This Accepted Manuscript has not been copyedited and formatted. The final version may differ from this version.
Research Articles: Behavioral/Cognitive
Cooperation not competition: bihemispheric tDCS and fMRI show role for
ipsilateral hemisphere in motor learning
Sheena Waters
1,2,3
1
, Tobias Wiestler and Jörn Diedrichsen
1,4
1
Institute of Cognitive Neuroscience, University College London, London, WC1 3AR, UK
2
Institute of Neurology, University College London, London, WC1 3AR, UK
3
Sobell Department for Motor Neuroscience and Motor Disorders, University College London, London, WC1
3AR, UK
4
Brain and Mind Institute, Western University, London ON, N6A 5B7, Canada
DOI: 10.1523/JNEUROSCI.3414-16.2017
Received: 3 November 2016
Revised: 1 April 2017
Accepted: 17 April 2017
Published: 3 July 2017
Author contributions: S.W.-M., T.W., and J.D. designed research; S.W.-M., T.W., and J.D. performed
research; S.W.-M., T.W., and J.D. analyzed data; S.W.-M. and J.D. wrote the paper.
Conflict of Interest: The authors declare no competing financial interests.
The research was supported by grants from the Wellcome trust (094874/Z/10/Z) and James McDonnell
foundation, both to JD, and a PhD studentship from the Brain Research Trust (6CHB) to SW. The Wellcome
Trust Centre for Neuroimaging is supported by core funding from the Wellcome Trust 091593/Z/10/Z).
Corresponding author: Jörn Diedrichsen, Brain and Mind Institute, Western University, N6A 5B7, London, ON,
Canada, Email: [email protected]
Cite as: J. Neurosci ; 10.1523/JNEUROSCI.3414-16.2017
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Accepted manuscripts are peer-reviewed but have not been through the copyediting, formatting, or proofreading
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Copyright © 2017 Waters et al.
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Cooperation not competition: bihemispheric tDCS and fMRI show role for ipsilateral
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hemisphere in motor learning
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Short title: Role of ipsilateral hemisphere in motor learning
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Sheena
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Waters1,2,3,
Tobias Wiestler1 , Jörn Diedrichsen1,4
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Corresponding author:
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Acknowledgements
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ƒ‰—Ž› ‡– ƒŽǤǡ ʹͲͲͻȌ •–—†‹‡• Šƒ˜‡ †‡‘•–”ƒ–‡† –Šƒ– ƒ…–‹˜‹–› ‹ ͳȄ
ϵϲ
ƒŽ–Š‘—‰Š ‘ˆ–‡ •—’’”‡••‡† „‡Ž‘™ „ƒ•‡Ž‹‡Ȅ…‘–ƒ‹• ‹ˆ‘”ƒ–‹‘ ƒ„‘—– ‘‰‘‹‰ ‹’•‹Žƒ–‡”ƒŽ
ϵϳ
‘˜‡‡–•Ǥ Š‹• ƒ…–‹˜‹–› …‘—Ž† …‘–”‹„—–‡ –‘ ‘˜‡‡– …‘–”‘Ž ‡‹–Š‡” –Š”‘—‰Š †‹”‡…–Ž›
ϵϴ
†‡•…‡†‹‰ ‹’•‹Žƒ–‡”ƒŽ ’”‘Œ‡…–‹‘•ǡ ‘” „› •Šƒ’‹‰ ƒ…–‹˜ƒ–‹‘ ’ƒ––‡”• ‘ –Š‡ …‘–”ƒŽƒ–‡”ƒŽ •‹†‡
ϵϵ
ȋ‡”•–›‡‡–ƒŽǤǡʹͲͲͷȌǤ……‘”†‹‰–‘–Š‹•Ǯ‹–‡”Š‡‹•’Š‡”‹……‘‘’‡”ƒ–‹‘ǯ‘†‡Žǡ„‹Š‡‹•’Š‡”‹…
ϭϬϬ
– ‹• ‘”‡ ‡ˆˆ‡…–‹˜‡ –Šƒ —‹Š‡‹•’Š‡”‹… •–‹—Žƒ–‹‘ǡ ‘– „‡…ƒ—•‡ ‹– •‹Ž‡…‡• ‹’•‹Žƒ–‡”ƒŽ
ϭϬϭ
…‘”–‡šǡ„—–„‡…ƒ—•‡‹–‹…”‡ƒ•‡•‹’•‹Žƒ–‡”ƒŽ’Žƒ•–‹…‹–›Ǥ
ϯ
d,ZK>K&/W^/>dZ>KZdy/EDKdKZ>ZE/E' ϰ
ϭϬϮ
‘ƒ†Œ—†‹…ƒ–‡„‡–™‡‡–Š‡•‡‡š’Žƒƒ–‹‘•ǡ™‡–‡•–‡†–Š‡‡ˆˆ‡…–•‘ˆ”‡˜‡”•‹‰–Š‡’‘Žƒ”‹–›
ϭϬϯ
‘ˆ „‹Š‡‹•’Š‡”‹… •–‹—Žƒ–‹‘Ǥ ‡ –”ƒ‹‡† ͸Ͷ ’ƒ”–‹…‹’ƒ–• ‹ ƒ †‘—„Ž‡Ǧ„Ž‹† •–—†›ǡ ‡‹–Š‡” —•‹‰
ϭϬϰ
•Šƒǡ —‹Š‡‹•’Š‡”‹…ǡ …‘˜‡–‹‘ƒŽ „‹Š‡‹•’Š‡”‹…ǡ ‘” ”‡˜‡”•‡†Ǧ’‘Žƒ”‹–› „‹Š‡‹•’Š‡”‹… –
ϭϬϱ
ȋFig. 1aȌǤƒ”–‹…‹’ƒ–•’‡”ˆ‘”‡†ƒ—‹ƒ—ƒŽ•‡“—‡…‡–ƒ•™‹–Š‡‹–Š‡”Ž‡ˆ–‘””‹‰Š–Šƒ†‘˜‡”
ϭϬϲ
ˆ‘—” †ƒ›• ȋFig. 1bȌǤ ˆ–‡” –”ƒ‹‹‰ ƒ† ƒ „‡Šƒ˜‹‘”ƒŽ ’‘•–Ǧ–‡•– ™‹–Š‘—– –ǡ ’ƒ”–‹…‹’ƒ–•
ϭϬϳ
—†‡”™‡–ˆ–‘‡Ž—…‹†ƒ–‡–Š‡‡—”ƒŽ…Šƒ‰‡•‹†—…‡†„›•–‹—Žƒ–‹‘Ǥ
ϭϬϴ
Š‡ ‹–‡”Š‡‹•’Š‡”‹… …‘’‡–‹–‹‘ ‘†‡Ž ȋƒ••—‹‰ ’‘Žƒ”‹–›Ǧ•’‡…‹ˆ‹… – ‡ˆˆ‡…–•ǡ •‡‡
ϭϬϵ
Table 1 ˆ‘” ‘–Š‡” ƒ••—’–‹‘•Ȍ ’”‡†‹…–• –Šƒ– ”‡˜‡”•‡†Ǧ’‘Žƒ”‹–› „‹Š‡‹•’Š‡”‹… – •Š‘—Ž†
ϭϭϬ
‹’ƒ‹” ’‡”ˆ‘”ƒ…‡ …‘’ƒ”‡† –‘ –Š‡ •Šƒ ‰”‘—’Ǥ ‘– ‘Ž› •Š‘—Ž† …‘–”ƒŽƒ–‡”ƒŽ …ƒ–Š‘†ƒŽ
ϭϭϭ
•–‹—Žƒ–‹‘ •—’’”‡•• –Š‡ ‘–‘” ƒ”‡ƒ• ‹˜‘Ž˜‡† ‹ Ž‡ƒ”‹‰ǡ „—– ‹’•‹Žƒ–‡”ƒŽ ƒ‘†ƒŽ •–‹—Žƒ–‹‘
ϭϭϮ
•Š‘—Ž†ˆ—”–Š‡”‹…”‡ƒ•‡‹–‡”Š‡‹•’Š‡”‹…‹Š‹„‹–‹‘Ǥˆ•Š‘—Ž†”‡˜‡ƒŽ‘’’‘•‹–‡…Šƒ‰‡•‹–Š‡
ϭϭϯ
Š‡‹•’Š‡”‡–Šƒ–”‡…‡‹˜‡†ƒ‘†ƒŽƒ†…ƒ–Š‘†ƒŽ•–‹—Žƒ–‹‘Ǥ
ϭϭϰ
‘–”ƒ•–‹‰Ž›ǡ –Š‡ ‹–‡”Š‡‹•’Š‡”‹… …‘‘’‡”ƒ–‹‘ ‘†‡Ž ȋƒ••—‹‰ ’‘Žƒ”‹–› —•’‡…‹ˆ‹…
ϭϭϱ
– ‡ˆˆ‡…–•Ȍ ’”‡†‹…–• –Šƒ– ’Žƒ•–‹…‹–› ‹• ‹†—…‡† ”‡‰ƒ”†Ž‡•• ‘ˆ …—””‡– †‹”‡…–‹‘ǡ ƒ† –Šƒ– –Š‡
ϭϭϲ
„‡Šƒ˜‹‘”ƒŽ ƒ†˜ƒ–ƒ‰‡ ƒ”‹•‡• ˆ”‘ ’Žƒ•–‹…‹–› ‹ „‘–Š Š‡‹•’Š‡”‡•Ǥ Š‡”‡ˆ‘”‡ǡ ”‡˜‡”•‡†Ǧ’‘Žƒ”‹–›
ϭϭϳ
– •Š‘—Ž† „‡ ƒ• ‡ˆˆ‡…–‹˜‡ ƒ• …‘˜‡–‹‘ƒŽ „‹Š‡‹•’Š‡”‹… –ǡ ƒ† „‘–Š •Š‘—Ž† „‡ ‘”‡
ϭϭϴ
‡ˆˆ‡…–‹˜‡ –Šƒ –Š‡ —‹Š‡‹•’Š‡”‹… ‘–ƒ‰‡Ǥ ‹˜‡ –Š‡ „‹Žƒ–‡”ƒŽ ƒ–—”‡ ‘ˆ –Š‡ ’”‡†‹…–‡† ’Žƒ•–‹…
ϭϭϵ
…Šƒ‰‡•ǡ –Š‡ –Ǧ”‡Žƒ–‡† ƒ†˜ƒ–ƒ‰‡• •Š‘—Ž† ‰‡‡”ƒŽ‹œ‡ –‘ –Š‡ —–”ƒ‹‡† Šƒ†Ǥ ‹ƒŽŽ›ǡ
ϭϮϬ
„‹Š‡‹•’Š‡”‹…–•Š‘—Ž†Ž‡ƒ†–‘ƒ…–‹˜‹–›…Šƒ‰‡•‹„‘–ŠŠ‡‹•’Š‡”‡•‹ƒ’‘Žƒ”‹–›Ǧ—•’‡…‹ˆ‹…
ϭϮϭ
ƒ‡”Ǥ
ϭϮϮ
Materials and methods
ϭϮϯ
ϭϮϰ
Participants
ϭϮϱ
‹š–›Ǧˆ‘—” Š‡ƒŽ–Š› ”‹‰Š–ǦŠƒ†‡† •—„Œ‡…–• ȋͷͶǤ͸ͻΨ ˆ‡ƒŽ‡•Ǣ ƒ˜‡”ƒ‰‡ ƒ‰‡ ʹʹǤͺͶ ά ͲǤͷ͸ ›‡ƒ”•Ȍ
ϭϮϲ
’ƒ”–‹…‹’ƒ–‡† ‹ –Š‹• •–—†›Ǥ ƒ”–‹…‹’ƒ–• …‘’Ž‡–‡† –Š‡ †‹„—”‰Š ƒ†‡†‡•• ˜‡–‘”›
ϭϮϳ
ȋŽ†ˆ‹‡Ž†ǡ ͳͻ͹ͳȌǡ ƒ• ™‡ŽŽ ƒ• ƒ •—”˜‡› ‘ˆ ‡†‹…ƒŽ Š‹•–‘”›ǡ —•‹…ƒŽ ƒ† …‘’—–‡” ‰ƒ‹‰
ϭϮϴ
‡š’‡”‹‡…‡ǡ ƒ† ’”‡˜‹‘—• ‡š’‘•—”‡ –‘ „”ƒ‹ •–‹—Žƒ–‹‘Ǥ š…Ž—•‹‘ …”‹–‡”‹ƒ ˆ‘” ’ƒ”–‹…‹’ƒ–‹‘
ϭϮϵ
™‡”‡ƒ•†‡–ƒ‹Ž‡†’”‡˜‹‘—•Ž›ȋƒ–‡”•Ǧ‡–‡‹‡”‡–ƒŽǤǡʹͲͳͶȌǤƒ”–‹…‹’ƒ–•‰ƒ˜‡™”‹––‡‹ˆ‘”‡†
ϭϯϬ
…‘•‡–‹ƒ……‘”†ƒ…‡–‘–Š‡‡…Žƒ”ƒ–‹‘‘ˆ‡Ž•‹‹ǡƒ†”‡…‡‹˜‡†ˆ‹ƒ…‹ƒŽ”‡—‡”ƒ–‹‘™‹–Š
ϭϯϭ
–Š‡’‘••‹„‹Ž‹–›‘ˆƒ††‹–‹‘ƒŽ„‘—•‡•ˆ‘”…‘’Ž‡–‹‰–Š‡•–—†›ƒ†’‡”ˆ‘”ƒ…‡‹’”‘˜‡‡–•Ǥ
ϭϯϮ
Š‡’”‘–‘…‘Ž™ƒ•ƒ’’”‘˜‡†„›–Š‡‡•‡ƒ”…Š–Š‹…•‘‹––‡‡Ǥ
ϭϯϯ
ϭϯϰ
Sequence task
ϭϯϱ
Š‡ ‡š’‡”‹‡– ”‡“—‹”‡† –Š‡ ˆƒ•– ’”‘†—…–‹‘ ‘ˆ †‹ˆˆ‡”‡– —‹ƒ—ƒŽ ͷǦ†‹‰‹– •‡“—‡…‡•Ǥ Š‡
ϭϯϲ
•‡“—‡…‡• ™‡”‡ ’‡”ˆ‘”‡† ‘ ƒ …—•–‘Ǧ„—‹Ž–ǡ Ǧ…‘’ƒ–‹„Ž‡ǡ ’‹ƒ‘ǦŽ‹‡ ‹•‘‡–”‹… ˆ‘”…‡
ϰ
d,ZK>K&/W^/>dZ>KZdy/EDKdKZ>ZE/E' ϱ
ϭϯϳ
‡›„‘ƒ”†Ǥ ‘”…‡• ‡š‡”–‡† ‘ ‡ƒ…Š ‡› ™‡”‡ ‡ƒ•—”‡† ‡˜‡”› ͷ • ˜‹ƒ –”ƒ•†—…‡”• ȋ‡•‹‰ Ƭ
ϭϯϴ
‘–”‘Ž‘‡›™‡ŽŽ…Ǥǡ
Ǧͳͷͳǡ†›ƒ‹…”ƒ‰‡‘ˆͲǦʹͷȌǤ
ϭϯϵ
Š‡ •‡“—‡…‡ –ƒ• ”‡“—‹”‡† ’ƒ”–‹…‹’ƒ–• –‘ ’”‡•• ‡ƒ…Š †‹‰‹– ‹ ƒ ’”‡Ǧ†‡ˆ‹‡† ‘”†‡”Ǥ ϭϰϬ
…‘’—–‡”•…”‡‡•Š‘™‡†‡ƒ…Š•‡“—‡…‡ƒ•ƒ•’‡…‹ˆ‹…‘”†‡”‹‰‘ˆ–Š‡—„‡”•ͳǦͷǤǮͳǯ”‡ˆ‡””‡†–‘
ϭϰϭ
Ž‡ˆ–Ǧ‘•– ƒ† Ǯͷǯ –‘ ”‹‰Š–Ǧ‘•– ‡›ǡ ƒ†–Š‡ •‡“—‡…‡ ™ƒ•‡š‡…—–‡†ˆ”‘ Ž‡ˆ––‘”‹‰Š–Ǥ ‘–Š‡”
ϭϰϮ
™‘”†•ǡ •‡“—‡…‡• ™‡”‡ …—‡† ‹ ƒ ‡š–”‹•‹…ǡ •’ƒ–‹ƒŽ ”‡ˆ‡”‡…‡ ˆ”ƒ‡Ǥ ƒ•‡† ‘ ’‹Ž‘–
ϭϰϯ
‡š’‡”‹‡–ƒ–‹‘ ȋ‹‡•–Ž‡” ƒ† ‹‡†”‹…Š•‡ǡ ʹͲͳ͵Ȍǡ ™‡ •‡Ž‡…–‡† –™‡Ž˜‡ •‡“—‡…‡• ‘ˆ ƒ–…Š‡†
ϭϰϰ
†‹ˆˆ‹…—Ž–›ǡ ‡š…Ž—†‹‰ •‡“—‡…‡• –Šƒ– …‘–ƒ‹‡† ƒ ”— ‘ˆ ‘”‡ –Šƒ –Š”‡‡ ƒ†Œƒ…‡– †‹‰‹–•Ǥ ”‘
ϭϰϱ
–Š‡•‡ –™‡Ž˜‡ •‡“—‡…‡•ǡ ‡ƒ…Š ’ƒ”–‹…‹’ƒ– ™ƒ• ƒ••‹‰‡† ƒ •‡– ‘ˆ ˆ‘—” •‡“—‡…‡• –Šƒ– ™‘—Ž† „‡
ϭϰϲ
–”ƒ‹‡† ȋ‹‡•–Ž‡” ‡– ƒŽǤǡ ʹͲͳͶȌǤ Š‡ ’‘••‹„Ž‡ •‡–• ˆ‘” –”ƒ‹‹‰ ™‡”‡ ȋͶͳ͵ͷʹǡ ʹͷ͵ͳͶǡ ͳͷͶʹ͵ǡ
ϭϰϳ
ͷͳʹͶ͵ȌǡȋͶͷͳ͵ʹǡʹͳͷ͵Ͷǡ͵ͳͶʹͷǡ͵ͷʹͶͳȌǡƒ†ȋͷʹͳ͵ͶǡͳͶͷ͵ʹǡʹ͵ͷͶͳǡͶ͵ͳʹͷȌǤ
ϭϰϴ
•ƒŽŽȋͲǤͷ͵…šͲǤͷ͵…Ȍ‰”‡‡„‘šˆŽƒ‹‰–Š‡’”‡•‡–‡†•‡“—‡…‡™ƒ•†‹•’Žƒ›‡†‘
ϭϰϵ
–Š‡ Ž‡ˆ– ƒ† ”‹‰Š– –‘ …—‡ –Š‡ ”‡•’‡…–‹˜‡ Šƒ† –Šƒ– •Š‘—Ž† ‡š‡…—–‡ –Š‡ •‡“—‡…‡Ǥ ”‡† „‘š
ϭϱϬ
ƒ’’‡ƒ”‡†‘–Š‡‘–Š‡”•‹†‡‹†‹…ƒ–‹‰–Šƒ––Š‡Šƒ†•Š‘—Ž†”‡ƒ‹ƒ–”‡•–‘–Š‡‡›„‘ƒ”†ǤŠ‡
ϭϱϭ
‹•–”—…–‹‘ƒŽ •–‹—Ž—• ”‡ƒ‹‡† ‘ –Š‡ •…”‡‡ ˆ‘” ʹǤ͹•ǡ ƒˆ–‡” ™Š‹…Š ˆ‹˜‡ ™Š‹–‡ ƒ•–‡”‹•• ™‡”‡
ϭϱϮ
’”‡•‡–‡†ƒ•ƒǮ‰‘ǯ•‹‰ƒŽǤƒ…Š‹•–”—…–‡†•‡“—‡…‡™ƒ•‡š‡…—–‡†‡‹–Š‡”ˆ‘—”–‹‡•‹ƒ”‘™ȋ‹
ϭϱϯ
–Š‡ ’”‡Ǧ–‡•– ƒ† ’‘•–Ǧ–‡•–Ȍ ‘” –Š”‡‡ –‹‡• ȋ†—”‹‰ –”ƒ‹‹‰ ƒ† ˆ ƒ…“—‹•‹–‹‘Ȍǡ ƒ† ‡ƒ…Š
ϭϱϰ
‡š‡…—–‹‘ ™ƒ• ‹†‹˜‹†—ƒŽŽ› –”‹‰‰‡”‡† ™‹–Š ƒ Ǯ‰‘ǯ •‹‰ƒŽǤ Š‡”‡ ™ƒ• ƒ ͷͲͲ• ‹–‡”˜ƒŽ „‡–™‡‡
ϭϱϱ
…‘•‡…—–‹˜‡•‡“—‡…‡‡š‡…—–‹‘•Ǥ‘”–Š‡”‡ƒ‹†‡”‘ˆ–Š‡ƒ—•…”‹’–ǡ™‡†‡ˆ‹‡ƒ•‹‰Ž‡Ǯ–”‹ƒŽǯ
ϭϱϲ
ƒ•–Š‡•‡–‘ˆ–Š”‡‡‘”ˆ‘—”…‘•‡…—–‹˜‡‡š‡…—–‹‘•‘ˆ–Š‡•ƒ‡•‡“—‡…‡Ǥ
ϭϱϳ
Š‡ ‘„Œ‡…–‹˜‡ ‘ˆ –Š‡ •‡“—‡…‡ –ƒ• ™ƒ• –‘ ’‡”ˆ‘” –Š‡ ˆ‹˜‡ †‹‰‹– ’”‡••‡• ƒ• “—‹…Ž› ƒ•
ϭϱϴ
’‘••‹„Ž‡™‹–Š ‹‹ƒŽ ‡””‘”•Ǥ‘” ƒ †‹‰‹–’”‡••–‘ „‡ ”‡‰‹•–‡”‡†ǡ –Š‡ƒ…–‹˜‡†‹‰‹– Šƒ† –‘ ‡š…‡‡†
ϭϱϵ
‡š‡”–‹‰ƒˆ‘”…‡‘ˆʹǤͷǡ™Š‹Ž‡ƒŽŽ‘–Š‡”†‹‰‹–•Šƒ†–‘‰‡‡”ƒ–‡ˆ‘”…‡•‘ˆ„‡Ž‘™ʹǤʹǤˆ–‡”‡ƒ…Š
ϭϲϬ
†‹‰‹– ’”‡••ǡ –Š‡ …‘””‡•’‘†‹‰ ƒ•–‡”‹• …Šƒ‰‡† …‘Ž‘” –‘ ’”‘˜‹†‡ ˆ‡‡†„ƒ… ƒ„‘—– ™Š‡–Š‡” –Š‡
ϭϲϭ
‹†‹˜‹†—ƒŽ ’”‡•• ™ƒ•ǣ Ǯ…‘””‡…–ǯ ȋ‰”‡‡Ȍǡ Ǯ‹…‘””‡…–ǯ ȋ”‡†Ȍǡ ‘” Ǯ–‘‘ Šƒ”†ǯ ȋ›‡ŽŽ‘™Ǣ –Šƒ– ‹•ǡ ‰”‡ƒ–‡”
ϭϲϮ
–Šƒ–Š‡—’’‡”Ž‹‹–•‡–ˆ‘”–Š‡–ƒ•ǡͺǤͻȌǤš‡…—–‹‘–‹‡ȋȌ™ƒ•‡ƒ•—”‡†ƒ•–Š‡†—”ƒ–‹‘
ϭϲϯ
„‡–™‡‡‘•‡–‘ˆ–Š‡ˆ‹”•–’”‡••ƒ†”‡Ž‡ƒ•‡‘ˆ–Š‡Žƒ•–’”‡••ǡƒ†‡””‘””ƒ–‡™ƒ•†‡ˆ‹‡†ƒ•–Š‡
ϭϲϰ
’‡”…‡–ƒ‰‡ ‘ˆ •‡“—‡…‡• –Šƒ– …‘–ƒ‹‡† ‘‡ ‘” ‘”‡ ‹…‘””‡…– †‹‰‹– ’”‡••‡•Ǥ Š”‘—‰Š‘—–
ϭϲϱ
„‡Šƒ˜‹‘”ƒŽ–”ƒ‹‹‰ǡƒ…‘•–ƒ–‡””‘””ƒ–‡™ƒ•‡…‘—”ƒ‰‡†„›‹•–”—…–‹‰’ƒ”–‹…‹’ƒ–•–‘•’‡‡†
ϭϲϲ
—’‹ˆ‡””‘””ƒ–‡™ƒ•Ž‘™‡”–ŠƒʹͲΨƒ†•Ž‘™†‘™‹ˆ‹–™ƒ•Š‹‰Š‡”Ǥ
ϭϲϳ
ˆ–‡” ‡ƒ…Š •‡“—‡…‡ ‡š‡…—–‹‘ǡ ’ƒ”–‹…‹’ƒ–• •ƒ™ ƒ „”‹‡ˆ ˆ‡‡†„ƒ… ȋͲǤͺ•Ȍǣ ‘‡ ‰”‡‡
ϭϲϴ
ƒ•–‡”‹•ȋαͳ’‘‹–Ȍ‹†‹…ƒ–‡†–Šƒ–ƒŽŽˆ‹˜‡’”‡••‡•‘ˆ–Š‡•‡“—‡…‡™‡”‡‡š‡…—–‡†…‘””‡…–Ž›Ǣ–Š”‡‡
ϭϲϵ
‰”‡‡ ƒ•–‡”‹•• ȋα ͵ ’‘‹–•Ȍ ‡ƒ– –Šƒ– –Š‡ •‡“—‡…‡ ™ƒ• ‡š‡…—–‡† …‘””‡…–Ž› ƒ† ™‹–Š ηʹͲΨ
ϭϳϬ
ˆƒ•–‡”–Šƒ–Š‡ƒ˜‡”ƒ‰‡‘ˆ–Š‡’”‡˜‹‘—•”—Ǣ‘‡„Ž—‡ƒ•–‡”‹••’‡…‹ˆ‹‡†–Šƒ––Š‡•‡“—‡…‡™ƒ•
ϭϳϭ
’‡”ˆ‘”‡†ʹͲΨ•Ž‘™‡”–Šƒ–Š‡ƒ˜‡”ƒ‰‡‘ˆ–Š‡’”‡˜‹‘—•”—ȋαͲ’‘‹–•ȌǢƒ†‘‡”‡†ƒ•–‡”‹•
ϱ
d,ZK>K&/W^/>dZ>KZdy/EDKdKZ>ZE/E' ϲ
ϭϳϮ
•‹‰‹ˆ‹‡† –Šƒ– ‘‡ ‘” ‘”‡ ‡””‘”• ™‡”‡ ƒ†‡ ȋα Ǧͳ ’‘‹–ȌǤ ƒ”–‹…‹’ƒ–• ”‡…‡‹˜‡† ƒ ˆ‹ƒ…‹ƒŽ
ϭϳϯ
„‘—•ƒ……‘”†‹‰–‘–Š‡‹”’‘‹–•…‘”‡Ǥ
ϭϳϰ
ϭϳϱ
