Neural networks underlying vocal control in experienced classical singers [Elektronische Ressource] / vorgelegt von Boris Kleber

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Neural networks underlying vocal  
control in experienced  
classical singers 
Dissertation 
der Fakultät für Informa‐t uionds Kognitionswissenschaften   
der Eberha‐rKadrl‐sUniversität Tübinge n
zur Erlangung des Grades ein es 
Doktors der Naturwissesnchafte n 
(Dr. rer. na t.)
vorgelegt von 
Dipl. ‐Psych. Boris Klebe r
aus Stuttga rt
Tübingen 
2009  Tag der mündlichen Qualifikation:   15.07.2009 
Dekan:   Prof. Dr.‐Ing. Oliver Kohlbacher 
1. Berichterstatter:    Prof. Dr. Martin Hautzinger  
2. Berichterstatter:    Prof. Dr. Niels Birbaumer  
  2   
 
 
Meinen Eltern, Heidemarie und Karl Kleber  
  3 Acknowledgements 
First and foremost, I would like  to express my deep est gratitude to Prof. Niels r‐Bi
baumer. I will never forget when we first met at the Stuttgart Opera and he asked me to 
visit him at  his institute in Tübingen, because “we may have a project you could be inte r‐
ested in”. It all came differ ently though. He strongly supported my idea for a new project 
involving singing and allowed me to change over from the BCI  group, when the DFG 
awarded the proposal.  He also  entrusted me with the organization of  a project  the con‐
tent of which  was way beyond  the scope  of this dissertation. I feel deeply honored by his 
faith in me! Ever since, Prof. Birbaumer has  been a great mentor whose door  was always 
open  when his support was needed.  He is not only  an  outstanding scientist but also a 
great  source of inspir ation  that never seem to run dry.  
I am deeply grateful to Prof.  Martin Hautzinger, who  has always been supportive du r‐
ing all stages of this dissertation. His expertise and understanding was indispensable  for 
accomplishing this work.  
Without Prof. John Gru zelier, I would most likely not be  where I am now . He stim u‐
lated and fostered my interest in neuroscience and show ed me  that the sciences and the 
arts can indeed celebrate a successful marriage. I am indebted  to you  for leading me on 
this path! Thank you ever so much!  
I would like to thank Dr. Ralf Veit and Prof. Martin Lotze for  sharing their knowledge 
on  fMRI with me, for  giving me  their invaluable expertise and friendship, and also for 
spending so many hours with me at the sc anner listening to “caro mio ben”! I am sure 
this song will never be forgotten!   
The Institute of Medical Psychology and Behavioral Neurobiology  is a special place 
and there are so many people who are responsible for that.  A big “ thanks” in particular 
goes to PD Dr. Ute Strehl for always being so sup portive and  to Lydia, Angela, Slavica, 
Hannelore, and  Maike.  There are many more, and it was  a privilege to work with  you 
guys!  
  4 Finally, I would like to thank my family!  Although I only had 10 years with my fat her, 
this was a very precious time. He was  of  a rare kind with his warm‐heartedness, his lov‐
ing being, his charisma and  his  unmatched musicality  as a widely respected concert i‐p
anist. He p laced a seed within me  and the plant is still growing. I will  thank you forever !  
There  are no words to express my gratitude to my mother. Your faith in me never failed 
and I  cannot say  enough  how much I thank you for this ! Last but defi nitely not least I 
would like to express my deeply felt thankfulness to Helga, to Mark and  also to Olga !  
 
  5  
 
Eidestattliche Erklärung 
 
Hiermit erkläre ich, dass ich unter Verwendung der im Literaturverzeichnis aufge‐
führten Quellen und unter fachlicher Betreuung diese Dissertation selbstständig verfasst 
habe. 
 
