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Research Articles: Behavioral/Cognitive

Eye-movement intervention enhances extinction via amygdala deactivation

Lycia D. de Voogd1, Jonathan W. Kanen1,3, David A. Neville1, Karin Roelofs1,2, Guillén Fernández1 and Erno J. Hermans1 1Donders Institute for Brain, Cognition and Behaviour, Radboud University and Radboud University Medical Center, 6500 HB, Nijmegen, The Netherlands 2Behavioural Science Institute, Radboud University, Nijmegen, The Netherlands 3Behavioural and Clinical Neuroscience Institute, Department of Psychology, University of Cambridge, Cambridge CB2 3EB, United Kingdom

DOI: 10.1523/JNEUROSCI.0703-18.2018

Received: 14 March 2018

Revised: 10 August 2018

Accepted: 16 August 2018

Published: 4 September 2018

Author contributions: L.D.d.V., K.R., G.F., and E.J.H. designed research; L.D.d.V. and J.K. performed research; L.D.d.V. and E.J.H. contributed unpublished reagents/analytic tools; L.D.d.V. analyzed data; L.D.d.V. wrote the first draft of the paper; L.D.d.V., J.K., D.A.N., K.R., G.F., and E.J.H. edited the paper; L.D.d.V. and E.J.H. wrote the paper.

Conflict of Interest: The authors declare no competing financial interests.

Correspondence to: Lycia D. de Voogd, Donders Institute for Brain, Cognition and Behaviour (Radboudumc), P.O. Box 9101, 6500 HB Nijmegen, The Netherlands, Ph: +31 (0)24 36 10878, Fx: +31 (0)24 36 10989, E-mail: [email protected]

Cite as: J. Neurosci ; 10.1523/JNEUROSCI.0703-18.2018

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Eye-movement intervention enhances extinction via amygdala deactivation Regular manuscript  › ‹ƒǤ†‡‘‘‰†ͳȗǡ ‘ƒ–ŠƒǤƒ‡ͳǡ͵ǡƒ˜‹†Ǥ‡˜‹ŽŽ‡ͳǡƒ”‹‘‡Ž‘ˆ•ͳǡʹǡ —‹ŽŽ± ‡”ž†‡œͳǡ ”‘ Ǥ ‡”ƒ•ͳ  ͳ‘†‡”• •–‹–—–‡ˆ‘””ƒ‹ǡ‘‰‹–‹‘ƒ†‡Šƒ˜‹‘—”ǡƒ†„‘—†‹˜‡”•‹–›ƒ†ƒ†„‘—†‹˜‡”•‹–› ‡†‹ ƒŽ‡–‡”ǡ͸ͷͲͲ ǡ‹Œ‡‰‡ǡŠ‡‡–Š‡”Žƒ†• ʹ‡Šƒ˜‹‘—”ƒŽ ‹‡ ‡ •–‹–—–‡ǡƒ†„‘—†‹˜‡”•‹–›ǡ‹Œ‡‰‡ǡŠ‡‡–Š‡”Žƒ†• ͵‡Šƒ˜‹‘—”ƒŽ ƒ† Ž‹‹ ƒŽ ‡—”‘• ‹‡ ‡ •–‹–—–‡ǡ ‡’ƒ”–‡– ‘ˆ •› Š‘Ž‘‰›ǡ ‹˜‡”•‹–› ‘ˆ ƒ„”‹†‰‡ǡƒ„”‹†‰‡ʹ͵ǡ‹–‡†‹‰†‘  ȗ‘””‡•’‘†‡ ‡–‘ǣ › ‹ƒǤ†‡‘‘‰† ‘†‡”• •–‹–—–‡ˆ‘””ƒ‹ǡ‘‰‹–‹‘ƒ†‡Šƒ˜‹‘—”ȋƒ†„‘—†— Ȍ ǤǤ‘šͻͳͲͳ ͸ͷͲͲ ‹Œ‡‰‡ǡŠ‡‡–Š‡”Žƒ†• ŠǣΪ͵ͳȋͲȌʹͶ͵͸ͳͲͺ͹ͺ šǣΪ͵ͳȋͲȌʹͶ͵͸ͳͲͻͺͻ Ǧƒ‹ŽǣŽǤ†‡˜‘‘‰†̷†‘†‡”•Ǥ”—ǤŽ 

Short title:›‡‘˜‡‡–•‡Šƒ ‡‡š–‹ –‹‘ Keywords: ›‰†ƒŽƒǢ ƒ˜Ž‘˜‹ƒ ˆ‡ƒ” ‘†‹–‹‘‹‰Ǣ š–‹ –‹‘ Ž‡ƒ”‹‰Ǣ ›‡ ‘˜‡‡– ‡•‡•‹–‹œƒ–‹‘ƒ†‡’”‘ ‡••‹‰ȋȌǢ‘‰‹–‹˜‡ ‘–”‘ŽǢˆ— –‹‘ƒŽ Ǥ Abstractǣͳͻͺ™‘”†• Introductionǣͷͻ͹™‘”†• Discussionǣͳ͸ͶͶ™‘”†• Number of figuresǣͷ Number of tablesǣ͸

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Abstract ’”‘˜‹‰ ‡š–‹ –‹‘ Ž‡ƒ”‹‰ ‹• ‡••‡–‹ƒŽ –‘ ‘’–‹‹œ‡ ’•› Š‘–Š‡”ƒ’› ˆ‘” ’‡”•‹•–‡– ˆ‡ƒ”Ǧ”‡Žƒ–‡† †‹•‘”†‡”•Ǥ  –™‘ ‹†‡’‡†‡– •–—†‹‡• ȋ„‘–Š αʹͶȌǡ ™‡ ˆ‘—† –Šƒ– ‰‘ƒŽǦ†‹”‡ –‡† ‡›‡ ‘˜‡‡–• ଶ ƒ –‹˜ƒ–‡ ƒ †‘”•ƒŽ ˆ”‘–‘Ǧ’ƒ”‹‡–ƒŽ ‡–™‘” ƒ† –”ƒ•‹‡–Ž› †‡ƒ –‹˜ƒ–‡ –Š‡ ƒ›‰†ƒŽƒ ȋߟ௣αǤͳ͹ȌǤ ‘‡ –‹˜‹–› ƒƒŽ›•‡• ”‡˜‡ƒŽ‡† –Šƒ– –Š‹• †‘™Ǧ”‡‰—Žƒ–‹‘ ’‘–‡–‹ƒŽŽ› ‡‰ƒ‰‡• ƒ ˜‡–”‘‡†‹ƒŽ ’”‡ˆ”‘–ƒŽ’ƒ–Š™ƒ›‘™–‘„‡‹˜‘Ž˜‡†‹ ‘‰‹–‹˜‡”‡‰—Žƒ–‹‘‘ˆ‡‘–‹‘Ǥ”‹–‹ ƒŽŽ›ǡ™Š‡‡›‡ ‘˜‡‡–•ˆ‘ŽŽ‘™‡†‡‘”›”‡ƒ –‹˜ƒ–‹‘†—”‹‰‡š–‹ –‹‘Ž‡ƒ”‹‰ǡ‹–”‡†— ‡†•’‘–ƒ‡‘—•ˆ‡ƒ” ଶ ”‡ ‘˜‡”›ʹͶŠ‘—”•Žƒ–‡”ȋߟ௣αǤʹͳȌǤ–”‘‰‡”ƒ›‰†ƒŽƒ†‡ƒ –‹˜ƒ–‹‘ˆ—”–Š‡”‘”‡’”‡†‹ –‡†ƒ•–”‘‰‡” ”‡†— –‹‘ ‹ •—„•‡“—‡– ˆ‡ƒ” ”‡ ‘˜‡”› ƒˆ–‡” ”‡‹•–ƒ–‡‡– ȋrαǤ͵ͻȌǤ  ‘ Ž—•‹‘ǡ ™‡ •Š‘™ –Šƒ– ‡š–‹ –‹‘Ž‡ƒ”‹‰ ƒ„‡‹’”‘˜‡†™‹–Šƒ‘Ǧ‹˜ƒ•‹˜‡‡›‡Ǧ‘˜‡‡–‹–‡”˜‡–‹‘–Šƒ––”‹‰‰‡”•ƒ –”ƒ•‹‡–•—’’”‡••‹‘‘ˆ–Š‡ƒ›‰†ƒŽƒǤ—”ˆ‹†‹‰–Šƒ–ƒ‘–Š‡”–ƒ•™Š‹ Š–ƒš‡•™‘”‹‰‡‘”› Ž‡ƒ†•–‘ƒ•‹‹Žƒ”ƒ›‰†ƒŽƒ•—’’”‡••‹‘ˆ—”–Š‡”‘”‡‹†‹ ƒ–‡•–Šƒ––Š‹•‡ˆˆ‡ –‹•Ž‹‡Ž›‘–•’‡ ‹ˆ‹  –‘‡›‡‘˜‡‡–•ǡ™Š‹ Š‹•‹Ž‹‡™‹–ŠƒŽƒ”‰‡„‘†›‘ˆ„‡Šƒ˜‹‘”ƒŽ•–—†‹‡•ǤŠ‹••–—†› ‘–”‹„—–‡•–‘ –Š‡—†‡”•–ƒ†‹‰‘ˆƒ™‹†‡Ž›—•‡†–”‡ƒ–‡–ˆ‘”–”ƒ—ƒ–‹ •›’–‘•„›’”‘˜‹†‹‰ƒ’ƒ”•‹‘‹‘—• ƒ ‘—–ˆ‘”Š‘™™‘”‹‰‡‘”›–ƒ••ƒ†‰‘ƒŽǦ†‹”‡ –‡†‡›‡‘˜‡‡–• ƒ‡Šƒ ‡‡š–‹ –‹‘Ǧ „ƒ•‡†’•› Š‘–Š‡”ƒ’›ǡƒ‡Ž›–Š”‘—‰Š‡—”ƒŽ ‹” —‹–•ȋe.g.ǡƒ›‰†ƒŽƒ†‡ƒ –‹˜ƒ–‹‘Ȍ•‹‹Žƒ”–‘–Š‘•‡ –Šƒ–•—’’‘”– ‘‰‹–‹˜‡ ‘–”‘Ž‘ˆ‡‘–‹‘Ǥ

