere, again supporting the view that absolute pitch is related to specialized lefthemispheric mechanisms located in the vicinity of the auditory cortex.
*Coauthor: Mathias Oechslin, Department of Neuropsychology, University of Zurich.
Musicians — Same or Different?
Speaker: Mari Tervaniemi
Cognitive Brain Research Unit, Department of Psychology, University of Helsinki
In the neurosciences of music, musicians have traditionally been treated as a unified group as if the demands set by their musical activities would be more or less equal in terms of perceptual, cognitive, and motor functions. However, obviously, their musical preferences differentiate them up to a high degree, for instance, in terms of the instrument they chose, the music genre they are mostly engaged with as well as their practicing style. This diversity in musicians' profiles has been recently taken into account in several empirical endeavors. This talk reviews the evidence available about the various neurocognitive profiles that these different musicians display.
Effects of Asymmetric Cultural Experiences on the Auditory Pathway:
Evidence from Music
Speaker: Patrick C.M. Wong
Departments of Communication, Sciences & Disorders and Otolaryngology, Head & Neck Surgery, Northwestern University
Cultural experiences come in many different forms, such as immersion in a particular linguistic community, exposure to faces of people with different racial backgrounds, or repeated encounters with music of a particular tradition. In most circumstances, these cultural experiences are asymmetric, meaning one type of experience occurs more frequently than other types (e.g., a person raised in India will likely encounter the Indian Todi scale more so than a Westerner). In this talk, I present recent findings from my laboratory that reveal the impact of short- and long-term asymmetric linguistic and musical experiences on how our nervous system responds to complex sounds.
I discuss experiments examining how musical experience may facilitate the learning of a tone language, how musicians develop neural circuitries that are sensitive to musical melodies played in their instrument of expertise, and how even non-musicians are particularly sensitive to music of their own culture(s). An understanding of these cultural asymmetries is useful in formulating a more comprehensive model of auditory perceptual expertise that considers how experiences shape auditory skill levels. Such a model has the potentials to aid in the development of rehabilitation programs for the effective treatment of neurological impairments.
Musical Memory — Music Is Memory
Coordinator: Barbara Tillmann
Introduction
Musical memory has remarkable features. However, current models of memory have been proposed mainly for verbal and visual stimuli and the functioning of non-verbal auditory memory is still not very well known. The goal of this symposium is to present the specific features of musical memory in comparison to language and other domains. The speakers review recent research investigating various aspects of musical memory and learning (short-term and working memory, implicit long-term memory and explicit long-term memory for episodic and semantic information) by covering developmental, cognitive and neurophysiological approaches in normal and brain-damaged participants. The contributions also address implications for medical rehabilitation for patients with memory disorders (epilepsy, Alzheimer or fronto-temporal dementia).
Matthew Schulkind address some basic questions about the specificity of musical memory: is it different from visual and verbal stimuli? Can it be measured using the same experimental manipulations and variables? Is music better remembered than other stimuli? Jennifer Saffran and Erik Thiessen investigate the role of melody in infant-directed-speech and song.
What is the minimum amount of information required to activate musical memory and to what extent this notion can be applied in Alzheimer patients is the focus of Emmanuel Bigand's talk, while Séverine Samson and her colleagues examine the neurobiological bases of learning and familiarity in music, and their implication in rehabilitation. Isabelle Peretz and Jennifer Saffran will conclude by presenting data on the neural correlates of the musical lexicon.
Is Memory for Music Special?
Speaker: Matthew Schulkind
Department of Psychology, Amherst College
Although psychologists since Ebbinghaus have studied memory, research in this area has focused on visual and verbal stimuli, with little attention paid to music. Although regrettable in some ways, the bias against studying memory for music presents researchers with an opportunity to examine whether memory principles derived from the study of visual and verbal stimuli apply to other domains. Three questions are addressed. First, is musical memory sensitive to the same kinds of experimental manipulations and variables as non-musical stimuli?
Obstacles that make direct comparison of musical and non-musical stimuli difficult are discussed. Second, is music remembered "better" than other kinds of stimuli? This claim abounds in the popular literature but has not been the subject of much empirical study. Third, is musical memory separate from other kinds of memory or does musical knowledge influence other forms of memory? This question is addressed by examining musical memory in dementia patients.
How the Melody Facilitates the Message, and Vice Versa, in Infant Learning and Memory
Speaker: Jenny Saffran*
Department of Psychology, University of Wisconsin, Madison
Melody is ubiquitous in infants' environments. In this talk, we consider recent studies investigating the role of melody in infant-directed-speech and song. Infants, like adults, remember more about word sequences when they are spoken melodiously or sung than when melody is absent. Conversely, infants remember more about melodies when they are paired with word sequences than when lyrics are absent. Melodies appear to enhance infants' encoding of non-musical sequential information, engaging attentional processes and providing correlated cues which facilitate learning. The fact that non-musical sequential information similarly enhances infants' encoding of melodies suggests the possibility of "virtuous circles": what infants learn and remember at one level of analysis facilitates subsequent discovery of structure at other levels, which may in turn reciprocally facilitate learning at the initial level.
*Coauthor: Erik Thiessen, Carnegie Mellon University.
Bootstrapping Memory for Music and Text
Speaker: Emmanuel Bigand
Institut Universitaire de France and Université de Bourgogne
We investigate the specificity of musical memory with a bootstrap procedure that consists of presenting participants with a randomly organized sample of short excerpts of musical pieces and texts. These pieces may be either famous (Vivaldi's "Four Seasons" for example) or unknown. The participant's task is to evaluate the familiarity of the excerpts.
The durations of these samples is manipulated from long duration (about 1 sec) to extremely short duration (50ms). At extremely short duration (below 250ms), it is basically impossible to explicitly recognize the nature of the excerpts. An additional manipulation consists of using famous music and famous texts (overlearned in childhood). The critical point of the study is to identify the minimum amount of time necessary to activate memory in music and language. In addition, this study is run with musically trained and untrained participants as well as Alzheimer patients and matched controls.
Current findings suggest that very short slices of information (50 ms) are processed easily enough to bootstrap memory for music in normal subjects. Memory is activated faster for famous music pieces than for linguistic texts. Implications for the dynamic qualities of human memory are also outlined.