 
(Boris Kleber)  
 
  6 Table of Contents  
Abb reviations..........................................................................................9 
Publications:.........................11 
Overview  12 
Chapter 1   Introduction ..................................................................13 
Chapter 2   Fundamentals of voice physiology........................15 
2.1 Anatomy of the larynx .......... 15 
2.2  Functional anatomy of the vocal system ...... 19 
2.3  Trained and untrained voices................................ ................................ ........................... 25 
Chapter 3   Expertise and deliberate practice.........................27 
Chapter 4   Principles of functional magnetic resonance imaging (fMRI) ......30  
4.1 Introduction................................ ................................ ................................ .............................. 30 
4.2  The physics of MRI ................. 31 
4.3 Functional MRI (fMRI) ................................ ......... 34  
4.4  Experimental designs of fMRI studies ................................ ........... 36 
4.5 Analysis of fMRI data................................ ............ 38 
4.5.1  Spatial and temporal processing ....... 40 
4.5.2  Statistical analyses ................................ ..43  
Chapter 5   Overt and imagined singing of an Italian aria...47 
5.1 Introduction................................ ................................ ................................ .............................. 47 
5.2 Methods ...... 52 
5.2.1  Participants: ............... 52 
5.2.2  Singing Task:................................ ................................ ................................ .............. 52 
5.2.3  fMRI technique: ........ 54 
5.3 Results................................ ......... 56 
5.3.1  Rating of imagery: ...56 
5.3.2  EMG ‐measures................................ ................................ ................................ .......... 56 
5.3.3  Results of fMRI analysis ........................ 56 
5.4 Discussion ................................ ..63 
5.4.1  Motor system: ............ 63 
  7 5.4.2  Areas involved in auditory control:................................ ................................ ..66 
5.4.3  Speech and so ng processing: ............... 68 
5.4.4  Emotional processing: ................................ ............................ 70 
5.5 Conclusions ................................ ............................... 71  
Chapter 6   Experience dependent neural adaptation in trained singers –  
an fMRI study ..............................................................73 
6.1 Introduction ................................ 73 
6.2 Methods ...... 75 
6.2.1  Participants: ................ 75 
6.2.2   Singing Experi ence: ................................ .76 
6.2.3  Singing Task: ................................ .............. 77 
6.2.4   fMRI Technique: ........ 77 
6.2.5  fMRI Data Analysis: .78 
6.3 Results................................ ................................ ................................ ......... 80 
6.3.1  Main effects: ................................ ................ 80 
6.3.2  Conjunction analysis: .............................. 80 
6.3.3  Vocal students versus laymen: ........... 81 
6.3.4  Opera singers versus laymen: ................................ ............. 81 
6.3.5  Op era singers versus vocal students: 81 
6.3.6  Singing practice effects: ................................ ......................... 85 
6.4 Discussion ................................ ..86 
6.4.1  Sensorimotor cortex ................................ ............................... 86 
6.4.2   Auditory corte x:................................ ........ 88 
6.4.3  Sensory association cortex ................... 89 
6.4.4   Performance monitoring ....................... 89 
6.4.5  Cerebellum, basal ganglia and the thalamus................................ ................ 90 
6.5 Conclusions ................................ ................................ ............................... 92  
Chapter 7   Summary........94 
Bibliography.........................................................98 
Appendix 1   ......................................................115  
Appendix 2  120  
 
  8 Abbreviations 
A1  primary auditory cort  ex
AAL  automatic anatomic labeling  
ACC   anterior cingulate cor tex
ACG  anterior cingulate gyr  us
BA  Brodmann area 
BOLD   blood oxygenation level dependent  
CBF  cerebral blood fl ow
cM1  contralateral primary motor cor  tex
df  degrees of freed om
DLPFC   dorsolaterparl efrontal cort ex
EMG   electromyograph y
EPI  echo planar imaging  
FDR   false discovery r ate
fMRI  functional Magnetic Resonance Imaging  
FWE   family wise error r ate
HRF  hemodynamic response function  
HDR   hemodynamic response  
Hipp   hippocampus  
IPC  inferiorpa  rietal cort ex
LI  lateralization inde x
M1  primary motor corte x
MNI  Montreal Neurological Institu te
MRI  magnetic resonance imaging  
MyHC   myosin heavy chain  
NMR   nuclear magnetic resonance  
p   probability 
PFC  prefrontal cort ex
PMC   premotor corte x
RF  radio frequenc y 
ROI  region of interes t
S1  primary somatosensory corte x
SD   standard deviati on
SFG  superior frontal gyr  us
SI I secondary somatosensory corte x
  9 SMA  supplementary motor cortex 
SPL  superior parietal cor tex
SPM  statistical parametrmiacappiln g  
SPS  samples per second  
SPSS  statistical package for the social sci ence
STF  slow twitch muscle fiber s