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Oxford University Press
Handbook of Music and Emotion
Theory, Research, Applications

CHAPTER 12
FUNCTIONAL NEUROIMAGING
Stefan Koelsch, Walter A. Siebel, and Thomas Fritz

Investigating Emotion With Music:
An fMRI Study
Stefan Koelsch,* Thomas Fritz, D. Yves v. Cramon, Karsten Müller,
and Angela D. Friederici
Max Planck Institute for Human Cognitive and Brain Sciences, Leipzig, Germany

Abstract: The present study used pleasant and unpleasant music to evoke emotion and functional magnetic resonance imaging (fMRI) to determine neural correlates of emotion processing. Unpleasant (permanently dissonant) music contrasted with pleasant (consonant) music showed activations of amygdala, hippocampus, parahippocampal gyrus, and temporal poles. These structures have previously been implicated in the emotional processing of stimuli with (negative) emotional valence; the present data show that a cerebral network comprising these structures can be activated during the perception of auditory (musical) information. Pleasant (contrasted to unpleasant) music showed activations of the inferior frontal gyrus (IFG, inferior Brodmann’s area (BA) 44, BA 45, and BA 46), the anterior superior
insula, the ventral striatum, Heschl’s gyrus, and the Rolandic operculum. IFG activations appear to reflect processes of music–syntactic analysis and working memory operations. Activations of Rolandic opercular areas possibly reflect the activation of mirror-function mechanisms during the perception of the pleasant tunes. Rolandic operculum, anterior superior insula, and ventral striatum may form a motor-related circuitry that serves the formation of (premotor) representations for vocal sound production during the perception of pleasant auditory information. In all of the mentioned structures, except the hippocampus, activations increased over time during the presentation of the musical stimuli, indicating that the effects of emotion processing have temporal dynamics; the temporal dynamics of emotion have so far mainly been neglected in the functional imaging literature. Hum Brain Mapp 27:239 –250, 2006.
© 2005 Wiley-Liss, Inc.

    * Correspondence to: Stefan Koelsch, Max-Planck-Institute of Human Cognitive and Brain Sciences, Stephanstr. 1a, 04103 Leipzig, Germany. E-mail: koelsch@cbs.mpg.de
    Received for publication 13 December 2004; Accepted 25 May 2005
    DOI: 10.1002/hbm.20180
    Published online 2 August 2005 in Wiley InterScience (www. interscience.wiley.com).

[ACKNOWLEDGMENTS
The project is part of the Research Group FOR 499 of the German Research Foundation “Acoustic communication of emotions in nonhuman mammals and man: production, perception and neural processing.” We thank W. A. Siebel for help in selecting the stimuli.]

Adults and children processing music:An fMRI study
Stefan Koelsch,a,b,* Thomas Fritz,b Katrin Schulze,b David Alsop,a and Gottfried Schlauga,*
aMusic and Neuroimaging Laboratory, Department of Neurology, Beth Israel Deaconess, Medical Center and Harvard Medical School,
Boston, MA 02215, USA
bMax Planck Institute for Human Cognitive and Brain Sciences, Leipzig, Germany
Received 8 July 2004; revised 25 October 2004; accepted 6 December 2004
Available online 2 March 2005
The present study investigates the functional neuroanatomy of music perception with functional magnetic resonance imaging (fMRI). Three different subject groups were investigated to examine developmental aspects and effects of musical training: 10-year-old children with varying degrees of musical training, adults without formal musical training (nonmusicians), and adult musicians. Subjects made judgements on sequences that ended on chords that were music-syntactically either regular or irregular. In adults, irregular chords activated the
inferior frontal gyrus, orbital frontolateral cortex, the anterior insula, ventrolateral premotor cortex, anterior and posterior areas of the superior temporal gyrus, the superior temporal sulcus, and the supramarginal gyrus. These structures presumably form different networks mediating cognitive aspects of music processing (such as processing of musical syntax and musical meaning, as well as auditory working memory), and possibly emotional aspects of music processing.
In the right hemisphere, the activation pattern of children was similar to that of adults. In the left hemisphere, adults showed larger activations than children in prefrontal areas, in the supramarginal gyrus, and in temporal areas. In both adults and children, musical training was correlated with stronger activations in the frontal operculum and the anterior portion of the superior temporal gyrus.
D 2005 Elsevier Inc. All rights reserved.