Learning and Familiarity in Music: Clinical Implications of Cognitive Neuroscience
Speaker: Séverine Samson*
University of Lille III, Neuropsychology and Auditory Cognition, Villeneuve d'Ascq & La Salpêtrière Hospital, Paris
By adapting methods of cognitive psychology to neuropsychology and brain imaging, we examined the neurobiological bases of learning and familiarity in music. We first present data obtained in brain pathologies of epileptic and degenerative origins as well as neuroimaging results to draw parallels between musical and language memory. Then, recent data illustrating how emotional content of stimuli influences musical memory are reported. These studies on musical memory as well as their relation to emotion open up interesting paths to devise new strategies of cognitive rehabilitation. We also present the benefit of music in rehabilitation of brain-damaged patients to trace the therapeutic implications of our results, thus strengthening the interest of examining the interactions between cognitive and clinical neurosciences.
*Coauthors: Delphine Dellacherie, University of Lille III, Neuropsychology and Auditory
Cognition, Villeneuve d'Ascq & La Salpêtrière Hospital, Paris; Hervé Platel, University of Caen & INSERM U923.
Neural Correlates of the Musical Lexicon
Speaker: Isabelle Peretz*
BRAMS, Université de Montreal
The musical lexicon is a representational system that contains all the representations of the specific musical phrases to which one has been exposed during one's lifetime. Accordingly, successful recognition of a familiar tune depends on a selection procedure that takes place in the musical lexicon. The goal of the study was to identify the neural correlates of the musical lexicon. Nine students with little musical training were scanned with functional magnetic resonance imaging while listening to familiar, unfamiliar and scrambled melodies. The familiar melodies were taken from instrumental pieces so as to avoid verbal associations; the unfamiliar melodies were retrogrades of the familiar melodies and the scrambled melodies contained the same tones in a random order. Listening to familiar melodies as compared to unfamiliar melodies was associated with activation in the left supplementary motor area (SMA) and the right superior temporal sulcus (STS). We suggest that the SMA involvement is likely to be related to inner singing. The STS may be more related to the retrieval of information from the musical lexicon. As expected, the results showed vast activation in auditory cortices during all conditions compared to silence. Additional activation in the left inferior parietal lobule (IPL) was also seen, which seems to be associated with processing of musical structure or expectancies.
*Coauthors: Nathalie Gosselin, BRAMS, Université de Montreal; Robert J. Zatorre and Pascal Belin, BRAMS, Montreal Neurological Institute, McGill University.
Emotions and Music: Normal and Disordered Development
Coordinators: Daniel J. Levitin and Mari Tervaniemi
Introduction
The symposium presents recent explorations and achievements in the fields of developmental auditory neuroscience, empirical music aesthetics, and music emotions in their normal and disordered forms. The aim of these studies is to illuminate the developmental time course of music perception and appreciation as well as the mental and neural determinants of aesthetic processes. Additional work examines responses to emotion and structure in music by typically developing individuals and individuals with Autistic Spectrum Disorders.
The symposium consists of four contributions which offer complementary view points and methodological solutions to probe this challenging but highly promising new area within the neurosciences of music. Maria Cristina Saccuman and her group, in collaboration with Stefan Koelsch, highlight mechanisms of music perception and demonstrate that specialization for processing music-like stimuli is present from the first days of life and that the dedicated systems are sensitive to violations of musical stimuli. Elvira Brattico talks about the aesthetic positive or negative emotions related to the subjective appraisal of music, the recognition and preference of the emotional content and their correlation to musical expertise. Daniel Levitin and Pamela Heaton address the issue of emotional content recognition in adolescents with Autistic Spectrum Disorders, while analyzing qualitative differences in this ability displayed by this particular population.
Music and the Infant Brain: a fMRI Study in Newborns
Speaker: Maria Cristina Saccuman*
Department of Psychology and Neuroscience, University Vita-Salute
Music has become a fruitful research field for cognitive neuroscience for the possibility it offers of addressing brain functional specialization, integration and plasticity.
Evidence from neurophysiological, electrophysiological, neuropsychological, and imaging studies, has converged on a specific network for music processing in adult non-musicians. However, it is still unclear to which extent these neural representations are due to adaptation of the brain resulting from passive exposure to the environment, or to genetic and neurobiological constraints. Very young infants are indeed surprisingly skilled at perceiving subtle aspects of musical stimuli, and ontogenetically it appears that infants' first steps into language are based on prosodic information, while musical communication in early childhood plays a major role for emotional, cognitive, and social development in children. Yet, very little is known about the early specialization of the brain for music. For the first time, we investigated the neural correlates of music processing in newborn infants with functional magnetic resonance imaging (fMRI).
18 healthy full-term non-sedated newborns within the first two days of life participated in the study. Subjects listened to musical stimuli alternating with silence. Three kinds of stimuli were used: (a) western tonal music excerpts; (b) the same excerpts altered so as to sound dissonant or (c) to include violations of tonal syntax, with continuous shifts between unrelated tonalities. Analyses show a specific activation for musical stimuli in the right hemisphere (superior temporal gyrus including A1, temporo-parietal junction, inferior parietal lobule), and more bilateral activation for the altered stimuli, with a decrease in signal in the right temporal region and additional activations in the left inferior frontal gyrus. Our results indicate that a specialization for processing music-like stimuli is present from the first days of life and that the dedicated systems are sensitive to violations of musical stimuli.
*Coauthors: Paola Scifo2,3, Guido Andreolli1, Danilo Spada5, Federica Navarra2,3, Cristina Baldoli3,4, Stefan Koelsch6, Daniela Perani1,2,3.
1 Department of Psychology and Neuroscience, University Vita-Salute San Raffaele, Milan.
2 Department of Nuclear Medicine Scientific Institute San Raffaele, Milan.
3 CERMAC San Raffaele Scientific Institute, Milan.
4 Department of Neuroradiology, Scientific Institute San Raffaele, Milan.
5 Psychology Institute School of Medicine, University of Milan.
6 Department of Psychology University of Sussex, Brighton, UK.
Subjective Appraisal of Music: Neuroimaging Evidence
Speaker: Elvira Brattico*
Cognitive Brain Research Unit, Department of Psychology, University of Helsinki
One fascination of music lies rightly in its power to induce at the same time contrasting emotions. Tears and joy may coexist during listening to music. This apparent paradox derives from the peculiar character of the musiclistening experience. Music is known to induce emotions, and most consistently few basic emotions (such as happiness and sadness) but more complex ones as well. Aesthetic positive or negative emotions related to the subjective appraisal of music are also typically triggered. A complete model of music listening should hence include both the initial emotional and perceptual processing of sounds and their subsequent conscious interpretation and subjective aesthetic appraisal. An aesthetic experience in general has been defined as an affective state determined by the interaction with an environmental stimulus to which we attribute positive or negative qualities (such as beauty or ugliness) as a result of cognitive and emotional evaluation.
