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PhD examination
Myeloarchitecture and Resting State Functional
Connectivity of Auditory Cortex in Musicians with
Absolute Pitch
Seung-Goo (ā€œSGā€) Kim
Max Planck Institute for Human Cognitive and Brain Sciences
5 Jan 2017
Institute for Psychologyā€Ø
Faculty of Biosciences, Pharmacy, and Psychologyā€Ø
University of Leipzig
1
Overview
ā€¢ Introduction: Absolute pitch (AP)
ā€¢ Study #1: In-vivo cortical myelin mapping
ā€¢ Study #2: Resting-state functional connectivity
ā€¢ Discussion: Distributed processing of AP
2
Introduction
Absolute pitch
3
What is absolute pitch (AP)?
ā€¢ Rare ability to recognize the name pitch without an external
reference [1].
ā€¢ Related to early musical experience (4~7 year-old [2]) and a
certain training method (ā€œļ¬xed-Doā€ than ā€œmovable-Doā€) [3].
ā€¢ Genetic factor: more APs in APā€™s family [2]; ā€Ø
Europeans (7%) vs. East-Asian (30%) in musicians [4].
ā€¢ Sometimes make mistakes in octave recognition [1,4]: ā€Ø
good at pitch chroma rather than pitch height.
[1] Miyazaki, 2004. [2] Baharloo et al., 1998. [3] Willson et al., 2012. [4] Miyazaki et al., 2012.
4
What is pitch chroma?
ā€¢ ā€œC3ā€
ā€¢ Pitch chroma:ā€Ø
Alphabetical label of 12
divisions of an octave
(C, C#, D, D#, ā€¦)
ā€¢ Pitch height: ā€Ø
Octave index
5
B2 C3 D3 E3 F3 G3 A3 B3 C4
C#3 D#3 F#3 G#3 A#3 C#4A#2
GarageBand, (C) Apple, Inc.
The 3rd octave The 4thThe 2nd
AP-test confusion matrix
Kim & Knƶsche, 2016. 6
ā€¢ Perceptual processā€Ø
Categorization of pitch chroma
ā€¢ Associative processā€Ø
Verbal/non-verbal labeling
Dichotomy of AP recognition
Levitin & Rogers, 2005. 7
Categorization
Labeling
ā€¢ Temporal lobe (perceptual proc.)ā€Ø
leftward asymmetry of planum temporale (PT)
area [1]; higher activation in the left PT [2,3].
ā€¢ Frontal lobe (associative proc.)ā€Ø
thicker inferior frontal gyrus (IFG) [4]; higher
activation in right dorsolateral prefrontal cortex
(DLPFC) [3-5].
ā€¢ Frontotemporal connectivityā€Ø
phase synchrony [6]; fractional anisotropy of
superior longitudinal fasiculus (SLF) [7].
Previous neuroimaging ļ¬ndings
DLPFC
HG
PT
http://clipartbest.com/
[1] Keenan et al., 2001. [2] Ohnishi et al., 2001. [3] Willson et al., 2009. [4] Dohn et al., 2015.ā€Ø
[5] Zatorre et al., 1998. [6] Elmer et al., 2015. [7] Oechslin et al., 2009.
8
HG: Heschlā€™s gyrus
But how does the AP work?
ā€¢ How does the cortical regions and the connectivity
between them contribute to the AP recognition?
> Multimodal imaging describing microscopic
architecture might give us better ideas to construct
a framework to explain how AP works.
9
Myeloarchitecture of auditory
cortex in musicians with AP
Study #1: Relaxometry at 7-T
10Kim & Knƶsche, 2016, Human Brain Mapping.
Intracortical myelination
ā€¢ Increases information transfer speed
(but not very much for short-range
connections in gray matter)
ā€¢ May reduce cross-talk between nearby
axons, thus increasing speciļ¬city
ā€¢ Prevents neuroplasticity after critical
period of neurodevelopment (McGee
et al., 2005)
Vogt, 1903
Cyto-
architecture
Myelo-
architecture
11
Research questions
ā€¢ Does the degree of cortical myelination in auditory
cortex (related to the perceptual subprocess of AP)
correlate with the acuity of AP?