Experimental Design
ϭϳϲ
ŽŽ ’ƒ”–‹…‹’ƒ–• …‘’Ž‡–‡† Ͷ •–—†› ’Šƒ•‡• ȋFig. 1Ȍǣ ƒ ’”‡Ǧ–‡•– ȋ†ƒ› ͳȌ ‹ ™Š‹…Š „ƒ•‡Ž‹‡
ϭϳϳ
’‡”ˆ‘”ƒ…‡ˆ‘”–™‡Ž˜‡ͷǦ†‹‰‹–•‡“—‡…‡•™ƒ•‡˜ƒŽ—ƒ–‡†ˆ‘”„‘–ŠŠƒ†•ǢƒͶǦ†ƒ›–”ƒ‹‹‰’Šƒ•‡
ϭϳϴ
ȋ†ƒ›•ʹǦͷȌ†—”‹‰™Š‹…Š’ƒ”–‹…‹’ƒ–•”‡’‡ƒ–‡†Ž›’”ƒ…–‹…‡†–Š‡•ƒ‡ˆ‘—”•‡“—‡…‡•™‹–Š‡‹–Š‡”
ϭϳϵ
–Š‡Ž‡ˆ–‘””‹‰Š–Šƒ†ˆ‘”ƒ’’”‘š‹ƒ–‡Ž›ƒŠ‘—”ȋ™Š‹…Š™ƒ•…‘—’Ž‡†™‹–Šʹͷ‹‘ˆ–ˆ”‘
ϭϴϬ
–Š‡‘•‡–‘ˆ–”ƒ‹‹‰ȌǢƒ’‘•–Ǧ–‡•–ȋ†ƒ›͸Ȍǡ™Š‹…Š™ƒ•…‘†—…–‡†‹–Š‡•ƒ‡ƒ‡”ƒ•–Š‡’”‡Ǧ
ϭϴϭ
–‡•–Ǣƒ†ƒ•‡••‹‘ȋ†ƒ›͹ȌǤ
ϭϴϮ
Š‡ ’”‡Ǧ–‡•– •–ƒ”–‡† ™‹–Š ƒ •Š‘”– ’”ƒ…–‹…‡ ”— ™‹–Š Ͷ –”‹ƒŽ• ‘” ͳ͸ ‡š‡…—–‹‘• ȋͶ
ϭϴϯ
‡š‡…—–‹‘•‘ˆ–™‘‡ƒ•›•‡“—‡…‡•’‡”Šƒ†Ȍ–‘ˆƒ‹Ž‹ƒ”‹œ‡’ƒ”–‹…‹’ƒ–•™‹–Š–Š‡–ƒ•Ǥ—”‹‰–Š‡
ϭϴϰ
’”‡Ǧ–‡•–ǡ ’ƒ”–‹…‹’ƒ–•’‡”ˆ‘”‡†ƒŽŽͳʹ •‡“—‡…‡•ȋͶ –‘Ǧ„‡Ǧ–”ƒ‹‡† ƒ† ͺ—–”ƒ‹‡†Ȍ ™‹–Š„‘–Š
ϭϴϱ
Ž‡ˆ–ƒ†”‹‰Š–Šƒ†•Ǥƒ…ŠŠƒ†™ƒ•”‡“—‹”‡†–‘’‡”ˆ‘”ʹ–”‹ƒŽ•’‡”•‡“—‡…‡™‹–ŠͶ‡š‡…—–‹‘•
ϭϴϲ
’‡” –”‹ƒŽ ȋ‹Ǥ‡Ǥ ͺ –‘–ƒŽ ‡š‡…—–‹‘•ȌǤ Š‡ ’”‡Ǧ–‡•– …‘•‹•–‡† ‘ˆ ‡‹‰Š– ”—• ™‹–Š ʹͶ –”‹ƒŽ• ’‡” Šƒ†ǡ
ϭϴϳ
™Š‹…Š ”‡•—Ž–‡† ‹ ƒ –‘–ƒŽ ‘ˆ ͻ͸ ‡š‡…—–‹‘• ’‡” Šƒ†Ǥ ‹–Š‹ –Š‡ ˆ‹”•– ˆ‘—” ”—•ǡ –Š‡ ‘”†‡” ‘ˆ
ϭϴϴ
•‡“—‡…‡•ƒ†Šƒ†•™ƒ•”ƒ†‘Ž›’‡”—–‡†ǡƒ†–Š‡‘”†‡”™ƒ•”‡˜‡”•‡†‹–Š‡•‡…‘†ŠƒŽˆ–‘
ϭϴϵ
…‘—–‡”„ƒŽƒ…‡’‘••‹„Ž‡Ž‡ƒ”‹‰‡ˆˆ‡…–•Ǥ
ϭϵϬ
‡ ƒ••‹‰‡† •—„Œ‡…–• ’•‡—†‘Ǧ”ƒ†‘Ž› –‘ ‘‡ ‘ˆ ˆ‘—” •–‹—Žƒ–‹‘ ‰”‘—’• ȋ•Šƒǡ
ϭϵϭ
—‹Š‡‹•’Š‡”‹…ǡ „‹Š‡‹•’Š‡”‹…ǡ ‘” ”‡˜‡”•‡†Ǧ’‘Žƒ”‹–› „‹Š‡‹•’Š‡”‹… – ‰”‘—’•Ȍ ƒ† –‘
ϭϵϮ
–”ƒ‹‹‰…‘Š‘”–ȋŽ‡ˆ–‘””‹‰Š–Šƒ†–”ƒ‹‹‰Ȍǡ•—…Š–Šƒ–‰”‘—’†‹ˆˆ‡”‡…‡•‹’”‡Ǧ–‡•–’‡”ˆ‘”ƒ…‡
ϭϵϯ
™‡”‡ ‹‹‹œ‡† ȋƒ–‡”•Ǧ‡–‡‹‡” ‡– ƒŽǤǡ ʹͲͳͶȌǤ ƒ…”‘•• – ‰”‘—’• ”‡˜‡ƒŽ‡† ‘
ϭϵϰ
†‹ˆˆ‡”‡…‡ ‹ „ƒ•‡Ž‹‡ –ƒ• ȋFȋ͵ǡ͸ͲȌ α ͲǤͳͷͺǡ p α ͲǤͻʹ͵Ȍ ȋ•‡‡ Table 2 ˆ‘” ‹†‹˜‹†—ƒŽ ‰”‘—’
ϭϵϱ
‡ƒ•Ȍǡ ƒ† –Š‡”‡ ™ƒ• ƒŽ•‘ ‘ •‹‰‹ˆ‹…ƒ– ’ƒ‹”Ǧ™‹•‡ †‹ˆˆ‡”‡…‡ „‡–™‡‡ ƒ› –™‘ •–‹—Žƒ–‹‘
ϭϵϲ
‰”‘—’•ȋƒŽŽtδͳǤͳͲǡp αͲǤʹ͹͵ȌǤ
ϭϵϳ
––Š‡„‡‰‹‹‰‘ˆ‡ƒ…Š•‡••‹‘ǡ’ƒ”–‹…‹’ƒ–•’”‘˜‹†‡†‹ˆ‘”ƒ–‹‘ƒ„‘—–•Ž‡‡’“—ƒŽ‹–›ǡ
ϭϵϴ
ƒŽ‡”–‡••ǡƒ––‡–‹‘ǡƒ†–ƒ•†‹ˆˆ‹…—Ž–›—•‹‰˜‹•—ƒŽƒƒŽ‘‰—‡•…ƒŽ‡•”ƒ‰‹‰ˆ”‘ͲȋŽ‘™‡•–Ȍ–‘
ϭϵϵ
ͳͲ ȋŠ‹‰Š‡•–ȌǤ Š‡”‡ ™‡”‡ ‘ •‹‰‹ˆ‹…ƒ– †‹ˆˆ‡”‡…‡• „‡–™‡‡ •–‹—Žƒ–‹‘ ‰”‘—’• ‹ –‡”• ‘ˆ
ϮϬϬ
–Š‡•‡’ƒ”ƒ‡–‡”•ȋTable 2ȌǤ
ϮϬϭ
‘‡•—”‡–Šƒ–„‘–Š’ƒ”–‹…‹’ƒ–•ƒ†‡š’‡”‹‡–‡”™‡”‡„Ž‹†–‘–ƒ••‹‰‡–ǡ–Š‡
ϮϬϮ
”ƒ†‘‹•ƒ–‹‘ ™ƒ• ’‡”ˆ‘”‡† „› ƒ ǡ ™Š‘ ™ƒ• ‘– ‹˜‘Ž˜‡† ‹ †ƒ–ƒ …‘ŽŽ‡…–‹‘Ǥ Š‡
ϮϬϯ
‡š’‡”‹‡–‡”ȋȌ‘Ž›‡™–Š‡Šƒ†–”ƒ‹‹‰…‘Š‘”–ƒ†–Š‡‡Ž‡…–”‘†‡ƒ””ƒ‰‡‡–ǡ„—–‘–
ϮϬϰ
™Š‡–Š‡” –Š‡ ’ƒ”–‹…‹’ƒ– ”‡…‡‹˜‡† ”‡ƒŽ ‘” •Šƒ •–‹—Žƒ–‹‘ǡ ™Š‹…Š ™ƒ• †‡–‡”‹‡† „› ƒ
ϮϬϱ
”ƒ†‘‹œ‡†…‘†‡‡–‡”‡†‹–‘–Š‡–ƒ…Š‹‡ƒ––Š‡„‡‰‹‹‰‘ˆ‡ƒ…Š•‡••‹‘Ǥ……‘”†‹‰Ž›ǡ
ϮϬϲ
͸͸Ǥ͹Ψ‘ˆ–Š‡•Šƒ’ƒ”–‹…‹’ƒ–•ȋͳͶȀʹͳȌŠƒ†„‹Š‡‹•’Š‡”‹…‡Ž‡…–”‘†‡ƒ””ƒ‰‡‡–ƒ†͵͵Ǥ͵Ψ
ϮϬϳ
Šƒ† —‹Š‡‹•’Š‡”‹… ‡Ž‡…–”‘†‡ ƒ””ƒ‰‡‡–Ǥ ‘ •‹‰‹ˆ‹…ƒ– „‡Šƒ˜‹‘”ƒŽ †‹ˆˆ‡”‡…‡• ™‡”‡ ˆ‘—†
ϲ
d,ZK>K&/W^/>dZ>KZdy/EDKdKZ>ZE/E' ϳ
ϮϬϴ
„‡–™‡‡–Š‡•‡–™‘•—„•‡–•‘ˆ–Š‡•Šƒ‰”‘—’‘’‘•–Ǧ–‡•–’‡”ˆ‘”ƒ…‡ȋFȋͳǡͳ͸ȌδͳǤͶͷͶǡpεͲǤʹͶͷǢ
ϮϬϵ
—•‹‰ ’”‡Ǧ–‡•– ƒ• …‘˜ƒ”‹ƒ–‡Ȍ ˆ‘” –”ƒ‹‡†Ȁ—–”ƒ‹‡† Šƒ† ƒ† –”ƒ‹‡†Ȁ—–”ƒ‹‡†
ϮϭϬ
•‡“—‡…‡Ǥ
Ϯϭϭ
—”‹‰–Š‡ˆ‘—”–”ƒ‹‹‰†ƒ›•ǡ’ƒ”–‹…‹’ƒ–•’”ƒ…–‹…‡†Ͷ‘ˆ–Š‡ ͳʹ•‡“—‡…‡•™‹–Š‡‹–Š‡”
ϮϭϮ
–Š‡‹” Ž‡ˆ– ‘” ”‹‰Š– Šƒ†Ǥ •‡••‹‘ …‘•‹•–‡† ‘ˆ ͳ͸ ”—• ™‹–Š ʹ –”‹ƒŽ• ’‡” •‡“—‡…‡ ‡ƒ…ŠǤ
Ϯϭϯ
Š‡”‡ˆ‘”‡ǡ ’ƒ”–‹…‹’ƒ–• ’‡”ˆ‘”‡† ͳʹͺ –”‹ƒŽ• ȋ͵ͺͶ •‡“—‡…‡ ‡š‡…—–‹‘•Ȍ ’‡” †ƒ›Ǥ  –Š‡ †ƒ›
Ϯϭϰ
ˆ‘ŽŽ‘™‹‰ –Š‡ ˆ‹ƒŽ •‡••‹‘ ‘ˆ –Ǧ…‘—’Ž‡† –”ƒ‹‹‰ǡ ƒ ’‘•–Ǧ–‡•– –Šƒ– Šƒ† ‡šƒ…–Ž› –Š‡ •ƒ‡
Ϯϭϱ
•–”—…–—”‡ƒ•–Š‡’”‡Ǧ–‡•–™ƒ•ƒ†‹‹•–‡”‡†Ǥ
Ϯϭϲ
Ϯϭϳ
Transcranial direct current stimulation
Ϯϭϴ
ƚ^ǁĂƐĂĚŵŝŶŝƐƚĞƌĞĚǀŝĂĂďŝŚĞŵŝƐƉŚĞƌŝĐŽƌƵŶŝŚĞŵŝƐƉŚĞƌŝĐŵŽŶƚĂŐĞ͘/ŶƚŚĞƵŶŝŚĞŵŝƐƉŚĞƌŝĐ
Ϯϭϵ
ŵŽŶƚĂŐĞ͕ǁĞƉůĂĐĞĚƚŚĞĂŶŽĚĞŽǀĞƌĐŽŶƚƌĂůĂƚĞƌĂůDϭĂŶĚĐĂƚŚŽĚĞŽǀĞƌŝƉƐŝůĂƚĞƌĂůƐƵƉƌĂŽƌďŝƚĂůƌŝĚŐĞ͘
ϮϮϬ
&ŽƌƚŚĞďŝŚĞŵŝƐƉŚĞƌŝĐŵŽŶƚĂŐĞ͕ǁĞƉŽƐŝƚŝŽŶĞĚƚŚĞĂŶŽĚĞŽǀĞƌĐŽŶƚƌĂůĂƚĞƌĂůĂŶĚƚŚĞĐĂƚŚŽĚĞĂďŽǀĞ
ϮϮϭ
ŝƉƐŝůĂƚĞƌĂůDϭ͕ǁŝƚŚƚŚĞƌĞǀĞƌƐĞĚͲƉŽůĂƌŝƚLJŵŽŶƚĂŐĞŝŶǀŽůǀŝŶŐƚŚĞŽƉƉŽƐŝƚĞƉŽůĂƌŝƚLJ͘dŚĞŚĂŶĚĂƌĞĂŽĨ
ϮϮϮ
DϭǁĂƐůŽĐĂůŝnjĞĚĂƐƚŚĞƉŽƐŝƚŝŽŶǁŚĞƌĞƐŝŶŐůĞͲƉƵůƐĞƐƵƉƌĂƚŚƌĞƐŚŽůĚdD^ĞǀŽŬĞĚĂǀŝƐŝďůĞƚǁŝƚĐŚŝŶ
ϮϮϯ
ƚŚĞĐŽŶƚƌĂůĂƚĞƌĂůĨŝƌƐƚĚŽƌƐĂůŝŶƚĞƌŽƐƐĞƵƐŵƵƐĐůĞ;ŽƌŽŽũĞƌĚŝĞƚĂů͕͘ϭϵϵϵͿ͘dŚŝƐǁĂƐŝŵƉůĞŵĞŶƚĞĚ
ϮϮϰ
ƵƐŝŶŐĂDĂŐƐƚŝŵŝ^ƚŝŵϮǁŝƚŚĂϱͲĐŵĨŝŐƵƌĞͲŽĨͲĞŝŐŚƚĐŽŝůƉŽƐŝƚŝŽŶĞĚƚĂŶŐĞŶƚŝĂůůLJƚŽƚŚĞŵĞĚŝĂůͲ
ϮϮϱ
ƐĂŐŝƚƚĂůƉůĂŶĞƐŬƵůůĂƚĂϰϱΣĂŶŐůĞĂŶĚǁŝƚŚƚŚĞŚĂŶĚůĞƉŽŝŶƚŝŶŐƉŽƐƚĞƌŝŽƌůLJĂŶĚĂŵŽŶŽƉŚĂƐŝĐƉƵůƐĞ
ϮϮϲ
ǁĂƐĚĞůŝǀĞƌĞĚ͘dŚŝƐĂŶŐůĞǁĂƐĐŚŽƐĞŶĂƐŝƚĞůŝĐŝƚƐƚŚĞƐƚƌŽŶŐĞƐƚƚŚĞƐƚƌŽŶŐĞƐƚƉĞƌƉĞŶĚŝĐƵůĂƌĨŝĞůĚƐ
ϮϮϳ
;:ĂŶƐƐĞŶĞƚĂů͕͘ϮϬϭϱͿǁŚŝĐŚŝƐŽƉƚŝŵĂůĨŽƌƐƚŝŵƵůĂƚŝŶŐĐŽƌƚŝĐŽƐƉŝŶĂůŶĞƵƌŽŶƐƚƌĂŶƐͲƐLJŶĂƉƚŝĐĂůůLJǀŝĂ
ϮϮϴ
ŚŽƌŝnjŽŶƚĂůĐŽƌƚŝĐŽͲĐŽƌƚŝĐĂůĐŽŶŶĞĐƚŝŽŶƐ;ŝ>ĂnjnjĂƌŽĞƚĂů͕͘ϮϬϬϴ͖ĞůǀĞŶĚĂŚůĞƚĂů͕͘ϮϬϭϰͿ͘ƚ^ǁĂƐ
ϮϮϵ
ĂĚŵŝŶŝƐƚĞƌĞĚŽǀĞƌĨŽƵƌĐŽŶƐĞĐƵƚŝǀĞƚƌĂŝŶŝŶŐĚĂLJƐĨŽƌƚŚĞĨŝƌƐƚϮϱŵŝŶŽĨĂŶĂƉƉƌŽdžŝŵĂƚĞůLJϲϬͲŵŝŶ
ϮϯϬ
ƐĞƐƐŝŽŶŽĨƐĞƋƵĞŶĐĞƚƌĂŝŶŝŶŐ͘ĐƵƌƌĞŶƚŽĨϮŵǁĂƐĚĞůŝǀĞƌĞĚƵƐŝŶŐĂŶĞƵƌŽŽŶŶͲƐƚŝŵƵůĂƚŽƌW>h^
Ϯϯϭ
;ŚƚƚƉ͗ͬͬǁǁǁ͘ŶĞƵƌŽĐŽŶŶ͘ĚĞͬĚĐͲƐƚŝŵƵůĂƚŽƌͺƉůƵƐͺĞŶͬͿƚŚƌŽƵŐŚƐĂůŝŶĞͲƐŽĂŬĞĚϯϱĐŵϮĞůĞĐƚƌŽĚĞƐ͘
ϮϯϮ
Ϯϯϯ
Behavioral analysis
Ϯϯϰ
Š‡”‡ ™ƒ• ‘ •‹‰‹ˆ‹…ƒ– †‹ˆˆ‡”‡…‡ „‡–™‡‡ –Š‡ – ‰”‘—’• ‹ –‡”• ‘ˆ ‡””‘” ”ƒ–‡
Ϯϯϱ
†—”‹‰ –Š‡ ’”‡Ǧ–‡•– ȋȋ͵ǡ͸ͲȌ α ͲǤʹͷʹǡ ’ α ͲǤͺ͸ͲǢ ƒ˜‡”ƒ‰‡† ƒ…”‘•• –”ƒ‹‹‰ ‰”‘—’ ƒ† Šƒ†Ȍ ‘”
Ϯϯϲ
–”ƒ‹‹‰ ȋȋ͵ǡ͸ͲȌ α ͳǤͲͺʹǡ ’ α ͲǤ͵͸ͶǢ ƒ˜‡”ƒ‰‡† ƒ…”‘•• †ƒ›ȌǤ ‡ ‘Ž› ˆ‘—† ƒ •‹‰‹ˆ‹…ƒ– ‡ˆˆ‡…– ‘ˆ
Ϯϯϳ
–‘‡””‘””ƒ–‡‹–Š‡’‘•–Ǧ–‡•–ǡ™‹–Š–”‡…‹’‹‡–•–‡†‹‰–‘„‡‘”‡ƒ……—”ƒ–‡–Šƒ•Šƒ
Ϯϯϴ
ȋȋ͵ǡ͸ͲȌα͵ǤͲͺ͸ǡ’αͲǤͲ͵͵ͻǢƒ˜‡”ƒ‰‡†ƒ…”‘••–”ƒ‹‹‰‰”‘—’ƒ†Šƒ†ȌǤ
Ϯϯϵ
ϮϰϬ
•‡“—‡…‡ǡ ƒ† Šƒ† ‘˜‡” ƒŽŽ ȋ…‘””‡…– ƒ† ‹…‘””‡…–Ȍ –”‹ƒŽ•Ǥ ‘” –Š‹• …ƒŽ…—Žƒ–‹‘ǡ ˆ‘” –Š‡
‘ ƒ†Œ—•– ˆ‘” –Š‡ †‹ˆˆ‡”‡– ‡””‘” ”ƒ–‡•ǡ ™‡ …ƒŽ…—Žƒ–‡† –Š‡ ‡†‹ƒ ˆ‘” ‡ƒ…Š ”—ǡ
Ϯϰϭ
‹…‘””‡…– –”‹ƒŽ ™ƒ• ”‡’Žƒ…‡† ™‹–Š –Š‡ ƒš‹— ‘ˆ –Šƒ– ‰”‘—’ ‘ˆ –”‹ƒŽ•ǡ –Š‡”‡„› ’‡ƒŽ‹œ‹‰
ϮϰϮ
‹ƒ……—”ƒ–‡’‡”ˆ‘”ƒ…‡Ǥ
ϳ
d,ZK>K&/W^/>dZ>KZdy/EDKdKZ>ZE/E' ϴ
Ϯϰϯ
‡…‘†—…–‡†’”‡Ǧ’Žƒ‡†•–ƒ–‹•–‹…ƒŽ…‘’ƒ”‹•‘•„‡–™‡‡‡ƒ…Š•–‹—Žƒ–‹‘‰”‘—’ƒ†
Ϯϰϰ
•Šƒǡ „‡–™‡‡ „‹Š‡‹•’‡”‹… ƒ† —‹Š‡‹•’Š‡”‹… ‰”‘—’•ǡ ƒ† „‡–™‡‡ –Š‡ –™‘ „‹Š‡‹•’Š‡”‹…
Ϯϰϱ
‰”‘—’• ˆ‘” –Š‡ Žƒ•– –”ƒ‹‹‰ †ƒ› ƒ† –Š‡ ’‘•–Ǧ–‡•–Ǥ ‘” –Š‡•‡ …‘’ƒ”‹•‘• „‡–™‡‡ ’ƒ‹”• ‘ˆ
Ϯϰϲ
‰”‘—’•ǡ™‡—•‡†ƒ‹™Š‹…Š–Š‡‡””‘”Ǧƒ†Œ—•–‡†ƒ–’”‡Ǧ–‡•–™ƒ•—•‡†ƒ•ƒ…‘˜ƒ”‹ƒ–‡
Ϯϰϳ
–‘ ƒ……‘—– ˆ‘” ’”‹‘” ‹–‡”Ǧ‹†‹˜‹†—ƒŽ †‹ˆˆ‡”‡…‡• ‹ ’‡”ˆ‘”ƒ…‡Ǥ Š‹• ’”‘…‡†—”‡ ‡ˆˆ‡…–‹˜‡Ž›
Ϯϰϴ
•—„–”ƒ…–•ˆ”‘‡ƒ…Š–”ƒ‹‹‰Ȁ’‘•–Ǧ–‡•–‡ƒ•—”‡‡––Š‡„‡•–’”‡†‹…–‹‘„ƒ•‡†‘–Š‡’”‡Ǧ–‡•–
Ϯϰϵ
‡ƒ•—”‡‡–Ǥ ‘’ƒ”‡† –‘ •‹’Ž› •—„–”ƒ…–‹‰ –Š‡ ’”‡Ǧ–‡•– ‡ƒ•—”‡‡–ǡ –Š‹• ‡–Š‘† ‹• Ž‡••
ϮϱϬ
•—•…‡’–‹„Ž‡ –‘ ‘‹•‡ ‹†—…‡† „› –Š‡ Žƒ”‰‡” ˜ƒ”‹ƒ„‹Ž‹–› ‹ –Š‡ ’”‡Ǧ–‡•– ‡ƒ•—”‡‡–•Ǥ Š‡
Ϯϱϭ
‹…Ž—†‡† Ǯ– ‰”‘—’ǯ ƒ† ǮŠƒ† –”ƒ‹‹‰ …‘Š‘”–ǯ ƒ• „‡–™‡‡Ǧ•—„Œ‡…– ˆƒ…–‘”•Ǥ Š‡
ϮϱϮ
–Š”‡•Š‘Ž†ˆ‘”ƒŽŽ•–ƒ–‹•–‹…ƒŽ…‘’ƒ”‹•‘•™ƒ•pδͲǤͲͷǤŽŽ†ƒ–ƒ’”‡•‡–‡†‹–Š‡–‡š–ƒ†ˆ‹‰—”‡•
Ϯϱϯ
ƒ”‡”‡’”‡•‡–‡†ƒ•‡ƒά•–ƒ†ƒ”†‡””‘”‘ˆ–Š‡‡ƒȋȌǤ
Ϯϱϰ
Ϯϱϱ
fMRI data acquisition
Ϯϱϲ
ŽŽ Ͷʹ„‹Š‡‹•’Š‡”‹…– ”‡…‹’‹‡–• ȋ•Šƒ ƒ† ”‡ƒŽȌ—†‡”™‡– ˆ •…ƒ‹‰ ‘‡ †ƒ› ƒˆ–‡”
Ϯϱϳ
–Š‡ ’‘•–Ǧ–‡•–Ǥ —‡ –‘ ”‡•‘—”…‡ Ž‹‹–ƒ–‹‘•ǡ ™‡ ™‡”‡ ‘– ƒ„Ž‡ –‘ ƒŽ•‘ •…ƒ –Š‡ —‹Š‡‹•’Š‡”‹…
Ϯϱϴ
‰”‘—’•ǤŠ‹Ž‡–Š‹•‰”‘—’™‘—Ž†Šƒ˜‡•—’’Ž‹‡†ƒ††‹–‹‘ƒŽ‹ˆ‘”ƒ–‹‘ǡ‹–™ƒ•‘–…”‹–‹…ƒŽ–‘–‡•–
Ϯϱϵ
‘—”ƒ‹Š›’‘–Š‡•‹•‘ˆ’‘Žƒ”‹–›Ǧ‹†‡’‡†‡–…Šƒ‰‡•‹„‘–ŠŠ‡‹•’Š‡”‡•ƒˆ–‡”„‹Š‡‹•’Š‡”‹…
ϮϲϬ
•–‹—Žƒ–‹‘Ǥ––Š‹•’‘‹–ǡƒ–Ž‡ƒ•–ͶͺŠŠƒ†‡Žƒ’•‡†•‹…‡–Š‡ˆ‹ƒŽ–ƒ’’Ž‹…ƒ–‹‘ǤŠ‡”‡ˆ‘”‡
Ϯϲϭ
ƒ› ‰”‘—’ †‹ˆˆ‡”‡…‡• •Š‘—Ž† „‡ †—‡ –‘ Ž‘‰Ǧ–‡” ‡—”‘’Žƒ•–‹… …Šƒ‰‡• ‹†—…‡† „› –Ǣ
ϮϲϮ
‹‡†‹ƒ–‡ ‘Ž‹‡ ‡ˆˆ‡…–• ‘ˆ – ȋ‡”‹‡”‹ ‡– ƒŽǤǡ ʹͲͳͲǢ ‹‡Ž‡ ‡– ƒŽǤǡ ʹͲͳ͵Ǣ –ƒŽ ‡– ƒŽǤǡ ʹͲͳͶȌ
Ϯϲϯ
•Š‘—Ž†Šƒ˜‡„‡‡™ƒ•Š‡†‘—–ǤŠ‡—‹Š‡‹•’Š‡”‹…–”‡…‹’‹‡–•™‡”‡‘–•…ƒ‡†Ǥ
Ϯϲϰ
Š‡ ˆ •‡••‹‘ …‘•‹•–‡† ‘ˆ ͺ ”—• …‘’”‹•‡† ‘ˆ ʹͶ ”ƒ†‘Ž› ‘”†‡”‡† –”‹ƒŽ• ȋ͵ ’‡”
Ϯϲϱ
–”‹ƒŽ –›’‡ȄͶ •‡“—‡…‡• š ʹ Šƒ†•Ȅ™‹–Š ͵ •‡“—‡…‡ ‡š‡…—–‹‘• ’‡” –”‹ƒŽǡ ›‹‡Ž†‹‰ ͹ʹ –‘–ƒŽ
Ϯϲϲ
‡š‡…—–‹‘• ’‡” ”—ȌǤ ƒ…Š –”‹ƒŽ …‘•‹•–‡† ‘ˆ ƒ …—‡‹‰ ’Šƒ•‡ ȋʹǤ͹ •ǡ ͳ Ȍǡ ˆ‘ŽŽ‘™‡† „› ͵
Ϯϲϳ
‡š‡…—–‹‘• ‘ˆ –Š‡ …—‡† •‡“—‡…‡ǡ –”‹‰‰‡”‡† ͵Ǥ͸• ƒ’ƒ”–Ǥ –”‹ƒŽǡ –Š‡”‡ˆ‘”‡ǡ Žƒ•–‡† ͳ͵Ǥͷ• ȋͷ •ȌǤ
Ϯϲϴ
ƒ…Š•‡“—‡…‡‡š‡…—–‹‘Šƒ†–‘„‡…‘’Ž‡–‡†™‹–Š‹ʹǤͺ•–‘ƒŽŽ‘™ˆ‘”ƒͲǤͺ•ˆ‡‡†„ƒ…’Šƒ•‡Ǥ
Ϯϲϵ
‘ ƒ–…Š „‡Šƒ˜‹‘”ƒŽ ’‡”ˆ‘”ƒ…‡ †—”‹‰ •…ƒ‹‰ǡ ’ƒ”–‹…‹’ƒ–• ™‡”‡ ‹•–”—…–‡† –‘
ϮϳϬ
’”‘†—…‡ –Š‡ •‡“—‡…‡ ƒ– ƒ ˆ‹š‡† •’‡‡† ‘ˆ ͳǤ͵• ƒ† ƒ• ƒ……—”ƒ–‡Ž› ƒ• ’‘••‹„Ž‡Ǥ Š‹• ™ƒ• –Š‡
Ϯϳϭ
ˆƒ•–‡•––Šƒ–•—„Œ‡…–•ƒ…”‘••ƒŽŽ‰”‘—’•…‘—Ž†ƒ…Š‹‡˜‡™‹–Š„‘–Š–”ƒ‹‡†ƒ†—–”ƒ‹‡†Šƒ†•™‹–Š
ϮϳϮ
Š‹‰Šƒ……—”ƒ…›ȋ̱ͻͲΨ…‘””‡…–ȌǤ††‹–‹‘ƒŽŽ›ǡ–Š”‘—‰Š‘—––”ƒ‹‹‰ǡ ˆ‘”…‡Ž‡˜‡Ž•‘ˆ–Š‡•‡“—‡…‡•
Ϯϳϯ
™‡”‡‡’–•‹‹Žƒ”„›‹’‘•‹‰ƒƒš‹ƒŽˆ‘”…‡–Š”‡•Š‘Ž†Ǥ‘ˆ‘”…‡Ž‡˜‡Žˆ‡‡†„ƒ…™ƒ•’”‘˜‹†‡†
Ϯϳϰ
†—”‹‰ˆ•…ƒ‹‰Ǥ
Ϯϳϱ
ƒ•‡Ž‹‡ƒ…–‹˜ƒ–‹‘™ƒ•‡ƒ•—”‡††—”‹‰ͺ”ƒ†‘Ž›‹–‡”•’‡”•‡†”‡•–’Šƒ•‡•‘ˆ
Ϯϳϲ
ͳ͵Ǥͷ•Ǥ ‘ ‘‹–‘” ˆ‘” ‹””‘” ƒ…–‹˜‹–› ‘ –Š‡ ‘Ǧ‘˜‹‰ Šƒ†ǡ ’ƒ”–‹…‹’ƒ–• ™‡”‡ ”‡“—‹”‡† –‘
Ϯϳϳ
‡‡’ƒŽŽ–‡†‹‰‹–•‘–Š‡‡›„‘ƒ”†ƒ†–‘‰‡‡”ƒ–‡ƒ•ƒŽŽ„ƒ•‡Ž‹‡ˆ‘”…‡‘ˆ̱ͲǤͷƒ–ƒŽŽ–‹‡•Ǥ
ϴ
d,ZK>K&/W^/>dZ>KZdy/EDKdKZ>ZE/E' ϵ
Ϯϳϴ
—…–‹‘ƒŽ‹ƒ‰‡•™‡”‡ƒ…“—‹”‡†—•‹‰ƒ͵‹‡‡•”‹‘ƒ…Š‹‡ǡ™‹–Šƒ͵ʹǦ…Šƒ‡ŽŠ‡ƒ†
Ϯϳϵ
…‘‹ŽǤʹ‡…Š‘Ǧ’Žƒƒ”•‡“—‡…‡™‹–Šƒ‘ˆʹǤ͹ʹ•™ƒ•—•‡†–‘ƒ…“—‹”‡–Š‡ˆ—…–‹‘ƒŽ˜‘Ž—‡•
ϮϴϬ
ȋͺ ”—•ǡ ͳͷͻ ˜‘Ž—‡• ’‡” ”—ǡ ͵ʹ ‹–‡”Ž‡ƒ˜‡† •Ž‹…‡• ™‹–Š ʹǤ͹  –Š‹…‡••Ȍ ‹ ƒ ‹–‡”Ž‡ƒ˜‡†
Ϯϴϭ
ƒ‡” ȋ͵ ‰ƒ’ǡ ƒ† ʹǤ͵šʹǤ͵ ʹ ‹Ǧ’Žƒ‡ ”‡•‘Ž—–‹‘ȌǤ Š‡ ˜‘Ž—‡• ™‡”‡ ƒ…“—‹”‡† ‹ ƒ
ϮϴϮ
‘„Ž‹“—‡ ‘”‹‡–ƒ–‹‘ ™‹–Š ƒ Ͷͷι–‹Ž–ƒ‰Ž‡ ˆ”‘ –Š‡ǦŽ‹‡Ȅ–Š‹••Ž‹…‡ ’”‡•…”‹’–‹‘’”‘˜‹†‡†
Ϯϴϯ
…‘˜‡”ƒ‰‡‘ˆ‘–‘””‡‰‹‘•‘–Š‡†‘”•ƒŽ•—”ˆƒ…‡‘ˆ–Š‡…‘”–‡šǡƒ•™‡ŽŽƒ•–Š‡•—’‡”‹‘”…‡”‡„‡ŽŽ—
Ϯϴϰ
ƒ†„ƒ•ƒŽ‰ƒ‰Ž‹ƒǡ„—–‡š…Ž—†‡†‹ˆ‡”‹‘”’”‡ˆ”‘–ƒŽƒ†‹ˆ‡”‹‘”ƒ†ƒ–‡”‹‘”–‡’‘”ƒŽŽ‘„‡•Ǥ‘”
Ϯϴϱ
ˆ—ŽŽ†‡–ƒ‹Ž•‘ˆ–Š‡ˆƒ…“—‹•‹–‹‘ǡ•‡‡ȋ‹‡•–Ž‡”‡–ƒŽǤǡʹͲͳͶȌǤ
Ϯϴϲ
Ϯϴϳ
fMRI analysis
Ϯϴϴ
Š‡ƒƒŽ›•‹•‘ˆ–Š‡ˆ†ƒ–ƒ‹•†‡•…”‹„‡†‹†‡–ƒ‹Ž‹‹‡•–Ž‡”‡–ƒŽǤȋʹͲͳͶȌǡ™Š‹…Š”‡’‘”–•–Š‡
Ϯϴϵ
”‡•—Ž–• ˆ”‘ –Š‡ •Šƒ ‰”‘—’Ǥ ˆ–‡” •–ƒ†ƒ”† ’”‡Ǧ’”‘…‡••‹‰ ȋ…‘””‡…–‹‘ ˆ‘” •Ž‹…‡ ƒ…“—‹•‹–‹‘ǡ
ϮϵϬ
‘–‹‘ ”‡ƒŽ‹‰‡–ǡ ƒ† …‘”‡‰‹•–”ƒ–‹‘ –‘ –Š‡ ‹†‹˜‹†—ƒŽ ƒƒ–‘‹…ƒŽ ‹ƒ‰‡Ȍǡ ™‡ —•‡† ƒ ˆ‹”•–Ǧ