Aesthetic experience of music may hence represent for system neuroscientists a useful model of the interaction between emotional, cognitive and evaluative self-referential processes evoked by an object. In a series of neuroimaging studies, we analyzed the main aspects of the aesthetic experience of music. The first study investigated the neural substrates of evaluative preference and descriptive correctness ratings of the same musical cadences by measuring the event-related potentials (ERPs).
The second study used a similar paradigm on subjects varying in their degree of musical expertise, to define the role of long-term knowledge on aesthetic listening to music. The third study aimed at disentangling the brain structures involved in basic emotion recognition and in subjective preference of musical excerpts. The current findings lead to a revised framework of music listening in which the subjective aspects of musical appraisal along with their neural determinants are integrated.
*Coauthor: Thomas Jacobsen, Cognitive Brain Research Unit, Department of Psychology, University of Helsinki.
Interpretation of Musical Emotion by Adolescents with
Autistic Spectrum Disorders (ASD)
Speaker: Daniel J. Levitin*
Department of Psychology, McGill University, Montreal
We will review recent research from our laboratory on emotional processing in music in children with autism spectrum disorders (including PDD-NOS, Autism, and Asperger's Syndrome). First, to test their recognition of emotions in music, We asked teenagers with and without ASD to describe musical excerpts using one of the four following emotions: happy, sad, scared or peaceful. Although the ASD individuals were less accurate at these identifications overall, they still performed significantly better than chance. Contrary to Baron-Cohen et al.'s (2000) "amygdala theory of autism," that individuals with ASD should be impaired at fear recognition, our ASD group identified fear significantly above chance.
Second, participants with and without ASD listened in random order to specially prepared examples of solo piano works in which the expressivity was manipulated (by parametrically altering the variability in timing and amplitude). The participants rated the level of expressivity they thought was contained in these excerpts on a continuous rating scale.
The judgments of control participants recreated the correct ranking order of the stimuli and showed monotonicity. In contrast, the judgments of ASD participants failed to distinguish between different levels of expressivity. We take this as evidence that individuals with ASD are attracted to the structural features of music rather than the expressive or emotional features (or at least if they appreciate music for its expressive/emotional content, they are not as sensitive to those same cues as are typically developing normal participants).
Does Autism Provide a Model for Studying Musical Emotion Recognition
in Typical Children?
Speaker: Pamela Heaton
Department of Psychology, Goldsmiths College, University of London
Autism is characterised by a deficit in emotional understanding within social domains. Whilst it may be predicted that such difficulties would generalise to music, anecdotal evidence suggests a strong orientation to music listening and experimental evidence has shown preserved sensitivity to the affective qualities of major and minor mode.
In order to extend these findings, children with autism, Down syndrome and typical development were compared on a task measuring perception of affective and movement states in music. The findings showed that discrimination performance depended on verbal mental age and there was no significant effect of diagnosis. In a second study carried out with high-functioning adults with autism, the nature of their personal experiences of music was probed. Analysis of verbal reports showed that participants exploited music for a wide range of social, emotional and cognitive purposes. However, whilst this provided evidence for spared musical appreciation in autism, it was noted that their descriptions of mood change in response to music were internally (arousal) rather than externally (emotive) focused.
Listening To and Making Music Facilitates Brain Recovery Processes
Coordinator: Gottfried Schlaug
Introduction
Emerging research over the last decade has shown that long-term music training and skill learning can be a strong stimulator for neuroplastic changes in the brain. Making music places unique demands on our nervous system, leading to strong coupling of perception and action mediated by sensory, motor, and integrative regions distributed throughout the brain. Furthermore, listening to music and making music provokes motion, improves and increases betweensubject communication and interaction, and is considered to be a pleasant and rewarding activity.
Several reports have now shown that listening to music and making music may have the power to make rehabilitation processes not only more enjoyable, but possibly more effective. Why is music so special and how does music listening/making achieve its rehabilitative effects? Music is a strong multimodal stimulus that simultaneously transmits visual, auditory, and motoric information to a specialized network consisting of fronto-temporo-parietal brain regions, whose components are also part of the putative human mirror neuron system. Among other functions, this system might support the coupling between a perception (visual or auditory) and motor action (leg, arm/hand or vocal). Music might be a special vehicle to involve the components of this system. Furthermore, music might provide an alternative entry point to remediate impaired neural processes by engaging and linking up brain centers that would otherwise not be engaged.
This symposium demonstrates several music-based therapeutic methods whose effectiveness has been demonstrated in clinical populations and whose underlying neuroscientific mechanisms we are beginning to understand. The session starts with an overview of the putative human mirror neuron system and how it relates to music perception and making. We suggest that music listening and music making, if used appropriately, can serve as a powerful and engaging treatment modality for enhancing brain recovery processes and neuroplasticity in general.
Brain Correlates of Mirror Function in Music
Speaker: Stefan Koelsch
Department of Psychology, University of Sussex, Brighton
In 2002, Kohler and colleagues reported neurons in the monkey premotor cortex that discharged when the animal performed a specific action, as well as when it heard the related sound, suggesting the presence of mirror neurons in the auditory domain. In 2005, Koelsch et al. reported premotor-like activation of the larynx during the perception of pleasant music, presumably reflecting the activation of an auditory perception-execution matching system that is engaged during the perception of vocalizable (melodic) auditory information. In 2004, a very similar activation was reported in a study on speech prosody (intonation contour) by Meyer at al.
Corroborating these findings, a recent study from Callan et al. (2006) revealed a remarkable overlap of brain structures serving the perception as well as the (covert) production of both speech and singing, again showing an overlap of brain structures involved in the perception and the production of musical stimuli. Other studies have shown premotor activation in piano players while listening to piano music (Haueisen & Knoesche, 2001) and in nonmusicians learning to play a piano piece (Lahav et al., 2007).
The combined findings suggest that different brain structures (such as the ventrolateral premotor cortex, Rolandic operculum, Broca's area, the insula, and inferior parietal regions) are involved in different aspects of perception-action mediation (like the translation of auditory signals into motor codes of different effectors and modulations of such translations by emotion, as well as by experience). These mechanisms are important for both the processing of music and of speech prosody.