ā€¢ Is myelo-correlate of AP diļ¬€erentiable from that of
the other psychoacoustic characteristics (e.g.,
frequency discrimination threshold)?
12
Participants & imaging
ā€¢ Participants
ā€¢ 8 AP musicians & 9 non-AP musicians categorized
based on behavioral test of AP
ā€¢ Cortical myelin mapping (7-T)
ā€¢ MP2RAGE sequence for quantitative mapping of
longitudinal relaxation rate (qR1), which strongly
correlates with myelin content in the cortex
13
In-vivo cortical myelin mapping
Kim & Knƶsche, 2016. 14
General linear model
ā€¢ x was either:
ā€¢ Absolute pitch group index
ā€¢ Absolute pitch score (APS) = 1 - scaled ([0,1]) error,
ā€¢ Frequency discrimination threshold (FDT) = -log10(Fm/F0)
15
qR1
= Ī²0
+ageĪ²1
+ sexĪ²2
+ethnicityĪ²3
+ xĪ²4
+Īµ
Absolute pitch score
HG
PT
STS
PP
STG
Frequency ā€Ø
discrimination threshold
STG
HG
PT
STS
PP
Kim & Knƶsche, 2016. 16
Concave
Convex
Pitch chroma vs. pitch height
Warren et al., 2003.17
Planum
polare
Planum
temporale
Primary auditory
cortex (overlap)
Chroma
Height
Kim & Knƶsche, 2016.
Dual pathway hypothesis (Rauschecker et al., 1995)
ā€¢ Ventral (ā€œwhatā€): vs. dorsal (ā€œwhereā€) auditory pathways
ā€¢ Spatially separated sensitivity to non-spatial (i.e. pitch) vs.
spatial (i.e. location) information
Barret & Hall, 2006. 18
ā€¢ Absolute recognition of pitch
chroma (rather than height)
ā€¢ Cortical myelin can increase
speciļ¬city and suppress
neuroplasticity
ā€¢ Dual pathway hypothesis: non-
spatial information (e.g.
phoneme) via ventral pathway.
ā€¢ Conjecture: Categorization of
pitch chroma along the ventral
pathway?
Categorical processing & ventral pathway
DLPFC
PT
HG
http://clipartbest.com/
19
PP
VLPFC
Summary of Study #1
ā€¢ We found: greater cortical myelin in the right
planum polare (PP) in musicians with than without
absolute pitch.
ā€¢ We think: the right PP and its connection (ventral
pathway) may be important in chromatic
categorization.
20
Resting-state functional connectivity of
the ventral pathway in musicians with AP
Study #2: Resting-state fMRI at 3-T
21Kim & Knƶsche, 2016, In preparation.
Spontaneity of AP
ā€¢ Unintended recognition of pitch chroma of
environmental sound (e.g., cluttering of glasses, siren
of ambulance) [1]
ā€¢ Uncontrollable and instant perception (e.g., Stroop-
like eļ¬€ect for incongruent pitch and pitch name [2,3])
ā€¢ Spontaneous & automatic processing of AP could be
reļ¬‚ected in BOLD signal during resting
[1] Miyazaki, 2004. [2] Itoh et al., 2005. [3] Schultz et al., 2013.
22
Research questions
ā€¢ Is the planum polare (PP) strongly
coupled with other cortical regions
that belong to the ventral pathway
in musicians with AP?
ā€¢ What about the planum temporale
(PT) and the dorsal pathway?