Ϯϵϭ
Ž‡˜‡ŽŽ‹‡ƒ”‘†‡Ž‹’Ž‡‡–‡†‹ͺȋ”‹•–‘‡–ƒŽǤǡʹͲͲ͹Ȍ–‘‡•–‹ƒ–‡–Š‡ƒ…–‹˜ƒ–‹‘ˆ‘”‡ƒ…Š
ϮϵϮ
‘ˆ–Š‡ˆ‘—”•‡“—‡…‡•ˆ‘”‡ƒ…ŠŠƒ†ǤŠ‡†‡•‹‰ƒ–”‹š…‘•‹•–‡†‘ˆƒ”‡‰”‡••‘”ˆ‘”‡ƒ…ŠŠƒ†
Ϯϵϯ
ƒ† •‡“—‡…‡ –›’‡ ƒ† ƒ ‹–‡”…‡’– ˆ‘” ‡ƒ…Š ”—Ǥ Š‡ ”‡‰”‡••‘” ™ƒ• ‘†‡ŽŽ‡† ƒ• ƒ „‘š…ƒ”
Ϯϵϰ
ˆ—…–‹‘‘ˆͳͲǤͺ•„‡‰‹‹‰™‹–Š–Š‡ˆ‹”•–‰‘…—‡‘ˆ–Š‡–”‹ƒŽǡ™Š‹…Š™ƒ•–Š‡…‘˜‘Ž˜‡†™‹–Šƒ
Ϯϵϱ
•–ƒ†ƒ”† Š‡‘†›ƒ‹… ”‡•’‘•‡ ˆ—…–‹‘Ǥ ”‘ –Š‡ ‡•–‹ƒ–‡† ”‡‰”‡••‹‘ ™‡‹‰Š–•ǡ ™‡
Ϯϵϲ
…‘’—–‡†–Š‡’‡”…‡–•‹‰ƒŽ…Šƒ‰‡…‘’ƒ”‡†–‘”‡•–ˆ‘”‡ƒ…ŠŠƒ†ǡƒ˜‡”ƒ‰‡†ƒ…”‘••–Š‡ˆ‘—”
Ϯϵϳ
•‡“—‡…‡•Ǥ
Ϯϵϴ
‘” –Š‡ ‰”‘—’ ƒƒŽ›•‹•ǡ ™‡ ”‡…‘•–”—…–‡† –Š‡ …‘”–‹…ƒŽ •—”ˆƒ…‡ ‘ˆ ‡ƒ…Š ‹†‹˜‹†—ƒŽ
Ϯϵϵ
’ƒ”–‹…‹’ƒ–—•‹‰”‡‡•—”ˆ‡”ȋƒŽ‡‡–ƒŽǤǡͳͻͻͻȌǡ™Š‹…Š’‡”‹–•–Š‡‡š–”ƒ…–‹‘‘ˆ–Š‡™Š‹–‡Ǧ‰”‡›
ϯϬϬ
ƒ––‡”•—”ˆƒ…‡ƒ†’‹ƒŽ•—”ˆƒ…‡ˆ”‘ƒƒ–‘‹…ƒŽ‹ƒ‰‡•ǤŠ‡ˆ—…–‹‘ƒŽ†ƒ–ƒ™ƒ•–Š‡’”‘Œ‡…–‡†
ϯϬϭ
‘–‘‡ƒ…Š‹†‹˜‹†—ƒŽ•—”ˆƒ…‡„›ƒ˜‡”ƒ‰‹‰ˆ‘”‡ƒ…Š•—”ˆƒ…‡‘†‡–Š‡˜‘š‡Ž•–Šƒ–Žƒ›„‡–™‡‡–Š‡
ϯϬϮ
™Š‹–‡Ǧ‰”‡›ƒ––‡”ƒ†’‹ƒŽ•—”ˆƒ…‡•Ǥ
ϯϬϯ
Š‡ ‹†‹˜‹†—ƒŽ •—”ˆƒ…‡• ™‡”‡ –Š‡ ƒŽ‹‰‡† –‘ ƒ •Šƒ”‡† •’Š‡”‹…ƒŽ –‡’Žƒ–‡ ˆ‘” Ž‡ˆ– ƒ†
ϯϬϰ
”‹‰Š–Š‡‹•’Š‡”‡•ǤŠ‹•ƒŽŽ‘™‡†—•–‘ˆŽ‹’–Š‡”‡•—Ž–•ˆ‘”–Š‡”‹‰Š–Šƒ†–”ƒ‹‡†…‘Š‘”–ǡ•—…Š–Šƒ–
ϯϬϱ
†ƒ–ƒ ˆ”‘ –Š‡ –”ƒ‹‡† Š‡‹•’Š‡”‡Ȅ–Š‡ Š‡‹•’Š‡”‡ …‘–”ƒŽƒ–‡”ƒŽ –‘ –Š‡ –”ƒ‹‡† Šƒ†Ȅ™ƒ•
ϯϬϲ
†‹•’Žƒ›‡† ‘–Š‡”‹‰Š–‰”‘—’Š‡‹•’Š‡”‡ǡƒ†–Š‡†ƒ–ƒˆ”‘–Š‡—–”ƒ‹‡†Š‡‹•’Š‡”‡‘–Š‡
ϯϬϳ
Ž‡ˆ–Ǥ‡–Š‡…‘†—…–‡†tǦ–‡•–•„‡–™‡‡•Šƒƒ†–‰”‘—’•ˆ‘”‡ƒ…Š•—”ˆƒ…‡˜‡”–‡šǡ—•‹‰ƒ
ϯϬϴ
—…‘””‡…–‡† Š‡‹‰Š– –Š”‡•Š‘Ž† ‘ˆ Tȋͳǡ͵ͻȌ α ʹǤ͹ͳǡ p α ͲǤͲͲͷǤ ƒ‹Ž›Ǧ™‹•‡ ‡””‘” ™ƒ• …‘–”‘ŽŽ‡† „›
ϯϬϵ
…ƒŽ…—Žƒ–‹‰ –Š‡ …”‹–‹…ƒŽ •‹œ‡ ‘ˆ –Š‡ Žƒ”‰‡•– •—’”ƒ–Š”‡•Š‘Ž† …Ž—•–‡” –Šƒ– ™‘—Ž† „‡ ‡š’‡…–‡† „›
ϯϭϬ
…Šƒ…‡ǡ —•‹‰ ƒ—••‹ƒ ‹‡Ž† –Š‡‘”› ƒ• ‹’Ž‡‡–‡† ‹ –Š‡ ˆ”‹•–ƒ– ’ƒ…ƒ‰‡ ȋ‘”•Ž‡› ‡– ƒŽǤǡ
ϯϭϭ
ͳͻͻ͸ȌǤ ‡•—Ž–• ™‡”‡ †‹•’Žƒ›‡† —•‹‰ –Š‡ ͵Ǧ˜‹•—ƒŽ‹•ƒ–‹‘ •‘ˆ–™ƒ”‡ ƒ”‡– ȋƒ ••‡ ‡– ƒŽǤǡ
ϯϭϮ
ʹͲͲͳȌǤ
ϵ
d,ZK>K&/W^/>dZ>KZdy/EDKdKZ>ZE/E' ϭϬ
ϯϭϯ
‘” –Š‡ ’”‘ˆ‹Ž‡ ’Ž‘–• •Š‘™ ‹ Fig. 3a,bǡ ™‡ †‡ˆ‹‡† ƒ Ž‹‡ ”—‹‰ ˆ”‘ –Š‡ ’‘•–‡”‹‘”
ϯϭϰ
’ƒ”‹‡–ƒŽ …‘”–‡š –Š”‘—‰Š –Š‡ Šƒ† ƒ”‡ƒ ‘ˆ ͳ –‘ –Š‡ ƒ–‡”‹‘” –‹’ ‘ˆ ”‘†ƒ ƒ”‡ƒ ͸ ȋ’”‡‘–‘”
ϯϭϱ
…‘”–‡šȌǤ‡–Š‡ƒ˜‡”ƒ‰‡†ƒ…–‹˜‹–›‘˜‡”–Š‡ƒ”‡ƒͳǤͷ…ƒ„‘˜‡ƒ†„‡Ž‘™–Šƒ–Ž‹‡ȋ’—”’Ž‡ƒ”‡ƒ
ϯϭϲ
‹Fig. 3dȌǤ
ϯϭϳ
‡ ƒŽ•‘ …‘†—…–‡† ƒ ”‡‰‹‘ ‘ˆ ‹–‡”‡•– ȋȌ ƒƒŽ›•‹• ˆ‘” –Š‡ …‘”–‹…ƒŽ ‘–‘” ”‡‰‹‘•Ǥ
ϯϭϴ
Š‡› ‹…Ž—†‡† –Š‡ Šƒ† ”‡‰‹‘ ‘ˆ ’”‹ƒ”› ‘–‘” …‘”–‡š ȋͳǡ ”‘†ƒ ƒ”‡ƒ ͶȌǢ ’”‹ƒ”›
ϯϭϵ
•‘ƒ–‘•‡•‘”›…‘”–‡šȋͳǡ”‘†ƒƒ”‡ƒͳǦ͵ȌǢ†‘”•ƒŽ’”‡‘–‘”…‘”–‡šȋ†ȌǢ•—’’Ž‡‡–ƒ”›
ϯϮϬ
‘–‘”ƒ”‡ƒ•ȋȀ’”‡ǦȌǢƒ†•—’‡”‹‘”’ƒ”‹‡–ƒŽŽ‘„‡ǡ†‹˜‹†‡†‹–‘ƒƒ–‡”‹‘”ȋ‹–”ƒǦ’ƒ”‹‡–ƒŽ
ϯϮϭ
•—Ž…—•ǡ Ȍ ƒ† ’‘•–‡”‹‘” ȋ‘……‹’‹–‘Ǧ’ƒ”‹‡–ƒŽ Œ—…–‹‘ǡ Ȍ ƒ•’‡…–Ǥ ‡ †‡ˆ‹‡† ƒŽŽ • ‘ –Š‡
ϯϮϮ
•›‡–”‹… ‰”‘—’ –‡’Žƒ–‡ ƒ† •—„•‡“—‡–Ž› ’”‘Œ‡…–‡† –Š‹• ‹–‘ ‹†‹˜‹†—ƒŽ †ƒ–ƒ •’ƒ…‡ ˜‹ƒ –Š‡
ϯϮϯ
”‡•’‡…–‹˜‡‹†‹˜‹†—ƒŽ•—”ˆƒ…‡Ǥ
ϯϮϰ
Š‡ ƒ˜‡”ƒ‰‡ ƒ…–‹˜‹–› ‹ –Š‡•‡ • ™ƒ• •—„‹––‡† –‘ ƒ –‘ …ƒŽ…—Žƒ–‡ †‹ˆˆ‡”‡…‡•
ϯϮϱ
„‡–™‡‡ – ‰”‘—’• ȋˆƒ…–‘” Ǯ–ǯȌ ƒ† ‹–‡”ƒ…–‹‘• ™‹–Š –”ƒ‹‹‰ Šƒ† ȋˆƒ…–‘” ǮŠƒ†ǯȌ ƒ†
ϯϮϲ
Š‡‹•’Š‡”‡ ȋˆƒ…–‘” ǮŠ‡‹•’Š‡”‡ǯȌǤ ‡…ƒ—•‡ –Š‡”‡ ™‡”‡ ‘ •‹‰‹ˆ‹…ƒ– ˆ—…–‹‘ƒŽ †‹ˆˆ‡”‡…‡•
ϯϮϳ
„‡–™‡‡Ž‡ˆ–ƒ†”‹‰Š–Šƒ†–”ƒ‹‹‰…‘Š‘”–•ȋȋͳǡ͵͸ȌδʹǤʹͳǡ’εͲǤͳͷǡˆ‘”ƒŽŽ͸•ǡ‡˜‡™‹–Š‘—–
ϯϮϴ
…‘””‡…–‹‘Ȍǡ‘”‹–‡”ƒ…–‹‘•™‹–Šƒ›‘–Š‡”ˆƒ…–‘”ǡ™‡ƒƒŽ›œ‡†–Š‡ˆ—…–‹‘ƒŽ†ƒ–ƒƒ˜‡”ƒ‰‡†‘˜‡”
ϯϮϵ
Ž‡ˆ–ƒ†”‹‰Š–Šƒ†–”ƒ‹‹‰…‘Š‘”–•Ǥ
ϯϯϬ
Results
ϯϯϭ
ϯϯϮ
Bihemispheric tDCS increases motor learning more than unihemispheric tDCS
ϯϯϯ
‡ˆ‹”•– †‡–‡”‹‡† ™Š‡–Š‡”…‘˜‡–‹‘ƒŽ „‹Š‡‹•’Š‡”‹… – ‹• ‘”‡ ‡ˆˆ‡…–‹˜‡ ‹ ’”‘‘–‹‰
ϯϯϰ
Ž‡ƒ”‹‰–Šƒ—‹Š‡‹•’Š‡”‹…•–‹—Žƒ–‹‘Ǥ‡—•‡†–Š‡‡””‘”Ǧƒ†Œ—•–‡†‡š‡…—–‹‘–‹‡ȋǡ•‡‡
ϯϯϱ
‡–Š‘†•Ȍ ƒ• ƒ ‘˜‡”ƒŽŽ ’‡”ˆ‘”ƒ…‡ ‡ƒ•—”‡Ǥ › –Š‡ ‡† ‘ˆ Ͷ †ƒ›• ‘ˆ •‡“—‡…‡ –”ƒ‹‹‰ǡ
ϯϯϲ
—‹Š‡‹•’Š‡”‹…–”‡…‹’‹‡–•‡š‡…—–‡†•‡“—‡…‡•ͲǤʹͲ͵άͲǤͳͲͳ•ȋͳ͸Ǥ͸ΨȌˆƒ•–‡”–Šƒ•Šƒ
ϯϯϳ
ȋFȋͳǡ͵ͳȌ αͶǤͶ͵ǡp αͲǤͲͶ͵ǢFig.2a/bǡblack vs. blue linesȌǤ•‹’”‡˜‹‘—•”‡’‘”–•ȋ‡‹•ƒ†”‹–•…Šǡ
ϯϯϴ
ʹͲͳͳȌǡ –Š‹• ƒ†˜ƒ–ƒ‰‡ ’‡”•‹•–‡† †—”‹‰ –Š‡ ’‘•–Ǧ–‡•–ǡ ™Š‹…Š ™ƒ• …‘†—…–‡† ™‹–Š‘—– – ‘‡
ϯϯϵ
†ƒ›ƒˆ–‡”–Š‡ˆ‹ƒŽ–”ƒ‹‹‰•‡••‹‘ȋFȋͳǡ͵ͳȌ αͺǤͳ͵ǡp αͲǤͲͲͺȌǤŠ‡”‡™ƒ•‘•‹‰‹ˆ‹…ƒ–†‹ˆˆ‡”‡…‡
ϯϰϬ
„‡–™‡‡–Š‡Ž‡ˆ–ƒ†”‹‰Š–Šƒ†–”ƒ‹‡†‰”‘—’•‹–Š‡•‹œ‡‘ˆ–Š‹•‡ˆˆ‡…–ȋFȋͳǡ͵ͳȌαʹǤͲ͵ǡp αͲǤͳ͸ͶȌǤ
ϯϰϭ
ƒ”–‹…‹’ƒ–•™Š‘”‡…‡‹˜‡†…‘˜‡–‹‘ƒŽ„‹Š‡‹•’Š‡”‹…–™‡”‡ƒadditionalͲǤʹ͸Ͳά
ϯϰϮ
ͲǤͳͳͶ• ˆƒ•–‡” –Šƒ –Š‘•‡ ™Š‘ ”‡…‡‹˜‡† —‹Š‡‹•’Š‡”‹… •–‹—Žƒ–‹‘Ǥ Š‡ †‹ˆˆ‡”‡…‡ „‡–™‡‡
ϯϰϯ
„‹Š‡‹•’Š‡”‹… ƒ† —‹Š‡‹•’Š‡”‹… ‰”‘—’• ™ƒ• ”‡Ž‹ƒ„Ž‡ ȋFȋͳǡʹͶȌ α ͳʹǤͶͶǡ p α ͲǤͲͲʹȌǡ ƒ• ™ƒ• –Š‡
ϯϰϰ
†‹ˆˆ‡”‡…‡ „‡–™‡‡ „‹Š‡‹•’Š‡”‹… ƒ† •Šƒ ‰”‘—’• ȋFȋͳǡ͵ͲȌ α ͸ͶǤͲʹǡ p α ͸ǤʹͶǦͲͻȌǤ Š‡•‡
ϯϰϱ
†‹ˆˆ‡”‡…‡• ™‡”‡ ƒ‹–ƒ‹‡† †—”‹‰ –Š‡ ’‘•–Ǧ–‡•– ȋ”‡•’‡…–‹˜‡Ž›ǡ FȋͳǡʹͶȌ α ͳͲǤͻͺ ǡ p α ͲǤͲͲ͵ ƒ†
ϯϰϲ
Fȋͳǡ͵ͲȌ α ͷͷǤ͵Ͳʹǡ p α ʹǤ͹ͷ‡ǦͲͺȌǤ ‡…‡ǡ ‘˜‹‰ –Š‡ …ƒ–Š‘†‡ ˆ”‘ –Š‡ •—’”ƒ‘”„‹–ƒŽ ”‹†‰‡
ϯϰϳ
ȋ—‹Š‡‹•’Š‡”‹… –Ȍ –‘ ‹’•‹Žƒ–‡”ƒŽ ͳ ȋ„‹Š‡‹•’Š‡”‹… –Ȍ ›‹‡Ž†‡† ‡ƒ”Ž› –™‹…‡ –Š‡
ϭϬ
d,ZK>K&/W^/>dZ>KZdy/EDKdKZ>ZE/E' ϭϭ
ϯϰϴ
’‡”ˆ‘”ƒ…‡‰ƒ‹ȋ͵͹ǤͺΨ”‡Žƒ–‹˜‡–‘•ŠƒȌǤŠ‡”‡ˆ‘”‡ǡ™‡”‡’Ž‹…ƒ–‡Š‡”‡Š‹‰Š‡”‡ˆˆ‡…–‹˜‡‡••
ϯϰϵ
ˆ‘”„‹Š‡‹•’Š‡”‹…–”‡Žƒ–‹˜‡–‘—‹Š‡‹•’Š‡”‹…•–‹—Žƒ–‹‘ȋ‹‡•‡–ƒŽǤǡʹͲͲͺǢƒŠ‘—†‹‡–
ϯϱϬ
ƒŽǤǡʹͲͳͳǢƒ”‘ƒ†‹–‡›ǡʹͲͳ͵Ǣ‹†‡„‡”‰‡–ƒŽǤǡʹͲͳ͵Ǣ‡Š‡–ƒŽǤǡʹͲͳ͵Ǣƒ”‘•‡–ƒŽǤǡʹͲͳ͸Ȍ
ϯϱϭ
‹–Š‡…‘–‡š–‘ˆƒ—Ž–‹’Ž‡Ǧ†ƒ›Ž‡ƒ”‹‰•–—†›Ǥ
ϯϱϮ
ϯϱϯ
Reversed-polarity bihemispheric tDCS increases motor learning
ϯϱϰ
Š‡‹–‡”Š‡‹•’Š‡”‹……‘’‡–‹–‹‘‘†‡Ž•—’’‘•‡•–Šƒ––Š‡ƒ†˜ƒ–ƒ‰‡‘ˆ„‹Š‡‹•’Š‡”‹…”‡Žƒ–‹˜‡
ϯϱϱ
–‘ —‹Š‡‹•’Š‡”‹… – ƒ”‹•‡• †—‡ –‘ …ƒ–Š‘†ƒŽ •—’’”‡••‹‘ ‘ˆ ‹’•‹Žƒ–‡”ƒŽ …‘”–‡šǤ Š‹• ‹†‡ƒ
ϯϱϲ
’”‡†‹…–•–Šƒ––Š‡”‡˜‡”•‡†Ǧ’‘Žƒ”‹–›‘–ƒ‰‡•Š‘—Ž†ƒ––‡—ƒ–‡‘–‘”Ž‡ƒ”‹‰”‡Žƒ–‹˜‡–‘•Šƒǡ
ϯϱϳ
ƒ• ‹– †‡…”‡ƒ•‡• …‘–”ƒŽƒ–‡”ƒŽ ͳ ‡š…‹–ƒ„‹Ž‹–› „‘–Š †‹”‡…–Ž› –Š”‘—‰Š …ƒ–Š‘†ƒŽ ƒ† ‹†‹”‡…–Ž›
ϯϱϴ
–Š”‘—‰Š‹’•‹Žƒ–‡”ƒŽƒ‘†ƒŽ•–‹—Žƒ–‹‘Ǥ
ϯϱϵ
—”†ƒ–ƒǡŠ‘™‡˜‡”ǡ•Š‘™‡†–Š‡…‘˜‡”•‡ǣ”‡˜‡”•‡†Ǧ’‘Žƒ”‹–›„‹Š‡‹•’Š‡”‹…–Ž‡†–‘
ϯϲϬ
•‹‰‹ˆ‹…ƒ–Ž›ˆƒ•–‡”•–Šƒ•Šƒ•–‹—Žƒ–‹‘ǡ„‘–Š†—”‹‰–Š‡ˆ‹ƒŽ–”ƒ‹‹‰†ƒ›ȋFȋͳǡ͵ͳȌ αʹ͹Ǥͷͷǡ
ϯϲϭ
p αͳǤͳͷͶ‡ǦͲͷȌƒ††—”‹‰–Š‡’‘•–Ǧ–‡•–ȋFȋͳǡ͵ͲȌαʹͷǤͲǡp αʹǤ͵ʹ‡ǦͲͷȌǤ–ƒ–‹•–‹…ƒŽŽ›ǡ–Š‡’‡”ˆ‘”ƒ…‡
ϯϲϮ
‘ˆ –Š‡ ”‡˜‡”•‡† ’‘Žƒ”‹–› ‰”‘—’ ™ƒ• ‹†‹•–‹‰—‹•Šƒ„Ž‡ ˆ”‘ …‘˜‡–‹‘ƒŽ „‹Š‡‹•’Š‡”‹… –
ϯϲϯ
ȋŽƒ•– –”ƒ‹‹‰ †ƒ›ǣ Fȋͳǡʹ͵Ȍ α ͳǤͺͳǡ p α ͲǤͳͻͳǡ ’‘•–Ǧ–‡•–ǣ Fȋͳǡʹ͵Ȍ α ͳǤͻ͵ǡ p α ͲǤͳ͹ͺȌǤ Š‡ • ˆ‘” –Š‡
ϯϲϰ
…‘˜‡–‹‘ƒŽƒ†”‡˜‡”•‡†Ǧ’‘Žƒ”‹–›„‹Š‡‹•’Š‡”‹…‰”‘—’•™‡”‡ƒŽ•‘•›‡–”‹…ƒ…”‘••Ž‡ˆ–ƒ†
ϯϲϱ
”‹‰Š–Šƒ†–”ƒ‹‡†‰”‘—’•ǣ–Š‡–‰”‘—’šŠƒ†…‘Š‘”–‹–‡”ƒ…–‹‘™ƒ•‘–•‹‰‹ˆ‹…ƒ–ȋFȋͳǡʹ͵Ȍ
ϯϲϲ
αͲǤͳͳǡpαͲǤ͹ͶͷͻȌǤ˜‡–Š‘—‰Š”‡˜‡”•‡Ǧ’‘Žƒ”‹–›„‹Š‡‹•’Š‡”‹…–Ž‡†—‡”‹…ƒŽŽ›–‘„‡––‡”
ϯϲϳ
‘—–…‘‡• –Šƒ —‹Š‡‹•’Š‡”‹… –ǡ –Š‹• ‡ˆˆ‡…– †‹† ‘– ”‡ƒ…Š •–ƒ–‹•–‹…ƒŽ •‹‰‹ˆ‹…ƒ…‡ ȋŽƒ•–
ϯϲϴ
–”ƒ‹‹‰ †ƒ›ǣ FȋͳǡʹͶȌ α ʹǤͻ͹ǡ p α ͲǤͲͻ͹ǡ ’‘•–Ǧ–‡•–ǣ FȋͳǡʹͶȌ α ʹǤͲʹǡ p α ͲǤͳ͸ͺȌǤ ‘™‡˜‡”ǡ –Š‡ –™‘
ϯϲϵ
„‹Š‡‹•’Š‡”‹… ‰”‘—’• …‘„‹‡† ™‡”‡ •‹‰‹ˆ‹…ƒ–Ž› ˆƒ•–‡” –Šƒ –Š‡ —‹Š‡‹•’Š‡”‹… ‰”‘—’ ȋŽƒ•–
ϯϳϬ
–”ƒ‹‹‰†ƒ›ǣFȋͳǡ͵ͺȌαͺǤ͸ͳǡp αͲǤͲͲ͸ǡ’‘•–Ǧ–‡•–ǣFȋͳǡʹͶȌα͸Ǥ͸͸ǡp αͲǤͲͳͶȌǤ
ϯϳϭ
‘ŽŽ‡…–‹˜‡Ž›ǡ –Š‡•‡ ”‡•—Ž–• ‹†‹…ƒ–‡ –Šƒ– „‹Š‡‹•’Š‡”‹… •–‹—Žƒ–‹‘ ™ƒ• ‘”‡ ‡ˆˆ‡…–‹˜‡
ϯϳϮ
–Šƒ —‹Š‡‹•’Š‡”‹… •–‹—Žƒ–‹‘Ǥ Š‹• ƒ††‹–‹‘ƒŽ „‡‡ˆ‹–ǡ Š‘™‡˜‡”ǡ ™ƒ• ‘– …‘ˆ‡””‡† „›
ϯϳϯ
‹’•‹Žƒ–‡”ƒŽ •—’’”‡••‹‘ǡ ƒ• ™‡ †‹† ‘– ˆ‹† ƒ ’‘Žƒ”‹–›Ǧ•’‡…‹ˆ‹… ‡ˆˆ‡…– ˆ‘” „‹Š‡‹•’Š‡”‹… –Ǥ
ϯϳϰ
•–‡ƒ†ǡany •–‹—Žƒ–‹‘‘ˆ–Š‡‹’•‹Žƒ–‡”ƒŽ‘–‘”ƒ”‡ƒ•ƒ……‡Ž‡”ƒ–‡†‘–‘”Ž‡ƒ”‹‰Ǥ
ϯϳϱ
Š‡•‡ ”‡•—Ž–•ǡ –Š‡”‡ˆ‘”‡ǡ ˆƒ˜‘” –Š‡ „‹Š‡‹•’Š‡”‹… …‘‘’‡”ƒ–‹‘ ‘†‡Žǡ „› ™Š‹…Š –Š‡
ϯϳϲ
ƒ††‹–‹‘ƒŽ Ž‡ƒ”‹‰ ƒ†˜ƒ–ƒ‰‡ ‘ˆ „‹Š‡‹•’Š‡”‹… ”‡Žƒ–‹˜‡ –‘ —‹Š‡‹•’Š‡”‹… – ƒ”‹•‡• ˆ”‘
ϯϳϳ
’Žƒ•–‹……Šƒ‰‡•‹„‘–ŠŠ‡‹•’Š‡”‡•–Šƒ–™‘—Ž†’”‘‘–‡’‡”ˆ‘”ƒ…‡ˆ‘”„‘–ŠŠƒ†•ǤŠ‹•‹†‡ƒ
ϯϳϴ
ƒ‡• –™‘ –‡•–ƒ„Ž‡ ’”‡†‹…–‹‘•Ǥ ‹”•–ǡ ƒ …‘•‹†‡”ƒ„Ž‡ ’ƒ”– ‘ˆ –Š‡ „‡Šƒ˜‹‘”ƒŽ – ƒ†˜ƒ–ƒ‰‡
ϯϳϵ
•Š‘—Ž† ‰‡‡”ƒŽ‹œ‡ –‘ –Š‡ —–”ƒ‹‡† Šƒ†Ǥ ‡…‘†Ž›ǡ ‡—”ƒŽ …Šƒ‰‡• •Š‘—Ž† ‘……—” ‹ „‘–Š
ϯϴϬ
Š‡‹•’Š‡”‡• ‹ ƒ ’‘Žƒ”‹–›Ǧ—•’‡…‹ˆ‹… ƒ‡”Ǥ  –Š‡ ”‡ƒ‹†‡” ‘ˆ –Š‡ ’ƒ’‡”ǡ ™‡ –‡•– –Š‡•‡
ϯϴϭ
’”‡†‹…–‹‘•Ǥ
ϯϴϮ
ϭϭ
d,ZK>K&/W^/>dZ>KZdy/EDKdKZ>ZE/E' ϭϮ
ϯϴϯ
Behavioral tDCS effects generalize to the untrained hand
ϯϴϰ
–Š‡’”‡Ǧ–‡•–ƒ†’‘•–Ǧ–‡•–ǡ’ƒ”–‹…‹’ƒ–•Š‹•‰‡‡”ƒŽ‹œƒ–‹‘™ƒ•‡˜‡‘”‡’”‘‘—…‡†‹–Š‡
ϯϴϱ
– ‰”‘—’•ǣ ”‡Žƒ–‹˜‡ –‘ •Šƒǡ —‹Š‡‹•’Š‡”‹… ’‡”ˆ‘”‡† –”ƒ‹‡† •‡“—‡…‡• ™‹–Š –Š‡‹”
ϯϴϲ
—–”ƒ‹‡† Šƒ†ǡ ƒŽŽ‘™‹‰ —• –‘ ƒ••‡•• ‹–‡”Ǧƒ—ƒŽ ‰‡‡”ƒŽ‹œƒ–‹‘Ǥ ‘•‹•–‡– ™‹–Š ’”‡˜‹‘—•
ϯϴϳ
”‡•—Ž–• ȋƒ–‡”•Ǧ‡–‡‹‡” ‡– ƒŽǤǡ ʹͲͳͶǢ ‹‡•–Ž‡” ‡– ƒŽǤǡ ʹͲͳͶȌǡ ‡˜‡ –Š‡ •Šƒ ‰”‘—’ •Š‘™‡†
ϯϴϴ
…‘•‹†‡”ƒ„Ž‡’‡”ˆ‘”ƒ…‡‹’”‘˜‡‡–•‘–Š‡—–”ƒ‹‡†Šƒ†ȋFig.2c/dȌǤȋFȋͳǡ͵ͳȌ αͳʹǤͶ͵ǡp α
ϯϴϵ
ͲǤͲͲͳȌǡ„‹Š‡‹•’Š‡”‹…ȋFȋͳǡʹͶȌ αʹ͸Ǥͻͻǡ’αͳǤ͵Ͷ‡ǦͲͷȌǡƒ†”‡˜‡”•‡†Ǧ’‘Žƒ”‹–›„‹Š‡‹•’Š‡”‹…‰”‘—’•
ϯϵϬ
ȋFȋͳǡʹͶȌ α ʹͷǤͻͺǡ ’ α ͳǤ͹ͺ‡ǦͲͷȌ ƒŽŽ ’‡”ˆ‘”‡† •‹‰‹ˆ‹…ƒ–Ž› „‡––‡” ‘ –Š‡ —–”ƒ‹‡† Šƒ†Ǥ
ϯϵϭ
††‹–‹‘ƒŽŽ›ǡ ’‡”ˆ‘”ƒ…‡ ˆ‘” –Š‡ —–”ƒ‹‡† Šƒ† ™ƒ• „‡––‡” ‹ „‹Š‡‹•’Š‡”‹… ”‡Žƒ–‹˜‡ –‘
ϯϵϮ
—‹Š‡‹•’Š‡”‹…–”‡…‹’‹‡–•ȋFȋͳǡ͵ͳȌ αͶǤʹ͸ǡp αͲǤͲͶ͹Ȍǡƒ†–Š‡”‡™ƒ•‘†‹ˆˆ‡”‡…‡„‡–™‡‡
ϯϵϯ
–Š‡„‹Š‡‹•’Š‡”‹…–‰”‘—’•ȋFȋͳǡʹ͵ȌαͲǤͲͷǡp αͲǤͺʹʹȌǤ
ϯϵϰ
’‘”–ƒ–Ž›ǡ –Š‡ ‡Šƒ…‡‡– ‘ˆ —–”ƒ‹‡† Šƒ† ’‡”ˆ‘”ƒ…‡ ™ƒ• ‡˜‡ Žƒ”‰‡” –Šƒ
ϯϵϱ
™Šƒ– ™‘—Ž† Šƒ˜‡ „‡‡ ‡š’‡…–‡† ‹ˆ ‰‡‡”ƒŽ‹œƒ–‹‘ ™‡”‡ •‹’Ž› ’”‘’‘”–‹‘ƒŽ –‘ –Š‡
ϯϵϲ
‹’”‘˜‡‡–• ‘ –Š‡ –”ƒ‹‡† Šƒ†ǣ ™Š‡ ‡š’”‡••‹‰ ’”‡ǦȀ’‘•–Ǧ–‡•– †‹ˆˆ‡”‡…‡ ˆ‘” –Š‡
ϯϵϳ
—–”ƒ‹‡†Šƒ†”‡Žƒ–‹˜‡–Š‡Ž‡ƒ”‹‰ ‰ƒ‹• ˆ‘”–Š‡–”ƒ‹‡†Šƒ†ǡ™‡ˆ‘—† –Šƒ––‹…”‡ƒ•‡†
ϯϵϴ
–Š‡’”‘’‘”–‹‘‘ˆ‰‡‡”ƒŽ‹œƒ–‹‘Ǥ‘”•Šƒ”‡…‹’‹‡–•ǡ–Š‡—–”ƒ‹‡†Šƒ†‰ƒ‹‡†ͷͺǤͷΨ‘ˆ–Š‡
ϯϵϵ
‹’”‘˜‡‡–‘ˆ–Š‡–”ƒ‹‡†Šƒ†ǡ™Š‡”‡ƒ•ǡˆ‘”„‘–Š„‹Š‡‹•’Š‡”‹…–‰”‘—’•ǡ–Š‡’‡”…‡–ƒ‰‡
ϰϬϬ
‘ˆ‹–‡”Ǧƒ—ƒŽ‰‡‡”ƒŽ‹œƒ–‹‘™ƒ•‰”‡ƒ–‡”ȋͺ͸ǦͺͻǤʹΨǢtȋͳǡ͵͵Ȍ εʹǤ͸͸ʹǡp δͲǤͲͳʹȌǤŠ‡•‡”‡•—Ž–•
ϰϬϭ
•—‰‰‡•– –Šƒ– „‹Š‡‹•’Š‡”‹… – ‹ˆŽ—‡…‡† ƒ‹Ž› ‡ˆˆ‡…–‘”Ǧ‹†‡’‡†‡– ”‡’”‡•‡–ƒ–‹‘• ‹
ϰϬϮ
„‘–ŠŠ‡‹•’Š‡”‡•Ǥ
ϰϬϯ
ϰϬϰ
Behavioural tDCS effects generalize to untrained sequences
ϰϬϱ
ŽŽ’ƒ”–‹…‹’ƒ–•ƒŽ•‘‹’”‘˜‡†‘–Š‡—–”ƒ‹‡†•‡“—‡…‡ˆ”‘’”‡Ǧ–‡•––‘’‘•–Ǧ–‡•–ȋ•‡‡Fig.