Thus, disruption of these mirror mechanisms can lead to both amusia and aphasia.
From Singing to Speaking: Why Does Singing Help Aphasic Patients?
Speaker: Gottfried Schlaug
Beth Israel Deaconess Medical Center & Harvard Medical School, Boston
The neural processes that underlie post-stroke language recovery remain largely unknown and thus have not been specifically targeted by aphasia therapies. Two possible pathways to recovery exist; one is through reactivation of perilesional areas in the left hemisphere and/or homologous language regions in the right. Because of its potential to engage/unmask language-capable brain regions in the unaffected right hemisphere, Melodic Intonation Therapy (MIT) is well-suited for facilitating language recovery in non-fluent aphasic patients, particularly those with left-hemisphere lesions encompassing large portions of the left frontal and superior temporal lobes. Patients enrolled in this study underwent an intense therapy program consisting of 75 treatment sessions. Across all patients, post-treatment evaluations showed significant improvements in behavioral measures of speech output (e.g., more meaningful words/min and increased phrase length) that correlated significantly with functional imaging changes found in a right hemispheric fronto-temporal network during overt speech in post- vs. pretreatment fMRI comparisons.
These data suggest that intensive MIT treatment leads to significant gains in speech production that can be maintained after therapy. And further, that these gains are supported by functional brain changes involving primarily right-hemispheric, languagecapable brain regions. We hypothesize that MIT's unique elements (melodic intonation, rhythmic tapping, and continuous voicing) play a critical role in facilitating this recovery from nonfluent aphasia.
Learning to Play Piano Supports Fine Motor Rehabilitation After Stroke
Speaker: Eckart Altenmüller
Institute of Music Physiology and Musicians' Medicine, University of Music and Drama, Hannover
In previous studies, it was shown that just three weeks of piano training can induce neuronal representations of skilled finger movements activated by auditory stimulation. In this study, we examined whether or not this kind of cross-modal mechanism for auditory-sensorimotor integration can be employed in the rehabilitation of motor functions following a stroke. For this purpose, we evaluated a music-supported training program designed to induce an auditory sensorimotor co-representation of movements in 32 stroke patients (17 affected in the left and 15 in the right upper extremity). Patients without any previous musical experience participated in an intensive step-by-step training, first of the paretic extremity, followed by training of both extremities. Training was applied 15 times over 3 weeks in addition to conventional treatment. Fine as well as gross motor skills were addressed by using either a MIDIpiano or electronic drum pads.
As a control, 30 stroke patients (15 affected left and 15 right) undergoing exclusively conventional therapies were recruited. Behavioral pre- and post-treatment motor functions were monitored using a computerized movement analysis system (Zebris) and an established array of motor tests (e.g. Action Research Arm Test, Box and Block Test). To investigate the activity of cortical regions in the control of movement, we studied event-related desynchronization/synchronization (ERD/ERS) and event-related coherences from all 62 subjects performing self-paced movements of the right index finger (MIDI-piano) and of the hole arm (drum pads). Patients showed significant improvement after treatment with respect to speed, precision and smoothness of movements, as shown by 3D movement analysis and clinical motor tests. Furthermore, compared to the control subjects, motor control in everyday activities improved significantly. Neurophysiological data showed a significantly larger decrease of EEG signal (power) before time of movement onset in the music supported training group in the posttraining register, which is associated with increased corticospinal excitability, whereas there were almost no differences in the control group. The music-supported training group presented a most pronounced enhancement of the coherences after the training compared to the control group, especially in the drums conditions.
This innovative therapeutic strategy is an effective approach for the motor skills neurorehabilitation in stroke patients.
*Coauthors: Sabine Schneider, Paul-Walter Schönle, and Thomas Münte, Institute of Music Physiology and Musicians' Medicine, University of Music and Drama, Hannover.
Rhythmic Auditory Stimulation Improves Gait in Stroke Patients
Speaker: Michael H. Thaut
Department of Music-Theatre-Dance, School of the Arts, Colorado State University & Center for Biomedical Research in Music, Fort Collins
The effectiveness of 2 different types of gait training in stroke rehabilitation, rhythmic auditory stimulation (RAS) versus neurodevelopmental therapy (NDT)/Bobath-based training, was compared in 2 groups of hemiparetic stroke patients over a 3-week period of daily training. Mean entry date into the study was 21.3 days post-stroke for the RAS group and 22.3 days for the control group. Patients entered the study as soon as they were able to complete 5 stride cycles with handheld assistance. Patients were closely equated by age, gender, and lesion site. Motor function in both groups was pre-assessed by the Barthel Index and the Fugl-Meyer Scales.
Pre- to post-test measures showed a significant improvement in the RAS group for velocity (P = .006), stride length (P = .0001), cadence (P = .0001) and symmetry (P = .0049) over the NDT/Bobath group. Effect sizes for RAS over NDT/Bobath training were 13.1 m/min for velocity, 0.18 m for stride length, and 19 steps/min for cadence. The data show that after 3 weeks of gait training, RAS is an effective therapeutic method to enhance gait training in hemiparetic stroke rehabilitation. Gains were significantly higher for RAS compared to NDT/Bobath training.
Music, Prosody, and Motor Programming: a Common Neural Organization?
Coordinators: Giuliano Avanzini and Katie Overy
Introduction
The human species is expert at communicating information, ideas and emotional states. Such communication takes place in various forms and involves infinitely complex arrangements of motor movements, from the vibrations of vocal chords to the bowing of a violin. Musical communication is particularly abstracted, but investigations into the neural basis of musical processing have revealed shared networks with both speech and motor processing, suggesting that similar organizational mechanisms may be involved. Impairments in musical skills, language skills and motor skills have also been found to co-occur. This symposium outlines a number of new approaches to the question of potentially shared networks between music and language.
Luciano Fadiga begins with a discussion of the neurophysiology of action representation and will present data showing that the motor cortex is activated while "listening to actions", ending with a new theoretical framework for the role of Broca's area. Aniruddh Patel presents recent data from two independent studies suggesting that individuals with amusia can have difficulties discriminating a statement from a question, based on the linguistic intonation contour. William Thompson presents data suggesting that individuals with amusia can have difficulties classifying certain types of emotional prosody, either in the presence or absence of linguistic information. Finally, Caroline Palmer presents new ERP data suggesting that listeners' behavioral and neural responses to musical structure depend very much on the sequential context, in a similar way to contextual influences in prosodic processing of language.