23
HG
DLPFC
PT
http://clipartbest.com/
VLPFC
PP
HG: Heschlā€™s gyrus
Participants & imaging
ā€¢ Same participants: 8 AP musicians & 9 non-AP musicians
ā€¢ Whole-brain functional images at 3-T
ā€¢ Echo-planer imaging; 2.3-mm isovoxel; TR = 1.4 s; for
9.8-min resting
ā€¢ Anatomical image at 3-T
ā€¢ MPRAGE; 1-mm isovoxel
24
Zero-lag cross-correlation
Kim & Knƶsche, In prep.25
Kim & Knƶsche, In prep.26
ā€¢ Frequency-
dependent
connectivity
(regardless of
time lag)
ā€¢ Averaged into ā€Ø
9 frequency bins
with 50% overlap:
[0, 20], ā€Ø
[10, 30], ā€Ø
[20, 40], ā€¦,ā€Ø
[90, 110] mHz
Cross-coherence
General linear model
ā€¢ FC was either:
ā€¢ cross-correlation
ā€¢ cross-coherenceā€Ø
ā€Ø
ā€¢ x was either:
ā€¢ AP-group index (AP)
ā€¢ AP score (APS)
27
FC = Ī²0
+ motionĪ²1
+ ethnicityĪ²2
+ xĪ²3
+ Īµ
AP/APS eļ¬€ect in correlation
28
Kim&Knƶsche,inprep.
AP/APS eļ¬€ect in coherence
Kim&Knƶsche,inprep.
29
Greater RSFC of the right PP
ā€¢ Ipsilateral STG, contralateral PP/STG: ā€Ø
adjacent/homologous regions via bilateral
ventral streams [1]
ā€¢ Bilateral STS: specialized auditory object
recognition; e.g. voice recognition [2]
ā€¢ Left IFG: verbal labeling of pitch chroma [3]
ā€¢ Bilateral ACC, medial SFG: default-mode
network [4]
[1] Upadhyay et al., 2008 [2] Belin et al., 2004. [3] Wengenroth et al., 2013. [4] Smith et al., 2009.
30
What about the left dorsal pathway?
31
ā€¢ Leftward asymmetry of planum temporale (PT) [1]
ā€¢ Greater rCBF/BOLD in the left PT during passive
listening [2,3]
ā€¢ Phase synchrony between PT and dorsal lateral
prefrontal cortex (DLPFC) [4], superior
longitudinal fasiculus [5]
ā˜…But not conclusive: the other studies also found
volume/thickness/FA increase in the right STG/
ILF [6,7].
[1] Schlaug et al., 1995. [2] Ohnishi et al., 2001. [3] Wilson et al., 2009. [4] Elmer et al., 2015.ā€Ø
[5] Oechslin et al., 2009. [6] Wengenroth et al., 2013. [7] Dohn et al., 2015.
-log10minP
RSFC seeding from planum temporale
Kim & Knƶsche, In prep.32
Summary of Study #2
ā€¢ We found: greater resting-state functional
connectivity (RSFC) of the the right planum polare
(PP) with the anteroventral auditory network and
the anteromedial network in musicians with
absolute pitch (AP).
ā€¢ We think: the ventral auditory pathway via the right
PP may be crucial in AP processing (presumably in
chromatic categorization).
33
Discussion
Distributed processing of AP
34
Main ļ¬ndings
ā€¢ Greater cortical myelin in the right planum
polare (PP) in musicians with absolute pitch
ā€¢ Greater RSFC of the right PP in
ā€¢ the right STG, the left PP/STG,
ā€¢ the bilateral STS,
ā€¢ the left IFG, and
ā€¢ the bilateral ACC/SFG
ā€¢ No diļ¬€erence in the left planum temporale.
35
Distributed processing of AP
Left dorsal pathway?
(spatial processing?)
Kim & Knƶsche, In prep.36
Future directions
ā€¢ Topological organization for pitch chroma in APs
ā€¢ Model-based encoding fMRI analysis
ā€¢ Dynamics of frontotemporal network
ā€¢ Eļ¬€ective connectivity in multimodal data
(fMRI+MEG)
37
Thank you for attention!