ϰϬϲ
2dȌǤŠ‹•‡ˆˆ‡…–™ƒ•’”‘‘–‡†„›–ǡ•—…Š–Šƒ–ǡˆ‘”—–”ƒ‹‡†•‡“—‡…‡•ǡ—‹Š‡‹•’Š‡”‹…
ϰϬϳ
–”‡…‹’‹‡–•™‡”‡ͲǤ͵ͳͲάͲǤͳͲͷ•ȋͳͻǤͶΨȌˆƒ•–‡”–Šƒ•Šƒƒ†„‹Š‡‹•’Š‡”‹…”‡…‹’‹‡–•
ϰϬϴ
ȋ…‘˜‡–‹‘ƒŽƒ†”‡˜‡”•‡†Ǧ’‘Žƒ”‹–›‰”‘—’•…‘„‹‡†Ȍ™‡”‡ͲǤʹ͵ͳάͲǤͳ͵͹•ȋͳ͹Ǥ͸ΨȌˆƒ•–‡”
ϰϬϵ
–Šƒ—‹Š‡‹•’Š‡”‹…ƒ‘†ƒŽ–”‡…‹’‹‡–•ȋƒ†„‘–Š‘ˆ–Š‡•‡†‹ˆˆ‡”‡…‡•™‡”‡•‹‰‹ˆ‹…ƒ–Ǣ
ϰϭϬ
”‡•’‡…–‹˜‡Ž›ȋͳǡ͵ͳȌα͸Ǥ͹͵ǡ’αͲǤͲͳͶƒ†ȋʹǡ͵͸ȌαͺǤͳ͵ǡ’αͲǤͲͲͺȌǤ††‹–‹‘ƒŽŽ›ǡ™Š‡™‡
ϰϭϭ
“—ƒ–‹ˆ‹‡†–Š‡’”‘’‘”–‹‘‘ˆ–”ƒ•ˆ‡”‘ˆ•’‡‡†ƒ†˜ƒ–ƒ‰‡•–‘—–”ƒ‹‡†•‡“—‡…‡•ˆ‘”–Š‡
ϰϭϮ
–”ƒ‹‡†Šƒ†ǡ™‡ˆ‘—†–Šƒ–ǡƒ…”‘••ƒŽŽ‰”‘—’•ǡ–‹…”‡ƒ•‡†–Š‡’”‘’‘”–‹‘‘ˆ–”ƒ•ˆ‡””‡Žƒ–‹˜‡
ϰϭϯ
–‘•Šƒȋȋ͵ǡ͸ͲȌαͶǤͻͲǡ’αͲǤͲͲͶͳȌǤŠ‡”‡ˆ‘”‡ǡ–Š‡‡ˆˆ‡…–‘ˆ–‹–Š‹••–—†›‹•‡š‡”–‡†ƒ–ƒ
ϰϭϰ
Žƒ”‰‡Ž›‡ˆˆ‡…–‘”Ǧƒ†•‡“—‡…‡Ǧ—•’‡…‹ˆ‹…ˆƒ•Š‹‘ǡƒ•™‡Šƒ˜‡”‡’‘”–‡†’”‡˜‹‘—•Ž›ȋƒ–‡”•Ǧ
ϰϭϱ
‡–‡‹‡”‡–ƒŽǤǡʹͲͳͶȌǤ
ϰϭϲ
ϭϮ
d,ZK>K&/W^/>dZ>KZdy/EDKdKZ>ZE/E' ϭϯ
ϰϭϳ
Bihemispheric tDCS causes activation increases in both hemispheres
ϰϭϴ
‘‡Ž—…‹†ƒ–‡–Š‡‡—”ƒŽ…‘•‡“—‡…‡•‘ˆ–•–‹—Žƒ–‹‘ǡ™‡‡ƒ•—”‡†ˆƒ…–‹˜ƒ–‹‘
ϰϭϵ
™Š‹Ž‡ ’ƒ”–‹…‹’ƒ–• ‡š‡…—–‡† –Š‡ ˆ‘—” •‡“—‡…‡• ™‹–Š ‡‹–Š‡” –Š‡ –”ƒ‹‡† ‘” —–”ƒ‹‡† Šƒ†Ǥ
ϰϮϬ
ƒ”–‹…‹’ƒ–• ™‡”‡ •…ƒ‡† –™‘ †ƒ›• ƒˆ–‡” –Š‡‹” ˆ‹ƒŽ –Ǧ…‘—’Ž‡† –”ƒ‹‹‰ •‡••‹‘ǡ •—…Š –Šƒ–
ϰϮϭ
‘—” ‡ƒ•—”‡ ™‘—Ž† ”‡ˆŽ‡…– Ž‡ƒ”‹‰Ǧ”‡Žƒ–‡† ’Žƒ•–‹…‹–›ǡ ”ƒ–Š‡” –Šƒ ‹‡†‹ƒ–‡ ‡ˆˆ‡…–• ‘”
ϰϮϮ
ƒˆ–‡”‡ˆˆ‡…–•‘ˆ–‘‡—”ƒŽ‡š…‹–ƒ„‹Ž‹–›‘”Š‡‘†›ƒ‹…•™Š‹…Š‹‰Š–‘–„‡†‹”‡…–Ž›”‡Žƒ–‡†
ϰϮϯ
–‘ •‡“—‡…‡ Ž‡ƒ”‹‰ ȋ‡”‹‡”‹ ‡– ƒŽǤǡ ʹͲͳͲǢ ‹‡Ž‡ ‡– ƒŽǤǡ ʹͲͳ͵Ǣ –ƒŽ ‡– ƒŽǤǡ ʹͲͳͶȌǤ ‘ ’”‡˜‡–
ϰϮϰ
ƒ…–‹˜‹–› †‹ˆˆ‡”‡…‡• ƒ––”‹„—–ƒ„Ž‡ –‘ •‹’Ž‡ „‡Šƒ˜‹‘—”ƒŽ †‹ˆˆ‡”‡…‡•ǡ ™‡ …Ž‘•‡Ž› ƒ–…Š‡† –Š‡
ϰϮϱ
’‡”ˆ‘”ƒ…‡ ‹ –Š‡ •…ƒ‡” ‹ –‡”• ‘ˆ ‘˜‡‡– •’‡‡† ƒ† ˆ‘”…‡ ƒ…”‘•• „‘–Š ‰”‘—’• ƒ†
ϰϮϲ
Šƒ†•Ǥ Š‡Š‡‹•’Š‡”‹……‘‘’‡”ƒ–‹‘‘†‡Ž’”‡†‹…–•–Šƒ–•‹‹Žƒ”‡—”ƒŽ…Šƒ‰‡••Š‘—Ž†‘……—”‹
ϰϮϳ
„‘–ŠŠ‡‹•’Š‡”‡•ǡ‹†‡’‡†‡–Ž›‘ˆ’‘Žƒ”‹–›Ǥ•‡š’‡…–‡†ǡˆ‘”–”ƒ‹‡†Šƒ†‡š‡…—–‹‘•ǡ–Š‡•Šƒ
ϰϮϴ
‰”‘—’‡šŠ‹„‹–‡†…‘–”ƒŽƒ–‡”ƒŽƒ…–‹˜ƒ–‹‘ƒ†‹’•‹Žƒ–‡”ƒŽ†‡ƒ…–‹˜ƒ–‹‘‹ͳƒ†ͳȋFig. 3a, Fig.
ϰϮϵ
4Ȍǡ ƒ• ‘ˆ–‡ ‘„•‡”˜‡† †—”‹‰ •‹’Ž‡ —‹ƒ—ƒŽ Šƒ† ‘˜‡‡–• •—…Š ƒ• –Š‘•‡ –Šƒ– ™‡ •–—†›
ϰϯϬ
Š‡”‡ȋ‹‡†”‹…Š•‡‡–ƒŽǤǡʹͲͳ͵Ǣ‹‡•–Ž‡”ƒ†‹‡†”‹…Š•‡ǡʹͲͳ͵ȌǤ…‘–”ƒ•–ǡ„‘–Š„‹Š‡‹•’Š‡”‹…
ϰϯϭ
– ‰”‘—’•Ȅ‹†‡’‡†‡–Ž› ‘ˆ ’‘Žƒ”‹–›Ȅ•Š‘™‡† ‰”‡ƒ–‡” …‘–”ƒŽƒ–‡”ƒŽ ƒ…–‹˜ƒ–‹‘ ƒ† ‘
ϰϯϮ
‹’•‹Žƒ–‡”ƒŽ†‡ƒ…–‹˜ƒ–‹‘Ǥ
ϰϯϯ
ϰϯϰ
ƒ •‹‰Ž‡ Ǯ–ǯ ‰”‘—’ǡ ƒ• ™‡ †‹† ‘– ˆ‹† ƒ› •‹‰‹ˆ‹…ƒ– …Ž—•–‡”• ‘ˆ †‹ˆˆ‡”‡–‹ƒŽ ƒ…–‹˜ƒ–‹‘
ϰϯϱ
„‡–™‡‡…‘˜‡–‹‘ƒŽƒ†”‡˜‡”•‡†Ǧ’‘Žƒ”‹–›–‰”‘—’•Ǥ•—”ˆƒ…‡Ǧ„ƒ•‡†‰”‘—’ƒƒŽ›•‹•ȋ•‡‡
ϰϯϲ
ƒ–‡”‹ƒŽ•ƒ†‡–Š‘†•Ȍ•Š‘™‡†–Šƒ–„‘–Š…‘–”ƒŽƒ–‡”ƒŽƒ†‹’•‹Žƒ–‡”ƒŽ’”‹ƒ”›•‘ƒ–‘•‡•‘”›
ϰϯϳ
ȋͳȌ ƒ† ’”‹ƒ”› ‘–‘” ȋͳȌ …‘”–‡š Šƒ† ‰”‡ƒ–‡” ƒ…–‹˜ƒ–‹‘ ‹ – ”‡…‹’‹‡–• ȋTable 3Ȍ ˆ‘”
ϰϯϴ
‘˜‡‡–• ‘ˆ –Š‡ –”ƒ‹‡† Šƒ†Ǥ —…–‹‘ƒŽ †‹ˆˆ‡”‡…‡• ˆ‘” –Š‡ —–”ƒ‹‡† Šƒ† ™‡”‡ ˜‹•—ƒŽŽ›
‘”•–ƒ–‹•–‹…ƒŽ…‘’ƒ”‹•‘ǡ™‡…‘„‹‡†–Š‡–™‘„‹Š‡‹•’Š‡”‹…•–‹—Žƒ–‹‘‰”‘—’•‹–‘
ϰϯϵ
•‹‹Žƒ”ǡ„—–•–ƒ–‹•–‹…ƒŽŽ›Ž‡••’”‘‘—…‡†ȋFig. 3b; Fig. 4b; Table 3ȌǤ
ϰϰϬ
ϰϰϭ
‹‡†”‹…Š•‡ǡʹͲͳ͵Ȍǡ…‘ˆ‹”‡†–Š‡•‡”‡•—Ž–•ȋFig. 3eȌǤ …”‘••„‘–ŠŠ‡‹•’Š‡”‡•ƒ†Šƒ†•ǡ™‡
ϰϰϮ
‘„•‡”˜‡†ƒ…–‹˜ƒ–‹‘‹…”‡ƒ•‡•‹ͳȋFȋͳǡͶͲȌαͻǤ͵Ͷǡ p αͲǤͲͲͶȌƒ†ͳȋFȋͳǡͶͲȌαͳ͵ǤͷͷǡpαͲǤͲͲͲ͹ȌǤ
ϰϰϯ
Š‡”‡™ƒ••‘‡‡˜‹†‡…‡ˆ‘”ƒ‹–‡”ƒ…–‹‘‘ˆ–šŠ‡‹•’Š‡”‡šŠƒ†‹ͳȋFȋͳǡͶͲȌαͶǤͳ͸ǡpα
ƒŽ›•‹• —•‹‰ ƒƒ–‘‹…ƒŽŽ› ’”‡†‡ˆ‹‡† ”‡‰‹‘• ‘ˆ ‹–‡”‡•–• ȋ•Ȍ ȋ‹‡•–Ž‡” ƒ†
ϰϰϰ
ͲǤͲͶͺȌǡƒŽ–Š‘—‰Š–Š‹•†‹†‘–“—‹–‡”‡ƒ…Š•‹‰‹ˆ‹…ƒ…‡‹ͳǣFȋͳǡͶͲȌα͵Ǥ͹ʹǡpαͲǤͲ͸ͳǤŠ‹•”‡ˆŽ‡…–•
ϰϰϱ
–Šƒ– „‹Š‡‹•’Š‡”‹… – ‡•’‡…‹ƒŽŽ› ‹…”‡ƒ•‡† ƒ…–‹˜ƒ–‹‘ ƒ••‘…‹ƒ–‡† ™‹–Š –Š‡ –”ƒ‹‡† Šƒ† ƒ†
ϰϰϲ
‹’•‹Žƒ–‡”ƒŽȋ—–”ƒ‹‡†ȌŠ‡‹•’Š‡”‡Ǥ
ϰϰϳ
ϰϰϴ
‹ –Š‡ Ž‡ˆ– Š‡‹•’Š‡”‡ †—”‹‰ Ž‡ˆ–ǦŠƒ† ‘˜‡‡–• ƒ• …‘’ƒ”‡† –‘ ƒ…–‹˜‹–› ‹ –Š‡ ”‹‰Š–
ϰϰϵ
Š‡‹•’Š‡”‡ †—”‹‰ ”‹‰Š–ǦŠƒ† ‘˜‡‡–• ȋ‡”•–›‡ ‡– ƒŽǤǡ ʹͲͲͷȌǤ Š‹• Š‡‹•’Š‡”‹…
ϰϱϬ
ƒ•›‡–”›™ƒ••‹‰‹ˆ‹…ƒ–ˆ‘”ͳȋȋͳǡ͵ͻȌα͵ͳǤͳ͵͹ǡ’αͳǤͻͺ͵‡Ǧ͸Ȍƒ†ͳȋȋͳǡ͵ͻȌαʹͳǤ͹͵ǡ’α͵Ǥ͸ʹͳ‡ǦͷȌǡ
ϰϱϭ
„—–†‹†‘–‹–‡”ƒ…–•‹‰‹ˆ‹…ƒ–Ž›™‹–Š–Š‡–‡ˆˆ‡…–Ǥ
‡ƒŽ•‘‘„•‡”˜‡†–Š‡‡š’‡…–‡†Š‡‹•’Š‡”‹…ƒ•›‡–”‹‡•™‹–Š‘”‡‹’•‹Žƒ–‡”ƒŽƒ…–‹˜‹–›
ϭϯ
d,ZK>K&/W^/>dZ>KZdy/EDKdKZ>ZE/E' ϭϰ
ϰϱϮ
ϰϱϯ
„—–•‹‰‹ˆ‹…ƒ–†‹ˆˆ‡”‡…‡•„‡–™‡‡–Š‡‰”‘—’•ȋTable 4ȌǤ‘™‡˜‡”ǡ–Š‡†‹ˆˆ‡”‡…‡•‹™‡”‡
ϰϱϰ
˜‡”› •ƒŽŽ ȋ‘ –Š‡ ‘”†‡” ‘ˆ ͶͲǦ͸Ͳ •Ȍǡ ƒ† –Š‡ †‹ˆˆ‡”‡…‡• ‹ ˆ‘”…‡ ”‡Žƒ–‹˜‡ –‘ •Šƒ ™‡”‡ ‘–
ϰϱϱ
…‘•‹•–‡–ƒ…”‘••–Š‡–™‘„‹Š‡‹•’Š‡”‹…–‰”‘—’•ȋˆ‘”…‡™ƒ••Ž‹‰Š–Ž›Ž‘™‡”–Šƒ•Šƒ‹–Š‡
ϰϱϲ
…‘˜‡–‹‘ƒŽ„‹Š‡‹•’Š‡”‹…–‰”‘—’ƒ†•Ž‹‰Š–Ž›Š‹‰Š‡”–Šƒ•Šƒ‹–Š‡”‡˜‡”•‡†Ǧ’‘Žƒ”‹–›
ϰϱϳ
– ‰”‘—’ȌǤ ‡˜‡”–Š‡Ž‡••ǡ –‘ ‡•—”‡ –Šƒ– –Š‡ ‘„•‡”˜‡† ‹…”‡ƒ•‡• ‹ ƒ…–‹˜ƒ–‹‘ …‘—Ž† ‘– „‡
ϰϱϴ
ƒ––”‹„—–ƒ„Ž‡ –‘ –Š‡•‡ •ƒŽŽ „‡Šƒ˜‹‘”ƒŽ †‹ˆˆ‡”‡…‡•ǡ ™‡ ‹…Ž—†‡† ‡š‡…—–‹‘ –‹‡ǡ ‡””‘” ”ƒ–‡ǡ ƒ†
ϰϱϵ
ˆ‘”…‡ƒ•…‘˜ƒ”‹ƒ–‡•‹ƒƒŽ›•‡•ƒ†ˆ‘—†–Šƒ–ǡˆ‘”ƒŽŽ…‘’ƒ”‹•‘•ǡ–Š‡‡ˆˆ‡…–‘ˆ–
ϰϲϬ
”‡ƒ‹‡†•‹‰‹ˆ‹…ƒ–ˆ‘”„‘–Šͳȋ”‡•’‡…–‹˜‡Ž›ǡ Fȋͳǡ͵ͻȌ α͹Ǥʹ͸ǡ p αͲǤͲͳǢFȋͳǡ͵ͻȌ αͳͶǤͲ͵ǡ p αͲǤͲͲͳǢ
ϰϲϭ
Fȋͳǡ͵ͻȌαͺǤʹͻǡ p αͲǤͲͲ͸Ȍƒ†ͳȋFȋͳǡ͵ͻȌαͺǤͲʹǡ p αͲǤͲͲ͹ǢFȋͳǡ͵ͻȌαͳͲǤͻͻǡ p αͲǤͲͲʹǢFȋͳǡ͵ͻȌαͳʹǤ͹ͺǡ p
ϰϲϮ
αͲǤͲͲͳȌǤ
ϰϲϯ
‡•’‹–‡‘—”„‡•–‡ˆˆ‘”–•–‘ƒ–…Š„‡Šƒ˜‹‘”ƒŽ’‡”ˆ‘”ƒ…‡†—”‹‰ˆǡ–Š‡”‡™‡”‡•Ž‹‰Š–ǡ
‘–Š‡” ’‘–‡–‹ƒŽ …‘ˆ‘—† –Šƒ– …‘—Ž† Ž‡ƒ† –‘ ‹…”‡ƒ•‡† ‹’•‹Žƒ–‡”ƒŽ ƒ…–‹˜ƒ–‹‘ ‹•
ϰϲϰ
‹””‘”‹‰Ǥ ‹””‘”‹‰ ”‡ˆ‡”• –‘ –Š‡ ’Š‡‘‡‘ ™Š‡”‡„›—•…Ž‡•‘ˆ –Š‡ ‘Ǧ‘˜‹‰ Šƒ†ƒ”‡
ϰϲϱ
ƒ…–‹˜ƒ–‡†•‹—Ž–ƒ‡‘—•Ž›™‹–Š–Š‘•‡‘ˆ–Š‡‘˜‹‰Šƒ†ȋ‡ƒ—Ž‡‡–ƒŽǤǡʹͲͳ͵ȌǤ—…Š‘˜‡‡–•
ϰϲϲ
ƒ”‡–›’‹…ƒŽŽ›˜‹•‹„Ž‡‹’ƒ–Š‘Ž‘‰‹…ƒŽ•–ƒ–‡•ȋ‡Ǥ‰Ǥ•–”‘‡Ȍǡ„—–ƒ”‡ƒŽ•‘’”‡•‡–ƒ†‡ƒ•—”ƒ„Ž‡‹
ϰϲϳ
Š‡ƒŽ–Š› ’‘’—Žƒ–‹‘•Ǥ —”‹‰ ˆǡ ’ƒ”–‹…‹’ƒ–• ™‡”‡ ”‡“—‹”‡†–‘ ”‡•– –Š‡’ƒ••‹˜‡Šƒ† ‘ –Š‡
ϰϲϴ
‡›„‘ƒ”† ™Š‹Ž‡ –Š‡ ƒ…–‹˜‡ Šƒ† ™ƒ• ‡š‡…—–‹‰ •‡“—‡…‡•Ǥ ‹””‘”‹‰ ™ƒ• ’ƒ”ƒ‡–‡”‹œ‡† —•‹‰
ϰϲϵ
–Š‡”ƒ‰‡‘ˆˆ‘”…‡•‘–Š‡’ƒ••‹˜‡Šƒ†ƒ…”‘••–Š‡–‹‡…‘—”•‡‘ˆƒ–”‹ƒŽǡ–Š‡ƒ••‘…‹ƒ–‡†•–ƒ†ƒ”†
ϰϳϬ
†‡˜‹ƒ–‹‘ǡ ƒ† –Š‡ …‘””‡Žƒ–‹‘ „‡–™‡‡ –Š‡ ˆ‘”…‡ –”ƒ…‡• ˆ‘” ƒ–…Š‹‰ †‹‰‹–• ‘ˆ –Š‡ ’ƒ••‹˜‡ ƒ†
ϰϳϭ
ƒ…–‹˜‡ Šƒ†Ǥ ˜‡ –Š‘—‰Š –Š‡ •‹‰‹ˆ‹…ƒ– ’‘•‹–‹˜‡ …‘””‡Žƒ–‹‘• ‹†‹…ƒ–‡† –Šƒ– ™‡ …‘—Ž†
ϰϳϮ
•—……‡••ˆ—ŽŽ›†‡–‡…––Š‡˜‡”›•—„–Ž‡‹””‘”‹‰‹‘—”Š‡ƒŽ–Š›…‘–”‘Ž’ƒ”–‹…‹’ƒ–•ǡ–Š‡”‡™‡”‡‘
ϰϳϯ
•‹‰‹ˆ‹…ƒ–†‹ˆˆ‡”‡…‡„‡–™‡‡–‰”‘—’•ˆ‘”ƒ›‘ˆ–Š‡•‡‡ƒ•—”‡•ȋTable 5ȌǤ
ϰϳϰ
ƒ‡ –‘‰‡–Š‡”ǡ –Š‡•‡ †ƒ–ƒ †‡‘•–”ƒ–‡ –Šƒ– „‹Š‡‹•’Š‡”‹… –Ȅ‹””‡•’‡…–‹˜‡ ‘ˆ –Š‡
ϰϳϱ
’‘Žƒ”‹–›Ȅ™ƒ• ƒ••‘…‹ƒ–‡† ™‹–Š ‹…”‡ƒ•‡• ‹ ƒ˜‡”ƒ‰‡ ƒ…–‹˜ƒ–‹‘ ‹ „‘–Š ‹’•‹Žƒ–‡”ƒŽ ƒ†
ϰϳϲ
…‘–”ƒŽƒ–‡”ƒŽ Š‡‹•’Š‡”‡•Ǥ Š‹• †‹ˆˆ‡”‡…‡ …‘—Ž† ‘– „‡ ‡š’Žƒ‹‡† „› „‡Šƒ˜‹‘”ƒŽ †‹ˆˆ‡”‡…‡•
ϰϳϳ
„‡–™‡‡–Š‡‰”‘—’•†—”‹‰–Š‡•…ƒǤ
ϰϳϴ
ϰϳϵ
Quantification of subject blindedness and perceptual side effects of tDCS
ϰϴϬ
—„•‡“—‡– –‘ ‡ƒ…Š •‡••‹‘ ‘ˆ – ƒ†‹‹•–”ƒ–‹‘ǡ ’ƒ”–‹…‹’ƒ–• —†‡”™‡– ƒ ’”‡˜‹‘—•Ž›
ϰϴϭ
†‡•‹‰‡†„ƒ––‡”›–‘…Šƒ”ƒ…–‡”‹œ‡–‡ˆˆ‡…–•ȋƒ–‡”•Ǧ‡–‡‹‡”‡–ƒŽǤǡʹͲͳͶȌǡ™‹–Š–Š‡ƒ††‹–‹‘‘ˆ
ϰϴϮ
“—‡•–‹‘• ƒ„‘—– ’‡”…‡’–—ƒŽ •‹†‡ ‡ˆˆ‡…–•Ǥ ’‡…‹ˆ‹…ƒŽŽ›ǡ ’ƒ”–‹…‹’ƒ–• ™‡”‡ ƒ•‡† –‘ ”ƒ–‡ –Š‡
ϰϴϯ
‡š’‡”‹‡…‡ ‘ˆ –‹‰Ž‹‰ǡ ’ƒ‹ǡ „—”‹‰ǡ ‹–…Š‹‰ǡ †‹œœ‹‡••ǡ ƒ† ‡–ƒŽ ˆƒ–‹‰—‡ ȋͳǣͳͲȌ ƒ† –Š‡ –‘
ϰϴϰ
†‡•…”‹„‡ȋ‹‹Ȍ–Š‡†—”ƒ–‹‘‘ˆ–Š‡‡ˆˆ‡…–Ǥ•…ƒ„‡•‡‡‹Fig. 5aǡ–”‡…‹’‹‡–•–‡†‡†–‘
ϰϴϱ
‡šŠ‹„‹– •Ž‹‰Š–Ž› Š‹‰Š‡” ‹–‡•‹–‹‡• ‘ˆ ’‡”…‡’–—ƒŽ ’”‘’‡”–‹‡• ‘ˆ –Ȅ‡•’‡…‹ƒŽŽ› ‹ –‡”• ‘ˆ
ϰϴϲ
–‹‰Ž‹‰ǡ„—”‹‰ƒ†‹–…Š‹‰ȄŠ‘™‡˜‡”ǡ‘‡‘ˆ–Š‡•‡˜ƒ”‹ƒ„Ž‡••Š‘™‡†ƒ•‹‰‹ˆ‹…ƒ–„‡–™‡‡Ǧ
ϭϰ
d,ZK>K&/W^/>dZ>KZdy/EDKdKZ>ZE/E' ϭϱ
ϰϴϳ
‰”‘—’†‹ˆˆ‡”‡…‡ȋƒŽŽtȋ͸ʹȌ δͳǤͶͶ͸ǡp εͲǤͳͷ͵Ȍǡ‘”‡š…‡‡†‡†Ž‡˜‡ŽͶȋ‘—–‘ˆƒƒš‹ƒŽ”ƒ–‹‰‘ˆͳͲȌ
ϰϴϴ
ˆ‘”ƒ›‘ˆ–Š‡͸Ͷ•—„Œ‡…–•Ǥ
ϰϴϵ
‘™‡˜‡”ǡ –Š‡ ‰”‘—’• –‡†‡† –‘ †‹ˆˆ‡” ‘ Š‘™ Ž‘‰ –Š‡› ‡š’‡”‹‡…‡† –Š‡•‡ •‹†‡ ‡ˆˆ‡…–•
ϰϵϬ
ȋFig. 5b)Ǥ ˜‡”ƒ‰‡† ƒ…”‘•• –Š‡ –Š”‡‡ – ‰”‘—’• ȋ—‹Š‡‹•’Š‡”‹…ǡ „‹Š‡‹•’Š‡”‹… ƒ†
ϰϵϭ
”‡˜‡”•‡†Ǧ’‘Žƒ”‹–› „‹Š‡‹•’Š‡”‹…Ȍǡ •—„Œ‡…–• ”‡’‘”–‡† •‹‰‹ˆ‹…ƒ–Ž› Ž‘‰‡” †—”ƒ–‹‘• ‘ˆ –‹‰Ž‹‰
ϰϵϮ
ȋtȋ͸ʹȌ αʹǤͷͷ͵ǡp αͲǤͲͳ͵Ȍǡ„—”‹‰ȋtȋ͸ʹȌ αʹǤͶͻʹǡp αͲǤͲͳͷȌǡƒ†‹–…Š‹‰ȋtȋ͸ʹȌ αʹǤͺ͵͵ǡp αͲǤͲͲ͸Ȍǡ
ϰϵϯ
ƒ†ƒ–‡†‡…›ˆ‘”Ž‘‰‡”†—”ƒ–‹‘‘ˆ’ƒ‹ȋtȋ͸ʹȌ αͳǤͺͲ͸ǡp αͲǤͲ͹͸ȌǤŠ‡”‡™‡”‡‘†‹ˆˆ‡”‡…‡•