Significant statement ‡ƒ”Ǧ”‡Žƒ–‡††‹•‘”†‡”•”‡’”‡•‡–ƒ•‹‰‹ˆ‹ ƒ–„—”†‡‘‹†‹˜‹†—ƒŽ•—ˆˆ‡”‡”•ƒ†•‘ ‹‡–›ǤŠ‡”‡‹•ƒ Š‹‰Š ‡‡† –‘ ‘’–‹‹œ‡ –”‡ƒ–‡–ǡ ‹ ’ƒ”–‹ —Žƒ” ˜‹ƒ non-invasive ‡ƒ•Ǥ ‡ ’‘–‡–‹ƒŽŽ› ‡ˆˆ‡ –‹˜‡ ‹–‡”˜‡–‹‘ ‹• ‡š‡ —–‹‘ ‘ˆ ‡›‡ ‘˜‡‡–• ˆ‘ŽŽ‘™‹‰ –”ƒ—ƒ ”‡ ƒŽŽǤ ‘™‡˜‡”ǡ ƒ ‡—”‘„‹‘Ž‘‰‹ ƒŽ —†‡”•–ƒ†‹‰‘ˆŠ‘™‡›‡‘˜‡‡–•”‡†— ‡–”ƒ—ƒ–‹ •›’–‘•‹•Žƒ ‹‰Ǥ‡†‡‘•–”ƒ–‡–Šƒ– ‰‘ƒŽǦ†‹”‡ –‡†‡›‡Ǧ‘˜‡‡–•ǡŽ‹‡™‘”‹‰‡‘”›–ƒ••ǡ†‡ƒ –‹˜ƒ–‡–Š‡ƒ›‰†ƒŽƒǡ–Š‡ ‘”‡‡—”ƒŽ •—„•–”ƒ–‡ ‘ˆ ˆ‡ƒ” Ž‡ƒ”‹‰Ǥ ˆˆ‡ –‹˜‡ ‘‡ –‹˜‹–› ƒƒŽ›•‡• ”‡˜‡ƒŽ‡† ƒ›‰†ƒŽƒ †‡ƒ –‹˜ƒ–‹‘ ’‘–‡–‹ƒŽŽ› ‡‰ƒ‰‡† †‘”•‘Žƒ–‡”ƒŽ ƒ† ˜‡–”‘‡†‹ƒŽ ’”‡ˆ”‘–ƒŽ ’ƒ–Š™ƒ›•Ǥ Š‡ ƒ’’Ž‹‡† †—”‹‰ •ƒˆ‡–›Ž‡ƒ”‹‰ǡ–Š‹•†‡ƒ –‹˜ƒ–‹‘’”‡†‹ –•ƒ”‡†— –‹‘‹Žƒ–‡”ˆ‡ƒ””‡ ‘˜‡”›ǤŠ‡•‡ˆ‹†‹‰•’”‘˜‹†‡ ƒ’ƒ”•‹‘‹‘—•ƒ†‡ Šƒ‹•–‹ ƒ ‘—–‘ˆŠ‘™„‡Šƒ˜‹‘”ƒŽƒ‹’—Žƒ–‹‘•–ƒš‹‰™‘”‹‰‡‘”› ƒ†•—’’”‡••‹‰ƒ›‰†ƒŽƒƒ –‹˜‹–› ƒƒŽ–‡””‡–‡–‹‘‘ˆ‡‘–‹‘ƒŽ‡‘”‹‡•Ǥ 

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Introduction

1 š–‹ –‹‘Ž‡ƒ”‹‰‹• ‘”‡–‘‘•–‡ˆˆ‡ –‹˜‡–Š‡”ƒ’‹‡•ˆ‘”†‹•‘”†‡”•‘ˆˆ‡ƒ”ƒ†ƒš‹‡–›ȋ‹••‘‡–ƒŽǤǡ 2 ʹͲͳ͵ȌǤ š’‘•—”‡ –Š‡”ƒ’›ǡ ˆ‘” ‹•–ƒ ‡ǡ ”‡•—Ž–• ‹ –Š‡ ˆ‘”ƒ–‹‘ ‘ˆ ƒ ‡š–‹ –‹‘ ‡‘”› –Šƒ– 3 •—’’”‡••‡• ˆ‡ƒ” ‡š’”‡••‹‘Ǥ ‡Žƒ’•‡ ‘ˆ ’ƒ–Š‘Ž‘‰‹ ƒŽ ˆ‡ƒ” ‹• ‡˜‡”–Š‡Ž‡•• ‘‘ ȋƒ”‡ǡ ʹͲͳͳǢ 4 —•‘‘” ‡– ƒŽǤǡ ʹͲͳͷ„ȌǤ ’”‘˜‹‰ ‡š–‹ –‹‘ Ž‡ƒ”‹‰ ‹• –Š‡”‡ˆ‘”‡ ƒ ‹’‘”–ƒ– ‰‘ƒŽ ‘ˆ 5 –”ƒ•Žƒ–‹‘ƒŽ ”‡•‡ƒ” Š ‹–‘ ˆ‡ƒ”Ǧ”‡Žƒ–‡† †‹•‘”†‡”• ȋ—•‘‘” ‡– ƒŽǤǡ ʹͲͳͷ„ȌǤ Šƒ”ƒ ‘Ž‘‰‹ ƒŽ 6 –”‡ƒ–‡–•Šƒ˜‡’”‘˜‡‡ˆˆ‡ –‹˜‡‹’”‡˜‡–‹‰ˆ‡ƒ””‡ ‘˜‡”›‹ƒ‹ƒŽ‘†‡Ž•ȋƒ†‡”‡–ƒŽǤǡʹͲͲͲȌǡ 7 „—– –Š‡•‡ ‡–Š‘†• ƒ”‡ ‘ˆ–‡ ‘– ƒ’’Ž‹ ƒ„Ž‡ ‹ Š—ƒ• ȋƒ†‡” ‡– ƒŽǤǡ ʹͲͲͲȌ ‘” Šƒ˜‡ ›‹‡Ž†‡† 8 ‹ ‘•‹•–‡–”‡•—Ž–•‹‡š’‡”‹‡–ƒŽ‘†‡Ž•™‹–ŠŠ—ƒ•ȋ‹†–‡–ƒŽǤǡʹͲͲͻǢ‘•‡–ƒŽǤǡʹͲͳʹȌǤ‡™ 9 ‘Ǧ‹˜ƒ•‹˜‡–‡ Š‹“—‡•Šƒ˜‡„‡‡†‡˜‡Ž‘’‡†–Šƒ––ƒ”‰‡–”‡ ‘•‘Ž‹†ƒ–‹‘‘ˆ–Š‡‘”‹‰‹ƒŽ‡‘”› 10 ”ƒ–Š‡”–Šƒ‡Šƒ ‡•–ƒ†ƒ”†‡š–‹ –‹‘Ž‡ƒ”‹‰ȋ Š‹ŽŽ‡”‡–ƒŽǤǡʹͲͳͲȌǤŽ–Š‘—‰Š–Š‡•‡”‡•—Ž–•ƒ”‡ 11 ’”‘‹•‹‰ǡ–Š‡‹” Ž‹‹ ƒŽ—–‹Ž‹–›‹••‘ˆƒ”— Ž‡ƒ”Ǥ 12 Ž‹‹ ƒŽŽ›‡ˆˆ‡ –‹˜‡–”‡ƒ–‡–•ƒ”‡‘–ƒŽ™ƒ›•†‡”‹˜‡†ˆ”‘•— Š‡š’‡”‹‡–ƒŽ‘†‡Ž•Ǥ‡ 13 ‡šƒ’Ž‡ ‹• ›‡ ‘˜‡‡– ‡•‡•‹–‹œƒ–‹‘ ƒ† ‡’”‘ ‡••‹‰ ȋǢ ‹••‘ ‡– ƒŽǤǡ ʹͲͳ͵Ȍǡ ƒ 14 ‡˜‹†‡ ‡Ǧ„ƒ•‡†–Š‡”ƒ’›ƒ†’ƒ”–‘ˆ‡–ƒŽŠ‡ƒŽ–Š ƒ”‡‰—‹†‡Ž‹‡•‹ƒ› ‘—–”‹‡•ȋ‹••‘‡–ƒŽǤǡ 15 ʹͲͳ͵Ǣ‡‡ƒ†—‹Œ’‡”•ǡʹͲͳ͵ȌǤ‡•’‹–‡‹–•™‹†‡—•‡ǡƒ‡ Šƒ‹•–‹ ǡ‡—”‘„‹‘Ž‘‰‹ ƒŽ—†‡”•–ƒ†‹‰ 16 ‘ˆ‹•Žƒ ‹‰Ǥ—”‹‰–”‡ƒ–‡–ǡ’ƒ–‹‡–•†‹˜‹†‡–Š‡‹”ƒ––‡–‹‘„‡–™‡‡”‡ ƒŽŽ‹‰–”ƒ—ƒ–‹  17 ‡‘”‹‡•ƒ†ƒ‹‰Žƒ–‡”ƒŽ‡›‡‘˜‡‡–•†‹”‡ –‡†„›–Š‡–Š‡”ƒ’‹•–̵•Šƒ†Ǥ›‡‘˜‡‡–•ƒ”‡ 18 ‡–”ƒŽ–‘–Š‡’”‘ ‡†—”‡ǡ„—–‹–‹•— Ž‡ƒ”‹ˆ–Š‡›’Žƒ›any”‘Ž‡‹–Š‡–Š‡”ƒ’‡—–‹ ‘—– ‘‡ƒ„‘˜‡ 19 ‘”ƒŽ‡š–‹ –‹‘ȋ‡‡ƒ†—‹Œ’‡”•ǡʹͲͳ͵ȌǤ •‹‰Š–‹–‘–Š‡’‘–‡–‹ƒŽ”‘Ž‡‘ˆ‡›‡‘˜‡‡–•ƒ† 20 –Š‡ ‡—”‘„‹‘Ž‘‰‹ ƒŽ ‡ Šƒ‹•• —†‡”Ž›‹‰ –Š‹• ƒ‹’—Žƒ–‹‘ ‹• ‘– ‘Ž› ”— ‹ƒŽ –‘ ˆ—”–Š‡” 21 ‘’–‹‹œ‡ –Š‹• –Š‡”ƒ’› „—– ™‘—Ž† ƒŽ•‘ ‹’‘”–ƒ–Ž› ƒ†˜ƒ ‡ ‘—” ˆ—†ƒ‡–ƒŽ —†‡”•–ƒ†‹‰ ‘ˆ 22 ‡š–‹ –‹‘Ž‡ƒ”‹‰Ǥ 23 ‡ Ž‡ƒ† ‹–‘ ƒ ‡—”ƒŽ ‡ Šƒ‹•–‹  ƒ ‘—– ‹• –Šƒ– ‰‘ƒŽǦ†‹”‡ –‡† ‡›‡ ‘˜‡‡–• ƒ”‡ 24 ƒ••‘ ‹ƒ–‡†™‹–Šƒ –‹˜ƒ–‹‘•‹–Š‡†‘”•ƒŽˆ”‘–‘Ǧ’ƒ”‹‡–ƒŽ‡–™‘”ǡ‹ Ž—†‹‰ˆ”‘–ƒŽ‡›‡ˆ‹‡Ž†•ȋ  Ǣ 25 ‘”„‡––ƒƒ†Š—ŽƒǡʹͲͲʹȌǡ•‹‹Žƒ”–‘™‘”‹‰‡‘”›–ƒ••–Šƒ–”‡“—‹”‡‰‘ƒŽǦ†‹”‡ –‡†ƒ––‡–‹‘Ǥ 26 ”‹–‹ ƒŽŽ›ǡ ™‘”‹‰ ‡‘”› –ƒ•• ƒ”‡ ƒ ‘’ƒ‹‡† „› ”‘„—•– †‡ƒ –‹˜ƒ–‹‘• ‹ ƒ ’‘•–‡”‹‘”Ǧ‡†‹ƒŽ 27 ‡–™‘”ȋ‹‡–ƒŽǤǡʹͲͲͻȌǡ‹ Ž—†‹‰–Š‡ƒ›‰†ƒŽƒǤ –‡”‡•–‹‰Ž›ǡƒ•‹‹Žƒ”ƒ –‹˜ƒ–‹‘‘ˆ–Š‡†‘”•ƒŽ 28 ˆ”‘–‘Ǧ’ƒ”‹‡–ƒŽ ‡–™‘” ƒ† †‡ƒ –‹˜ƒ–‹‘ ‘ˆ –Š‡ ƒ›‰†ƒŽƒ Šƒ• „‡‡ ˆ‘—† †—”‹‰ ‘‰‹–‹˜‡ 29 ”‡‰—Žƒ–‹‘‘ˆ‡‘–‹‘ƒ•™‡ŽŽȋ Š•‡”ƒ† ”‘••ǡʹͲͲͷȌǤŠ‹•‹•‹’‘”–ƒ–„‡ ƒ—•‡–ƒ”‰‡–‹‰–Š‡ 30 ƒ›‰†ƒŽƒˆ‘ŽŽ‘™‹‰‡‘”›”‡ƒ –‹˜ƒ–‹‘ǡ„›„Ž‘ ‹‰’”‘–‡‹•›–Š‡•‹•ǡ’”‡˜‡–•ˆ‡ƒ””‡ ‘˜‡”›‹ 31 ”‘†‡–• ȋƒ†‡” ‡– ƒŽǤǡ ʹͲͲͲȌǤ ‹‹Žƒ”Ž›ǡ •›•–‡‹  ƒ†‹‹•–”ƒ–‹‘ ‘ˆ ’”‘’”ƒ‘Ž‘Žǡ ƒ βǦƒ†”‡‘ ‡’–‘” 32 ƒ–ƒ‰‘‹•–ǡ’”‡•—ƒ„Ž›‡š‡”–•‹–•‡ˆˆ‡ –•‘ˆ‡ƒ””‡ ‘˜‡”›˜‹ƒ‹Š‹„‹–‹‘‘ˆ’”‘–‡‹•›–Š‡•‹•‹–Š‡