The Supramodal Syntax: a Missing Link Between Action Representation and
Human Communication?
Speaker: Luciano Fadiga
DSBTA Section of Human Physiology, Faculty of Medicine, Ferrara
In the last decade new evidence is growing in favour of an additional, more cognitive, role played by motor and premotor centers. Clear motor activation is evident when one simply imagines a motor act and it has been shown that during observation of others' actions a temporo-parietalfrontal circuit becomes active in the observer's brain. Which is the function of this "high-level" motor involvement? Wh does our brain look inside its own motor representations to understand how other individuals act? Is this motor involvement functional to perception or does it represent a mere epiphenomenon?
In the first part of my talk I review the neurophysiology of action representation in monkey premotor cortex, then describe some TMS experiments on action perception in humans. The second part of my talk shows some recent results demonstrating that not only observing actions but also "listening to actions" enhances the excitability of the motor cortex. During passive listening of speech, normal subjects motorically "resonate" by internally re-acting the listened words. If on one hand these data give support to Liberman's motor theory of speech perception, on the other hand they suggest that "visuomotor" and "acoustic-motor" matching systems may represent two particular aspects of a more general mechanism, used by the brain to map sensory information on its own motor repertoire.
Finally, I present data and theoretical framework supporting a new interpretation of the role played by Broca's area. Recent brain imaging studies report that, in addition to speech-related tasks, Broca's area is significantly involved also during tasks devoid of verbal content, such as arithmetic calculation and listening to music. Taking into account the large variety of experimental paradigms inducing such activation, I present neurophysiological and brain imaging data trying to integrate, on a common ground, these apparently different competencies.
Impairments of Linguistic Prosody Among Individuals with Amusia
Speaker: Aniruddh D. Patel
The Neurosciences Institute, San Diego
To what extent do music and language share neural mechanisms for processing pitch patterns? Musical tone-deafness (amusia) provides important evidence on this question. Amusics have problems with musical melody perception, yet early work suggested that they had no problems 29 with the perception of speech intonation (Ayotte, Peretz, & Hyde, 2002). However, recent data indicate that about 30% of amusics from independent studies (British and French-Canadian) have difficulty discriminating a statement from a question on the basis of a final pitch fall or rise. This suggests that pitch direction perception deficits in amusia (known from previous psychophysical work) can extend to speech. For British amusics, the direction deficit is related to the rate of change of the final pitch glide in statements/questions, with increased discrimination difficulty when rates are relatively slow.
These findings suggest that amusia provides a useful window on the neural relations between melodic processing in language and music.
Impairments of Emotional Prosody Among Individuals with Amusia
Speaker: William Forde Thompson
Department of Psychology, Macquarie University, Sydney
Most congenital amusic individuals can distinguish questions and statements at normal levels, implying that musical and linguistic skills have different processing requirements. However, while statements and questions have linguistic connotations, emotional communication occurs in both music and speech prosody. Moreover, the same acoustic cues convey similar emotional messages in both domains, suggesting that the acoustic code for emotional communication is shared by music and speech prosody.
Thus, if musical impairments extend to speech prosody, they are likely to occur for tests of sensitivity to emotional prosody. In this study, groups of amusic and non-impaired individuals were evaluated on a test of emotional prosody. The task involved classifying the emotion conveyed in short spoken phrases. Phrases consisted of emotionally neutral words (e.g., "The broom is in the cupboard") spoken to convey each of five emotions: disgust, fear, happiness, sadness, and irritation.
The amusic group classified happiness and sadness at normal levels, but were abnormally poor at classifying fear and irritation. This selective difficulty with certain emotions occurred for normal speech, or when prosodic stimuli were presented in the absence of linguistic information. The results will be discussed in view of mixed and often fragile data on the relation between musical ability and sensitivity to emotional cues in speech prosody. Implications for understanding amusia will also be discussed.
Perceiving Music in Context
Speakers:
Caroline Palmer*, Department of Psychology, McGill University, Montreal
Karsten Steinhauer*, School of Communication Sciences and Disorders, Montreal
Listeners' aesthetic and emotional responses to music typically occur in the context of long musical passages that contain structures defined in terms of the events that precede them. We describe behavioral and ERP studies investigating listeners' responses to musical accents that arise in longer musical sequences. Musically trained listeners performed a timbre change detection task. A single-tone timbre change was positioned within 4-bar melodies to coincide with or without melodic contour accents and rhythmic accents (temporal gaps).
Responses were slower to timbre changes near a contour change, but faster for timbre changes near a temporal gap, suggesting that temporal gaps were processed more quickly and signaled more closure than contour changes. Participants also responded faster to timbre changes later in the melody (likely due to increased predictability). An ERP experiment was conducted with the same melodies (350 ms between tone onsets). ERP responses to ( task-relevant) attended timbre changes elicited a Mismatch negativity (MMN) around 200ms and a late positive component around 400 ms (P300), reflecting updating of the timbre change in working memory. In contrast, (task-irrelevant) temporal gaps elicited an MMN only, with no subsequent P300 (indicating that temporal gaps were not updated). Specific to the musical context, the peak amplitudes of the components changed systematically across the sequence.
Listeners' behavioral and neural responses to musical structure depend on the sequential context, and change systematically as sequential predictability and listeners' expectations change.
*Coauthors: Lisa Jewett and Christine Capota, Department of Psychology, McGill University, Montreal.
New Directions: Cochlear Implants
Coordinator: Nina Kraus
Introduction
The improvement of new technologies on the care and rehabilitation in hearing-impaired population has fostered research on fine perception in both speech and music. In particular, the aim of this symposium is to review the current findings and understand the mutual interactions between music and language in children and adults with cochlear implants. Robert Shannon explores in his talk the contribution of spectral and temporal envelope and fine structure to various aspects of speech and music perception. Sandra Trehub discusses the different aspects of music perception in congenitally deaf children, who have only heard music electrically (via cochlear implants) rather than acoustically. Finally Nina Kraus presents a series of experiments demonstrating effects of training on music perception in cochlear implants and in a simulated cochlear implant model.