ϰϵϰ
‹–Š‡†—”ƒ–‹‘‘ˆ†‹œœ‹‡••ȋtȋ͸ʹȌ αͳǤͲͷͳǡp αͲǤʹͻͺȌ‘”‡–ƒŽˆƒ–‹‰—‡ȋtȋ͸ʹȌ αͲǤ͸͹ͳǡp αͲǤͷͲͷȌ
ϰϵϱ
„‡–™‡‡ •Šƒ ƒ† – ‰”‘—’•Ǥ • ‘„•‡”˜‡† ’”‡˜‹‘—•Ž› ȋƒ–‡”•Ǧ‡–‡‹‡” ‡– ƒŽǤǡ ʹͲͳͶȌǡ
ϰϵϲ
’ƒ”–‹…‹’ƒ–• ‡š’‡”‹‡…‡† ‘ •‹‰‹ˆ‹…ƒ– †‹ˆˆ‡”‡…‡• ‹ ‘˜‡”ƒŽŽ †‹•…‘ˆ‘”–ǡ ’‡”…‡‹˜‡† –
ϰϵϳ
‹–‡•‹–›ǡ‘”†‹•–”ƒ…–‹‘†—‡–‘–ȋTable 2ȌǤ‘”‡‘˜‡”ǡ χ Ǧ‰‘‘†‡••‘ˆˆ‹––‡•–••Š‘™‡†–Šƒ–
ϰϵϴ
–Š‡”‡ ™‡”‡ ‘ †‹ˆˆ‡”‡…‡• ‹ –Š‡ †‡–‡…–ƒ„‹Ž‹–› ‘ˆ – ƒ••‹‰‡– „‡–™‡‡ – ‰”‘—’• ƒ†
ϰϵϵ
•ŠƒȋTable 2ȌǤŠ‡•‡ˆ‹†‹‰•…‘ŽŽ‡…–‹˜‡Ž›•—‰‰‡•––Šƒ–ǡ†‡•’‹–‡•Ž‹‰Š–’‡”…‡’–—ƒŽ†‹ˆˆ‡”‡…‡•ǡ
ϱϬϬ
–„Ž‹†‹‰‡–Š‘†•™‡”‡•—ˆˆ‹…‹‡–ǤŠ‹•‹•…‘‰”—‡–™‹–Š‘–Š‡””‡…‡–‡˜‹†‡…‡ȋ‡••Ž‡”‡–
ϱϬϭ
ƒŽǤǡʹͲͳʹȌǡ‹…Ž—†‹‰ƒ•–—†›–Šƒ–‹˜‡•–‹‰ƒ–‡†‘˜‡”–™‹…‡ƒ•ƒ›•—„Œ‡…–•ƒ–ʹȋ—••‘‡–ƒŽǤǡ
ϱϬϮ
ʹͲͳ͵ȌǤ
ϱϬϯ
ϱϬϰ
Assessment of behavioral side effects of tDCS
ϱϬϱ
 ƒ††‹–‹‘ –‘ –Š‡ ƒ‹ –”ƒ‹‹‰ –ƒ• ȋ–Š‡ •‡“—‡…‡ –ƒ• †‡•…”‹„‡† ƒ„‘˜‡Ȍǡ ™‡ –‡•–‡† ƒŽŽ
ϱϬϲ
„‹Š‡‹•’Š‡”‹…–ƒ†•Šƒ”‡…‹’‹‡–•‘–™‘ƒ††‹–‹‘ƒŽ–ƒ••‘ˆƒ—ƒŽ•‹ŽŽ†—”‹‰–Š‡’”‡Ǧ
ϱϬϳ
–‡•– ƒ† ’‘•–Ǧ–‡•–ǣ ‹†‹˜‹†—ƒ–‹‘ ȋ–Š‡ ƒ„‹Ž‹–› –‘ ‘˜‡ –Š‡ †‹‰‹–• •‡’ƒ”ƒ–‡Ž›Ȍ ƒ† …‘ˆ‹‰—”ƒ–‹‘
ϱϬϴ
‡š‡…—–‹‘ ȋ–Š‡ •‹ŽŽ ‘ˆ ’”‡••‹‰ …‡”–ƒ‹ †‹‰‹–• ƒ– –Š‡ •ƒ‡ –‹‡ ™Š‹Ž‡ ‡‡’‹‰ …‘ƒ…–‹˜ƒ–‹‘ ‘ˆ
ϱϬϵ
—‹–‡†‡† †‹‰‹–• ‹‹ƒŽȌȄˆ‘” ˆ—ŽŽ †‡–ƒ‹Ž• ‘ˆ –Š‡•‡ –ƒ••ǡ •‡‡ ƒ–‡”•Ǧ‡–‡‹‡” ‡– ƒŽǤ ȋʹͲͳͶȌǤ
ϱϭϬ
‡‹–Š‡” „‹Š‡‹•’Š‡”‹… – ‰”‘—’ ‡šŠ‹„‹–‡† ƒ› •‹†‡ ‡ˆˆ‡…–• ˆ”‘ –Ǧ…‘—’Ž‡† •‡“—‡…‡
ϱϭϭ
–”ƒ‹‹‰‘‡‹–Š‡”‘ˆ–Š‡•‡•‹ŽŽ•ȋFig. 5c, dȌǡƒ†–Š‡”‡™‡”‡‘•‹‰‹ˆ‹…ƒ–†‹ˆˆ‡”‡…‡•„‡–™‡‡
ϱϭϮ
–Š‡ –™‘ „‹Š‡‹•’Š‡”‹… ‰”‘—’• ƒ† •Šƒ ȋƒŽŽ Fȋʹǡ͵ͷȌδʹǤͲͻʹǡ ’εͲǤͳ͵ͻǢ —•‹‰ …‘””‡…–‹‘
ϱϭϯ
™‹–Š ’”‡Ǧ–‡•– ’‡”ˆ‘”ƒ…‡ȌǤ Š‡”‡ˆ‘”‡ǡ ƒ• ‹ ‘—” ’”‡˜‹‘—• ™‘”ǡ ™‡ †‹† ‘– ˆ‹† ƒ› ƒ†˜‡”•‡
ϱϭϰ
–”ƒ†‡Ǧ‘ˆˆ• „‡–™‡‡ —•‹‰ – –‘ ˆƒ…‹Ž‹–ƒ–‡ ƒ—ƒŽ ‘–‘” •‹ŽŽ Ž‡ƒ”‹‰ ƒ† ’‡”ˆ‘”ƒ…‡ ‘
ϱϭϱ
—–”ƒ‹‡†–ƒ••ȋƒ–‡”•Ǧ‡–‡‹‡”‡–ƒŽǤǡʹͲͳͶȌǤ
ϱϭϲ
Discussion
ϱϭϳ
ϱϭϴ
—”•–—†›’”‘˜‹†‡•‡˜‹†‡…‡ˆ‘”ƒƒ…–‹˜‡”‘Ž‡‘ˆ‹’•‹Žƒ–‡”ƒŽ‘–‘””‡‰‹‘•‹—‹ƒ—ƒŽ‘–‘”
ϱϭϵ
•‹ŽŽ Ž‡ƒ”‹‰Ǥ ‡ ”‡’Ž‹…ƒ–‡† –Š‡ …Žƒ••‹… ‘„•‡”˜ƒ–‹‘ –Šƒ– „‹Š‡‹•’Š‡”‹… – ‹• ‘”‡ ‡ˆˆ‡…–‹˜‡
ϱϮϬ
–Šƒ •–‹—Žƒ–‹‰ ‘Ž› ‘‡ Š‡‹•’Š‡”‡ ȋ‹‡• ‡– ƒŽǤǡ ʹͲͲͺȌǤ ”‹–‹…ƒŽŽ›ǡ „‹Š‡‹•’Š‡”‹… – ™‹–Š
ϱϮϭ
–Š‡ ƒ‘†‡ ‘˜‡” ‹’•‹Žƒ–‡”ƒŽ ‘–‘” …‘”–‡š Ž‡† –‘ •‹‹Žƒ” Ž‡ƒ”‹‰ ƒ†˜ƒ–ƒ‰‡• ƒ• ™‹–Š –Š‡ ƒ‘†‡
ϭϱ
d,ZK>K&/W^/>dZ>KZdy/EDKdKZ>ZE/E' ϭϲ
ϱϮϮ
‘˜‡” …‘–”ƒŽƒ–‡”ƒŽ ‘–‘” …‘”–‡šǤ ‹ƒŽŽ›ǡ „‘–Š ‘–ƒ‰‡• Ž‡† –‘ Ž‘‰ǦŽƒ•–‹‰ ‹…”‡ƒ•‡• ‘ˆ
ϱϮϯ
ˆ—…–‹‘ƒŽ ƒ…–‹˜‹–› ‹ „‹Žƒ–‡”ƒŽ •‡•‘”‹‘–‘” ƒ”‡ƒ• ƒ† –Š‡ –Ǧ‹†—…‡† Ž‡ƒ”‹‰ ƒ†˜ƒ–ƒ‰‡
ϱϮϰ
‰‡‡”ƒŽ‹œ‡†–‘–Š‡—–”ƒ‹‡†Šƒ†Ǥ
ϱϮϱ
—” ”‡•—Ž–• …Ž‡ƒ”Ž› ƒ”‰—‡ ƒ‰ƒ‹•– –Š‡ ‹–‡”Š‡‹•’Š‡”‹…Ǧ…‘’‡–‹–‹‘ ‘†‡Ž ƒ• ƒ
ϱϮϲ
‡š’Žƒƒ–‹‘ ˆ‘” –Š‡ ƒ†˜ƒ–ƒ‰‡ ‘ˆ „‹Š‡‹•’Š‡”‹… ‘˜‡” —‹Š‡‹•’Š‡”‹… –Ǥ ……‘”†‹‰ –‘ –Š‹•
ϱϮϳ
‹†‡ƒǡ …ƒ–Š‘†ƒŽ •–‹—Žƒ–‹‘ •—’’”‡••‡• –Š‡ ‹’•‹Žƒ–‡”ƒŽ Š‡‹•’Š‡”‡ǡ •—„•‡“—‡–Ž› ”‡Ž‡ƒ•‹‰ –Š‡
ϱϮϴ
…‘–”ƒŽƒ–‡”ƒŽ Š‡‹•’Š‡”‡ ˆ”‘ ‹–‡”Š‡‹•’Š‡”‹… •—’’”‡••‹‘Ǥ —” ”‡•—Ž–• ˆ‘” –Š‡ ”‡˜‡”•‡†Ǧ
ϱϮϵ
’‘Žƒ”‹–›„‹Š‡‹•’Š‡”‹…‰”‘—’ƒ”‡ƒ–‘††•™‹–Š–Š‹•‡š’Žƒƒ–‹‘ǡ„‡…ƒ—•‡‡š…‹–ƒ–‘”›•–‹—Žƒ–‹‘
ϱϯϬ
‘ˆ–Š‡‹’•‹Žƒ–‡”ƒŽ…‘”–‡š•Š‘—Ž†Šƒ˜‡Ž‡†–‘ƒ––‡—ƒ–‡†ǡ”ƒ–Š‡”–Šƒƒ……‡Ž‡”ƒ–‡†ǡ‘–‘”Ž‡ƒ”‹‰
ϱϯϭ
”‡Žƒ–‹˜‡–‘—‹Š‡‹•’Š‡”‹…ȋ‘”‡˜‡•ŠƒȌ•–‹—Žƒ–‹‘Ǥ
ϱϯϮ
•–‡ƒ†ǡ–Š‡ˆ—ŽŽ’ƒ––‡”‘ˆ‘—””‡•—Ž–•…ƒ„‡‡š’Žƒ‹‡†—†‡”–Š‡ƒ••—’–‹‘–Šƒ––
ϱϯϯ
‹…”‡ƒ•‡† ’Žƒ•–‹…‹–› ‹ „‘–Š Š‡‹•’Š‡”‡• ‹†‡’‡†‡–Ž› ‘ˆ ’‘Žƒ”‹–›ǡ ƒ† –Šƒ– –Š‡ –™‘ ‘–‘”
ϱϯϰ
…‘”–‹…‡•…‘‘’‡”ƒ–‡‹’”‘†—…‹‰Š‹‰ŠŽ‡˜‡Ž•‘ˆ•‹ŽŽȋ•‡‡Table 1ȌǤ‡Ž‹‹–ƒ–‹‘‘ˆ‘—”•–—†›‹•
ϱϯϱ
–Šƒ– ™‡ †‹† ‘– †‹”‡…–Ž› ‡ƒ•—”‡ …Šƒ‰‡• ‹ „‹Žƒ–‡”ƒŽ • „‡ˆ‘”‡ ƒ† ƒˆ–‡” –Ǥ Š—•ǡ ™‡
ϱϯϲ
…ƒ‘– ƒ‡ •–”‘‰ ‹ˆ‡”‡…‡ ƒ„‘—– ™Š‡–Š‡” –Š‡ …ƒ–Š‘†ƒŽ •–‹—Žƒ–‹‘ ‹ –Š‡ „‹Š‡‹•’Š‡”‹…
ϱϯϳ
‘–ƒ‰‡ ‹…”‡ƒ•‡† ‘” †‡…”‡ƒ•‡† •Š‘”– –‡” ‡š…‹–ƒ„‹Ž‹–› ‹ ͳǤ ‘” ‡šƒ’Ž‡ǡ –Š‡”‡ ‹• •‘‡
ϱϯϴ
‡˜‹†‡…‡ –Šƒ– …ƒ–Š‘†ƒŽ —‹Š‡‹•’Š‡”‹… – •–‹—Žƒ–‹‘ ƒ– ʹ  ‹…”‡ƒ•‡•ǡ ”ƒ–Š‡” –Šƒ
ϱϯϵ
†‡…”‡ƒ•‡•ǡ • ȋ‘”†‹ŽŽ‘Ǧƒ–‡‘• ‡– ƒŽǤǡ ʹͲͳʹǢ ƒ–•‹ƒ†œ‡ ‡– ƒŽǤǡ ʹͲͳ͵Ǣ „—– •‡‡ǡ ‡‰‹œ ‡– ƒŽǤǡ
ϱϰϬ
ʹͲͳ͵ȌǤ ’‘”–ƒ–Ž›ǡ ’”‡˜‹‘—• •–—†‹‡• ‘ˆ –Š‡ ‡ˆˆ‡…– ‘ˆ „‹Š‡‹•’Š‡”‹… – ‘ • Šƒ˜‡ ‘–
ϱϰϭ
‡šŠ‹„‹–‡† …‘•‹•–‡– ‡š…‹–ƒ„‹Ž‹–› …Šƒ‰‡• —•‹‰ ƒ „‹Š‡‹•’Š‡”‹… ‘–ƒ‰‡ǣ ‹ …‘–”ƒ•– –‘
ϱϰϮ
—‹Š‡‹•’Š‡”‹…–ǡ„‹Š‡‹•’Š‡”‹…•–‹—Žƒ–‹‘Šƒ•„‡‡ˆ‘—†–‘‡‹–Š‡”’”‘†—…‡‘•‹‰‹ˆ‹…ƒ–
ϱϰϯ
…Šƒ‰‡•‹•ȋ̵Š‡ƒ‡–ƒŽǤǡʹͲͳͶȌǡ‘”…Šƒ‰‡•–Šƒ–™‡”‡•–ƒ–‹•–‹…ƒŽŽ›Ž‡••”‘„—•–ȋ‘”†‹ŽŽ‘Ǧ
ϱϰϰ
ƒ–‡‘• ‡– ƒŽǤǡ ʹͲͳʹȌǤ Š‡•‡ ”‡•—Ž–• ”ƒ‹•‡ –Š‡ ’‘••‹„‹Ž‹–› –Šƒ– –Š‡ …Šƒ‰‡• ‹ ‘–‘” ’Žƒ•–‹…‹–›
ϱϰϱ
•Š‘™ „‡Šƒ˜‹‘”ƒŽŽ› ‹ –Š‹• ƒ† ’”‡˜‹‘—• ȋ‹‡• ‡– ƒŽǤǡ ʹͲͲͺȌ •–—†‹‡•ǡ ”‡Ž› ‘ †‹ˆˆ‡”‡–
ϱϰϲ
‡…Šƒ‹•• –Šƒ –Š‘•‡ ”‡ˆŽ‡…–‡† ‹ –Š‡ ’‘Žƒ”‹–›Ǧ•’‡…‹ˆ‹… …Šƒ‰‡• ‹ •Ǥ ’ƒ”•‹‘‹‘—•
ϱϰϳ
‡š’Žƒƒ–‹‘ ˆ‘” ‘—” ”‡•—Ž–• ‹• –Šƒ– –Š‡ „‡Šƒ˜‹‘”ƒŽ – ‡ˆˆ‡…–• ƒ”‡ ”‡Žƒ–‡† –‘ –Š‡ •’ƒ–‹ƒŽ
ϱϰϴ
†‹•–”‹„—–‹‘ ‘ˆ ‡Ž‡…–”‹…ƒŽ …—””‡–•ǡ ”ƒ–Š‡” –Šƒ –‘ –Š‡ …—””‡– †‹”‡…–‹‘Ǥ ‹‘’Š›•‹…ƒŽ …—””‡–
ϱϰϵ
‘†‡ŽŽ‹‰ ‘ˆ – ȋ”—‘‰ ‡– ƒŽǤǡ ʹͲͳͶǢ ƒ”‘• ‡– ƒŽǤǡ ʹͲͳ͸Ȍ †‡‘•–”ƒ–‡• –Šƒ– –Š‡
ϱϱϬ
—‹Š‡‹•’Š‡”‹… ‘–ƒ‰‡ ’”‹ƒ”‹Ž› •‡†• …—””‡– –Š”‘—‰Š …‘–”ƒŽƒ–‡”ƒŽ ’”‡‘–‘” ƒ†
ϱϱϭ
‹’•‹Žƒ–‡”ƒŽ ’”‡ˆ”‘–ƒŽ ”‡‰‹‘•ǡ ™Š‡”‡ƒ• „‹Š‡‹•’Š‡”‹… •–‹—Žƒ–‹‘ –ƒ”‰‡–• ‘–‘” ƒ† ’”‡‘–‘”
ϱϱϮ
”‡‰‹‘•„‹Žƒ–‡”ƒŽŽ›Ǥ’‘”–ƒ–Ž›ǡ–Š‡™‡ƒ”ƒ†‹ƒŽ…—””‡–•–Šƒ–‰‹˜‡”‹•‡–‘–Š‡’‘Žƒ”‹–›Ǧ•’‡…‹ˆ‹…
ϱϱϯ
‡ˆˆ‡…–• ‘ • •™‹–…Š †‹”‡…–‹‘• ‘ ‘’’‘•‹–‡ •‹†‡• ‘ˆ –Š‡ ‰›”—• ȋƒŠƒ ‡– ƒŽǤǡ ʹͲͳ͵ȌǤ
ϱϱϰ
Š‡”‡ˆ‘”‡ǡ ’”‡‘–‘” ƒ”‡ƒ• ‹ „‘–Š Š‡‹•’Š‡”‡• …‘—Ž† ‡š’‡”‹‡…‡ ‡‹–Š‡” •—’’”‡••‹‘ ‘”
ϱϱϱ
‡š…‹–ƒ–‹‘ †‡’‡†‹‰ ‘ –Š‡‹” ˆ‘Ž†‹‰ ‰‡‘‡–”›Ǥ Š‡ ‡ˆˆ‡…–• ‘ˆ – ‘ ‡—”‘’Žƒ•–‹…‹–›ǡ
ϱϱϲ
–Š‡”‡ˆ‘”‡ǡ …‘—Ž† „‡ ‡†‹ƒ–‡† „› –Š‡ …‘•‹†‡”ƒ„Ž› •–”‘‰‡” –ƒ‰‡–‹ƒŽ …—””‡–•ǡ ™Š‹…Šǡ ‹
ϱϱϳ
…‘–”ƒ•––‘–Š‡”ƒ†‹ƒŽ…—””‡–•ǡ†‘‘–Šƒ˜‡’‘Žƒ”‹–›Ǧ•’‡…‹ˆ‹…‡ˆˆ‡…–•ȋƒŠƒ‡–ƒŽǤǡʹͲͳ͵ȌǤ†‡”
ϭϲ
d,ZK>K&/W^/>dZ>KZdy/EDKdKZ>ZE/E' ϭϳ
ϱϱϴ
–Š‹•ƒ••—’–‹‘ǡ„‹Š‡‹•’Š‡”‹…– ȋ‹†‡’‡†‡–Ž› ‘ˆ’‘Žƒ”‹–›Ȍ ™‘—Ž†Šƒ˜‡ ‹…”‡ƒ•‡†‡—”ƒŽ
ϱϱϵ
’Žƒ•–‹…‹–›‹„‘–ŠŠ‡‹•’Š‡”‡•‹ƒƒ‡”—”‡Žƒ–‡†–‘–Š‡…Šƒ‰‡•‡ƒ•—”‡ƒ„Ž‡™‹–Š•Ǥ
ϱϲϬ
˜‡—†‡”–Š‡ƒ••—’–‹‘‘ˆ’‘Žƒ”‹–›Ǧ—•’‡…‹ˆ‹…–‡ˆˆ‡…–•‘’Žƒ•–‹…‹–›ǡ‘—””‡•—Ž–•
ϱϲϭ
”‡ƒ‹ ‹…‘’ƒ–‹„Ž‡ ™‹–Š –Š‡ Š‡‹•’Š‡”‹… …‘’‡–‹–‹‘ ‘†‡Žǣ ‹ˆ –Š‡ …ƒ–Š‘†‡ ’”‘‘–‡†
ϱϲϮ
’Žƒ•–‹…‹–›‹‹’•‹Žƒ–‡”ƒŽͳ†—”‹‰–”ƒ‹‹‰ǡ–Š‡–Š‡‹–‡”Š‡‹•’Š‡”‹……‘’‡–‹–‹‘‘†‡Ž™‘—Ž†
ϱϲϯ
Šƒ˜‡’”‡†‹…–‡†ƒ†‹•ƒ†˜ƒ–ƒ‰‡‘ˆ„‘–Š„‹Š‡‹•’Š‡”‹…‘–ƒ‰‡•”‡Žƒ–‹˜‡–‘–Š‡—‹Š‡‹•’Š‡”‹…
ϱϲϰ
‘–ƒ‰‡ǡ ƒ• „‹Š‡‹•’Š‡”‹… •–‹—Žƒ–‹‘ ™‘—Ž† ‹…”‡ƒ•‡ –Š‡ ’—–ƒ–‹˜‡Ž› Šƒ”ˆ—Ž ‹’•‹Žƒ–‡”ƒŽ
ϱϲϱ
ƒ…–‹˜ƒ–‹‘ȋ•‡‡Table 1ȌǤ
ϱϲϲ
•–‡ƒ†‘ˆ’”‘„‹‰‡š…‹–ƒ„‹Ž‹–›‘ˆ–Š‡’”‹ƒ”›‘–‘”…‘”–‡šƒˆ–‡”–•–‹—Žƒ–‹‘—•‹‰
ϱϲϳ
ǡ™‡‡˜ƒŽ—ƒ–‡†–ƒ•Ǧ”‡Žƒ–‡†ƒ…–‹˜‹–›—•‹‰ˆƒˆ–‡”–”ƒ•‹‡––‡ˆˆ‡…–•Šƒ†„‡‡™ƒ•Š‡†
ϱϲϴ
‘—–Ǥƒ†˜ƒ–ƒ‰‡‘ˆ–Š‹•ƒ’’”‘ƒ…Š‹•–Šƒ–™‡ƒ˜‘‹†‡†’‘••‹„Ž‡‹–‡”ˆ‡”‡…‡™‹–Š–Š‡’”‘…‡••‘ˆ
ϱϲϵ
‡‘”› …‘•‘Ž‹†ƒ–‹‘ –Š”‘—‰Š –Š‡ ‡…‡••ƒ”› •–‹—Žƒ–‹‘ –‘ ͳ ™Š‡ ‡ƒ•—”‹‰ •
ϱϳϬ
ȋ—‡ŽŽ„ƒ…Š‡” ‡– ƒŽǤǡ ʹͲͲʹȌǤ ‡ ‘„•‡”˜‡† –Šƒ– –Š‡ ƒ˜‡”ƒ‰‡ ƒ…–‹˜‹–› †—”‹‰ –”ƒ‹‡† Šƒ†
ϱϳϭ
‘˜‡‡–• ™ƒ• Žƒ”‰‡” ‹ „‹Š‡‹•’Š‡”‹… – ‰”‘—’• ‹ „‘–Š …‘–”ƒŽƒ–‡”ƒŽ ƒ† ‹’•‹Žƒ–‡”ƒŽ
ϱϳϮ
•‡•‘”‹‘–‘” ”‡‰‹‘•Ǥ Š‡ ”‡•—Ž–• ˆ‘” –Š‡ —–”ƒ‹‡† Šƒ† ™‡”‡ •‹‹Žƒ”ǡ ƒŽ„‡‹– Ž‡•• ”‘„—•–Ǥ
ϱϳϯ
”‡˜‹‘—• ™‘” Šƒ• †‡‘•–”ƒ–‡† „‹Žƒ–‡”ƒŽ ƒ…–‹˜‹–› ‹…”‡ƒ•‡• ‹ ͳ †—”‹‰ –Š‡ ƒ’’Ž‹…ƒ–‹‘ ‘ˆ
ϱϳϰ
„‹Š‡‹•’Š‡”‹…–ȋ‹†‡„‡”‰‡–ƒŽǤǡʹͲͳ͵ȌǤ‹‹Žƒ”‘Ž‹‡ȋ™‘ƒ†ƒ‰ǡʹͲͳͳǢ–ƒ‰‰‡–ƒŽǤǡ
ϱϳϱ
ʹͲͳʹȌ‘”•Š‘”–ǦŽƒ•–‹‰ȋƒ—†‡™‹‰‡–ƒŽǤǡʹͲͲͳǢ™‘‡–ƒŽǤǡʹͲͲͺǢƒ‰‡– ƒŽǤǡ ʹͲͲͻǢ –ƒ‰‰‡–ƒŽǤǡ
ϱϳϲ
ʹͲͲͻǢ ‹ ‡– ƒŽǤǡ ʹͲͳʹȌ …Šƒ‰‡• —†‡”‡ƒ–Š –Š‡ ƒ‘†‡ Šƒ˜‡ ƒŽ•‘ „‡‡ ”‡’‘”–‡† ˆ‘”
ϱϳϳ
—‹Š‡‹•’Š‡”‹… –Ǥ ’‘”–ƒ–Ž›ǡ ™‡ ‡ƒ•—”‡† ˆ—…–‹‘ƒŽ ƒ…–‹˜‹–› ƒ’’”‘š‹ƒ–‡Ž› Ͷͺ Š‘—”•
ϱϳϴ
ƒˆ–‡” –Š‡ ‡† ‘ˆ –Š‡ ˆ‹ƒŽ •–‹—Žƒ–‹‘Ǥ Š‡”‡ˆ‘”‡ǡ –Š‡ ƒ…–‹˜‹–› ‹…”‡ƒ•‡• ”‡’‘”–‡† Š‡”‡ ”‡ˆŽ‡…–
ϱϳϵ
Ž‘‰‡”ǦŽƒ•–‹‰…Šƒ‰‡•…ƒ—•‡†„›‡—”ƒŽ’Žƒ•–‹…‹–›ǡ”ƒ–Š‡”–Šƒƒ›‹‡†‹ƒ–‡‡ˆˆ‡…–•‘ˆ–‘
ϱϴϬ
–Š‡Š‡‘†›ƒ‹…”‡•’‘•‡Ǥ
ϱϴϭ
Š‡ˆƒ…––Šƒ––Š‡ƒ…–‹˜ƒ–‹‘…Šƒ‰‡•‹‘—”•–—†›™‡”‡”‡•–”‹…–‡†–‘’”‹ƒ”›‘–‘”ƒ†