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33 ƒ›‰†ƒŽƒȋᑣ„‹‡ ƒ†‡†‘—šǡʹͲͲͶȌǤ›‰†ƒŽƒ”‡ƒ –‹˜‹–›‡ƒ•—”‡†™‹–ŠǦˆ ‹Š—ƒ•‹• 34 ˆ—”–Š‡”‘”‡ †‡ ”‡ƒ•‡† ƒˆ–‡” ’”‘’”ƒ‘Ž‘Ž ƒ†‹‹•–”ƒ–‹‘ ȋ —”Ž‡ƒ ‡– ƒŽǤǡ ʹͲͳͲȌǤ †‡‡†ǡ ‹– Šƒ• 35 „‡‡•Š‘™–Šƒ–™‘”‹‰‡‘”›ǦŽ‹‡–ƒ••ǡ•— Šƒ•ƒ‰ƒ‡‘ˆ‡–”‹•ȋ ‘Ž‡•‡–ƒŽǤǡʹͲͲͻǢ ƒ‡•‡– 36 ƒŽǤǡʹͲͳͷȌ ƒƒˆˆ‡ ––Š‡‡‘–‹‘ƒŽ‹–›‘ˆ‡‘”‹‡•Ǥ‡–Š‡”‡ˆ‘”‡Š›’‘–Š‡•‹œ‡†ȋͳȌ–Šƒ–‰‘ƒŽǦ†‹”‡ –‡† 37 ‡›‡ ‘˜‡‡–• ‘—Ž† „‡ —•‡† ƒ• ƒ ‘-‹˜ƒ•‹˜‡ –‘‘Ž –‘ –”ƒ•‹‡–Ž› •—’’”‡•• ƒ›‰†ƒŽƒ ƒ –‹˜‹–›ǡ 38 ‘’ƒ”ƒ„Ž‡ –‘ ™‘”‹‰ ‡‘”› –ƒ••ǡ ƒ† ȋʹȌ –Šƒ– ƒ ™‡ŽŽǦ–‹‡† ƒ’’Ž‹ ƒ–‹‘ ‘ˆ –Š‹• †‡ƒ –‹˜ƒ–‹‘ 39 ˆ‘ŽŽ‘™‹‰‡‘”›”‡ƒ –‹˜ƒ–‹‘ ‘—Ž†”‡†— ‡ˆ‡ƒ””‡ ‘˜‡”›Ǥ 40 ‘ –‡•– ‘—” Š›’‘–Š‡•‡•ǡ ‹ ‡š’‡”‹‡– ‘‡ ȋ α ʹͶȌǡ ’ƒ”–‹ ‹’ƒ–• ’‡”ˆ‘”‡† ƒ –™‘Ǧ„ƒ  41 ™‘”‹‰ ‡‘”› –ƒ• ƒ† ‰‘ƒŽǦ†‹”‡ –‡† ‡›‡ ‘˜‡‡–• ‹ ƒ „Ž‘  †‡•‹‰ ™Š‹Ž‡ —†‡”‰‘‹‰ 42 ˆ— –‹‘ƒŽ Ǥ‡–‡•–‡†™Š‡–Š‡”„‘–Š–ƒ••™‘—Ž†•—’’”‡••ƒ›‰†ƒŽƒƒ –‹˜‹–›ƒ•™‡ŽŽƒ•ƒŽ–‡”–Š‡ 43 ‘—’Ž‹‰„‡–™‡‡–Š‡ƒ›‰†ƒŽƒƒ††‘”•ƒŽŽƒ–‡”ƒŽ’”‡ˆ”‘–ƒŽ ‘”–‡šǤ –Š‡•‡ ‘†‡š’‡”‹‡–ȋα 44 ʹͶȌǡ™‡‹–‡‰”ƒ–‡†‡›‡‘˜‡‡–•‹–‘ƒ‡•–ƒ„Ž‹•Š‡†ƒ˜Ž‘˜‹ƒˆ‡ƒ” ‘†‹–‹‘‹‰Ȁ‡š–‹ –‹‘Ȁ 45 ”‡ ƒŽŽ ’ƒ”ƒ†‹‰ ƒ† –‡•–‡† ™Š‡–Š‡” ‰‘ƒŽǦ†‹”‡ –‡† ‡›‡ ‘˜‡‡–• ’”‡˜‡– ˆ‡ƒ” ”‡ ‘˜‡”› ˜‹ƒ –Š‹• 46 ƒ›‰†ƒŽƒ†‡ƒ –‹˜ƒ–‹‘Ǥ 47  48 Materials and Methods 49 50  ͳ 51 52 Participants 53 ™‡–›Ǧˆ‘—” ”‹‰Š–ǦŠƒ†‡† Š‡ƒŽ–Š› ˜‘Ž—–‡‡”• ȋͳʹ ˆ‡ƒŽ‡•ǡ ͳʹ ƒŽ‡•Ǣ ʹ͵Ǧ͵͹ ›‡ƒ”• ȏαʹ͸Ǥͻͷǡ 54 α͵Ǥ͸ȐȌ ‘’Ž‡–‡† –Š‡ •–—†›Ǥ š Ž—•‹‘ ”‹–‡”‹ƒ ™‡”‡ ƒ› ‘–”ƒ‹†‹ ƒ–‹‘• ˆ‘”  Ǥ ŽŽ ‰ƒ˜‡ 55 ™”‹––‡ ‹ˆ‘”‡† ‘•‡–ƒ†™‡”‡’ƒ‹†ˆ‘”–Š‡‹”’ƒ”–‹ ‹’ƒ–‹‘ǤŠ‹••–—†›™ƒ•ƒ’’”‘˜‡†„›–Š‡ 56 Ž‘ ƒŽ‡–Š‹ ƒŽ”‡˜‹‡™„‘ƒ”†ȋ”‡‰‹‘”Š‡Ǧ‹Œ‡‰‡ȌǤ 57  58 Experimental design 59 Š‡–ƒ•• ‘•‹•–‡†‘ˆ͸„Ž‘ •‘ˆƒ–™‘Ǧ„ƒ ™‘”‹‰‡‘”›–ƒ•ȋ‹‡–ƒŽǤǡʹͲͲͻȌǡ͸„Ž‘ •‘ˆ 60 •‘‘–ŠǦ’—”•—‹–Žƒ–‡”ƒŽ‡›‡‘˜‡‡––ƒ•ǡƒ†ƒƒ††‹–‹‘ƒŽͺ„Ž‘ •‘ˆŽ‘™ǦŽ‡˜‡Žˆ‹šƒ–‹‘„ƒ•‡Ž‹‡Ǥ 61 Š‡†—”ƒ–‹‘‘ˆ‡ƒ Š„Ž‘ ™ƒ•ʹ͹•‡ ‘†•Ǥ‹–Š‹‡ƒ Š–™‘Ǧ„ƒ „Ž‘ ǡ’ƒ”–‹ ‹’ƒ–••ƒ™ƒ”ƒ†‘ 62 •‡“—‡ ‡ ‘•‹•–‹‰ ‘ˆ ͳͷ •‹‰Ž‡ †‹‰‹–•Ǥ ƒ Š †‹‰‹– ™ƒ• ’”‡•‡–‡† ˆ‘” ͶͲͲ •ǡ ˆ‘ŽŽ‘™‡† „› ƒ 63 ‹–‡”•–‹—Ž—•‹–‡”˜ƒŽȋ  Ȍ‘ˆͳͶͲͲ•Ǥƒ”–‹ ‹’ƒ–•™‡”‡ƒ•‡†–‘†‡–‡ –™Š‡–Š‡”–Š‡ —””‡–‹–‡ 64 Šƒ†ƒ’’‡ƒ”‡†–™‘’‘•‹–‹‘•„ƒ ‹–Š‡•‡“—‡ ‡ƒ†™‡”‡‹•–”— –‡†–‘ƒ‡ƒ„—––‘’”‡••™Š‡ 65 †‡–‡ –‹‰ƒ–ƒ”‰‡–Ǥ ‘”–Š‡‡›‡Ǧ‘˜‡‡–„Ž‘ •ǡ’ƒ”–‹ ‹’ƒ–•™‡”‡‹•–”— –‡†–‘ˆ‘ŽŽ‘™ƒŽƒ–‡”ƒŽŽ›