The Importance of Temporal Envelope and Fine Structure Cues for Speech and Music: Lessons from Cochlear Implants
Speaker: Robert V. Shannon*
Department of Auditory Implants and Perception, House Ear Institute, Los Angeles
Research and outcomes with cochlear implants have revealed a dichotomy in the cues necessary for speech and music recognition. Speech can be conveyed by a small number of noise bands each modulated slowly in time. While such a representation allows good speech recognition, it does not provide adequate information for speaker identification, voice quality, speech in noise, or melody recognition. We review experiments investigating the contribution of spectral and temporal envelope and fine structure to various aspects of speech and music perception. In general, coarse spectral and temporal resolution are adequate for speech recognition and for conveying prosodic and rhythmic properties of music. But voice quality and identification, as well as melody recognition, requires spectral fine structure cues. With fixed constraints on spectral resolution, such as it occurs with a hearing loss or cochlear implant, music training can produce a considerable improvement in performance.
*Coauthors: John Galvin and Qian-Jie Fu, Department of Auditory Implants and Perception, House Ear Institute, Los Angeles.
Music in the Lives of Deaf Children with Cochlear Implants
Speaker: Sandra Trehub
Department of Psychology, University of Toronto, Mississauga, ON
Cochlear implants provide reasonable information about timing but very coarse information about pitch. As a result, many adult implant users find music unpleasant or uninteresting, even if they were music lovers prior to becoming deaf. The situation seems to be different for congenitally deaf children, who have only heard music electrically (via cochlear implants) rather than acoustically. These children generally enjoy music listening and musicmaking even though what they hear is very different from what is heard by listeners with normal hearing. Like hearing children, child implant users recognize instrumental renditions of familiar pop songs that they usually hear with words.
Unlike hearing children, however, they cannot recognize those songs from the melody alone. Surprisingly, they remember more details of the theme music that accompanies their favorite television programs. It is likely that rhythm plays a larger role in music processing for child implant users than it does for hearing children. 24 Indeed, the songs that child implant users sing reveal poor reproduction of tunes but age-appropriate reproduction of rhythms. Finally, there are indications that deaf children with implants use the melody (i.e., prosody) and vocal timbre of speech to recognize familiar voices.
Training-induced Malleability in Neural Encoding of Pitch, Time and Timbre
Speaker: Nina Kraus
Department of Communication Sciences, Neurobiology and Physiology, Otolaryngology, Northwestern University
Speech and music sounds consist of three fundamental components: pitch, harmonics (timbre/formants) and timing (ranging from consonants to rhythm in music). It is possible to access the neural transcription of these elements objectively, non-invasively and with great fidelity with scalp electrodes in humans1. The acoustic cues relating to pitch, timing and harmonics perception have distinct subcortical representations which can be selectively enhanced or degraded in different populations.
Musically trained subjects have been found to have enhanced subcortical representation of pitch, timing and harmonics (timbre). The effects of musical experience on the nervous system's response to sound are pervasive and extend beyond music to the domains of language and emotion2,3. In impaired systems, these subcortical neural events provide a biological marker of specific aspects of deficient sound encoding. Children with reading disorders reveal deficiencies in encoding timing and harmonics, and this pattern is consistent with the phonological processing problems inherent in reading disorders4. The opposite pattern is seen for a subgroup of autistic children who show abnormal pitch representation, consistent with inability to perceive the "intention" of what is said (e.g. question vs. statement)5.
In the normal system, it appears that cortical functions like music and language fundamentally shape auditory processing that occurs early in the sensory processing stream2,3,6. This top-down influence is likely mediated by the extensive (corticofugal) circuitry of descending efferent fibers that synapse all along the auditory pathway7. Sensory shaping occurs not only with lifelong experience but can be effected by short-term training. I discuss a series of experiments demonstrating the lifelong and short-term8,9 malleability of the neural encoding of pitch, timing and timbre. In particular, I will discuss the effect of training in a simulated cochlear implant model and the implications for hearing music through a cochlear implant.
It appears that in order to facilitate sensory learning, the impaired system can capitalize on 1) the shared biological resources underlying the neural processing of language, music and emotion, 2) the apparent cognitive-sensory reciprocity, and 3) the seemingly independent neural representations of pitch, timing and harmonic cues.
References
1 Kraus, N. & Nicol, T. (2005) Brainstem origins for cortical "what" and "where" pathways in the auditory system. Trends in Neurosciences 28: 176-181.
2 Musacchia, G., Sams, M., Skoe, E.& Kraus, N. (2007) Musicians have enhanced subcortical auditory and audiovisual processing of speech and music. Proc. Nat. Acad. Sci. 104: 15894-158983.
3 Wong, P.C.M., Skoe, E., Russo, N.M.,Dees, T. & Kraus, N. (2007) Musical experience shapes human brainstem encoding of linguistic pitchpatterns. Nature Neurosci. 10: 420-422.
4 Banai, K., Nicol, T., Zecker,S. & Kraus, N. (2005) Brainstem timing: implications forcortical processing and literacy. J. Neurosci. 25 (43): 9850-9857.
5 Russo, N.M., Bradlow, A.R., Skoe,E., Trommer, B.L., Nicol., T., Zecker,S. & Kraus, N. (in press) Deficient brainstem encoding of pitch in children with Autism Spectrum Disorders. Clinical Neurophys.
6 Krishnan, A., Xu, Y.S., Gandour,J. & Cariani, P. (2005) Encoding of pitch in the human brainstem is sensitive to language experience. Cogn. Brain. Res. 25: 161-168.
7 Suga, N., Xiao, Z.J., Ma, X.F. & Ji,W.Q. (2002) Plasticity and corticofugal modulationfor hearing in adult animals. Neuron 36: 9-18.
8 Song, J.H., Skoe E., Wong,P.C.M. & Kraus, N. (in press) Plasticity in the adult human auditory brainstem following short-term linguistic training. J. Cogn. Neurosci.
9 Russo, N., Nicol, T., Zecker, S, Hayes,E. & Kraus, N. (2005) Auditory training improves neuraltiming in the human brainstem. Behav. Brain Res. 156: 95-103.
Supported by National Institutes of Health (R01DC001510) and National Science Foundation (NSF 0544846).
Web Sites
Pierfranco and Luisa Mariani Foundation
The Mariani Foundation was the lead organizer for the Neurosciences and Music III conference. This nonprofit organization's core meeting is to promote progress in child neurology by providing services, educational events, and funding for research. Go to their site for more information about neuroscience and music conferences and see their neuromusic news releases for information about recent papers in the field.