ϱϴϮ
•‡•‘”› …‘”–‡š •Š‘—Ž† ‘– ‡…‡••ƒ”‹Ž› „‡ –ƒ‡ ƒ• ‡˜‹†‡…‡ –Šƒ– –Š‡ ”‡Ž‡˜ƒ– ‡—”‘’Žƒ•–‹…
ϱϴϯ
…Šƒ‰‡• ‘Ž› ‘……—””‡† Š‡”‡Ǥ ƒ–Š‡”ǡ ‹– ‹• ‡“—ƒŽŽ› ’‘••‹„Ž‡ –Šƒ– – Ž‡† –‘ ‹…”‡ƒ•‡† ’Žƒ•–‹…‹–›
ϱϴϰ
„‹Žƒ–‡”ƒŽŽ› ‹ ’”‡‘–‘” ‘” •—’’Ž‡‡–ƒ”› ‘–‘” …‘”–‡šǡ ƒ† –Šƒ– –Š‡ ‹…”‡ƒ•‡† ƒ…–‹˜‹–› ‹ ͳ
ϱϴϱ
ƒ†ͳ”‡ˆŽ‡…–•–Š‡‹…”‡ƒ•‡†‘†—Žƒ–‘”›‹’—–ˆ”‘–Š‡•‡ƒ”‡ƒ•Ǥ
ϱϴϲ
…”‡ƒ•‡† ’Žƒ•–‹…‹–› ‹ ‘–‘” ƒ† ’”‡Ǧ‘–‘” ƒ”‡ƒ• ‘ˆ „‘–Š Š‡‹•’Š‡”‡• ™‘—Ž† ƒŽ•‘
ϱϴϳ
‡š’Žƒ‹ ‘—” ‘„•‡”˜ƒ–‹‘ –Šƒ– –Š‡ „‡Šƒ˜‹‘”ƒŽ ƒ†˜ƒ–ƒ‰‡• †—‡ –‘ – ‰‡‡”ƒŽ‹œ‡† –‘ –Š‡
ϱϴϴ
—–”ƒ‹‡† Šƒ†Ǥ  ƒ ’”‡˜‹‘—• •–—†›ǡ ™‡ —•‡† —Ž–‹Ǧ˜‘š‡Ž ’ƒ––‡” ƒƒŽ›•‹• –‘ ‹†‡–‹ˆ›
ϱϴϵ
…‘‘ƒŽ‹–‹‡•‹–Š‡‡—”ƒŽ‡…‘†‹‰‘ˆ•’‡…‹ˆ‹…‘–‘”•‡“—‡…‡•ˆ‘”–Š‡ Ž‡ˆ–ƒ†”‹‰Š–Šƒ†•
ϱϵϬ
ȋ‹‡•–Ž‡” ‡– ƒŽǤǡ ʹͲͳͶȌǤ ‡ ˆ‘—† ™‹†‡•’”‡ƒ†ǡ •Šƒ”‡† ”‡’”‡•‡–ƒ–‹‘• ‹ „‘–Š ’”‡‘–‘” ƒ†ǡ
ϱϵϭ
•—”’”‹•‹‰Ž›ǡ’”‹ƒ”›‘–‘”ƒ”‡ƒ•‘ˆ„‘–ŠŠ‡‹•’Š‡”‡•Ǥ
‹˜‡–Šƒ––‹…”‡ƒ•‡†–Š‡ƒ‘—–
ϱϵϮ
‘ˆ ‹–‡”ƒ—ƒŽ ‰‡‡”ƒŽ‹œƒ–‹‘ǡ ‹– ƒ’’‡ƒ”• Ž‹‡Ž› –Šƒ– ‡ˆˆ‡…–‘”Ǧ‹†‡’‡†‡– •‡“—‡…‡
ϱϵϯ
”‡’”‡•‡–ƒ–‹‘•™‡”‡’ƒ”–‹…—Žƒ”Ž›ˆƒ…‹Ž‹–ƒ–‡†Ǥ
ϭϳ
d,ZK>K&/W^/>dZ>KZdy/EDKdKZ>ZE/E' ϭϴ
ϱϵϰ
‘•—ƒ”‹œ‡ǡ™‡†‡‘•–”ƒ–‡–Šƒ–…‘˜‡–‹‘ƒŽƒ†”‡˜‡”•‡†Ǧ’‘Žƒ”‹–›„‹Š‡‹•’Š‡”‹…
ϱϵϱ
– •‹‹Žƒ”Ž› ‹…”‡ƒ•‡ ‘–‘” Ž‡ƒ”‹‰ ƒ† ƒ…–‹˜ƒ–‹‘ ‹ „‘–Š ƒ‘†‡Ǧ ƒ† …ƒ–Š‘†‡Ǧ
ϱϵϲ
‘†—Žƒ–‡†Š‡‹•’Š‡”‡•ǤŠ—•ǡ‘—”•–—†›’”‘˜‹†‡•‡˜‹†‡…‡ˆ‘”ƒƒ…–‹˜‡”‘Ž‡‘ˆ–Š‡‹’•‹Žƒ–‡”ƒŽ
ϱϵϳ
…‘”–‡š ‹ —‹ƒ—ƒŽ ‘–‘” …‘–”‘Ž ‹ ›‘—‰ǡ Š‡ƒŽ–Š› ‹†‹˜‹†—ƒŽ•ǡ …‘•‹•–‡– ™‹–Š ’”‡˜‹‘—•
ϱϵϴ
”‡’‘”–•‹‘Ž†‡”ƒ†—Ž–•‘”˜‹…–‹•‘ˆ•–”‘‡ȋ‘Šƒ•‡Ǧ‡”‰‡–ƒŽǤǡʹͲͲʹǢ‹‡”ƒ‡–ƒŽǤǡʹͲͳͶȌǤ
ϱϵϵ
– ‡ˆˆ‡…–• ‘ ’Žƒ•–‹…‹–› ƒ”‡ •–‹ŽŽ ‘ˆ–‡ …‘•–”—‡† ‹ –‡”• ‘ˆ ‡š…‹–ƒ–‹‘Ȁ‹Š‹„‹–‹‘ ‘ˆ ‡—”ƒŽ
ϲϬϬ
–‹••—‡—†‡”–Š‡‡Ž‡…–”‘†‡•ǡƒ†–Š‡‹–‡”Š‡‹•’Š‡”‹……‘’‡–‹–‹‘‘†‡Ž‹•…‘‘Ž›—•‡†–‘
ϲϬϭ
‡š’Žƒ‹ –Š‡ •—’‡”‹‘”‹–› ‘ˆ „‹Š‡‹•’Š‡”‹… –Ǥ Š‡ ‹†‡ƒ ‘ˆ ’”‘‘–‹‰ ‘–‘” Ž‡ƒ”‹‰ „›
ϲϬϮ
”‡†—…‹‰‹’•‹Žƒ–‡”ƒŽ‡š…‹–ƒ„‹Ž‹–›‹•’ƒ”–‹…—Žƒ”Ž›’‡”–‹‡––‘•–”‘‡‡—”‘”‡Šƒ„‹Ž‹–ƒ–‹‘ǡ™Š‡”‡‹–
ϲϬϯ
Šƒ• Ž‡† –‘ –Š‡ •—’’‘•‹–‹‘ –Šƒ– •—’’”‡••‹‰ ƒ…–‹˜‹–› ‹ –Š‡ Š‡ƒŽ–Š› Š‡‹•’Š‡”‡ –‘ ”‡Ž‡ƒ•‡
ϲϬϰ
‹Š‹„‹–‹‘‘ˆŽ‡•‹‘‡†…‘”–‡šƒ›ˆƒ…‹Ž‹–ƒ–‡”‡…‘˜‡”›ȋ—‡Žƒ†‘Š‡ǡʹͲͲ͸Ǣ‹ŽŽ‹ƒ•‡–ƒŽǤǡ
ϲϬϱ
ʹͲͳͲǢ‘Ž‘‰‹‹‡–ƒŽǤǡʹͲͳͳǢ‹–•…Š‡ƒ†ƒ—Ž—•ǡʹͲͳͳǢƒ‡—…Š‹ƒ†œ—‹ǡʹͲͳʹǢ‹‡”ƒ‡–
ϲϬϲ
ƒŽǤǡʹͲͳʹǢ”ƒ—•‡‡–ƒŽǤǡʹͲͳ͵Ǣ‡ˆ‡„˜”‡‡–ƒŽǤǡʹͲͳͶȌǤŠ‹Ž‡–Š‡Š‡‹•’Š‡”‹……‘‘’‡”ƒ–‹‘‘†‡Ž
ϲϬϳ
’”‘’‘•‡† Š‡”‡ —•– „‡ –‡•–‡† ‹ ‡Ž†‡”Ž› ’ƒ”–‹…‹’ƒ–• ƒ† ‹†‹˜‹†—ƒŽ• ™‹–Š •–”‘‡ǡ ‘—” †ƒ–ƒ
ϲϬϴ
•—‰‰‡•– –Šƒ– ƒ •‹’Ž‡ ‡š…‹–ƒ–‹‘Ȁ‹Š‹„‹–‹‘ ‘†‡Ž ƒ› „‡ –‘‘ •‹’Ž‹•–‹… ƒ† •Š‘—Ž† „‡
ϲϬϵ
ƒ„ƒ†‘‡†‹ˆƒ˜‘”‘ˆƒˆ”ƒ‡™‘”–Šƒ–ƒ…‘™Ž‡†‰‡•–Š‡„”‘ƒ†‡ˆˆ‡…–•‘ˆ–…—””‡–ƒ†–Š‡
ϲϭϬ
”‘Ž‡•–Šƒ–„‘–ŠŠ‡‹•’Š‡”‡•’Žƒ›‹–Š‡‡…‘†‹‰‘ˆ‹ˆ‘”ƒ–‹‘†—”‹‰‘–‘”Ž‡ƒ”‹‰Ǥ
ϲϭϭ
ϲϭϮ
ϲϭϯ
ϲϭϰ
References
ϲϭϱ
ϲϭϲ
ϲϭϳ
ϲϭϴ
ϲϭϵ
ϲϮϬ
ϲϮϭ
ϲϮϮ
ϲϮϯ
ϲϮϰ
ϲϮϱ
ϲϮϲ
ϲϮϳ
ϲϮϴ
ϲϮϵ
ϲϯϬ
ϲϯϭ
ϲϯϮ
ϲϯϯ
ϲϯϰ
ϲϯϱ
ŶƚĂů͕ŝŬƐŽŶD͕ĂƚƚĂ͕>ĂĨŽŶ͕ĞĐŚĞŶƚW͕WĂƌƌĂ>͕WĂƵůƵƐt;ϮϬϭϰͿ/ŵĂŐŝŶŐĂƌƚŝĨĂĐƚƐŝŶĚƵĐĞĚ
ďLJĞůĞĐƚƌŝĐĂůƐƚŝŵƵůĂƚŝŽŶĚƵƌŝŶŐĐŽŶǀĞŶƚŝŽŶĂůĨDZ/ŽĨƚŚĞďƌĂŝŶ͘EĞƵƌŽŝŵĂŐĞϴϱWƚϯ͗ϭϬϰϬͲ
ϭϬϰϳ͘
ĂƚƐŝŬĂĚnjĞ'͕DŽůŝĂĚnjĞs͕WĂƵůƵƐt͕<ƵŽD&͕EŝƚƐĐŚĞD;ϮϬϭϯͿWĂƌƚŝĂůůLJŶŽŶͲůŝŶĞĂƌƐƚŝŵƵůĂƚŝŽŶ
ŝŶƚĞŶƐŝƚLJͲĚĞƉĞŶĚĞŶƚĞĨĨĞĐƚƐŽĨĚŝƌĞĐƚĐƵƌƌĞŶƚƐƚŝŵƵůĂƚŝŽŶŽŶŵŽƚŽƌĐŽƌƚĞdžĞdžĐŝƚĂďŝůŝƚLJŝŶ
ŚƵŵĂŶƐ͘:WŚLJƐŝŽůϱϵϭ͗ϭϵϴϳͲϮϬϬϬ͘
ĂƵĚĞǁŝŐ:͕EŝƚƐĐŚĞD͕WĂƵůƵƐt͕&ƌĂŚŵ:;ϮϬϬϭͿZĞŐŝŽŶĂůŵŽĚƵůĂƚŝŽŶŽĨK>DZ/ƌĞƐƉŽŶƐĞƐƚŽ
ŚƵŵĂŶƐĞŶƐŽƌŝŵŽƚŽƌĂĐƚŝǀĂƚŝŽŶďLJƚƌĂŶƐĐƌĂŶŝĂůĚŝƌĞĐƚĐƵƌƌĞŶƚƐƚŝŵƵůĂƚŝŽŶ͘DĂŐŶZĞƐŽŶDĞĚ
ϰϱ͗ϭϵϲͲϮϬϭ͘
ĞĂƵůĞs͕dƌĞŵďůĂLJ^͕dŚĞŽƌĞƚ,;ϮϬϭϯͿ/ŶƚĞƌŚĞŵŝƐƉŚĞƌŝĐĐŽŶƚƌŽůŽĨƵŶŝůĂƚĞƌĂůŵŽǀĞŵĞŶƚ͘EĞƵƌĂů
WůĂƐƚϮϬϭϮ͗ϲϮϳϴϭϲ͘
ŽůŽŐŶŝŶŝE͕sĂůůĂƌ'͕ĂƐĂƚŝ͕>ĂƚŝĨ>͕ůͲEĂnjĞƌZ͕tŝůůŝĂŵƐ:͕ĂŶĐŽ͕DĂĐĞĂ͕dĞƐŝŽ>͕ŚĞƐƐĂ͕
&ƌĞŐŶŝ&;ϮϬϭϭͿEĞƵƌŽƉŚLJƐŝŽůŽŐŝĐĂůĂŶĚďĞŚĂǀŝŽƌĂůĞĨĨĞĐƚƐŽĨƚ^ĐŽŵďŝŶĞĚǁŝƚŚĐŽŶƐƚƌĂŝŶƚͲ
ŝŶĚƵĐĞĚŵŽǀĞŵĞŶƚƚŚĞƌĂƉLJŝŶƉŽƐƚƐƚƌŽŬĞƉĂƚŝĞŶƚƐ͘EĞƵƌŽƌĞŚĂďŝůEĞƵƌĂůZĞƉĂŝƌϮϱ͗ϴϭϵͲϴϮϵ͘
ŽƌŽŽũĞƌĚŝ͕&ŽůƚLJƐ,͕<ƌŝŶŐƐd͕^ƉĞƚnjŐĞƌh͕dŚƌŽŶ͕dŽƉƉĞƌZ;ϭϵϵϵͿ>ŽĐĂůŝnjĂƚŝŽŶŽĨƚŚĞŵŽƚŽƌŚĂŶĚ
ĂƌĞĂƵƐŝŶŐƚƌĂŶƐĐƌĂŶŝĂůŵĂŐŶĞƚŝĐƐƚŝŵƵůĂƚŝŽŶĂŶĚĨƵŶĐƚŝŽŶĂůŵĂŐŶĞƚŝĐƌĞƐŽŶĂŶĐĞŝŵĂŐŝŶŐ͘
ůŝŶEĞƵƌŽƉŚLJƐŝŽůϭϭϬ͗ϲϵϵͲϳϬϰ͘
ĞŶŐŝnj͕DƵƌĂƐĞE͕ZŽƚŚǁĞůů:;ϮϬϭϯͿKƉƉŽƐŝƚĞĞĨĨĞĐƚƐŽĨǁĞĂŬƚƌĂŶƐĐƌĂŶŝĂůĚŝƌĞĐƚĐƵƌƌĞŶƚ
ƐƚŝŵƵůĂƚŝŽŶŽŶĚŝĨĨĞƌĞŶƚƉŚĂƐĞƐŽĨƐŚŽƌƚŝŶƚĞƌǀĂůŝŶƚƌĂĐŽƌƚŝĐĂůŝŶŚŝďŝƚŝŽŶ;^//Ϳ͘džƉƌĂŝŶZĞƐ͘
ŚĞŶZ;ϮϬϬϰͿ/ŶƚĞƌĂĐƚŝŽŶƐďĞƚǁĞĞŶŝŶŚŝďŝƚŽƌLJĂŶĚĞdžĐŝƚĂƚŽƌLJĐŝƌĐƵŝƚƐŝŶƚŚĞŚƵŵĂŶŵŽƚŽƌĐŽƌƚĞdž͘
džƉƌĂŝŶZĞƐϭϱϰ͗ϭͲϭϬ͘
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d,ZK>K&/W^/>dZ>KZdy/EDKdKZ>ZE/E' ϲϯϲ
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ĂůĞD͕&ŝƐĐŚů͕^ĞƌĞŶŽD/;ϭϵϵϵͿŽƌƚŝĐĂůƐƵƌĨĂĐĞͲďĂƐĞĚĂŶĂůLJƐŝƐ͘/͘^ĞŐŵĞŶƚĂƚŝŽŶĂŶĚƐƵƌĨĂĐĞ
ƌĞĐŽŶƐƚƌƵĐƚŝŽŶ͘EĞƵƌŽŝŵĂŐĞϵ͗ϭϳϵͲϭϵϰ͘
ĂƐŬĂůĂŬŝƐ:͕ŚƌŝƐƚĞŶƐĞŶ<͕&ŝƚnjŐĞƌĂůĚW͕ZŽƐŚĂŶ>͕ŚĞŶZ;ϮϬϬϮͿdŚĞŵĞĐŚĂŶŝƐŵƐŽĨ
ŝŶƚĞƌŚĞŵŝƐƉŚĞƌŝĐŝŶŚŝďŝƚŝŽŶŝŶƚŚĞŚƵŵĂŶŵŽƚŽƌĐŽƌƚĞdž͘:WŚLJƐŝŽůϱϰϯ͗ϯϭϳͲϯϮϲ͘
ĞůǀĞŶĚĂŚů/͕'ĂƚƚŝŶŐĞƌE͕ĞƌŐĞƌd͕'ůĞŝĐŚ͕^ŝĞďŶĞƌ,Z͕DĂůůs;ϮϬϭϰͿdŚĞZŽůĞŽĨWƵůƐĞ^ŚĂƉĞŝŶ
DŽƚŽƌŽƌƚĞdždƌĂŶƐĐƌĂŶŝĂůDĂŐŶĞƚŝĐ^ƚŝŵƵůĂƚŝŽŶhƐŝŶŐ&ƵůůͲ^ŝŶĞ^ƚŝŵƵůŝ͘W>Ž^KŶĞ
ϵ͗ĞϭϭϱϮϰϳ͘
ŝ>ĂnjnjĂƌŽs͕ŝĞŵĂŶŶh͕>ĞŵŽŶZE;ϮϬϬϴͿ^ƚĂƚĞŽĨƚŚĞĂƌƚ͗WŚLJƐŝŽůŽŐLJŽĨƚƌĂŶƐĐƌĂŶŝĂůŵŽƚŽƌĐŽƌƚĞdž
ƐƚŝŵƵůĂƚŝŽŶ͘ƌĂŝŶ^ƚŝŵƵůϭ͗ϯϰϱͲϯϲϮ͘
ŝWŝŶŽ'͕WĞůůĞŐƌŝŶŽ'͕ƐƐĞŶnjĂ'͕ĂƉŽŶĞ&͕&ĞƌƌĞƌŝ&͕&ŽƌŵŝĐĂ͕ZĂŶŝĞƌŝ&͕dŽŵďŝŶŝD͕ŝĞŵĂŶŶh͕
ZŽƚŚǁĞůů:͕ŝ>ĂnjnjĂƌŽs;ϮϬϭϰͿDŽĚƵůĂƚŝŽŶŽĨďƌĂŝŶƉůĂƐƚŝĐŝƚLJŝŶƐƚƌŽŬĞ͗ĂŶŽǀĞůŵŽĚĞůĨŽƌ
ŶĞƵƌŽƌĞŚĂďŝůŝƚĂƚŝŽŶ͘EĂƚZĞǀEĞƵƌŽůϭϬ͗ϱϵϳͲϲϬϴ͘
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ŝŶĚŝǀŝĚƵĂƚĞĚĨŝŶŐĞƌŵŽǀĞŵĞŶƚƐ͘ĞƌĞďŽƌƚĞdžϮϯ͗ϭϯϲϮͲϭϯϳϳ͘
&ĞƌďĞƌƚ͕WƌŝŽƌŝ͕ZŽƚŚǁĞůů:͕ĂLJ>͕ŽůĞďĂƚĐŚ:'͕DĂƌƐĚĞŶ;ϭϵϵϮͿ/ŶƚĞƌŚĞŵŝƐƉŚĞƌŝĐŝŶŚŝďŝƚŝŽŶ
ŽĨƚŚĞŚƵŵĂŶŵŽƚŽƌĐŽƌƚĞdž͘:WŚLJƐŝŽůϰϱϯ͗ϱϮϱͲϱϰϲ͘
&ƌŝƐƚŽŶ<:͕ƐŚďƵƌŶĞƌ:͕<ŝĞďĞů^:͕EŝĐŚŽůƐd͕WĞŶŶLJt͕ĞĚƐ;ϮϬϬϳͿ^ƚĂƚŝƐƚŝĐĂůWĂƌĂŵĞƚƌŝĐDĂƉƉŝŶŐ͗
dŚĞŶĂůLJƐŝƐŽĨ&ƵŶĐƚŝŽŶĂůƌĂŝŶ/ŵĂŐĞƐ͗ĐĂĚĞŵŝĐWƌĞƐƐ͘
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Zd͕ĂƌŵĞŶĂ:D;ϮϬϬϵͿŽƌƚŝĐĂůƌĞƉƌĞƐĞŶƚĂƚŝŽŶŽĨŝƉƐŝůĂƚĞƌĂůĂƌŵŵŽǀĞŵĞŶƚƐŝŶŵŽŶŬĞLJĂŶĚ
ŵĂŶ͘:EĞƵƌŽƐĐŝϮϵ͗ϭϮϵϰϴͲϭϮϵϱϲ͘
,ƵŵŵĞů&͕ŽŚĞŶ>';ϮϬϬϲͿEŽŶͲŝŶǀĂƐŝǀĞďƌĂŝŶƐƚŝŵƵůĂƚŝŽŶ͗ĂŶĞǁƐƚƌĂƚĞŐLJƚŽŝŵƉƌŽǀĞ
ŶĞƵƌŽƌĞŚĂďŝůŝƚĂƚŝŽŶĂĨƚĞƌƐƚƌŽŬĞ͍>ĂŶĐĞƚEĞƵƌŽůϱ͗ϳϬϴͲϳϭϮ͘
:ĂŶŐ^,͕ŚŶ^,͕LJƵŶtD͕<ŝŵ^͕>ĞĞDz͕<ǁŽŶz,;ϮϬϬϵͿdŚĞĞĨĨĞĐƚŽĨƚƌĂŶƐĐƌĂŶŝĂůĚŝƌĞĐƚĐƵƌƌĞŶƚ
ƐƚŝŵƵůĂƚŝŽŶŽŶƚŚĞĐŽƌƚŝĐĂůĂĐƚŝǀĂƚŝŽŶďLJŵŽƚŽƌƚĂƐŬŝŶƚŚĞŚƵŵĂŶďƌĂŝŶ͗ĂŶĨDZ/ƐƚƵĚLJ͘
EĞƵƌŽƐĐŝ>ĞƚƚϰϲϬ͗ϭϭϳͲϭϮϬ͘
:ĂŶƐƐĞŶD͕KŽƐƚĞŶĚŽƌƉd&͕^ƚĞŐĞŵĂŶ&;ϮϬϭϱͿdŚĞĐŽŝůŽƌŝĞŶƚĂƚŝŽŶĚĞƉĞŶĚĞŶĐLJŽĨƚŚĞĞůĞĐƚƌŝĐ
ĨŝĞůĚŝŶĚƵĐĞĚďLJdD^ĨŽƌDϭĂŶĚŽƚŚĞƌďƌĂŝŶĂƌĞĂƐ͘:EĞƵƌŽĞŶŐZĞŚĂďŝůϭϮ͗ϰϳ͘
:ŽŚĂŶƐĞŶͲĞƌŐ,͕ZƵƐŚǁŽƌƚŚD&͕ŽŐĚĂŶŽǀŝĐD͕<ŝƐĐŚŬĂh͕tŝŵĂůĂƌĂƚŶĂ^͕DĂƚƚŚĞǁƐWD;ϮϬϬϮͿ
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ϵϵ͗ϭϰϱϭϴͲϭϰϱϮϯ͘
<ĂƌŽŬ^͕tŝƚŶĞLJ';ϮϬϭϯͿŶŚĂŶĐĞĚŵŽƚŽƌůĞĂƌŶŝŶŐĨŽůůŽǁŝŶŐƚĂƐŬͲĐŽŶĐƵƌƌĞŶƚĚƵĂůƚƌĂŶƐĐƌĂŶŝĂů
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<ĞƐƐůĞƌ^<͕dƵƌŬĞůƚĂƵďW͕ĞŶƐŽŶ:'͕,ĂŵŝůƚŽŶZ,;ϮϬϭϮͿŝĨĨĞƌĞŶĐĞƐŝŶƚŚĞĞdžƉĞƌŝĞŶĐĞŽĨĂĐƚŝǀĞĂŶĚ
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ϮϬ
>ŝŶĚĞŶďĞƌŐZ͕EĂĐŚƚŝŐĂůů>͕DĞŝŶnjĞƌD͕^ŝĞŐDD͕&ůŽĞů;ϮϬϭϯͿŝĨĨĞƌĞŶƚŝĂůĨĨĞĐƚƐŽĨƵĂůĂŶĚ
hŶŝŚĞŵŝƐƉŚĞƌŝĐDŽƚŽƌŽƌƚĞdž^ƚŝŵƵůĂƚŝŽŶŝŶKůĚĞƌĚƵůƚƐ͘:EĞƵƌŽƐĐŝϯϯ͗ϵϭϳϲͲϵϭϴϯ͘
DĂŚŵŽƵĚŝ,͕ŽƌŚĂŶŝ,ĂŐŚŝŐŚŝ͕WĞƚƌĂŵĨĂƌW͕:ĂŚĂŶƐŚĂŚŝ^͕^ĂůĞŚŝ͕&ƌĞŐŶŝ&;ϮϬϭϭͿdƌĂŶƐĐƌĂŶŝĂů
ĚŝƌĞĐƚĐƵƌƌĞŶƚƐƚŝŵƵůĂƚŝŽŶ͗ĞůĞĐƚƌŽĚĞŵŽŶƚĂŐĞŝŶƐƚƌŽŬĞ͘ŝƐĂďŝůZĞŚĂďŝůϯϯ͗ϭϯϴϯͲϭϯϴϴ͘
DŝĞůŬĞ͕tƌĞĚĞ͕^ĐŚƵůnjͲ^ĐŚĂĞĨĨĞƌt͕dĂŐŚŝnjĂĚĞŚͲtĂŐŚĞĨŝ͕EŝƚƐĐŚĞD͕ZŽŚĚĞs͕>ŝĞďĞƚĂŶnj
;ϮϬϭϯͿĂƚŚŽĚĂůƚƌĂŶƐĐƌĂŶŝĂůĚŝƌĞĐƚĐƵƌƌĞŶƚƐƚŝŵƵůĂƚŝŽŶŝŶĚƵĐĞƐƌĞŐŝŽŶĂů͕ůŽŶŐͲůĂƐƚŝŶŐ
ƌĞĚƵĐƚŝŽŶƐŽĨĐŽƌƚŝĐĂůďůŽŽĚĨůŽǁŝŶƌĂƚƐ͘EĞƵƌŽůZĞƐ͘
DŽƌĚŝůůŽͲDĂƚĞŽƐ>͕dƵƌƉŝŶͲ&ĞŶŽůů>͕DŝůůĂŶͲWĂƐĐƵĂů:͕EƵŶĞnjͲWĞƌĞnjE͕WĂŶLJĂǀŝŶ/͕'ŽŵĞnjͲƌŐƵĞůůĞƐ
:D͕ŽƚŝĂͲWĂŶŝĂŐƵĂ͕&ŽĨĨĂŶŝ'͕>ĂŶŐE͕KůŝǀŝĞƌŽ;ϮϬϭϮͿĨĨĞĐƚƐŽĨƐŝŵƵůƚĂŶĞŽƵƐďŝůĂƚĞƌĂů