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66 ‘˜‹‰ †‘– ™‹–Š –Š‡‹” ‡›‡•Ǥ Š‡ •’‡‡† ‘ˆ –Š‡ ‡›‡ ‘˜‡‡–• ™ƒ• ̱ͳ œǡ „ƒ•‡† ‘ ’”‡˜‹‘—• 67 Žƒ„‘”ƒ–‘”›‘†‡Ž•‘ˆȋ˜ƒ†‡ ‘—–‡–ƒŽǤǡʹͲͳ͵ȌǤ ‘”ͳͷ’ƒ”–‹ ‹’ƒ–•ǡ‡›‡Ǧ–”ƒ ‹‰†ƒ–ƒ™‡”‡ 68 ”‡ ‘”†‡†ƒ†˜‡”‹ˆ‹‡†’ƒ”–‹ ‹’ƒ–• ‘’Ž‹‡†™‹–Š–Š‡‹•–”— –‹‘•Ǥ 69  70 MRI data acquisition 71  • ƒ•™‡”‡ƒ “—‹”‡†—•‹‰ƒ‹‡‡•ȋ”Žƒ‰‡ǡ ‡”ƒ›Ȍ ›”ƒ͵• ƒ‡”Ǥ 72 ʹȗǦ™‡‹‰Š–‡† „Ž‘‘† ‘š›‰‡ƒ–‹‘ Ž‡˜‡ŽǦ†‡’‡†‡– ȋȌ ‹ƒ‰‡• ™‡”‡ ”‡ ‘”†‡† —•‹‰ ƒ 73 —•–‘‹œ‡†  •‡“—‡ ‡™‹–Šƒ• ‡†‹‰•Ž‹ ‡ƒ “—‹•‹–‹‘ȋ͵͹ƒš‹ƒŽ•Ž‹ ‡•ǢǡͳǤͺͻ•Ǣǡʹͷ •Ǣ 74 ‡‡”ƒŽ‹œ‡† —–‘ ƒŽ‹„”ƒ–‹‰ ƒ”–‹ƒŽŽ› ƒ”ƒŽŽ‡Ž  “—‹•‹–‹‘• ȋ Ǣ ”‹•™‘Ž† ‡– ƒŽǤǡ ʹͲͲʹȌ 75 ƒ ‡Ž‡”ƒ–‹‘ˆƒ –‘”ʹǢˆŽ‹’ƒ‰Ž‡ǡͻͲιǢ•Ž‹ ‡ƒ–”‹š•‹œ‡ǡ͸Ͷš͸ͶǢ•Ž‹ ‡–Š‹ ‡••ǡ͵Ǥ͵Ǣ•Ž‹ ‡‰ƒ’ǡͲǤ͵ 76 Ǣ ǡʹͳʹšʹͳʹǢ„ƒ†™‹†–Šǣͳ͹͹͸ œȀ’šǢ‡ Š‘•’ƒ ‹‰ǣͲǤ͸ͷ•ȌǤ•–”— –—”ƒŽ‹ƒ‰‡ȋͳ 77 ‹•‘–”‘’‹ Ȍ™ƒ•ƒ “—‹”‡†—•‹‰ƒͳǦ™‡‹‰Š–‡†͵ƒ‰‡–‹œƒ–‹‘Ǧ’”‡’ƒ”‡†”ƒ’‹†‰”ƒ†‹‡–Ǧ‡ Š‘ 78 •‡“—‡ ‡ȋǦ ǢǡʹǤ͹͵•ǢǡʹǤͻͷ•ǢˆŽ‹’ƒ‰Ž‡ǡ͹ιǢ ǡʹͷ͸šʹͷ͸šͳ͹͸ȌǤ 79 80 MRI data preprocessing and statistical analyses 81  †ƒ–ƒ™‡”‡’”‡Ǧ’”‘ ‡••‡†‹•–ƒ†ƒ”†•–‡”‡‘–ƒ –‹ ȋ ͳͷʹȌ•’ƒ ‡ˆ‘”–Š‡’—”’‘•‡‘ˆ™Š‘Ž‡Ǧ 82 „”ƒ‹‰”‘—’ƒƒŽ›•‡•Ǥ—–—ƒŽ‹ˆ‘”ƒ–‹‘ƒš‹‹œƒ–‹‘Ǧ„ƒ•‡†”‹‰‹†Ǧ„‘†›”‡‰‹•–”ƒ–‹‘™ƒ•—•‡†–‘ 83 ”‡‰‹•–‡” •–”— –—”ƒŽ ƒ† ˆ— –‹‘ƒŽ ‹ƒ‰‡•Ǥ — –‹‘ƒŽ ‹ƒ‰‡• ™‡”‡ ‘–‹‘ ‘””‡ –‡† —•‹‰ ”‹‰‹†Ǧ 84 „‘†›–”ƒ•ˆ‘”ƒ–‹‘•Ǥ–”— –—”ƒŽ‹ƒ‰‡•™‡”‡•‡‰‡–‡†‹–‘‰”ƒ›ƒ––‡”ǡ™Š‹–‡ƒ––‡”ǡƒ†  85 ‹ƒ‰‡• —•‹‰ ƒ —‹ˆ‹‡† ’”‘„ƒ„‹Ž‹•–‹  –‡’Žƒ–‡ ”‡‰‹•–”ƒ–‹‘ ƒ† –‹••—‡ Žƒ••‹ˆ‹ ƒ–‹‘ ‡–Š‘† 86 ȋ•Š„—”‡” ƒ† ”‹•–‘ǡ ʹͲͲͷȌǤ ‹••—‡ ‹ƒ‰‡• ™‡”‡ –Š‡ ”‡‰‹•–‡”‡† ™‹–Š •‹–‡Ǧ•’‡ ‹ˆ‹  –‹••—‡ 87 –‡’Žƒ–‡•ȋ ”‡ƒ–‡†ˆ”‘͵ͺͶͳǦ™‡‹‰Š–‡†• ƒ•Ȍ—•‹‰ȋ•Š„—”‡”ǡʹͲͲ͹Ȍǡƒ†”‡‰‹•–‡”‡† 88 ȋ—•‹‰ƒƒˆˆ‹‡–”ƒ•ˆ‘”ƒ–‹‘Ȍ™‹–Š–Š‡ ͳͷʹ–‡’Žƒ–‡‹ Ž—†‡†‹ͺȋ ǣ̴ͲͲ͹Ͳ͵͹ȌǤ 89 †‡–‹ ƒŽ –”ƒ•ˆ‘”ƒ–‹‘• ™‡”‡ ƒ’’Ž‹‡† –‘ ƒŽŽ ˆ— –‹‘ƒŽ ‹ƒ‰‡•ǡ ™Š‹ Š ™‡”‡ ”‡•Ž‹ ‡† ‹–‘ ʹ  90 ‹•‘–”‘’‹ ˜‘š‡Ž•ƒ†•‘‘–Š‡†™‹–Šƒ͸   ƒ—••‹ƒ‡”‡ŽǤ 91  ‡•’‘•‡• –‘ –Š‡ –™‘Ǧ„ƒ  –ƒ• ƒ† Žƒ–‡”ƒŽ ‡›‡ ‘˜‡‡–• ™‡”‡ ‘†‡Ž‡† —•‹‰ „‘šǦ ƒ” 92 ”‡‰”‡••‘”•ȋ†—”ƒ–‹‘‘ˆʹ͹•ȌǤŠ‡•‡–™‘”‡‰”‡••‘”•™‡”‡–‡’‘”ƒŽŽ› ‘˜‘Ž˜‡†™‹–Š–Š‡ ƒ‘‹ ƒŽ 93 Š‡‘†›ƒ‹  ”‡•’‘•‡ ˆ— –‹‘ ȋ  Ȍ ‹ Ž—†‡† ‹ ͺ ȋ ǣ̴ͲͲ͹Ͳ͵͹ȌǤ ††‹–‹‘ƒŽŽ›ǡ •‹š 94 ‘˜‡‡– ’ƒ”ƒ‡–‡” ”‡‰”‡••‘”• ȋ͵ –”ƒ•Žƒ–‹‘•ǡ ͵ ”‘–ƒ–‹‘•Ȍ †‡”‹˜‡† ˆ”‘ ”‹‰‹†Ǧ„‘†› ‘–‹‘ 95 ‘””‡ –‹‘ǡ Š‹‰ŠǦ’ƒ•• ˆ‹Ž–‡”‹‰ ȋͳȀͳʹͺ œ —–Ǧ‘ˆˆȌǡ ƒ† ȋͳȌ •‡”‹ƒŽ ‘””‡Žƒ–‹‘ ‘””‡ –‹‘• ™‡”‡ 96 ‹ Ž—†‡† ‹ –Š‡ ‘†‡ŽǤ ‹‰Ž‡Ǧ•—„Œ‡ – ‘–”ƒ•– ƒ’• ‘ˆ –Š‡ –™‘Ǧ„ƒ  ƒ† ‡›‡Ǧ‘˜‡‡– „Ž‘ • 97 ƒ‰ƒ‹•–ˆ‹šƒ–‹‘™‡”‡‡–‡”‡†‹–‘•‡ ‘†ǦŽ‡˜‡Ž‘‡Ǧ•ƒ’Ž‡–Ǧ–‡•–•Ǥ