Brams (International Laboratory for Brain, Music, and Sound Research)
This research unit affiliated with McGill, the Université de Montreal, and the Montreal Neurological Research Institute, provides links to many music and brain science researchers in the Montreal region.
House Ear Institute
Information about ongoing research into the auditory system and technologies to improve hearing. Tigerspeech is a program designed to help people with cochlear implants learn to make use of their prosthetics.
Nature Magazine Web Focus: Science & Music
This recent nine-part series features essays on the interface between science and music, as well as a podcast.
This is Your Brain on Music
The promotional Web site for Daniel Levitin's book describes the role of various brain regions in music processing and includes references to the brain in popular songs.
Books
Brown S, Volgsten U, eds. 2005. Music and Manipulation: On the Social Uses and Social Control of Music. Berghahn Books, New York.
Blacking J. 1995. How Musical is Man? University of Washington Press, Seattle.
Levitin DJ. 2006. This Is Your Brain on Music: The Science of a Human Obsession. Plume, New York.
Mithen S. 2006. The Singing Neanderthals: The Origins of Music, Language, Mind and Body. Harvard University Press, Cambridge, MA.
Patel AD. 2007. Music, Language and the Brain. Oxford University Press, New York.
Peretz I, Zatorre R, eds. 2003. The Cognitive Neuroscience of Music. Oxford University Press, New York.
Sacks O. 2007. Musicophilia: Tales of Music and the Brain. Knopf, New York.
Thaut MH. 2005. Rhythm, Music, and the Brain: Scientific Foundations and Clinical Applications. Routledge, New York.
Wallin NL, Merker B, Brown S, eds. 2001. The Origins of Music. The MIT Press, Cambridge, MA.
Articles
Ayotte J, Peretz I, Hyde K. 2002. Congenital amusia: A group study of adults afflicted with a music-specific disorder. Brain 125: 238-251. (PDF, 385 KB) Full Text
Brown S, Martinez MJ. 2007. Activation of premotor vocal areas during musical discrimination. Brain Cogn. 63: 59-69. (PDF, 765 KB) Full Text
Brown S, Ngan E, Liotti M. 2008. A larynx area in the human motor cortex. Cereb. Cortex 18: 837-845. Full Text
Chen JL, Penhune VB, Zatorre RJ. 2008. Listening to musical rhythms recruits motor regions of the brain. Cereb. Cortex. [Epub ahead of print]
Chen JL, Penhune VB, Zatorre RJ. 2008. Moving on time: brain network for auditory-motor synchronization is modulated by rhythm complexity and musical training. J. Cogn. Neurosci. 20: 226-239.
Chen JL, Zatorre RJ, Penhune VB. 2006. Interactions between auditory and dorsal premotor cortex during synchronization to musical rhythms. Neuroimage 32: 1771-1781.
Dalla Bella S. 2008. Singing out of tune: Disturbances of vocal performance in the general population. J. Acoust. Soc. Am. 123: 3379.
Grahn JA, Brett M. 2007. Rhythm and beat perception in motor areas of the brain. J. Cogn. Neurosci. 19: 893-906.
Gunji A, Ishii R, Chau W, et al. 2007. Rhythmic brain activities related to singing in humans. Neuroimage 34: 426-434.
Hyde KL, Zatorre RJ, Griffiths TD, et al. 2006. Morphometry of the amusic brain: a two-site study. Brain 129: 2562-2570. (PDF, 424 KB) Full Text
Large EW. (in press). Resonating to musical rhythm: theory and experiment. Grondin S, ed. The Psychology of Time. Elsevier, New York.
Mandell J, Schulze K, Schlaug G. 2007. Congenital amusia: an auditory-motor feedback disorder? Restor. Neurol. Neurosci. 25: 323-334.
Marmel F, Tillmann B, Dowling WJ. 2008. Tonal expectations influence pitch perception. Percept. Psychophys. 70: 841-852.
Patel AD. 2003. Language, music, syntax and the brain. Nat. Neurosci. 6: 674-681.
Patel AD, Iversen JR. 2007. The linguistic benefits of musical abilities. Trends Cogn Sci. 11: 369-72.
Patel AD, Iversen JR, Chen Y, Repp BH. 2005. The influence of metricality and modality on synchronization with a beat. Exp. Brain Res. 163: 226-238.
Peretz I, Zatorre RJ.2005. Brain organization for music processing. Annu. Rev. Psychol. 56: 89-114.
Ruiz MH, Koelsch S, Bhattacharya J. 2008. Decrease in early right alpha band phase synchronization and late gamma band oscillations in processing syntax in music. Hum. Brain Mapp. [Epub ahead of print]
Schön D, Boyer M, Moreno S, et al. 2008. Songs as an aid for language acquisition. Cognition 106: 975-983.
Shannon RV. 2005. Speech and music have different requirements for spectral resolution. Int. Rev. Neurobiol. 70: 121-134.
Stegemöller EL, Skoe E, Nicol T, et al. 2008. Musical training and vocal production of speech and song. Music Perception 25: 419-428.
Wong PCM, Skoe E, Russo NM, et al. 2007. Musical experience shapes human brainstem encoding of linguistic pitch patterns. Nat. Neurosci. 10: 420-422.
Volkova A, Trehub SE, Schellenberg EG. 2006. Infants' memory for musical performances. Dev. Sci. 9: 583-589.
Zatorre RJ, Chen JL, Penhune VB. 2007. When the brain plays music: auditory–motor interactions in music perception and production. Nat. Rev. Neuro. 8: 547-558.
Organizers
Isabelle Peretz, PhD
Université de Montréal
International Laboratory for Brain, Music, and Sound Research (Brams)
e-mail | web site | publications
Isabelle Peretz is professor of psychology at the Université de Montréal, and codirector of the International Laboratory for Brain, Music, and Sound Research (BRAMS). She also holds the positions of Canada Research Chair in Neurocognition in Music, Casavant Chair in Neurocognition of Music, and associate professor in the School of Hearing and Audiology at the University of Montreal. Her research interests include the biological foundations of music, how the brain is organized for music, music-specific impairments, neural correlates of musical emotions, speech prosody, music and speech in singing, and the neural correlates of pitch-related deficits.