ƚ^ŽĨƚŚĞŚƵŵĂŶŵŽƚŽƌĐŽƌƚĞdž͘ƌĂŝŶ^ƚŝŵƵůϱ͗ϮϭϰͲϮϮϮ͘
DƵĞůůďĂĐŚĞƌt͕ŝĞŵĂŶŶh͕tŝƐƐĞů:͕ĂŶŐE͕<ŽĨůĞƌD͕&ĂĐĐŚŝŶŝ^͕ŽƌŽŽũĞƌĚŝ͕WŽĞǁĞt͕,ĂůůĞƚƚD
;ϮϬϬϮͿĂƌůLJĐŽŶƐŽůŝĚĂƚŝŽŶŝŶŚƵŵĂŶƉƌŝŵĂƌLJŵŽƚŽƌĐŽƌƚĞdž͘EĂƚƵƌĞϰϭϱ͗ϲϰϬͲϲϰϰ͘
DƵƌĂƐĞE͕ƵƋƵĞ:͕DĂnjnjŽĐĐŚŝŽZ͕ŽŚĞŶ>';ϮϬϬϰͿ/ŶĨůƵĞŶĐĞŽĨŝŶƚĞƌŚĞŵŝƐƉŚĞƌŝĐŝŶƚĞƌĂĐƚŝŽŶƐŽŶ
ŵŽƚŽƌĨƵŶĐƚŝŽŶŝŶĐŚƌŽŶŝĐƐƚƌŽŬĞ͘ŶŶEĞƵƌŽůϱϱ͗ϰϬϬͲϰϬϵ͘
EĂƌŽƐ'͕'ĞLJĞƌD͕<ŽĐŚ^͕DĂLJƌ>͕ůůŝŶŐĞƌd͕'ƌŝŵŵ&͕'ŚĂƌĂďĂŐŚŝ;ϮϬϭϲͿŶŚĂŶĐĞĚŵŽƚŽƌ
ůĞĂƌŶŝŶŐǁŝƚŚďŝůĂƚĞƌĂůƚƌĂŶƐĐƌĂŶŝĂůĚŝƌĞĐƚĐƵƌƌĞŶƚƐƚŝŵƵůĂƚŝŽŶ͗/ŵƉĂĐƚŽĨƉŽůĂƌŝƚLJŽƌĐƵƌƌĞŶƚ
ĨůŽǁĚŝƌĞĐƚŝŽŶ͍ůŝŶEĞƵƌŽƉŚLJƐŝŽůϭϮϳ͗ϮϭϭϵͲϮϭϮϲ͘
Eŝ͕'ƵŶƌĂũ͕EĞůƐŽŶ:͕zĞŚ/:͕ĂƐƚŝůůŽ'͕,ŽƋƵĞd͕ŚĞŶZ;ϮϬϬϵͿdǁŽƉŚĂƐĞƐŽĨŝŶƚĞƌŚĞŵŝƐƉŚĞƌŝĐ
ŝŶŚŝďŝƚŝŽŶďĞƚǁĞĞŶŵŽƚŽƌƌĞůĂƚĞĚĐŽƌƚŝĐĂůĂƌĞĂƐĂŶĚƚŚĞƉƌŝŵĂƌLJŵŽƚŽƌĐŽƌƚĞdžŝŶŚƵŵĂŶ͘
ĞƌĞďŽƌƚĞdžϭϵ͗ϭϲϱϰͲϭϲϲϱ͘
EŝƚƐĐŚĞD͕WĂƵůƵƐt;ϮϬϬϬͿdžĐŝƚĂďŝůŝƚLJĐŚĂŶŐĞƐŝŶĚƵĐĞĚŝŶƚŚĞŚƵŵĂŶŵŽƚŽƌĐŽƌƚĞdžďLJǁĞĂŬ
ƚƌĂŶƐĐƌĂŶŝĂůĚŝƌĞĐƚĐƵƌƌĞŶƚƐƚŝŵƵůĂƚŝŽŶ͘:WŚLJƐŝŽůϱϮϳWƚϯ͗ϲϯϯͲϲϯϵ͘
EŝƚƐĐŚĞD͕WĂƵůƵƐt;ϮϬϭϭͿdƌĂŶƐĐƌĂŶŝĂůĚŝƌĞĐƚĐƵƌƌĞŶƚƐƚŝŵƵůĂƚŝŽŶͲͲƵƉĚĂƚĞϮϬϭϭ͘ZĞƐƚŽƌEĞƵƌŽů
EĞƵƌŽƐĐŝϮϵ͗ϰϲϯͲϰϵϮ͘
EŝƚƐĐŚĞD͕EŝƚƐĐŚĞD^͕<ůĞŝŶ͕dĞƌŐĂƵ&͕ZŽƚŚǁĞůů:͕WĂƵůƵƐt;ϮϬϬϯͿ>ĞǀĞůŽĨĂĐƚŝŽŶŽĨĐĂƚŚŽĚĂů
ƉŽůĂƌŝƐĂƚŝŽŶŝŶĚƵĐĞĚŝŶŚŝďŝƚŝŽŶŽĨƚŚĞŚƵŵĂŶŵŽƚŽƌĐŽƌƚĞdž͘ůŝŶEĞƵƌŽƉŚLJƐŝŽůϭϭϰ͗ϲϬϬͲ
ϲϬϰ͘
KΖ^ŚĞĂ:͕ŽƵĚƌŝĂƐD,͕^ƚĂŐŐ:͕ĂĐŚƚŝĂƌs͕<ŝƐĐŚŬĂh͕ůŝĐŚĞƌ:h͕:ŽŚĂŶƐĞŶͲĞƌŐ,;ϮϬϭϰͿWƌĞĚŝĐƚŝŶŐ
ďĞŚĂǀŝŽƵƌĂůƌĞƐƉŽŶƐĞƚŽd^ŝŶĐŚƌŽŶŝĐŵŽƚŽƌƐƚƌŽŬĞ͘EĞƵƌŽŝŵĂŐĞϴϱWƚϯ͗ϵϮϰͲϵϯϯ͘
KůĚĨŝĞůĚZ;ϭϵϳϭͿdŚĞĂƐƐĞƐƐŵĞŶƚĂŶĚĂŶĂůLJƐŝƐŽĨŚĂŶĚĞĚŶĞƐƐ͗ƚŚĞĚŝŶďƵƌŐŚŝŶǀĞŶƚŽƌLJ͘
EĞƵƌŽƉƐLJĐŚŽůŽŐŝĂϵ͗ϵϳͲϭϭϯ͘
WĞƌĞnjD͕ŽŚĞŶ>';ϮϬϬϵͿ/ŶƚĞƌŚĞŵŝƐƉŚĞƌŝĐŝŶŚŝďŝƚŝŽŶďĞƚǁĞĞŶƉƌŝŵĂƌLJŵŽƚŽƌĐŽƌƚŝĐĞƐ͗ǁŚĂƚŚĂǀĞ
ǁĞůĞĂƌŶĞĚ͍:WŚLJƐŝŽůϱϴϳ͗ϳϮϱͲϳϮϲ͘
ZĂŚŵĂŶ͕ZĞĂƚŽ͕ƌůŽƚƚŝD͕'ĂƐĐĂ&͕ĂƚƚĂ͕WĂƌƌĂ>͕ŝŬƐŽŶD;ϮϬϭϯͿĞůůƵůĂƌĨĨĞĐƚƐŽĨĐƵƚĞ
ŝƌĞĐƚƵƌƌĞŶƚ^ƚŝŵƵůĂƚŝŽŶ͗^ŽŵĂƚŝĐĂŶĚ^LJŶĂƉƚŝĐdĞƌŵŝŶĂůĨĨĞĐƚƐ͘:WŚLJƐŝŽů͘
ZĞŝƐ:͕&ƌŝƚƐĐŚ;ϮϬϭϭͿDŽĚƵůĂƚŝŽŶŽĨŵŽƚŽƌƉĞƌĨŽƌŵĂŶĐĞĂŶĚŵŽƚŽƌůĞĂƌŶŝŶŐďLJƚƌĂŶƐĐƌĂŶŝĂůĚŝƌĞĐƚ
ĐƵƌƌĞŶƚƐƚŝŵƵůĂƚŝŽŶ͘ƵƌƌKƉŝŶEĞƵƌŽůϮϰ͗ϱϵϬͲϱϵϲ͘
ZƵƐƐŽZ͕tĂůůĂĐĞ͕&ŝƚnjŐĞƌĂůĚW͕ŽŽƉĞƌEZ;ϮϬϭϯͿWĞƌĐĞƉƚŝŽŶŽĨŽŵĨŽƌƚƵƌŝŶŐĐƚŝǀĞĂŶĚ^ŚĂŵ
dƌĂŶƐĐƌĂŶŝĂůŝƌĞĐƚƵƌƌĞŶƚ^ƚŝŵƵůĂƚŝŽŶ͗ŽƵďůĞůŝŶĚ^ƚƵĚLJ͘ƌĂŝŶ^ƚŝŵƵůĚŽŝ͗
ϭϬ͘ϭϬϭϲͬũ͘ďƌƐ͘ϮϬϭϯ͘Ϭϱ͘ϬϬϵ͘΀ƉƵďĂŚĞĂĚŽĨƉƌŝŶƚ΁͘
^ĞŚŵ͕<ŝƉƉŝŶŐ:͕^ĐŚĂĨĞƌ͕sŝůůƌŝŶŐĞƌ͕ZĂŐĞƌƚW;ϮϬϭϯͿŽŵƉĂƌŝƐŽŶďĞƚǁĞĞŶhŶŝͲĂŶĚŝůĂƚĞƌĂů
ƚ^ĨĨĞĐƚƐŽŶ&ƵŶĐƚŝŽŶĂůŽŶŶĞĐƚŝǀŝƚLJŽĨƚŚĞ,ƵŵĂŶDŽƚŽƌŽƌƚĞdž͘&ƌŽŶƚ,ƵŵEĞƵƌŽƐĐŝ
ϳ͗ϭϴϯ͘
^ƚĂŐŐ:͕KΖ^ŚĞĂ:͕<ŝŶĐƐĞƐd͕tŽŽůƌŝĐŚD͕DĂƚƚŚĞǁƐWD͕:ŽŚĂŶƐĞŶͲĞƌŐ,;ϮϬϬϵͿDŽĚƵůĂƚŝŽŶŽĨ
ŵŽǀĞŵĞŶƚͲĂƐƐŽĐŝĂƚĞĚĐŽƌƚŝĐĂůĂĐƚŝǀĂƚŝŽŶďLJƚƌĂŶƐĐƌĂŶŝĂůĚŝƌĞĐƚĐƵƌƌĞŶƚƐƚŝŵƵůĂƚŝŽŶ͘Ƶƌ:
EĞƵƌŽƐĐŝϯϬ͗ϭϰϭϮͲϭϰϮϯ͘
^ƚĂŐŐ:͕ĂĐŚƚŝĂƌs͕KΖ^ŚĞĂ:͕ůůŵĂŶ͕ŽƐŶĞůůZ͕<ŝƐĐŚŬĂh͕DĂƚƚŚĞǁƐWD͕:ŽŚĂŶƐĞŶͲĞƌŐ,
;ϮϬϭϮͿŽƌƚŝĐĂůĂĐƚŝǀĂƚŝŽŶĐŚĂŶŐĞƐƵŶĚĞƌůLJŝŶŐƐƚŝŵƵůĂƚŝŽŶͲŝŶĚƵĐĞĚďĞŚĂǀŝŽƵƌĂůŐĂŝŶƐŝŶ
ĐŚƌŽŶŝĐƐƚƌŽŬĞ͘ƌĂŝŶϭϯϱ͗ϮϳϲͲϮϴϰ͘
dĂŬĞƵĐŚŝE͕/njƵŵŝ^;ϮϬϭϮͿDĂůĂĚĂƉƚŝǀĞƉůĂƐƚŝĐŝƚLJĨŽƌŵŽƚŽƌƌĞĐŽǀĞƌLJĂĨƚĞƌƐƚƌŽŬĞ͗ŵĞĐŚĂŶŝƐŵƐĂŶĚ
ĂƉƉƌŽĂĐŚĞƐ͘EĞƵƌĂůWůĂƐƚϮϬϭϮ͗ϯϱϵϳϮϴ͘
ϮϬ
d,ZK>K&/W^/>dZ>KZdy/EDKdKZ>ZE/E' Ϯϭ
ϳϯϳ
ϳϯϴ
ϳϯϵ
ϳϰϬ
ϳϰϭ
ϳϰϮ
ϳϰϯ
ϳϰϰ
ϳϰϱ
ϳϰϲ
ϳϰϳ
ϳϰϴ
ϳϰϵ
ϳϱϬ
ϳϱϭ
ϳϱϮ
ϳϱϯ
ϳϱϰ
ϳϱϱ
ϳϱϲ
ϳϱϳ
ϳϱϴ
ϳϱϵ
ϳϲϬ
ϳϲϭ
ϳϲϮ
ϳϲϯ
ϳϲϰ
ϳϲϱ
ϳϲϲ
ϳϲϳ
ϳϲϴ
ϳϲϵ
dĂŬĞƵĐŚŝE͕KŽƵĐŚŝĚĂz͕/njƵŵŝ^;ϮϬϭϮͿDŽƚŽƌĐŽŶƚƌŽůĂŶĚŶĞƵƌĂůƉůĂƐƚŝĐŝƚLJƚŚƌŽƵŐŚŝŶƚĞƌŚĞŵŝƐƉŚĞƌŝĐ
ŝŶƚĞƌĂĐƚŝŽŶƐ͘EĞƵƌĂůWůĂƐƚϮϬϭϮ͗ϴϮϯϮϴϱ͘
dƌƵŽŶŐY͕,ƵďĞƌD͕yŝĞy͕ĂƚƚĂ͕ZĂŚŵĂŶ͕WĂƌƌĂ>͕ŵŽĐŚŽǁƐŬŝ:W͕ŝŬƐŽŶD;ϮϬϭϰͿůŝŶŝĐŝĂŶ
ĐĐĞƐƐŝďůĞdŽŽůƐĨŽƌ'h/ŽŵƉƵƚĂƚŝŽŶĂůDŽĚĞůƐŽĨdƌĂŶƐĐƌĂŶŝĂůůĞĐƚƌŝĐĂů^ƚŝŵƵůĂƚŝŽŶ͗
KE^/ĂŶĚ^W,Z^͘ƌĂŝŶ^ƚŝŵƵů͘
sĂŶƐƐĞŶ͕>ĞǁŝƐ:t͕ƌƵƌLJ,͕,ĂĚũŝŬŚĂŶŝE͕dŽŽƚĞůůZ͕ĂŬŝƌĐŝŽŐůƵD͕DŝůůĞƌD/;ϮϬϬϭͿDĂƉƉŝŶŐ
ǀŝƐƵĂůĐŽƌƚĞdžŝŶŵŽŶŬĞLJƐĂŶĚŚƵŵĂŶƐƵƐŝŶŐƐƵƌĨĂĐĞͲďĂƐĞĚĂƚůĂƐĞƐ͘sŝƐŝŽŶƌĞƐĞĂƌĐŚϰϭ͗ϭϯϱϵͲ
ϭϯϳϴ͘
sĞƌŶŝĞƌŝ&͕ƐƐĞŶnjĂ'͕DĂŐŐŝŽW͕dŝďƵnjnjŝ&͕ĂƉƉĂƐŽĚŝ&͕ůƚĂŵƵƌĂ͕ŽƌďĞƚƚŽD͕dƌŽƚƚĂ>͕WĂůĂnjnjŽW͕
ƌĐŽůĂŶŝD͕dĞĐĐŚŝŽ&͕ZŽƐƐŝŶŝWD;ϮϬϭϬͿŽƌƚŝĐĂůŶĞƵƌŽŵŽĚƵůĂƚŝŽŶŵŽĚŝĨŝĞƐĐĞƌĞďƌĂů
ǀĂƐŽŵŽƚŽƌƌĞĂĐƚŝǀŝƚLJ͘^ƚƌŽŬĞϰϭ͗ϮϬϴϳͲϮϬϵϬ͘
sĞƌƐƚLJŶĞŶd͕ŝĞĚƌŝĐŚƐĞŶ:͕ůďĞƌƚE͕ƉĂƌŝĐŝŽW͕/ǀƌLJZ;ϮϬϬϱͿ/ƉƐŝůĂƚĞƌĂůŵŽƚŽƌĐŽƌƚĞdžĂĐƚŝǀŝƚLJ
ĚƵƌŝŶŐƵŶŝŵĂŶƵĂůŚĂŶĚŵŽǀĞŵĞŶƚƐƌĞůĂƚĞƐƚŽƚĂƐŬĐŽŵƉůĞdžŝƚLJ͘:EĞƵƌŽƉŚLJƐŝŽůϵϯ͗ϭϮϬϵͲϭϮϮϮ͘
sŝŶĞƐt͕ĞƌƌƵƚŝ͕^ĐŚůĂƵŐ';ϮϬϬϴͿƵĂůͲŚĞŵŝƐƉŚĞƌĞƚ^ĨĂĐŝůŝƚĂƚĞƐŐƌĞĂƚĞƌŝŵƉƌŽǀĞŵĞŶƚƐĨŽƌ
ŚĞĂůƚŚLJƐƵďũĞĐƚƐΖŶŽŶͲĚŽŵŝŶĂŶƚŚĂŶĚĐŽŵƉĂƌĞĚƚŽƵŶŝͲŚĞŵŝƐƉŚĞƌĞƐƚŝŵƵůĂƚŝŽŶ͘D
EĞƵƌŽƐĐŝϵ͗ϭϬϯ͘
tĂƚĞƌƐͲDĞƚĞŶŝĞƌ^͕,ƵƐĂŝŶD͕tŝĞƐƚůĞƌd͕ŝĞĚƌŝĐŚƐĞŶ:;ϮϬϭϰͿŝŚĞŵŝƐƉŚĞƌŝĐdƌĂŶƐĐƌĂŶŝĂůŝƌĞĐƚ
ƵƌƌĞŶƚ^ƚŝŵƵůĂƚŝŽŶŶŚĂŶĐĞƐĨĨĞĐƚŽƌͲ/ŶĚĞƉĞŶĚĞŶƚZĞƉƌĞƐĞŶƚĂƚŝŽŶƐŽĨDŽƚŽƌ^LJŶĞƌŐLJĂŶĚ
^ĞƋƵĞŶĐĞ>ĞĂƌŶŝŶŐ͘:ŽƵƌŶĂůŽĨEĞƵƌŽƐĐŝĞŶĐĞϯϰ;ϯͿ͘
tŝĞƐƚůĞƌd͕ŝĞĚƌŝĐŚƐĞŶ:;ϮϬϭϯͿ^ŬŝůůůĞĂƌŶŝŶŐƐƚƌĞŶŐƚŚĞŶƐĐŽƌƚŝĐĂůƌĞƉƌĞƐĞŶƚĂƚŝŽŶƐŽĨŵŽƚŽƌ
ƐĞƋƵĞŶĐĞƐ͘Ğ>ŝĨĞϮ͗ĞϬϬϴϬϭ͘
tŝĞƐƚůĞƌd͕tĂƚĞƌƐͲDĞƚĞŶŝĞƌ^͕ŝĞĚƌŝĐŚƐĞŶ:;ϮϬϭϰͿĨĨĞĐƚŽƌͲŝŶĚĞƉĞŶĚĞŶƚŵŽƚŽƌƐĞƋƵĞŶĐĞ
ƌĞƉƌĞƐĞŶƚĂƚŝŽŶƐĞdžŝƐƚŝŶĞdžƚƌŝŶƐŝĐĂŶĚŝŶƚƌŝŶƐŝĐƌĞĨĞƌĞŶĐĞĨƌĂŵĞƐ͘:EĞƵƌŽƐĐŝϯϰ͗ϱϬϱϰͲϱϬϲϰ͘
tŝůůŝĂŵƐ:͕WĂƐĐƵĂůͲ>ĞŽŶĞ͕&ƌĞŐŶŝ&;ϮϬϭϬͿ/ŶƚĞƌŚĞŵŝƐƉŚĞƌŝĐŵŽĚƵůĂƚŝŽŶŝŶĚƵĐĞĚďLJĐŽƌƚŝĐĂů
ƐƚŝŵƵůĂƚŝŽŶĂŶĚŵŽƚŽƌƚƌĂŝŶŝŶŐ͘WŚLJƐdŚĞƌϵϬ͗ϯϵϴͲϰϭϬ͘
tŽƌƐůĞLJ<:͕DĂƌƌĞƚƚ^͕EĞĞůŝŶW͕sĂŶĚĂů͕&ƌŝƐƚŽŶ<:͕ǀĂŶƐ;ϭϵϵϲͿƵŶŝĨŝĞĚƐƚĂƚŝƐƚŝĐĂůĂƉƉƌŽĂĐŚ
ĨŽƌĚĞƚĞƌŵŝŶŝŶŐƐŝŐŶŝĨŝĐĂŶƚƐŝŐŶĂůƐŝŶŝŵĂŐĞƐŽĨĐĞƌĞďƌĂůĂĐƚŝǀĂƚŝŽŶ͘,ƵŵƌĂŝŶDĂƉƉϰ͗ϱϴͲϳϯ͘
ŝŵĞƌŵĂŶD͕,ĞŝƐĞ<&͕'ĞƌůŽĨĨ͕ŽŚĞŶ>'͕,ƵŵŵĞů&;ϮϬϭϰͿŝƐƌƵƉƚŝŶŐƚŚĞŝƉƐŝůĂƚĞƌĂůŵŽƚŽƌ
ĐŽƌƚĞdžŝŶƚĞƌĨĞƌĞƐǁŝƚŚƚƌĂŝŶŝŶŐŽĨĂĐŽŵƉůĞdžŵŽƚŽƌƚĂƐŬŝŶŽůĚĞƌĂĚƵůƚƐ͘ĞƌĞďŽƌƚĞdž
Ϯϰ͗ϭϬϯϬͲϭϬϯϲ͘
ŝŵĞƌŵĂŶD͕,ĞŝƐĞ<&͕,ŽƉƉĞ:͕ŽŚĞŶ>'͕'ĞƌůŽĨĨ͕,ƵŵŵĞů&;ϮϬϭϮͿDŽĚƵůĂƚŝŽŶŽĨƚƌĂŝŶŝŶŐďLJ
ƐŝŶŐůĞͲƐĞƐƐŝŽŶƚƌĂŶƐĐƌĂŶŝĂůĚŝƌĞĐƚĐƵƌƌĞŶƚƐƚŝŵƵůĂƚŝŽŶƚŽƚŚĞŝŶƚĂĐƚŵŽƚŽƌĐŽƌƚĞdžĞŶŚĂŶĐĞƐ
ŵŽƚŽƌƐŬŝůůĂĐƋƵŝƐŝƚŝŽŶŽĨƚŚĞƉĂƌĞƚŝĐŚĂŶĚ͘^ƚƌŽŬĞϰϯ͗ϮϭϴϱͲϮϭϵϭ͘
ϳϳϬ
ϳϳϭ
Ϯϭ
d,ZK>K&/W^/>dZ>KZdy/EDKdKZ>ZE/E' ϮϮ
Figure captions
ϳϳϮ
ϳϳϯ
ϳϳϰ
Figure 1: Experimental Design.
ϳϳϱ
(a) – ‘–ƒ‰‡•ǣ •Šƒ •–‹—Žƒ–‹‘ ȋ„Ž—‡ȌǢ —‹Š‡‹•’Š‡”‹… ȋǮ‹ǯȌ – ȋ„Žƒ…ȌǢ
ϳϳϲ
„‹Š‡‹•’Š‡”‹…ȋǮ‹ǯȌ–ȋ”‡†ȌǢƒ†”‡˜‡”•‡†Ǧ’‘Žƒ”‹–›„‹Š‡‹•’Š‡”‹…ȋǮ‹ǦǯȌ–ȋ‰”‡‡ȌǤŠ‡
ϳϳϳ
Ž‘…ƒ–‹‘ ‘ˆ –Š‡ ƒ‘†‡ ‹• ‹†‹…ƒ–‡† „› ƒ ǮΪǯ ƒ† –Š‡ Ž‘…ƒ–‹‘ ‘ˆ –Š‡ …ƒ–Š‘†‡ „› ƒ ǮǦǯǤ ‡”‡ǡ –Š‡
ϳϳϴ
‡Ž‡…–”‘†‡’Žƒ…‡‡–ˆ‘”–Š‡Ž‡ˆ–ǦŠƒ†–”ƒ‹‡†‰”‘—’‹••Š‘™ȋˆ‘”™Š‹…Š–Š‡”‹‰Š–Š‡‹•’Š‡”‡
ϳϳϵ
™ƒ• –Š‡ Ǯ–”ƒ‹‡† Š‡‹•’Š‡”‡ǯȌǤ ‘” ”‹‰Š–ǦŠƒ† –”ƒ‹‡† ’ƒ”–‹…‹’ƒ–•ǡ –Š‡ ‡Ž‡…–”‘†‡ Ž‘…ƒ–‹‘ ™ƒ•
ϳϴϬ
”‡˜‡”•‡†ˆ‘”ƒŽŽ•–‹—Žƒ–‹‘‰”‘—’•Ǥ‘”’—”’‘•‡•‘ˆ†‘—„Ž‡Ǧ„Ž‹†‹‰ǡʹȀ͵‘ˆ–Š‡•Šƒ‰”‘—’Šƒ†
ϳϴϭ
ƒ„‹Š‡‹•’Š‡”‹…ƒ†ͳȀ͵Šƒ†ƒ—‹Š‡‹•’Š‡”‹…‘–ƒ‰‡Ǥ
ϳϴϮ
(b) ”‘…‡†—”‡ǣ —”‹‰ –Š‡ ’”‡Ǧ–‡•–ǡ ’ƒ”–‹…‹’ƒ–• ’‡”ˆ‘”‡† ͳʹ ͷǦ†‹‰‹– •‡“—‡…‡• ™‹–Š ‡‹–Š‡”
ϳϴϯ
Šƒ†Ǥ—„•‡“—‡–Ž›ǡ’ƒ”–‹…‹’ƒ–•™‡”‡ƒ••‹‰‡†–‘‘‡‘ˆ–Š‡ˆ‘—”–‰”‘—’•ƒ†–”ƒ‹‡†ˆ‘”Ͷ
ϳϴϰ
†ƒ›•™‹–Š‡‹–Š‡”–Š‡Ž‡ˆ–‘”–Š‡”‹‰Š–Šƒ†ǡ”‡•—Ž–‹‰‹ͺ†‹ˆˆ‡”‡–‰”‘—’•ȋˆ‘”†‡–ƒ‹Ž••‡‡Table
ϳϴϱ
1ȌǤ —”‹‰ –Š‡ ’‘•–Ǧ–‡•–ǡ ’ƒ”–‹…‹’ƒ–• ™‡”‡ –‡•–‡† ƒ‰ƒ‹ ȋƒ• ‹ –Š‡ ’”‡Ǧ–‡•–Ȍ ™‹–Š‘—– –Ǥ
ϳϴϲ
‹ƒŽŽ›ǡƒŽŽ’ƒ”–‹…‹’ƒ–•™‹–Šƒ„‹Š‡‹•’Š‡”‹…‘–ƒ‰‡ȋ„—–‘–™‹–Š—‹Š‡‹•’Š‡”‹…‘–ƒ‰‡Ȍ
ϳϴϳ
—†‡”™‡–ˆ—…–‹‘ƒŽ•…ƒ‹‰‘†ƒ›͹Ǥ
ϳϴϴ
ϳϴϵ
Figure 2: Bihemispheric tDCS accelerates learning and generalization in a polarity-
ϳϵϬ
independent manner.
ϳϵϭ
ȋaȌ˜‡”ƒ‰‡‹–Š‡’”‡Ǧ–‡•–ǡ–”ƒ‹‹‰ǡƒ†’‘•–Ǧ–‡•–ˆ‘”•Šƒȋ„Ž—‡Ȍǡ—‹Š‡‹•’Š‡”‹…ȋ„Žƒ…Ȍǡ
ϳϵϮ
„‹Š‡‹•’Š‡”‹…ȋ”‡†Ȍǡƒ†”‡˜‡”•‡†Ǧ’‘Žƒ”‹–›„‹Š‡‹•’Š‡”‹…ȋ‰”‡‡Ȍ–‰”‘—’•Ǥ—„Œ‡…–•–”ƒ‹‡†
ϳϵϯ
™‹–Š ‡‹–Š‡” –Š‡ Ž‡ˆ– Šƒ† ‘” ȋbȌ ”‹‰Š– Šƒ†Ǥ ȋcȌ ”‡Ǧ ƒ† ’‘•–Ǧ–‡•– †ƒ–ƒ ˆ‘” –Š‡ –”ƒ‹‡† ƒ† ȋdȌ
ϳϵϰ
—–”ƒ‹‡†•‡“—‡…‡•Ǥ‡•—Ž–•ƒ”‡•Š‘™ˆ‘”–Š‡–”ƒ‹‡†ȋȌƒ†—–”ƒ‹‡†ȋȌŠƒ†•ǡ•‡’ƒ”ƒ–‡†
ϳϵϱ
„›‰”‘—’ǤŠ‡’”‡Ǧ–‡•–”‡•—Ž–•ȋ‰”‡›†ƒ•Š‡†Ž‹‡Ȍƒ”‡ƒ˜‡”ƒ‰‡†ƒ…”‘••ƒŽŽˆ‘—”‰”‘—’•Ǥ””‘”„ƒ”•
ϳϵϲ
ƒ†•Šƒ†‡†”‡‰‹‘‹†‹…ƒ–‡„‡–™‡‡Ǧ•—„Œ‡…–Ǥ
ϳϵϳ
ϳϵϴ
ϳϵϵ
Figure 3: Bihemispheric tDCS recipients show greater average activation than sham in
ϴϬϬ
bilateral sensorimotor areas.