www.elsevier.com/locate/ynimg
YNIMG-02951; No. of pages: 9; 4C:
NeuroImage 25 (2005) 1068– 1076

Amygdala activity can be modulated by unexpected chord functions during music listening
Stefan Koelsch(a,b), Thomas Fritz (a) and Gottfried Schlaug (c)
aJunior Research GroupNeurocognition of Music,Max Planck Institute for Human Cognitive and Brain Science, Leipzig,Germany,
bDepartment of Psychology,University of Sussex, Brighton,UK and cDepartment of Neurology,Music andNeuroimaging Laboratory,
Beth Israel Deaconess Medical Center and HarvardMedical School, Boston,USA
Correspondence to Stefan Koelsch, PhD, Leader Independent Junior Research Group,Max-Planck-Institute for Human Cognitive and Brain Sciences, Stephanstr.1a 04103 Leipzig,Germany
Tel: +49 341355 21711; fax: +49 341355 217 30; e-mail: koelsch@cbs.mpg.de
Received14 September 2008; accepted 23 September 2008
DOI:10.1097/WNR.0b013e32831a8722
Numerous earlier studies have investigated the cognitive
processing of musical syntax with regular and irregular chord sequences.
However, irregular sequences may also be perceived as unexpected, and therefore have a di¡erent emotional valence than regular sequences.We provide behavioral data showing that irregular chord functions presented in chord sequence paradigms are perceived as less pleasant than regular sequences. A reanalysis of functional MRI data showed increased blood oxygen leveldependent signal changes bilaterally in the amygdala in response to music-syntactically irregular (compared with regular) chord functions.The combineddata indicate thatmusic-syntactically irregular events elicit brain activity related to emotional processes, and that, in addition to intensely pleasurable music or highly unpleasant music, single chord functions can also modulate amygdala activity.

NeuroReport 19:1815^1819 c 2008 Wolters Kluwer Health | Lippincott Williams &Wilkins.

Towards a neural basis of music-evoked emotions
Stefan Koelsch
Cluster of Excellence ‘‘Languages of Emotion’’, Freie Universität Berlin, Habelschwerdter Allee 45, 14195 Berlin, Germany
Music is capable of evoking exceptionally strong emotions and of reliably affecting the mood of individuals. Functional neuroimaging and lesion studies show that music-evoked emotions can modulate activity in virtually all limbic and paralimbic brain structures. These structures are crucially involved in the initiation, generation, detection, maintenance, regulation and termination of emotions that have survival value for the individual and the species. Therefore, at least some music-evoked emotions involve the very core of evolutionarily adaptive neuroaffective mechanisms. Because dysfunctions in these structures are related to emotional disorders, a better understanding of music-evoked emotions and their neural correlates can lead to amore systematic and effective use of music in therapy.

Trends in Cognitive Sciences Vol.14 No. 3, 131-137

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A Neuroscientific Perspective on Music Therapy
Stefan Koelsch
Department of Psychology, University of Sussex, Falmer, Brighton, United Kingdom
During the last years, a number of studies demonstrated thatmusic listening (and even more so music production) activates a multitude of brain structures involved in cognitive, sensorimotor, and emotional processing. For example, music engages sensory processes, attention, memory-related processes, perception-actionmediation (“mirror neuron system” activity), multisensory integration, activity changes in core areas of emotional processing, processing of musical syntax and musical meaning, and social cognition. It is likely that the engagement of these processes by music can have beneficial effects on the psychological and physiological health of individuals, although the mechanisms underlying such effects are currently not well understood. This article gives a brief overview of factors contributing to the effects of music-therapeutic work. Then, neuroscientific studies using music to investigate emotion, perception-action mediation (“mirror function”), and social cognition are reviewed, including illustrations of the relevance of these domains for music therapy.

The Neurosciences and Music III—Disorders and Plasticity: Ann. N.Y. Acad. Sci. 1169: 374–384 (2009).
doi: 10.1111/j.1749-6632.2009.04592.x c 2009 New York Academy of Sciences.

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