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98  ‹ƒŽŽ›ǡ™‡ ‘†— –‡†ƒ ’•› Š‘’Š›•‹‘Ž‘‰‹ ƒŽ ‹–‡”ƒ –‹‘ȋ ȌƒƒŽ›•‹•™‹–Š–Š‡ƒ›‰†ƒŽƒ 99 ȋŽ‡ˆ– ƒ† ”‹‰Š– •‡’ƒ”ƒ–‡Ž›Ȍ ƒ• •‡‡†• ˆ‘” „‘–Š –Š‡ ‡›‡Ǧ‘˜‡‡– ‘†‹–‹‘ ƒ† –Š‡ –™‘Ǧ„ƒ  100 ‘†‹–‹‘Ǥ‡’‡”ˆ‘”‡†–Š‡ ƒƒŽ›•‹•‹š’‡”‹‡–ͳǡ„‡ ƒ—•‡Š‡”‡™‡—•‡†ƒ„Ž‘ ‡††‡•‹‰ǡ 101 ™Š‹ Š ‹• ‘’–‹ƒŽ ˆ‘” ‹˜‡•–‹‰ƒ–‹‰ –ƒ•Ǧ†”‹˜‡ ‘‡ –‹˜‹–› Šƒ‰‡• ‘ –‘’ ‘ˆ –ƒ• ƒ –‹˜ƒ–‹‘ 102 ȋ ”‹•–‘‡–ƒŽǤǡͳͻͻ͹ȌǤŠ‡•‡ȋˆ‘—”Ȍˆ‹”•–Ž‡˜‡Ž‘†‡Ž•™‡”‡‹†‡–‹ ƒŽ–‘–Š‡‘†‡Ž†‡• ”‹„‡†ƒ„‘˜‡ǡ 103 „—–‡ƒ Š‹ Ž—†‡†–™‘ƒ††‹–‹‘ƒŽ”‡‰”‡••‘”•ƒ‡Ž›ȏͳȐ–Š‡–‹‡•‡”‹‡•ˆ‘”–Š‡ˆ‹”•–‡‹‰‡˜ƒ”‹ƒ–‡‘ˆ 104 –Š‡ƒ›‰†ƒŽƒ•‡‡†ȋ‡‹–Š‡”Ž‡ˆ–‘””‹‰Š–Ȍǡƒ†ȏʹȐ–Š‡’”‘†— –‘ˆ–Š‹•–‹‡•‡”‹‡•ȏƒˆ–‡”Š‡‘†›ƒ‹  105 ”‡•’‘•‡ˆ— –‹‘ȋ  Ȍ†‡ ‘˜‘Ž—–‹‘Ȑ™‹–Š–ƒ•”‡‰”‡••‘”ȋ‡‹–Š‡”–Š‡–™‘Ǧ„ƒ „Ž‘ •‘”–Š‡‡›‡ 106 ‘˜‡‡–„Ž‘ •˜‡”•—•ˆ‹šƒ–‹‘Ȍ–Šƒ–™ƒ•–‡’‘”ƒŽŽ› ‘˜‘Ž˜‡†™‹–Š–Š‡ ƒ‘‹ ƒŽ  ‹ Ž—†‡† 107 ‹ ͺ ȋ ǣ̴ͲͲ͹Ͳ͵͹ȌǤ ‹‰Ž‡Ǧ•—„Œ‡ – ‘–”ƒ•– ƒ’• ‘ˆ –Š‡ –™‘Ǧ„ƒ  ƒ† ‡›‡Ǧ‘˜‡‡– 108 „Ž‘ • ƒ‰ƒ‹•– ˆ‹šƒ–‹‘ ™‡”‡ ‡–‡”‡† ‹–‘ •‡ ‘†ǦŽ‡˜‡Ž ˆ—ŽŽ ˆƒ –‘”‹ƒŽ ”‡’‡ƒ–‡† ‡ƒ•—”‡• • 109 ™‹–ŠŠ‡‹•’Š‡”‡ƒ•ƒ™‹–Š‹Ǧ•—„Œ‡ –˜ƒ”‹ƒ„Ž‡Ǥ 110 ƒ•‡†‘‘—”ƒ’”‹‘”‹Š›’‘–Š‡•‡•ǡ”‡•—Ž–•ˆ‘”–Š‡ƒ›‰†ƒŽƒƒ†˜‡–”ƒŽ‡†‹ƒŽ’”‡ˆ”‘–ƒŽ 111 ‘”–‡šȋ˜ Ȍ™‡”‡ ‘””‡ –‡†ˆ‘””‡†— ‡†•‡ƒ” Š˜‘Ž—‡•—•‹‰•ƒŽŽ˜‘Ž—‡ ‘””‡ –‹‘•ȋȌ 112 ƒ†™‡”‡ˆƒ‹Ž›Ǧ™‹•‡‡””‘”ȋ Ȍ ‘””‡ –‡†—•‹‰˜‘š‡ŽǦŽ‡˜‡Ž•–ƒ–‹•–‹ •Ǥ‘ˆ–Š‡ƒ›‰†ƒŽƒ™ƒ• 113 „ƒ•‡†‘ƒ‰”‘—’ƒ•–Šƒ–™ƒ• ”‡ƒ–‡†„›ƒ˜‡”ƒ‰‹‰‹†‹˜‹†—ƒŽƒ›‰†ƒŽƒ•‡‰‡–ƒ–‹‘•ȋαʹͶȌ‘ˆ 114 ͳǦ™‡‹‰Š–‡†‹ƒ‰‡•—•‹‰  ȋ ǣ̴ͲͲʹͺʹ͵ǡ•‡‡ 115 Š––’•ǣȀȀˆ•ŽǤˆ”‹„Ǥ‘šǤƒ Ǥ—Ȁˆ•ŽȀˆ•Ž™‹‹Ȁ Ȍǡ™Š‹ Š™‡”‡™ƒ”’‡†‹–‘ •’ƒ ‡—•‹‰Ǥ 116 Š‡˜ ™ƒ•†‡ˆ‹‡†ƒ•ͳͲ•’Š‡”‡ƒ”‘—†–Š‡ͲͶͲǦ͵ ‘‘”†‹ƒ–‡„ƒ•‡†‘ƒ’”‡˜‹‘—••–—†› 117 ȋ Š‹ŽŽ‡”ƒ†‡Ž‰ƒ†‘ǡʹͲͳͲȌǤ 118 119 Eye tracking 120 ‘”ͳͷ’ƒ”–‹ ‹’ƒ–•ǡ‡›‡–”ƒ ‹‰™ƒ•”‡ ‘”†‡†—•‹‰ƒǦ ‘’ƒ–‹„Ž‡‡›‡Ǧ–”ƒ ‹‰•›•–‡ȋ›‡ 121 ”ƒ Ǧ ƒ‡”ƒ —‹–ǡ  ǡ ‡•‘‘–‘”‹  •–”—‡–•ȌǤ ƒ–ƒ ™‡”‡ ’”‡’”‘ ‡••‡† —•‹‰ ‹ǦŠ‘—•‡ 122 •‘ˆ–™ƒ”‡ ȋ ‡”ƒ• ‡– ƒŽǤǡ ʹͲͳ͵Ȍ ‹’Ž‡‡–‡† ‹ ƒ–Žƒ„ ͹ǤͳͶ ȋƒ–Š‘”•ǡ  ǣ̴ͲͲͳ͸ʹʹȌǤ 123 Ž‹•™‡”‡”‡‘˜‡†ˆ”‘–Š‡•‹‰ƒŽ—•‹‰Ž‹‡ƒ”‹–‡”’‘Žƒ–‹‘Ǥ›‡Ǧ–”ƒ ‹‰†ƒ–ƒ™‡”‡‘”ƒŽ‹œ‡† 124 „ƒ•‡† ‘ ƒ ƒŽ‹„”ƒ–‹‘ ƒ– –Š‡ •–ƒ”– ‘ˆ –Š‡ ‡š’‡”‹‡–Ǥ ‹•—ƒŽ ‹•’‡ –‹‘ ”‡˜‡ƒŽ‡† ’ƒ”–‹ ‹’ƒ–• 125 ‘’Ž‹‡†™‹–Š–Š‡‹•–”— –‹‘•‘ˆ–Š‡–ƒ•Ǥ 