Robert J. Zatorre, PhD
Montreal Neurological Institute, McGill University
e-mail | web site | publications
Robert Zatorre is a professor in the Department of Neurology and Neurosurgery at the Montreal Neurological Institute at McGill University. He heads the Auditory Processing Laboratory, which conducts basic research to understand the function of complex auditory processes, especially the processing of musical sounds and speech. He also works on auditory spatial processes and cross-modal plasticity, as well as anatomical measures of auditory cortex and its relation to hemispheric asymmetries.
Virginia Penhune, PhD
Concordia University
e-mail | web site | publications
Virginia Penhune is professor of psychology at Concordia University in Montreal. She heads the Laboratory for Motor Learning and Neural Plasticity, which investigates changes in the human brain that occur due to motor learning and performance. Her team is working to identify brain regions that control changes in movement kinematics through learning, specific kinematic parameters that change as a skill is acquired, and how musical training affects the ability to learn.
Giuliano Avanzini, PhD
Fondazione IRCCS, Istituto Neurologico "C. Besta", Milan
e-mail | web site | publications
Giuliano Avanzini is emeritus chairman of the Department of Neurosciences at the Institute Neurologico "C. Besta" in Milan, Italy and professor of neurology at the University of Ferrara, Italy. Since the 1970s he has conducted research in neurophysiology, in particular epilepsy. He is editor-in-chief of Neurological Sciences and a member of the editorial boards of the European Journal of Neurology, Epilepsy, Epilepsy Research, and Neurological Acta Scandinavia. He is president of the International School of Neurological Sciences of Venice, director of the Summer School of Epileptology, and past-president of the International League Against Epilepsy.
Luisa Lopez, MD, PhD
University of Rome "Tor Vergata"
e-mail
Luisa Lopez is a neurophysiologist with a PhD in child neurology. She heads the child neurology unit in "Eugenio Litta" Rehabilitation Center in Grottaferrata, Rome. She also consults and teaches in the Child Neurology Department at University of Rome, Tor Vergata. Her combined interests in child neurology and music have driven her toward the Mariani Foundation, where she has consulted in its neuroscience and music project since 2000.
Maria Majno, PhD
Pierfranco and Luisa Mariani Foundation
e-mail | web site | publications
Maria Majno is executive director of the Fondazione Pierfranco e Luisa Mariani for Child Neurology (Milan, Italy), a nonprofit organization dedicated to care and services, research, and continuous specialized education in the field of developmental disabilities. Ever since her initial involvement with the Mariani Foundation in 1987, she has been in charge of general management and programming of training and international meetings, and of related publishing activities. Her background in the humanities and music has steered her interest toward the relationship between music and the neurosciences, which in recent years has become a main focus of the Foundation's activities.
Keynote Speaker
Steven Mithen, PhD
University of Reading
e-mail | web site
Stephen Mithen is an archeologist whose work centers on four themes: late Pleistocene and early Holocene hunter-gatherers and farmers; computational archeology; the evolution of the human mind, language, and music; and water, life, and civilization. He completed his PhD in archaeology at Cambridge University. Between 1987 and 1992 he was a research fellow at Trinity Hall and then lecturer in archaeology at Cambridge. After moving to the University of Reading, he was promoted to senior lecturer (1996), reader (1998), and then professor of early prehistory (2000). In August 2002 he was appointed as the first head of the School of Human & Environmental Sciences, formed by the Departments of Archaeology, Geography, Soil Science and the Postgraduate Institute of Sedimentology. He was elected as a Fellow of the British Academy in 2004. He is the author of The Singing Neanderthals: The Origins of Music, Language, Mind, and Body.
Speakers
Eckart Altenmüller, MD
University of Music and Drama, Hannover
e-mail | web site | publications
Emmanuel Bigand, PhD
Institut Universitaire de France
e-mail | web site | publications
Elvira Brattico, PhD
University of Helsinki
e-mail | web site | publications
Steven Brown, PhD
Simon Fraser University
e-mail | web site | publications
Joyce L. Chen
McGill University, BRAMS
e-mail | publications
Simone Dalla Bella, PhD
University of Finance and Management, Warsaw
e-mail | web site | publications
Luciano Fadiga, PhD
University of Ferrara
e-mail | web site | publications
Jessica A. Grahn, PhD
Cambridge University
e-mail | web site | publications
Pamela Heaton, PhD
Goldsmiths, University of London
e-mail | web site | publications
John R. Iversen, PhD
The Neurosciences Institute
e-mail | web site | publications
Lutz Jäncke, PhD
University of Zurich
e-mail | web site
Stefan Koelsch, PhD
University of Sussex
e-mail | web site | publications
Nina Kraus, PhD
Northwestern University
e-mail | web site | publications
Edward W. Large, PhD
Florida Atlantic University
e-mail | web site | publications
Daniel J. Levitin, PhD
McGill University
e-mail | web site | publications
Mathias Oechslin
University of Zurich
e-mail | web site
Caroline Palmer, PhD
McGill University, BRAMS
e-mail | web site | publications
Christo Pantev, PhD
University of Münster
e-mail | web site | publications
Aniruddh D. Patel, PhD
The Neurosciences Institute
e-mail | web site | publications
Maria Cristina Saccuman, PhD
University Vita-Salute San Raffaele
publications
Jenny Saffran, PhD
University of Wisconsin
e-mail | web site | publications
Séverine Samson
University of Lille 3
Villeneuve d'Ascq & La Salpêtrière Hospital, Paris
e-mail | web site | publications
Gottfried Schlaug, MD, PhD
Beth Israel Deaconess Medical Center
Harvard Medical School
e-mail | web site | publications
Matthew Schulkind, PhD
Amherst College
e-mail | web site | publications
Robert V. Shannon, PhD
House Ear Institute
e-mail | web site | publications
Joel S. Snyder, PhD
University of Nevada, Las Vegas
e-mail | web site | publications
Karsten Steinhauer, PhD
McGill University
e-mail | web site | publications
Mari Tervaniemi, PhD
University of Helsinki
e-mail | web site | publications
Michael H. Thaut, PhD
Colorado State University
e-mail | web site | publications
William Forde Thompson, PhD
Macquarie University
e-mail | web site | publications
Laurel J. Trainor, PhD
McMaster University & Rotman Research Institute
e-mail | web site | publications
Sandra Trehub, PhD
University of Toronto
e-mail | web site | publications
Patrick C.M. Wong, PhD
Northwestern University
e-mail | web site | publications
Kathleen McGowan
Kathleen McGowan is a freelance magazine writer specializing in science and medicine.
This conference and eBriefing promoted by:
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