38

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Myeloarchitecture and Intrinsic Functional Connectivity of Auditory Cortex in Musicians with Absolute Pitch

  • 1. PhD examination Myeloarchitecture and Resting State Functional Connectivity of Auditory Cortex in Musicians with Absolute Pitch Seung-Goo (ā€œSGā€) Kim Max Planck Institute for Human Cognitive and Brain Sciences 5 Jan 2017 Institute for Psychologyā€Ø Faculty of Biosciences, Pharmacy, and Psychologyā€Ø University of Leipzig 1
  • 2. Overview ā€¢ Introduction: Absolute pitch (AP) ā€¢ Study #1: In-vivo cortical myelin mapping ā€¢ Study #2: Resting-state functional connectivity ā€¢ Discussion: Distributed processing of AP 2
  • 4. What is absolute pitch (AP)? ā€¢ Rare ability to recognize the name pitch without an external reference [1]. ā€¢ Related to early musical experience (4~7 year-old [2]) and a certain training method (ā€œļ¬xed-Doā€ than ā€œmovable-Doā€) [3]. ā€¢ Genetic factor: more APs in APā€™s family [2]; ā€Ø Europeans (7%) vs. East-Asian (30%) in musicians [4]. ā€¢ Sometimes make mistakes in octave recognition [1,4]: ā€Ø good at pitch chroma rather than pitch height. [1] Miyazaki, 2004. [2] Baharloo et al., 1998. [3] Willson et al., 2012. [4] Miyazaki et al., 2012. 4
  • 5. What is pitch chroma? ā€¢ ā€œC3ā€ ā€¢ Pitch chroma:ā€Ø Alphabetical label of 12 divisions of an octave (C, C#, D, D#, ā€¦) ā€¢ Pitch height: ā€Ø Octave index 5 B2 C3 D3 E3 F3 G3 A3 B3 C4 C#3 D#3 F#3 G#3 A#3 C#4A#2 GarageBand, (C) Apple, Inc. The 3rd octave The 4thThe 2nd
  • 6. AP-test confusion matrix Kim & Knƶsche, 2016. 6
  • 7. ā€¢ Perceptual processā€Ø Categorization of pitch chroma ā€¢ Associative processā€Ø Verbal/non-verbal labeling Dichotomy of AP recognition Levitin & Rogers, 2005. 7 Categorization Labeling
  • 8. ā€¢ Temporal lobe (perceptual proc.)ā€Ø leftward asymmetry of planum temporale (PT) area [1]; higher activation in the left PT [2,3]. ā€¢ Frontal lobe (associative proc.)ā€Ø thicker inferior frontal gyrus (IFG) [4]; higher activation in right dorsolateral prefrontal cortex (DLPFC) [3-5]. ā€¢ Frontotemporal connectivityā€Ø phase synchrony [6]; fractional anisotropy of superior longitudinal fasiculus (SLF) [7]. Previous neuroimaging ļ¬ndings DLPFC HG PT http://clipartbest.com/ [1] Keenan et al., 2001. [2] Ohnishi et al., 2001. [3] Willson et al., 2009. [4] Dohn et al., 2015.ā€Ø [5] Zatorre et al., 1998. [6] Elmer et al., 2015. [7] Oechslin et al., 2009. 8 HG: Heschlā€™s gyrus
  • 9. But how does the AP work? ā€¢ How does the cortical regions and the connectivity between them contribute to the AP recognition? > Multimodal imaging describing microscopic architecture might give us better ideas to construct a framework to explain how AP works. 9
  • 10. Myeloarchitecture of auditory cortex in musicians with AP Study #1: Relaxometry at 7-T 10Kim & Knƶsche, 2016, Human Brain Mapping.