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126  ʹ 127 128 Participants 129 ™‡–›Ǧˆ‘—””‹‰Š–ǦŠƒ†‡†Š‡ƒŽ–Š›˜‘Ž—–‡‡”•ȋͳʹˆ‡ƒŽ‡•ǡͳʹƒŽ‡•ǢʹͲǦ͵Ͷ›‡ƒ”•ȏαʹͶǤͺǡα͵Ǥ͸ȐȌ 130 ‘’Ž‡–‡†–Š‡•–—†›Ǥƒ††‹–‹‘ƒŽͷ’ƒ”–‹ ‹’ƒ–•†‹†‘– ‘’Ž‡–‡–Š‡‡–‹”‡‡š’‡”‹‡–†—‡‘Ǧ 131 ‘’Ž‹ƒ ‡ ™‹–Š ‹•–”— –‹‘• ȋe.g., ˆƒŽŽ‹‰ ƒ•Ž‡‡’ȌǤ š Ž—•‹‘ ”‹–‡”‹ƒ ™‡”‡ǣ —””‡– ‘” Ž‹ˆ‡–‹‡ 132 Š‹•–‘”› ‘ˆ ’•› Š‹ƒ–”‹ ǡ ‡—”‘Ž‘‰‹ ƒŽǡ ‘” ‡†‘ ”‹‡ ‹ŽŽ‡••ǡ —””‡– –”‡ƒ–‡– ™‹–Š ƒ› ‡†‹ ƒ–‹‘ 133 –Šƒ– ƒˆˆ‡ –• ‡–”ƒŽ ‡”˜‘—• •›•–‡ ‘” ‡†‘ ”‹‡ •›•–‡•ǡ ƒ˜‡”ƒ‰‡ —•‡ ‘ˆ ‘”‡ –Šƒ ͵ ƒŽ ‘Š‘Ž‹  134 „‡˜‡”ƒ‰‡•†ƒ‹Ž›ǡƒ˜‡”ƒ‰‡—•‡‘ˆ”‡ ”‡ƒ–‹‘ƒŽ†”—‰•™‡‡Ž›‘”‘”‡ǡŠƒ„‹–—ƒŽ•‘‹‰ǡ’”‡†‘‹ƒ– 135 Ž‡ˆ–ǦŠƒ†‡†‡••ǡ— ‘””‡ –‡†˜‹•‹‘ǡ‹–‡•‡†ƒ‹Ž›’Š›•‹ ƒŽ‡š‡” ‹•‡ǡƒ†ƒ› ‘–”ƒ‹†‹ ƒ–‹‘•ˆ‘” 136  Ǥƒ”–‹ ‹’ƒ–•‰ƒ˜‡™”‹––‡‹ˆ‘”‡† ‘•‡–ƒ†™‡”‡’ƒ‹†ˆ‘”–Š‡‹”’ƒ”–‹ ‹’ƒ–‹‘ǤŠ‹••–—†› 137 ™ƒ•ƒ’’”‘˜‡†„›–Š‡Ž‘ ƒŽ‡–Š‹ ƒŽ”‡˜‹‡™„‘ƒ”†ȋ”‡‰‹‘”Š‡Ǧ‹Œ‡‰‡ȌǤ 138  139 Experimental design 140 ƒ”–‹ ‹’ƒ–• ™‡”‡ –‡•–‡† ‹ ƒ †‹ˆˆ‡”‡–‹ƒŽ †‡Žƒ› ˆ‡ƒ” ‘†‹–‹‘‹‰ ’ƒ”ƒ†‹‰ ȋ Š‹ŽŽ‡” ‡– ƒŽǤǡ ʹͲͳͲǡ 141 ʹͲͳ͵Ȍ ‘ –Š”‡‡ ‘•‡ —–‹˜‡ †ƒ›• ™‹–Š ʹͶŠ ‹ „‡–™‡‡Ǥ Š‡ ˆ‹”•– †ƒ› ‘’”‹•‡† ƒ ƒ “—‹•‹–‹‘ 142 •‡••‹‘ǡ–Š‡•‡ ‘††ƒ›ƒ‡š–‹ –‹‘•‡••‹‘ǡƒ†–Š‡–Š‹”††ƒ›ƒ”‡ ƒŽŽ•‡••‹‘ǤŠ‡•–‹—Ž—••‡– 143 ƒ ”‘••–Š‡–Š”‡‡†ƒ›• ‘•‹•–‡†‘ˆˆ‘—”•“—ƒ”‡•ƒ• ‘†‹–‹‘‡†•–‹—Ž‹ȋȌ™‹–Šƒ†‹ˆˆ‡”‡– ‘Ž‘”Ǥ 144 Š‡ Ž—‹ƒ ‡ ‘ˆ –Š‡ •–‹—Ž‹ǡ „ƒ ‰”‘—†ǡ ƒ†   • ”‡‡ ™ƒ• ‡“—ƒŽ‹œ‡†Ǥ  †ƒ› ‘‡ǡ –™‘ —‡• 145 ȋΪ•ǡ Ͷ• †—”ƒ–‹‘Ȍ ™‡”‡ ’ƒ”–‹ƒŽŽ› ”‡‹ˆ‘” ‡† ȋ͵͹Ǥͷ Ψ ”‡‹ˆ‘” ‡‡– ”ƒ–‡Ȍ ™‹–Š ƒ ‹Ž† ‡Ž‡ –”‹ ƒŽ 146 •Š‘ –‘–Š‡ˆ‹‰‡”•ȋi.e.ǡ–Š‡— ‘†‹–‹‘‡†•–‹—Ž—•ǢȌǤŠ‡–™‘‘–Š‡” —‡•ȋǦ•ǡͶ•†—”ƒ–‹‘Ȍ 147 ™‡”‡‡˜‡””‡‹ˆ‘” ‡†Ǥ –‘–ƒŽǡ–Š‡”‡™‡”‡͸Ͷ–”‹ƒŽ•ȋͳ͸–”‹ƒŽ•’‡”Ȍǡ‹ Ž—†‹‰–Š‡Ǧ–”‹ƒŽ•Ǥ 148 Š‡Ϊ•”‡‹ˆ‘” ‡†ǡΪ•—”‡‹ˆ‘” ‡†ǡƒ†Ǧ•™‡”‡’”‡•‡–‡†‹ƒ’•‡—†‘”ƒ†‘‘”†‡”ǤŠ‡   149 ™ƒ•Œ‹––‡”‡†„‡–™‡‡Ͷ•ƒ†ͺ•™‹–Šƒƒ˜‡”ƒ‰‡‘ˆ͸•Ǥ 150  †ƒ› –™‘ǡ ‡š–‹ –‹‘ ‹ Ž—†‡† Ͷͺ  –”‹ƒŽ• ȋͳʹ –”‹ƒŽ• ’‡” ǡ Ͷ• †—”ƒ–‹‘Ȍ ƒ† ʹͶ ‡›‡Ǧ

151 ‘˜‡‡–„Ž‘ •ȋͳͲ•†—”ƒ–‹‘ȌǤ‡ΪȋΪ‡›‡Ȍƒ†‘‡ǦȋǦ‡›‡Ȍ™‡”‡ƒŽ™ƒ›•ˆ‘ŽŽ‘™‡†„›ƒ

152 ‡›‡Ǧ‘˜‡‡– „Ž‘  ™Š‹Ž‡ –Š‡ ‘–Š‡” Ϊ ȋڐ‘Ǧ‡›‡Ȍ ƒ† –Š‡ ‘–Š‡” Ǧ ȋǦ‘Ǧ‡›‡Ȍ ™‡”‡ ƒŽ™ƒ›• 153 ˆ‘ŽŽ‘™‡†„›ƒˆ‹šƒ–‹‘„Ž‘ ǤŠ‡  „‡–™‡‡ƒ†‡›‡‘˜‡‡–„Ž‘ ™ƒ•Œ‹––‡”‡†„‡–™‡‡ͲǤͷ• 154 ƒ†ͳǤͷ•™Š‹ Š™ƒ•†‘‡–‘‹‹‹œ‡‡›‡Ǧ‘˜‡‡–ƒ–‹ ‹’ƒ–‹‘†—”‹‰–Š‡’”‡•‡–ƒ–‹‘Ǥ‹–Š 155 –Š‡†—”ƒ–‹‘‘ˆͳͲ•ǡ™‡•–ƒ›‡†‘–Š‡Ž‘™‡”‡†‘ˆ™Šƒ–‹•—•‡†‹–”‡ƒ–‡–ǡ‹™Š‹ Š–Š‡ 156 †—”ƒ–‹‘‘ˆ‡›‡‘˜‡‡–•˜ƒ”‹‡•„‡–™‡‡ͺƒ†ͻ͸•ȋ‡‡ƒ†—‹Œ’‡”•ǡʹͲͳ͵ȌǤŠ‹•ͳͲ•†—”ƒ–‹‘ 157 Ž‹‹–•–Š‡Ž‡‰–Š‘ˆ–Š‡‡š’‡”‹‡–ǡ™Š‹Ž‡•–‹ŽŽ‹ Ž—†‹‰–Š‡’‡ƒ‘ˆ–Š‡Ž‘‘†š›‰‡ƒ–‹‘‡˜‡Ž 158 ‡’‡†‡– ȋȌ ”‡•’‘•‡ ™‹–Š‹ –Š‡ ‡›‡Ǧ‘˜‡‡– „Ž‘ • ȋ ‡‡‰‡” ƒ† ‡••ǡ ʹͲͲʹȌǤ • ‹