ϴϬϭ
(a)”‘ˆ‹Ž‡’Ž‘–‘ˆΨ•‹‰ƒŽ…Šƒ‰‡”‡Žƒ–‹˜‡–‘”‡•–‹–Š‡–”ƒ‹‡†ƒ†—–”ƒ‹‡†Š‡‹•’Š‡”‡•ˆ‘”
ϴϬϮ
–”ƒ‹‡† Šƒ† ‡š‡…—–‹‘•Ǥ ‡•—Ž–• ƒ”‡ •Š‘™ ˆ‘” –Š‡ •Šƒǡ „‹Š‡‹•’Š‡”‹… ȋ‹ –Ȍǡ ƒ†
ϴϬϯ
”‡˜‡”•‡†Ǧ’‘Žƒ”‹–› – ‰”‘—’• ȋ‹Ǧ –ȌǤ Š‡ šǦ…‘‘”†‹ƒ–‡ ‹†‹…ƒ–‡• –Š‡ †‹•–ƒ…‡ ˆ”‘ –Š‡
ϴϬϰ
ˆ—†—• ‘ˆ –Š‡ …‡–”ƒŽ •—Ž…—• ƒŽ‘‰ –Š‡ …‘”–‹…ƒŽ •—”ˆƒ…‡ ‹ ǡ ”—‹‰ ˆ”‘ ‘……‹’‹–ƒŽǦ’ƒ”‹‡–ƒŽ
ϴϬϱ
Œ—…–‹‘ ȋ’‘•–‡”‹‘”Ȍ –‘ –Š‡ ”‘•–”ƒŽ –‹’ ‘ˆ ’”‡‘–‘” …‘”–‡š ȋƒ–‡”‹‘”ȌǤ (b) ”‘ˆ‹Ž‡ ’Ž‘– ˆ‘” –Š‡
ϴϬϲ
—–”ƒ‹‡†Šƒ†Ǥ(c)˜‡”ƒ‰‡•—Ž…ƒŽ†‡’–ŠƒŽ‘‰–Š‡’”‘ˆ‹Ž‡Ǥ(d)‘…ƒ–‹‘‘ˆƒ”‡ƒ•ƒ˜‡”ƒ‰‡†‹–Š‡
ϴϬϳ
’”‘ˆ‹Ž‡ ’Ž‘–• ‘ ƒ ‹ˆŽƒ–‡† „”ƒ‹ •—”ˆƒ…‡ ȋ’—”’Ž‡ ƒ”‡ƒȌ (e) ˜‡”ƒ‰‡ Ψ •‹‰ƒŽ …Šƒ‰‡ ‹ ͸
ϮϮ
d,ZK>K&/W^/>dZ>KZdy/EDKdKZ>ZE/E' Ϯϯ
ϴϬϴ
ƒƒ–‘‹…ƒŽŽ›Ǧ†‡ˆ‹‡† • ȋ‹‡•–Ž‡” ƒ† ‹‡†”‹…Š•‡ǡ ʹͲͳ͵Ǣ ‹‡•–Ž‡” ‡– ƒŽǤǡ ʹͲͳͶȌǤ ”ƒ…‡–•
ϴϬϵ
‹†‹…ƒ–‡ –Šƒ– –Š‡ –™‘ „‹Š‡‹•’Š‡”‹… ‰”‘—’• ™‡”‡ •‹‰‹ˆ‹…ƒ–Ž› †‹ˆˆ‡”‡– ˆ”‘ •Šƒ ȋȗȗǣ
ϴϭϬ
pδͲǤͲͲͺ͵ǡ•–ƒ–‹•–‹…ƒŽ–Š”‡•Š‘Ž†ˆ‘”—Ž–‹’Ž‡…‘’ƒ”‹•‘•ǡȗǣpδͲǤͲͷȌǤ„„”‡˜‹ƒ–‹‘•ǣα…‡–”ƒŽ
ϴϭϭ
•—Ž…—•Ǣዐ–”ƒ’ƒ”‹‡–ƒŽ•—Ž…—•Ǣ‘α’‘•–…‡–”ƒŽ•—Ž…—•Ǣα•—’‡”‹‘”ˆ”‘–ƒŽ•—Ž…—•Ǥ
ϴϭϮ
ϴϭϯ
Figure 4: Bihemispheric tDCS recipients exhibit more activity in bilateral sensorimotor
ϴϭϰ
areas relative to sham. ϴϭϱ
(a)…–‹˜ƒ–‹‘ƒ’•ˆ‘”–Š‡–”ƒ‹‡†ƒ†—–”ƒ‹‡†Š‡‹•’Š‡”‡•ˆ‘”–Š‡•Šƒǡ„‹Š‡‹•’Š‡”‹…ȋ‹
ϴϭϲ
–Ȍǡƒ†”‡˜‡”•‡†Ǧ’‘Žƒ”‹–›ȋ‹Ǧ–Ȍ‰”‘—’•ƒ˜‡”ƒ‰‡†ƒ…”‘••Šƒ†–”ƒ‹‹‰…‘Š‘”–ǤŠ‹Ž‡
ϴϭϳ
–Š‡•Šƒ‰”‘—’‡šŠ‹„‹–‡†‹’•‹Žƒ–‡”ƒŽ†‡ƒ…–‹˜ƒ–‹‘ȋ–›’‹…ƒŽŽ›ˆ‘—†‹ˆ•–—†‹‡•‘ˆ—‹Žƒ–‡”ƒŽ
ϴϭϴ
‘˜‡‡–•Ȍǡ –Š‹• ’ƒ––‡” ™ƒ• ‘– ˆ‘—† ˆ‘” ‡‹–Š‡” „‹Š‡‹•’Š‡”‹… – ‰”‘—’Ǥ ††‹–‹‘ƒŽŽ›ǡ
ϴϭϵ
– ‰”‘—’• ‡šŠ‹„‹–‡† Š‹‰Š‡” …‘–”ƒŽƒ–‡”ƒŽ ƒ…–‹˜ƒ–‹‘Ǥ (b) ‘””‡•’‘†‹‰ ƒ’• ˆ‘” •‡“—‡…‡•
ϴϮϬ
‡š‡…—–‡† ™‹–Š –Š‡ —–”ƒ‹‡† Šƒ†Ǥ (c, d) ‹ˆˆ‡”‡…‡ Ǧƒ’• ˆ‘” –Š‡ ƒ…–‹˜ƒ–‹‘ ‹ –Š‡ –
ϴϮϭ
‰”‘—’•”‡Žƒ–‹˜‡–‘•Šƒˆ‘”–Š‡–”ƒ‹‡†ȋcȌƒ†—–”ƒ‹‡†ȋdȌŠƒ†•Ǥ
ϴϮϮ
„„”‡˜‹ƒ–‹‘•ǣα…‡–”ƒŽ•—Ž…—•Ǣዐ–”ƒ’ƒ”‹‡–ƒŽ•—Ž…—•Ǣ‘α’‘•–…‡–”ƒŽ•—Ž…—•Ǣα
ϴϮϯ
•—’‡”‹‘”ˆ”‘–ƒŽ•—Ž…—•Ǥ
ϴϮϰ
ϴϮϱ
ϴϮϲ
ϴϮϳ
Figure 5: Online perceptual side effects of tDCS. ϴϮϴ
‡ ‘‹–‘”‡† –Š‡ ’‘–‡–‹ƒŽ ‘Ž‹‡ •‹†‡ ‡ˆˆ‡…–• ‘ˆ – ‹ ‘”†‡” –‘ ‡˜ƒŽ—ƒ–‡ ’‡”…‡’–—ƒŽ
ϴϮϵ
†‹ˆˆ‡”‡…‡• „‡–™‡‡ – ‰”‘—’• ƒ† •ŠƒǤ (a) ‘” –Š‡ ‹–‡•‹–› Ž‡˜‡Ž ‘ˆ ƒŽŽ •‹š ‡ˆˆ‡…– –›’‡•ǡ
ϴϯϬ
–Š‡”‡™‡”‡‘•‹‰‹ˆ‹…ƒ–†‹ˆˆ‡”‡…‡•ƒ…”‘••–‰”‘—’•Ǥ(b)‘™‡˜‡”ǡ–Š‡†‹ˆˆ‡”‡…‡•„‡–™‡‡
ϴϯϭ
– ƒ† •Šƒ –‡†‡† –‘ „‡ ‰”‡ƒ–‡” ˆ‘” –Š‡ †—”ƒ–‹‘ ‘ˆ –Š‡•‡ •‹†‡ ‡ˆˆ‡…–• ƒ† •‹‰‹ˆ‹…ƒ–
ϴϯϮ
†‹ˆˆ‡”‡…‡•™‡”‡‘„•‡”˜‡†ˆ‘”†—”ƒ–‹‘‘ˆ„—”‹‰ƒ†‹–…Š‹‰ȋindicated by asterisksȌǤ(c) ‘•–Ǧ
ϴϯϯ
–‡•–’‡”ˆ‘”ƒ…‡‘ˆ–Š‡…‘ˆ‹‰—”ƒ–‹‘–ƒ••’‡‡†ȋleft panelȌƒ†‡ƒ†‡˜‹ƒ–‹‘ȋright panelȌ
ϴϯϰ
ˆ‘” –Š‡ •Šƒ ȋblueȌǡ „‹Š‡‹•’Š‡”‹… ȋredȌǡ ƒ† ”‡˜‡”•‡†Ǧ’‘Žƒ”‹–› „‹Š‡‹•’Š‡”‹… ȋgreenȌ ‰”‘—’•Ǥ
ϴϯϱ
(d) ‘•–Ǧ–‡•– ’‡”ˆ‘”ƒ…‡ ‘ˆ ‹†‹˜‹†—ƒ–‹‘ Ǥ – ”‡…‹’‹‡–• ‡š’‡”‹‡…‡† ‘ ƒ†˜‡”•‡
ϴϯϲ
„‡Šƒ˜‹‘”ƒŽ‡ˆˆ‡…–•ƒ•ƒ”‡•—Ž–‘ˆ–Ǧ…‘—’Ž‡†•‡“—‡…‡–”ƒ‹‹‰Ǥ
ϴϯϳ
ϴϯϴ
Ϯϯ
a
Sham
Uni tDCS
-
- +
+
or
Untrained
hemisphere
Bi tDCS
Bi-RP tDCS
-
+
Trained
hemisphere
Only bihemispheric
b
Training + tDCS
( Sham, Uni, Bi, or Bi-RP tDCS )
Pre-test
Post-test
fMRI
12 Sequences
4 Sequences
6
7
or
12 Sequences
Day
1
4 Sequences
2
3
2 mA tDCS
0min
25min
tDCS
Training
60min
4
5
a
b
Left-hand trained
3
Right-hand trained
Sham
Uni tDCS
Bi tDCS
Execution Time (s)
2.5
Bi-RP tDCS
2
1.5
1
Pretest
c
1
2
3
4
Training day
Posttest
Trained sequences
2.4
Pretest
d
1
2
3
4
Training day
Posttest
Untrained sequences
Execution Time (s)
2.2
pre-test
pre-test
2
1.8
1.6
1.4
1.2
1.0
0.8
0.6
T
U
Left-hand
trained
U
T
Right-hand
trained
T
U
Left-hand
trained
U
T
Right-hand
trained
Trained Hemisphere
Untrained Hemisphere
3
Signal change [%]
Trained Hand
a
Sham
Bi tDCS
Bi-RP tDCS
2
1
0
-30
-40
-30
-20
-10
0
10
20
30
-40
-30
-20
-10
0
10
20
30
-20
-10
0
10
20
30
-40
-30
-20
-10
0
10
20
30
3
Signal change [%]
Untrained Hand
b
-40
2
1
0
Distance from Central Sulcus [mm]
c
Sulcal depth [au]
d
PoCS
1
CS
IPS
SFS
0
OPJ
-1
-40
IPS
-30
S1
-20
-10
M1
0
PMd
10
20
30
Distance from Central Sulcus [mm]
% signal change
e
2
*
*
2 **
1.5
1.5
1
1
0.5
*
Hand
trained untrained
**
Sham
Bi
Bi-RP
0.5
**
*
0
0
M1
S1
PMd
SMA
Untrained hemisphere
IPS
OPJ
M1
S1
PMd
SMA
Trained hemisphere
IPS
OPJ
Intensity [0:10]
a
6
Tingling
Pain
Burning
Itching
Dizziness
Fatigue
4
2
0
*
Duration [min]
b
6
*
Sham
Uni tDCS
Bi tDCS
Bi-RP tDCS
*
4
2
0
c
d
Configuration Task
Sham
Bi tDCS
Bi-RP tDCS
0.4
2.5
2
1.5
1.4
pre-test
1.3
pre-test
RMSE
pre-test
Mean Deviation
Execution Time (s)
3.5
3
Individuation Task
0.3
1.2
1.1
1
0.9
0.2
1
0.8
T
U
Left-hand
trained
U
T
Right-hand
trained
T
U
Left-hand
trained
U
T
Right-hand
trained
T
U
Left-hand
trained
U
T
Right-hand
trained
d,ZK>K&/W^/>dZ>KZdy/EDKdKZ>ZE/E' ϭ
Table 1: Experimental prediction of hemispheric competition and cooperation models,
assuming either polarity specific or polarity unspecific effects.
tĞŚĂǀĞůŝƐƚĞĚƚŚĞĐŽŶĚŝƚŝŽŶƐŝŶƚŚĞŽƌĚĞƌŽĨƚŚĞƉƌĞĚŝĐƚĞĚďĞŚĂǀŝŽƌĂůƉĞƌĨŽƌŵĂŶĐĞ;фŵĞĂŶƐůŽǁĞƌ
ŵŽǀĞŵĞŶƚƚŝŵĞ͕ŝ͘Ğ͘ďĞƚƚĞƌƉĞƌĨŽƌŵĂŶĐĞͿ͘dŚĞƉƌĞĚŝĐƚŝŽŶƐĨŽƌƚŚĞŚĞŵŝƐƉŚĞƌŝĐĐŽŵƉĞƚŝƚŝŽŶŵŽĚĞů
ĂƐƐƵŵŝŶŐƉŽůĂƌŝƚLJͲƐƉĞĐŝĨŝĐĞĨĨĞĐƚƐŽŶƉůĂƐƚŝĐŝƚLJĂŶĚĨŽƌƚŚĞŚĞŵŝƐƉŚĞƌŝĐĐŽŽƉĞƌĂƚŝŽŶŵŽĚĞůĂƐƐƵŵŝŶŐ
ƉŽůĂƌŝƚLJͲƵŶƐƉĞĐŝĨŝĐĞĨĨĞĐƚƐĂƌĞƉƌĞƐĞŶƚĞĚŝŶƚŚĞŝŶƚƌŽĚƵĐƚŝŽŶ͘hŶĚĞƌƚŚĞŚĞŵŝƐƉŚĞƌŝĐĐŽŵƉĞƚŝƚŝŽŶ
ŵŽĚĞůǁŝƚŚƉŽůĂƌŝƚLJͲƵŶƐƉĞĐŝĨŝĐƚ^ĞĨĨĞĐƚƐ͕ƐƚŝŵƵůĂƚŝŶŐĐŽŶƚƌĂůĂƚĞƌĂůDϭŽŶůLJǁŽƵůĚďĞĞdžƉĞĐƚĞĚƚŽ
ďĞŵŽƌĞĞĨĨĞĐƚŝǀĞƚŚĂŶƐƚŝŵƵůĂƚŝŶŐďŽƚŚ;ƐŝŶĐĞďŽƚŚDϭƐĂƌĞĐŽŵƉĞƚŝŶŐͿ͘,ŽǁĞǀĞƌ͕ďŽƚŚ
ďŝŚĞŵŝƐƉŚĞƌŝĐŵŽŶƚĂŐĞƐƐŚŽƵůĚƐƚŝůůďĞŵŽƌĞĞĨĨĞĐƚŝǀĞƚŚĂŶƐŚĂŵĂƐĐŽŶƚƌĂůĂƚĞƌĂůDϭŝƐďĞŝŶŐ
ƐƚŝŵƵůĂƚĞĚ͘hŶĚĞƌƚŚĞŚĞŵŝƐƉŚĞƌŝĐĐŽŽƉĞƌĂƚŝŽŶŵŽĚĞůǁŝƚŚƉŽůĂƌŝƚLJͲƐƉĞĐŝĨŝĐƚ^ĞĨĨĞĐƚƐ͕
ƵŶŝŚĞŵŝƐƉŚĞƌŝĐƚ^ŝƐĞdžƉĞĐƚĞĚƚŽŚĂǀĞƚŚĞŐƌĞĂƚĞƐƚĨĂĐŝůŝƚĂƚŽƌLJĞĨĨĞĐƚďĞĐĂƵƐĞĐŽŶƚƌĂůĂƚĞƌĂůDϭŝƐ
ďĞŝŶŐƐƚŝŵƵůĂƚĞĚĂŶĚŝƉƐŝůĂƚĞƌĂůDϭŝƐƵŶĂĨĨĞĐƚĞĚ͘dŚĞĞdžĂĐƚƉƌĞĚŝĐƚŝŽŶĨŽƌƚŚĞƌĞŵĂŝŶŝŶŐƚŚƌĞĞ
ĐŽŶĚŝƚŝŽŶĚĞƉĞŶĚƐŽŶƚŚĞƌĞůĂƚŝǀĞŝŵƉŽƌƚĂŶĐĞĨŽƌĞĂĐŚŚĞŵŝƐƉŚĞƌĞŝŶĚĞǀĞůŽƉŝŶŐƚŚĞƐŬŝůů͘
WŽůĂƌŝƚLJƐƉĞĐŝĨŝĐ
ĂŶŽĚĂůŝŶĐƌĞĂƐĞƐ͕
ĐĂƚŚŽĚĂůĚĞĐƌĞĂƐĞƐƉůĂƐƚŝĐŝƚLJ
WŽůĂƌŝƚLJƵŶƐƉĞĐŝĨŝĐ
ďŽƚŚĂŶŽĚĂůĂŶĚĐĂƚŚŽĚĂů
ŝŶĐƌĞĂƐĞƉůĂƐƚŝĐŝƚLJ
,ĞŵŝƐƉŚĞƌŝĐĐŽŵƉĞƚŝƚŝŽŶ
ŵŽĚĞů
ŝфhŶŝф^ŚĂŵфŝͲZW
,ĞŵŝƐƉŚĞƌŝĐĐŽŽƉĞƌĂƚŝŽŶŵŽĚĞů
hŶŝфŝсŝͲZWф^ŚĂŵ
ŝсŝͲZWфhŶŝф^ŚĂŵ
ϭ
hŶŝфŝчŝͲZWч^ŚĂŵ
d,ZK>K&/W^/>dZ>KZdy/EDKdKZ>ZE/E' ϭ
Table 2: Participant demographic and psychological variables
‘‡˜ƒŽ—ƒ–‡ ‰”‘—’ †‹ˆˆ‡”‡…‡†‹ˆˆ‡”‡…‡•‹•‡šƒ††‡–‡…–ƒ„‹Ž‹–›‘ˆ–•–ƒ–—•ǡƒŽŽ–Š”‡‡–
‰”‘—’• ™‡”‡ ‹†‹˜‹†—ƒŽŽ› …‘’ƒ”‡† –‘ •Šƒ —•‹‰ ƒ χ Ǧ ‰‘‘†‡•• ‘ˆ ˆ‹– –‡•–Ǥ ‘” ƒŽŽ ‘–Š‡”
’ƒ”ƒ‡–‡”•ǡ‹š‡†Ǧ‡ˆˆ‡…–•™ƒ•…ƒŽ…—Žƒ–‡†™‹–Š–Š‡„‡–™‡‡ˆƒ…–‘”Ǯ–ǯǤŽŽ˜ƒ”‹ƒ„Ž‡•
™‡”‡ƒ˜‡”ƒ‰‡†ƒ…”‘••Ž‡ˆ–ƒ†”‹‰Š–Šƒ†–”ƒ‹‹‰…‘Š‘”–•Ǥ
Group
Sham
Uni tDCS
Bi tDCS
Bi-RP tDCS
Over all 4 groups
N ȋǦ–”ƒ‹‡†ǢǦ–”ƒ‹‡†Ȍ
‡šȋ‡ƒŽ‡ǣƒŽ‡Ȍ
Detectability of tDCS status
–•–ƒ–—•ǣȋ‡•ǣ‘Ȍ
tDCS perception
Averaged across all training
days
‹•…‘ˆ‘”–ȋͲǣͳͲȌ
‡”…‡‹˜‡†–‡•‹–›ȋͲǣͳͲȌ
‹•–”ƒ…–‹‘†—‡–‘–
ȋͲǣͳͲȌ
Demographics
‰‡
ʹͳȋͳͳǢͳͲȌ
ͳ͵ǣͺ
ͳͷȋͺǢ͹Ȍ
͹ǣͺ
ͳͶȋ͹Ǣ͹Ȍ
͸ǣͺ
ͳͶȋ͹Ǣ͹Ȍ
ͻǣͷ
ȱ2(1)
ͲǤ͸͸
p
ͲǤͶʹ
ͳͷǣ͸
ͳͳǣͶ
ͳͲǣͶ
ͳͳǣ͵
ͳͷǣ͸
ȱ2(1)
ͲǤͲ͸
͵ʹǣͳͳ
p
ͲǤͺͲ
F(3,60)
p
ʹǤ͵ʹάͲǤͶ͸
ʹǤͺͳάͲǤͶͲ
ͳǤͲ͹άͲǤ͵ͳ
ʹǤ͹ʹάͲǤ͵ͻ
͵ǤʹͺάͲǤͶͶ
ͳǤͺͲάͲǤ͵ͺ
͵Ǥ͵ͲάͲǤͷͶ
ʹǤͺͶάͲǤ͸ʹ
ͳǤ͸͵άͲǤͶ͵
ʹǤͺͺάͲǤ͵͹
͵ǤͻͳάͲǤͶͳ
ͳǤͷ͹άͲǤ͵ͻ
ͲǤͺͶ͸
ͳǤͳ͸Ͷ
ͲǤͺʹͷ
ͲǤͶ͹Ͷ
ͲǤ͵͵ͳ
ͲǤͶͺͷ
ʹʹǤͳͻάͲǤͻͷ
ʹͶǤͲͲάͳǤͷͻ
ʹʹǤͶ͵άͲǤͺʹ
F(3,60)
ͲǤͷͳͺ
p
ͲǤ͸͹ͳ
ƒ†‡†‡••ȋ†‹„—”‰ŠȌ
”‡˜‹‘—•‘–‘”–”ƒ‹‹‰ȋŠ”•Ȍ
Baseline motor
performance
ƒ•‡Ž‹‡
Psychological measures
Averaged across all days
Ž‡‡’‘—”•
Ž‡‡’—ƒŽ‹–›ȋͲǣͳͲȌ
Ž‡”–‡••ȋͲǣͳͲȌ
––‡–‹‘ȋͲǣͳͲȌ
ƒ•‹ˆˆ‹…—Ž–›ȋͲǣͳͲȌ
Averaged across final
training day and post-test
‡ƒ–‹…‡…ƒŽŽȋΨ…‘””‡…–Ȍ
ͺͺǤͲͻάͳǤͺ͹
͸ͶͲάʹͻ͵
ͺͶǤ͵͵άͶǤͷ͹
ͺ͹ʹά͵ͻ͸
ʹʹǤͻ͵ά
ͲǤͻͻ
͹͹ǤͷάͶǤͳͷ
ͶͶ͹άʹͶ͸
ͺʹǤͳͶά͵Ǥͻͷ
Ͷͺ͸ά͵Ͳ͸
ͳǤ͸͸ͳ
ͲǤ͵͵͸
F(3,60)
ͲǤͳͺͷ
ͲǤͺͲͲ
p
ʹǤ͵͸άͲǤͲ͸ͷ
͹ǤʹͻάͲǤʹͳ
͹ǤʹͺάͲǤʹ͸
͸Ǥ͹͹άͲǤ͵ͷ
͹Ǥ͸ͷάͲǤʹͺ
ͶǤͷ͸άͲǤͶͷ
ʹǤ͵͵άͲǤͳͷ
͸ǤͻͶάͲǤʹʹ
͸Ǥͻ͹άͲǤͳͻ
͸Ǥ͸ʹάͲǤͷͳ
͹Ǥͷ͸άͲǤ͵Ͷ
ͷǤͶ͵άͲǤͶʹ
ʹǤͶͷάͲǤͳͲ
͹ǤͶͻάͲǤͳͻ
͹ǤͺͻάͲǤʹͶ
͹ǤʹͳάͲǤͷͷ
ͺǤͳͺάͲǤ͵ʹ
ͶǤ͹ͳάͲǤ͵͹
ʹǤ͵ʹάͲǤͲ͸ͻ
͹ǤͶͻάͲǤʹͳ
͹Ǥʹ͹άͲǤʹ͸
͹ǤͳʹάͲǤʹͶ
͹ǤͲͶάͲǤͷʹ
ͷǤͳ͸άͲǤ͵Ͳ
ͲǤͳͷͻ
F(3,60)
ͳǤ͵ͻ͵
ʹǤͳͷͷ
ͲǤͶͳ͵
ͳǤͶͻͲ
ͲǤͻͻ͵
ͲǤͻʹͶ
p
ͲǤʹͷͶ
ͲǤͳͲʹ
ͲǤ͹ͶͶ
ͲǤʹʹ͸
ͲǤͶͲ͵
͹͵ǤʹͳάͶǤͷͻ
ͺͺǤ͵͵ά͵Ǥ͵͵
ͻ͸ǤͶ͵άͳǤͷ͹
ͻ͵Ǥ͹ͷάʹǤͷͶ
͹Ǥ͵͸ʹ
ͲǤͲͲͲ͵
ϭ
d,ZK>K&/W^/>dZ>KZdy/EDKdKZ>ZE/E' ϭ
Table 3: Effect of tDCS on average activation using surface-based analysis.
‡–™‡‡Ǧ•—„Œ‡…– ƒƒŽ›•‹• ™‹–Š —…‘””‡…–‡† Š‡‹‰Š– –Š”‡•Š‘Ž† ‘ˆ Tȋͳǡ͵ͻȌ α ʹǤ͹ͳǡ p α ͲǤͲͲͷǤ ”‡ƒ
‹†‹…ƒ–‡• –Š‡ •‹œ‡ ‘ˆ –Š‡ •—’”ƒ–Š”‡•Š‘Ž† …Ž—•–‡”ǡ Tȋͳǡ͵ͻȌ –Š‡ ƒš‹ƒŽ tǦ˜ƒŽ—‡ǡ PǦ…Ž—•–‡” ‹• –Š‡
…‘””‡…–‡†’”‘„ƒ„‹Ž‹–›‘ˆ‘„•‡”˜‹‰ƒ…Ž—•–‡”‘ˆ–Š‹••‹œ‡‘”„‹‰‰‡”‘˜‡”–Š‡™Š‘Ž‡…‘”–‹…ƒŽ•—”ˆƒ…‡
„›”ƒ†‘…Šƒ…‡ȋ‘”•Ž‡›‡–ƒŽǤǡͳͻͻ͸ȌǤ‘‘”†‹ƒ–‡•”‡ˆŽ‡…––Š‡Ž‘…ƒ–‹‘‘ˆ–Š‡…Ž—•–‡”’‡ƒ‹
•’ƒ…‡Ǥ
Region (Brodmann
Area)
Area
(mm2)
P
(clust)
Peak
value
T(1,39)
MNI coordinates
x
y
z
AVERAGE ACTIVATION
tDCS>sham (% signal activation > rest)
Trained hand
Contralateral (Trained) hemisphere
‘•–…‡–”ƒŽȋͳǢ͵Ȍ
ͳͶͷǤ͵͵ ͵Ǥ͸Ͷ
‘•–…‡–ƒŽȋͳǢ͵Ȍ
ͳͳ͸Ǥʹͳ ͵Ǥͺͳ
”‡…‡–”ƒŽȋͳǢͶȌ
ͳͳͶǤͲͳ ͵Ǥ͹ͺ
Ipsilateral (Untrained) hemisphere
”‡…‡–”ƒŽȋͳǢͶȌ
ͷ͹ͶǤͲͳ ͷǤͳͳ
‘•–…‡–”ƒŽȋͳǢ͵Ȍ
ʹʹͻǤ͸ͷ ͶǤͶͷ
”„‹–ƒŽƒ”‡ƒȀƒ”•
ͳͺʹǤ͸͵ ͷǤͶͻ
–”‹ƒ‰—Žƒ”‹•ȋͶ͹Ȍ
Untrained hand
Contralateral (Untrained) hemisphere
”„‹–ƒŽƒ”‡ƒȀƒ”•
ͳͳ͸Ǥ͵ͳ ͶǤ͵ͻ
–”‹ƒ‰—Žƒ”‹•ȋͶ͹Ȍ
ͶͻǤͺͻ
ͳͺǤʹ͸
͵͹Ǥ͹͹
ǦʹʹǤʹʹ
Ǧ͵ͻǤͷͺ
ǦʹͲǤͳ͸
ͷͳǤ͹ͺ
͸ͷǤ͸ͳ
ͷ͹Ǥʹʹ
δͲǤͲͲͳ
δͲǤͲͲͳ
ͲǤͲͲʹ
Ǧ͵͹ǤͶͳ
Ǧͳ͹ǤͲ͵
ǦͶʹǤͶ͸
Ǧʹ͸Ǥ͵͸
Ǧ͵ͻǤ͵ͳ
ʹͻǤͻͺ
ͷͷǤͺͳ
͸ͻǤ͸ͳ
ǦͲǤ͹ͺ
ͲǤͲ͵ͳ
Ipsilateral (Trained) hemisphere
All non-significant
ͲǤͲͲ͹
ͲǤͲ͵Ͳ
ͲǤͲ͵͵
ϭ
ǦͶͳǤ͸ͺ
͵ͲǤͲͳ
ǦͲǤͶͶ
d,ZK>K&/W^/>dZ>KZdy/EDKdKZ>ZE/E' ϭ
Table 4: Execution time, error rate, and force during fMRI.
ƒ„Ž‡•Š‘™•‡ƒȋά•–ƒ†ƒ”†‡””‘”Ȍ‘ˆ„‡Šƒ˜‹‘”ƒŽ’ƒ”ƒ‡–‡”•ˆ‘”ƒŽŽ–Š”‡‡–‰”‘—’•†—”‹‰
ˆ •…ƒ‹‰ ȋƒ˜‡”ƒ‰‡† ƒ…”‘•• Šƒ† –”ƒ‹‹‰ …‘Š‘”–ȌǤ Š‡ Žƒ•– …‘Ž— ‹†‹…ƒ–‡• ƒ FǦ–‡•–
…‘’ƒ”‹‰–Š‡͵‰”‘—’•Ǥ
Trained Hand
š‡…—–‹‘–‹‡ȋ•Ȍ
””‘””ƒ–‡ȋΨȌ
‘”…‡ȋȌ
Untrained Hand
š‡…—–‹‘–‹‡ȋ•Ȍ
””‘””ƒ–‡ȋΨȌ
‘”…‡ȋȌ
Šƒ
‡ƒ
‹–
‡ƒ Ǧ‹–
‡ƒ ͳǤ͵͸
ȋͲǤͲ͵Ȍ ͳǤ͵Ͳ
ͳ͵Ǥ͹͸ ȋʹǤ͵͸Ȍ ͷǤͲͺ
ͷǤͻͻ
ȋͲǤ͵ͶȌ ͷǤ͸ͷ
ȋͲǤͲʹȌ ͳǤ͵ʹ
ȋͳǤͳͺȌ ͷǤ͸ͳ
ȋͲǤʹͻȌ ͸Ǥͺͷ
ȋͲǤͲʹȌ ʹǤ͸Ͳʹ
ȋͳǤ͹ͲȌ ͹ǤʹͲ͸
ȋͲǤʹʹȌ ͶǤ͸ʹ͹
ͲǤͲͺ͸ͻ
ͲǤͲͲʹʹ
ͲǤͲͳͷ͹
ͳǤͶͲ
ͳͷǤͶ͵
ͷǤ͸ͻ
ȋͲǤͲʹȌ ͳǤ͵ͳ
ȋͳǤʹ͵Ȍ ͷǤ͹ͷ
ȋͲǤʹ͹Ȍ ͸Ǥ͹ͳ
ȋͲǤͲͳȌ ͶǤͷͲ͸
ȋͳǤ͵ͲȌ ͳͳǤͺ͸Ͷ
ȋͲǤʹͻȌ ͷǤ͹ͳͺ
ͲǤͲͳ͹Ͷ
ͲǤͲͲͲͳ
ͲǤͲͲ͸͸
ȋͲǤͲͶȌ
ȋʹǤͲ͵Ȍ
ȋͲǤʹ͸Ȍ
ͳǤ͵ʹ
͸ǤͷͲ
ͷǤͷͲ
ƒ…”‘••‰”‘—’•
Fȋʹǡ͵ͻȌ
p
ϭ
d,ZK>K&/W^/>dZ>KZdy/EDKdKZ>ZE/E' ϭ
Table 5: Mirroring of digit forces during fMRI.
ƒ„Ž‡ •Š‘™• ‡ƒ ȋά•–ƒ†ƒ”† ‡””‘”Ȍ ‘ˆ –Š‡ ƒ˜‡”ƒ‰‡ ˆ‘”…‡ ”ƒ‰‡ ȋƒš‹— Ȃ ‹‹—Ȍ ƒ†
•–ƒ†ƒ”††‡˜‹ƒ–‹‘‘ˆ–Š‡’ƒ••‹˜‡ȋ‘Ǧ‘˜‹‰ȌŠƒ†Ǥ‡•—Ž–•™‡”‡•’Ž‹–†‡’‡†‹‰‘™Š‡–Š‡”
–Š‡ –”ƒ‹‡† Šƒ† ‘” –Š‡ —–”ƒ‹‡† Šƒ† ™ƒ• –Š‡ ’ƒ••‹˜‡ Šƒ†Ǥ Š‡ …‘””‡Žƒ–‹‘ „‡–™‡‡ –Š‡
ˆ‘”…‡• ™ƒ• …ƒŽ…—Žƒ–‡† „‡–™‡‡ –Š‡ ˆ‘”…‡ –‹‡ •‡”‹‡• ‘ˆ –Š‡ ƒ–…Š‹‰ †‹‰‹–• ‘ˆ –Š‡ ƒ…–‹˜‡ ƒ†
’ƒ••‹˜‡Šƒ†•ǤŠ‡Žƒ•–…‘Ž—‹†‹…ƒ–‡•–Š‡Ǧ–‡•–…‘’ƒ”‹‰–Š‡͵‰”‘—’•Ǥ
Trained Hand
‘”…‡”ƒ‰‡ȏȐ
–ƒ†ƒ”††‡˜‹ƒ–‹‘ȏȐ
‘””‡Žƒ–‹‘‘ˆˆ‘”…‡–”ƒ…‡•
Untrained Hand
‘”…‡”ƒ‰‡ȏȐ
–ƒ†ƒ”††‡˜‹ƒ–‹‘ȏȐ
‘””‡Žƒ–‹‘‘ˆˆ‘”…‡–”ƒ…‡•
Šƒ
‡ƒ ‹–
‡ƒ ƒ…”‘••‰”‘—’•
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ȋͲǤͲͲʹȌ ͲǤͲ͹
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ȋͲǤͲͲȌ ͲǤͲͶ
ȋͲǤͲͷȌ ͲǤͲ͵
ȋͲǤͲ͵Ȍ ͲǤ͵ͷ͸
ȋͲǤͲͳȌ ͲǤͶͻ͹
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