  • 11. Intracortical myelination ā€¢ Increases information transfer speed (but not very much for short-range connections in gray matter) ā€¢ May reduce cross-talk between nearby axons, thus increasing speciļ¬city ā€¢ Prevents neuroplasticity after critical period of neurodevelopment (McGee et al., 2005) Vogt, 1903 Cyto- architecture Myelo- architecture 11
  • 12. Research questions ā€¢ Does the degree of cortical myelination in auditory cortex (related to the perceptual subprocess of AP) correlate with the acuity of AP? ā€¢ Is myelo-correlate of AP diļ¬€erentiable from that of the other psychoacoustic characteristics (e.g., frequency discrimination threshold)? 12
  • 13. Participants & imaging ā€¢ Participants ā€¢ 8 AP musicians & 9 non-AP musicians categorized based on behavioral test of AP ā€¢ Cortical myelin mapping (7-T) ā€¢ MP2RAGE sequence for quantitative mapping of longitudinal relaxation rate (qR1), which strongly correlates with myelin content in the cortex 13
  • 14. In-vivo cortical myelin mapping Kim & Knƶsche, 2016. 14
  • 15. General linear model ā€¢ x was either: ā€¢ Absolute pitch group index ā€¢ Absolute pitch score (APS) = 1 - scaled ([0,1]) error, ā€¢ Frequency discrimination threshold (FDT) = -log10(Fm/F0) 15 qR1 = Ī²0 +ageĪ²1 + sexĪ²2 +ethnicityĪ²3 + xĪ²4 +Īµ
  • 16. Absolute pitch score HG PT STS PP STG Frequency ā€Ø discrimination threshold STG HG PT STS PP Kim & Knƶsche, 2016. 16 Concave Convex
  • 17. Pitch chroma vs. pitch height Warren et al., 2003.17 Planum polare Planum temporale Primary auditory cortex (overlap) Chroma Height Kim & Knƶsche, 2016.
  • 18. Dual pathway hypothesis (Rauschecker et al., 1995) ā€¢ Ventral (ā€œwhatā€): vs. dorsal (ā€œwhereā€) auditory pathways ā€¢ Spatially separated sensitivity to non-spatial (i.e. pitch) vs. spatial (i.e. location) information Barret & Hall, 2006. 18
  • 19. ā€¢ Absolute recognition of pitch chroma (rather than height) ā€¢ Cortical myelin can increase speciļ¬city and suppress neuroplasticity ā€¢ Dual pathway hypothesis: non- spatial information (e.g. phoneme) via ventral pathway. ā€¢ Conjecture: Categorization of pitch chroma along the ventral pathway? Categorical processing & ventral pathway DLPFC PT HG http://clipartbest.com/ 19 PP VLPFC
  • 20. Summary of Study #1 ā€¢ We found: greater cortical myelin in the right planum polare (PP) in musicians with than without absolute pitch. ā€¢ We think: the right PP and its connection (ventral pathway) may be important in chromatic categorization. 20
  • 21. Resting-state functional connectivity of the ventral pathway in musicians with AP Study #2: Resting-state fMRI at 3-T 21Kim & Knƶsche, 2016, In preparation.