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159 ‡š’‡”‹‡– ͳǡ –Š‡ •’‡‡† ‘ˆ –Š‡ ‘˜‹‰ †‘– ™ƒ• ̱ͳ œǡ „ƒ•‡† ‘ ’”‡˜‹‘—• Žƒ„‘”ƒ–‘”› ‘†‡Ž• ‘ˆ 160 ȋ˜ƒ†‡ ‘—–‡–ƒŽǤǡʹͲͳ͵ȌǤŠ‡˜‹•—ƒŽƒ‰Ž‡™ƒ•ƒ’’”‘š‹ƒ–‡Ž›ͳͳιǤŠ‡  ƒˆ–‡”–Š‡‡›‡Ǧ 161 ‘˜‡‡– „Ž‘  ˜ƒ”‹‡† „‡–™‡‡ Ͷ• ƒ† ͺ• ™‹–Š ƒ ƒ˜‡”ƒ‰‡ ‘ˆ ͸•Ǥ  †ƒ› –Š”‡‡ǡ –Š‡ ‡š’‡”‹‡– 162 •–ƒ”–‡†™‹–Šƒ”‡Ǧ‡š–‹ –‹‘•‡••‹‘ȋ”‡Ǧ‡š–‹ –‹‘ͳȌǡ™Š‹ Š‹ Ž—†‡†ʹͶ–”‹ƒŽ•ȋ͸–”‹ƒŽ•’‡”ǡͶ• 163 †—”ƒ–‹‘Ȍ ™‹–Š ƒ   Œ‹––‡”‡† „‡–™‡‡ Ͷ• ƒ† ͺ• ȋƒ˜‡”ƒ‰‡ ‘ˆ ͸•ȌǤ ˆ–‡” –Š‹• •‡••‹‘ –Š‡”‡ ™ƒ• ƒ 164 ”‡‹•–ƒ–‡‡–’”‘ ‡†—”‡ȋ ƒƒ‡”‡–ƒŽǤǡʹͲͳͶȌ ‘•‹•–‹‰‘ˆ͵—Ǧ•‹‰ƒŽ‡†’”‡•‡–ƒ–‹‘•ȋ  ǣ 165 ͳͲ•ȌǤ ‘ŽŽ‘™‹‰–Š‹•ǡ’ƒ”–‹ ‹’ƒ–•—†‡”™‡–ƒ•‡ ‘†”‡Ǧ‡š–‹ –‹‘•‡••‹‘ȋ”‡Ǧ‡š–‹ –‹‘ʹȌǡ™Š‹ Š 166 ‹ Ž—†‡† ʹͶ  –”‹ƒŽ• ȋ͸ –”‹ƒŽ• ’‡” ǡ Ͷ• †—”ƒ–‹‘ȌǤ   ™ƒ• Œ‹––‡”‡† „‡–™‡‡ Ͷ• ƒ† ͺ• ™‹–Š ƒ 167 ƒ˜‡”ƒ‰‡‘ˆ͸•Ǥ‡‡ ‹‰—”‡ʹˆ‘”ƒ‘˜‡”˜‹‡™Ǥ 168  169 Questionnaires and debriefing 170 ƒ”–‹ ‹’ƒ–• ‘’Ž‡–‡†–Š‡‡ ‡’”‡••‹‘ ˜‡–‘”›ȋ Ǣ‡ ‡–ƒŽǤǡͳͻͻ͸Ȍƒ†–Š‡–”ƒ‹–˜‡”•‹‘ 171 ‘ˆ–ƒ–‡Ǧ”ƒ‹–š‹‡–› ˜‡–‘”›ȋ Ǧ–Ǣƒ†‡”Ž‘‡‰ǡͳͻͺͲȌǤ • ‘”‡ƒ„‘˜‡ͳ͵™ƒ•—•‡†–‘ 172 ‡š Ž—†‡’ƒ”–‹ ‹’ƒ–•ˆ”‘–Š‡ƒƒŽ›•‡•ǡ„—–‘‡‘ˆ–Š‡’ƒ”–‹ ‹’ƒ–•Šƒ†ƒ• ‘”‡Š‹‰Š‡”–Šƒ–Š‡ —–Ǧ 173 ‘ˆˆǤ˜‡”ƒ‰‡ • ‘”‡™ƒ•͵Ǥͷȋ”ƒ‰‡ǣͲǦͳͲȌƒ† Ǧ–™ƒ•͵͵Ǥͷȋ”ƒ‰‡ǣʹͷǦͶͺȌǤƒ”–‹ ‹’ƒ–•™‡”‡ 174 †‡„”‹‡ˆ‡†ƒˆ–‡”–Š‡ ‘’Ž‡–‹‘‘ˆ–Š‡‡š’‡”‹‡–ƒ†ƒ•‡†ƒ„‘—––Š‡‹” ‘–‹‰‡ ›‘™Ž‡†‰‡‘ 175 –Š‡‘ —””‡ ‡‘ˆ‡Ž‡ –”‹ ƒŽ•Š‘ •ǡƒ•™‡ŽŽƒ•–Š‡”‡Žƒ–‹‘•Š‹’„‡–™‡‡–Š‡•ƒ†‡›‡Ǧ‘˜‡‡– 176 „Ž‘ •Ǥƒ”–‹ ‹’ƒ–•™‡”‡ˆ—”–Š‡”‘”‡ƒ•‡†ƒ„‘—––Š‡‹”‘™Ž‡†‰‡‘ˆƒ†™Š‡–Š‡”–Š‡›ƒ– 177 •‘‡–‹‡†—”‹‰–Š‡‡š’‡”‹‡––Š‘—‰Š–‘ˆ–Š‡‡š’‡”‹‡–‹–Š‡ ‘–‡š–‘ˆ–”‡ƒ–‡–Ǥ ‹˜‡ 178 ’ƒ”–‹ ‹’ƒ–•”‡’‘”–‡††‘‹‰•‘Ǥ‡–Š‡”‡ˆ‘”‡”‡†‹†–Š‡ƒƒŽ›•‡•‘ˆ–Š‡–™‘”‡Ǧ‡š–‹ –‹‘’Šƒ•‡•‘ 179 †ƒ› ͵ ‡š Ž—†‹‰ –Š‡•‡ ˆ‹˜‡ ’ƒ”–‹ ‹’ƒ–•Ǥ Š‡ ”‡•—Ž–• ƒ† ‘ Ž—•‹‘• ”‡ƒ‹‡† –Š‡ •ƒ‡ ƒ† 180 –Š‡”‡ˆ‘”‡–Š‡”‡•—Ž–•ƒ”‡”‡’‘”–‡†‹ Ž—†‹‰ƒŽŽ’ƒ”–‹ ‹’ƒ–•Ǥ 181  182 Peripheral stimulation 183 Ž‡ –”‹ ƒŽ•Š‘ •™‡”‡†‡Ž‹˜‡”‡†˜‹ƒ–™‘‰Ȁ‰Ž‡Ž‡ –”‘†‡•ƒ––ƒ Š‡†–‘–Š‡†‹•–ƒŽ’ŠƒŽƒ‰‡•‘ˆ–Š‡ 184 •‡ ‘† ƒ† –Š‹”† ˆ‹‰‡”• ‘ˆ –Š‡ ”‹‰Š– ‘” Ž‡ˆ– Šƒ† ȋ ‘—–‡”„ƒŽƒ ‡† „‡–™‡‡ •—„Œ‡ –•Ȍ —•‹‰ ƒ 185  ʹͲͲͲ ȋ‹‘Ǧ”‘–‡ ŠȌ †‡˜‹ ‡Ǥ Š‘  †—”ƒ–‹‘ ™ƒ• ʹͲͲ •ǡ ƒ† ‹–‡•‹–› ˜ƒ”‹‡† ‹ ͳͲ 186 ‹–‡•‹–› •–‡’• „‡–™‡‡ ͲǦͶͲȀͲǦͺͲǤ —”‹‰ ƒ •–ƒ†ƒ”†‹œ‡† •Š‘  ‹–‡•‹–› ƒ†Œ—•–‡– 187 ’”‘ ‡†—”‡ǡ‡ƒ Š’ƒ”–‹ ‹’ƒ–”‡ ‡‹˜‡†ƒ†•—„Œ‡ –‹˜‡Ž›”ƒ–‡†ˆ‹˜‡•Š‘ •ǡƒŽŽ‘™‹‰•Š‘ ‹–‡•‹–›–‘ 188 ‘˜‡”‰‡–‘ƒŽ‡˜‡Ž‡š’‡”‹‡ ‡†ƒ•— ‘ˆ‘”–ƒ„Ž‡ǡ„—–‘–’ƒ‹ˆ—ŽǤŠ‡”‡•—Ž–‹‰ƒ˜‡”ƒ‰‡‹–‡•‹–› 189 •–‡’™ƒ•ͶǤͺȋǣͳǤͺȌ‘ƒ• ƒŽ‡ˆ”‘ͳ–‘ͳͲǤŠ‡‹–‡•‹–›•–‡’™ƒ••‡–‘†ƒ›ͳƒ†”‡ƒ‹‡†–Š‡ 190 •ƒ‡‘†ƒ›͵ˆ‘”–Š‡”‡‹•–ƒ–‡‡–’”‘ ‡†—”‡Ǥ 191