  • 22. Spontaneity of AP ā€¢ Unintended recognition of pitch chroma of environmental sound (e.g., cluttering of glasses, siren of ambulance) [1] ā€¢ Uncontrollable and instant perception (e.g., Stroop- like eļ¬€ect for incongruent pitch and pitch name [2,3]) ā€¢ Spontaneous & automatic processing of AP could be reļ¬‚ected in BOLD signal during resting [1] Miyazaki, 2004. [2] Itoh et al., 2005. [3] Schultz et al., 2013. 22
  • 23. Research questions ā€¢ Is the planum polare (PP) strongly coupled with other cortical regions that belong to the ventral pathway in musicians with AP? ā€¢ What about the planum temporale (PT) and the dorsal pathway? 23 HG DLPFC PT http://clipartbest.com/ VLPFC PP HG: Heschlā€™s gyrus
  • 24. Participants & imaging ā€¢ Same participants: 8 AP musicians & 9 non-AP musicians ā€¢ Whole-brain functional images at 3-T ā€¢ Echo-planer imaging; 2.3-mm isovoxel; TR = 1.4 s; for 9.8-min resting ā€¢ Anatomical image at 3-T ā€¢ MPRAGE; 1-mm isovoxel 24
  • 25. Zero-lag cross-correlation Kim & Knƶsche, In prep.25
  • 26. Kim & Knƶsche, In prep.26 ā€¢ Frequency- dependent connectivity (regardless of time lag) ā€¢ Averaged into ā€Ø 9 frequency bins with 50% overlap: [0, 20], ā€Ø [10, 30], ā€Ø [20, 40], ā€¦,ā€Ø [90, 110] mHz Cross-coherence
  • 27. General linear model ā€¢ FC was either: ā€¢ cross-correlation ā€¢ cross-coherenceā€Ø ā€Ø ā€¢ x was either: ā€¢ AP-group index (AP) ā€¢ AP score (APS) 27 FC = Ī²0 + motionĪ²1 + ethnicityĪ²2 + xĪ²3 + Īµ
  • 28. AP/APS eļ¬€ect in correlation 28 Kim&Knƶsche,inprep.
  • 29. AP/APS eļ¬€ect in coherence Kim&Knƶsche,inprep. 29
  • 30. Greater RSFC of the right PP ā€¢ Ipsilateral STG, contralateral PP/STG: ā€Ø adjacent/homologous regions via bilateral ventral streams [1] ā€¢ Bilateral STS: specialized auditory object recognition; e.g. voice recognition [2] ā€¢ Left IFG: verbal labeling of pitch chroma [3] ā€¢ Bilateral ACC, medial SFG: default-mode network [4] [1] Upadhyay et al., 2008 [2] Belin et al., 2004. [3] Wengenroth et al., 2013. [4] Smith et al., 2009. 30
  • 31. What about the left dorsal pathway? 31 ā€¢ Leftward asymmetry of planum temporale (PT) [1] ā€¢ Greater rCBF/BOLD in the left PT during passive listening [2,3] ā€¢ Phase synchrony between PT and dorsal lateral prefrontal cortex (DLPFC) [4], superior longitudinal fasiculus [5] ā˜…But not conclusive: the other studies also found volume/thickness/FA increase in the right STG/ ILF [6,7]. [1] Schlaug et al., 1995. [2] Ohnishi et al., 2001. [3] Wilson et al., 2009. [4] Elmer et al., 2015.ā€Ø [5] Oechslin et al., 2009. [6] Wengenroth et al., 2013. [7] Dohn et al., 2015.
  • 32. -log10minP RSFC seeding from planum temporale Kim & Knƶsche, In prep.32
  • 33. Summary of Study #2 ā€¢ We found: greater resting-state functional connectivity (RSFC) of the the right planum polare (PP) with the anteroventral auditory network and the anteromedial network in musicians with absolute pitch (AP). ā€¢ We think: the ventral auditory pathway via the right PP may be crucial in AP processing (presumably in chromatic categorization). 33
  • 35. Main ļ¬ndings ā€¢ Greater cortical myelin in the right planum polare (PP) in musicians with absolute pitch ā€¢ Greater RSFC of the right PP in ā€¢ the right STG, the left PP/STG, ā€¢ the bilateral STS, ā€¢ the left IFG, and ā€¢ the bilateral ACC/SFG ā€¢ No diļ¬€erence in the left planum temporale. 35
  • 36. Distributed processing of AP Left dorsal pathway? (spatial processing?) Kim & Knƶsche, In prep.36
  • 37. Future directions ā€¢ Topological organization for pitch chroma in APs ā€¢ Model-based encoding fMRI analysis ā€¢ Dynamics of frontotemporal network ā€¢ Eļ¬€ective connectivity in multimodal data (fMRI+MEG) 37
  • 38. Thank you for attention! 38