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192 Peripheral measurements 193 Ž‡ –”‘†‡”ƒŽƒ –‹˜‹–›™ƒ•ƒ••‡••‡†—•‹‰–™‘‰Ȁ‰Ž‡Ž‡ –”‘†‡•ƒ––ƒ Š‡†–‘–Š‡†‹•–ƒŽ’ŠƒŽƒ‰‡• 194 ‘ˆ–Š‡ˆ‹”•–ƒ†•‡ ‘†ˆ‹‰‡”•‘ˆ–Š‡Ž‡ˆ–‘””‹‰Š–Šƒ†ȋ ‘—–‡”„ƒŽƒ ‡†„‡–™‡‡•—„Œ‡ –•Ȍ—•‹‰ƒ 195 ”ƒ‹’•›•–‡ƒ†”‡ ‘”†‡†—•‹‰”ƒ‹‹•‹‘‡ ‘”†‡”•‘ˆ–™ƒ”‡ȋ”ƒ‹”‘†— –• „ ǡ 196 —‹ Šǡ ‡”ƒ›ȌǤƒ–ƒ™‡”‡’”‡’”‘ ‡••‡†—•‹‰‹ǦŠ‘—•‡•‘ˆ–™ƒ”‡Ǣ”ƒ†‹‘ˆ”‡“—‡ ›ȋ Ȍƒ”–‹ˆƒ –• 197 ™‡”‡ ”‡‘˜‡† ƒ† ƒ Ž‘™Ǧ’ƒ•• ˆ‹Ž–‡” ™ƒ• ƒ’’Ž‹‡†Ǥ ‹ ‘†— –ƒ ‡ ”‡•’‘•‡• ȋȌ ™‡”‡ 198 ƒ—–‘ƒ–‹ ƒŽŽ› • ‘”‡† ™‹–Š ƒ††‹–‹‘ƒŽ ƒ—ƒŽ •—’‡”˜‹•‹‘ —•‹‰ —–‘‘ƒ–‡ ȋ ”‡‡ ‡– ƒŽǤǡ ʹͲͳͶȌ 199 ‹’Ž‡‡–‡† ‹ ƒ–Žƒ„ ͹ǤͳͶ ȋƒ–Š‘”•ǡ  ǣ̴ͲͲͳ͸ʹʹȌǤ ‡ ‘’–‡† –‘ —•‡ –Š‡ ƒ‰‹–—†‡ 200 ‡–Š‘†ǡ •‹ ‡ ‹– Šƒ• „‡‡ ‘•‹†‡”‡† –Š‡ •–ƒ†ƒ”† ‡–Š‘† ‘ˆ • ‘”‹‰ • ˆ‘” •‡˜‡”ƒŽ †‡ ƒ†‡• 201 ȋ†‡Ž„‡”‰ǡͳͻ͹ʹȌǤƒ’Ž‹–—†‡•ȋ‡ƒ•—”‡†‹P‹‡Ȍ™‡”‡†‡–‡”‹‡†ˆ‘”‡ƒ Š–”‹ƒŽ™‹–Š‹ƒ 202 ‘•‡–Žƒ–‡ ›™‹†‘™„‡–™‡‡ͲǤͷƒ†ͶǤͷ•ƒˆ–‡”•–‹—Ž—•‘•‡–ǡ™‹–Šƒ‹‹—”‹•‡–‹‡‘ˆͲǤͷ• 203 ƒ†ƒƒš‹—”‹•‡–‹‡‘ˆͷ•ƒˆ–‡””‡•’‘•‡‘•‡–Ǥ‡‹ˆ‘” ‡†–”‹ƒŽ•™‡”‡‘‹––‡†ƒ†ƒŽŽ‘–Š‡” 204 ”‡•’‘•‡ ƒ’Ž‹–—†‡• ™‡”‡ •“—ƒ”‡Ǧ”‘‘– –”ƒ•ˆ‘”‡† ’”‹‘” –‘ •–ƒ–‹•–‹ ƒŽ ƒƒŽ›•‹•Ǥ ‡ •—„Œ‡ – ™ƒ• 205 ‘‹––‡†ˆ”‘–Š‡ƒƒŽ›•‡•‘†ƒ›ͳ„‡ ƒ—•‡‘ˆˆƒ‹Ž‡†”‡ ‘”†‹‰•’”‡•—ƒ„Ž›†—‡–‘‘–‹‘‘ˆ 206 –Š‡ Šƒ†Ǥ ‘—” ”‡’‡ƒ–‡†Ǧ‡ƒ•—”‡• • ™‡”‡ ‘†— –‡†ǡ ‘‡ ˆ‘” ‡ƒ Š ‡š’‡”‹‡–ƒŽ ’Šƒ•‡ 207 ȋ “—‹•‹–‹‘ǡ š–‹ –‹‘ǡ ‡Ǧ‡š–‹ –‹‘ͳǡ ƒ† ‡Ǧ‡š–‹ –‹‘ʹȌǤ ƒ Š  ‹ Ž—†‡†  ȋΪǡ ǦȌ 208 ƒ†š–‹ –‹‘ƒ‹’—Žƒ–‹‘ȋ‡›‡ǡ‘Ǧ‡›‡Ȍƒ•™‹–Š‹Ǧ•—„Œ‡ –ˆƒ –‘”•Ǥ—”‹‰–Š‡‡š–‹ –‹‘ƒ†”‡Ǧ 209 ‡š–‹ –‹‘ ’Šƒ•‡•ǡ ƒ ƒ††‹–‹‘ƒŽ ™‹–Š‹Ǧ•—„Œ‡ – ˆƒ –‘” ™ƒ• ‹ Ž—†‡†ǡ ƒ‡Ž› ‹‡ ȋ‡ƒ”Ž›ǡ Žƒ–‡ȌǤ 210 —„•‡“—‡–Ž›ǡ†‹ˆˆ‡”‡–‹ƒŽ™‡”‡ ƒŽ —Žƒ–‡†ȋڏ‹—•ǦȌ–‘–‡•–ˆ‘”†‹ˆˆ‡”‡ ‡•„‡–™‡‡–Š‡ 211 –™‘ ‘†‹–‹‘•ȋ‡›‡‘˜‡‡–ƒ†‘Ǧ‡›‡‘˜‡‡–ȌǤ‘–‡•–ˆ‘”•’‘–ƒ‡‘—•”‡ ‘˜‡”›‘ˆˆ‡ƒ”ǡ 212 –Š‡ †‹ˆˆ‡”‡–‹ƒŽ ”‡•’‘•‡ ‘ –Š‡ Žƒ•– –”‹ƒŽ ‘ˆ ‡š–‹ –‹‘ ™ƒ• •—„–”ƒ –‡† ˆ”‘ –Š‡ ˆ‹”•– †‹ˆˆ‡”‡–‹ƒŽ 213 ”‡•’‘•‡†—”‹‰”‡Ǧ‡š–‹ –‹‘ͳȋ Š‹ŽŽ‡”‡–ƒŽǤǡʹͲͳͲǡʹͲͳ͵ȌǤŠ‡”‡‹•–ƒ–‡‡–”‡ ‘˜‡”›‹†‡š™ƒ• 214 ƒŽ —Žƒ–‡†‹ƒ•‹‹Žƒ”™ƒ›„›•—„–”ƒ –‹‰–Š‡Žƒ•–†‹ˆˆ‡”‡–‹ƒŽ”‡•’‘•‡†—”‹‰”‡Ǧ‡š–‹ –‹‘ͳˆ”‘ 215 –Š‡ ˆ‹”•– †‹ˆˆ‡”‡–‹ƒŽ ”‡•’‘•‡ †—”‹‰ ”‡Ǧ‡š–‹ –‹‘ʹ ȋ Š‹ŽŽ‡” ‡– ƒŽǤǡ ʹͲͳͲǡ ʹͲͳ͵ȌǤ ‘” –Š‡ 216 •’‘–ƒ‡‘—•”‡ ‘˜‡”›‹†‡šƒ†”‡‹•–ƒ–‡‡–”‡ ‘˜‡”›‹†‡šƒƒŽ›•‡•ǡ™‡ ‘˜ƒ”‹‡†–Š‡‘”†‡”‘ˆ

217 –Š‡ΪȋΪ‡›‡‘”ڐ‘Ǧ‡›‡Ȍ’”‡•‡–ƒ–‹‘Ǥƒ•–Ž›ǡ–Š‡ƒ‘—–‘ˆƒ›‰†ƒŽƒ•—’’”‡••‹‘–Šƒ–‘ —””‡† 218 ‘†ƒ›ʹ†—”‹‰–Š‡‡›‡Ǧ‘˜‡‡–„Ž‘ •™ƒ•ƒ††‡†ƒ•ƒ ‘˜ƒ”‹ƒ–‡–‘–Š‡”‡ ‘˜‡”›‹†‡šƒƒŽ›•‡• 219 ‘†ƒ›͵–‘–‡•–™Š‡–Š‡”ƒ›‰†ƒŽƒ†‡ƒ –‹˜ƒ–‹‘’”‡†‹ –‡†ˆ‡ƒ””‡ ‘˜‡”›Ǥ 220 ›‡ –”ƒ ‹‰ ™ƒ• ”‡ ‘”†‡† —•‹‰ ƒ Ǧ ‘’ƒ–‹„Ž‡ ‡›‡Ǧ–”ƒ ‹‰ •›•–‡ ȋ›‡ ”ƒ Ǧ 221 ƒ‡”ƒ —‹–ǡ  ǡ ‡•‘‘–‘”‹  •–”—‡–•ȌǤ ƒ–ƒ ™‡”‡ ’”‡’”‘ ‡••‡† —•‹‰ ‹ǦŠ‘—•‡ •‘ˆ–™ƒ”‡ 222 ȋ ‡”ƒ•‡–ƒŽǤǡʹͲͳ͵Ȍ‹’Ž‡‡–‡†‹ƒ–Žƒ„͹ǤͳͶȋƒ–Š‘”•ǡ ǣ̴ͲͲͳ͸ʹʹȌǤŽ‹•™‡”‡ 223 ”‡‘˜‡†ˆ”‘–Š‡•‹‰ƒŽ—•‹‰Ž‹‡ƒ”‹–‡”’‘Žƒ–‹‘Ǥ›‡Ǧ–”ƒ ‹‰†ƒ–ƒ†—”‹‰–Š‡‡›‡Ǧ‘˜‡‡–•

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224 „Ž‘ • ™‡”‡ ‘”ƒŽ‹œ‡† „ƒ•‡† ‘ ƒ ƒŽ‹„”ƒ–‹‘ ƒ– –Š‡ •–ƒ”– ‘ˆ –Š‡ ‡š’‡”‹‡–Ǥ ‹•—ƒŽ ‹•’‡ –‹‘ 225 ”‡˜‡ƒŽ‡†’ƒ”–‹ ‹’ƒ–• ‘’Ž‹‡†™‹–Š–Š‡‹•–”— –‹‘•‘ˆ–Š‡–ƒ•Ǥ‡‡ ‹‰—”‡ͶǤ 226 227 Physiological noise correction 228 ‹‰‡”’—Ž•‡™ƒ•”‡ ‘”†‡†—•‹‰ƒ’—Ž•‡‘š‹‡–‡”ƒˆˆ‹š‡†–‘–Š‡–Š‹”†ˆ‹‰‡”‘ˆ–Š‡Ž‡ˆ–‘””‹‰Š–Šƒ† 229 ȋ ‘—–‡”„ƒŽƒ ‡† „‡–™‡‡ •—„Œ‡ –•ȌǤ ‡•’‹”ƒ–‹‘ ™ƒ• ‡ƒ•—”‡† —•‹‰ ƒ ”‡•’‹”ƒ–‹‘ „‡Ž– ’Žƒ ‡† 230 ƒ”‘—† –Š‡ ’ƒ”–‹ ‹’ƒ–ǯ• ƒ„†‘‡Ǥ —Ž•‡ ƒ† ”‡•’‹”ƒ–‹‘ ‡ƒ•—”‡• ™‡”‡ —•‡† ˆ‘” ”‡–”‘•’‡ –‹˜‡ 231 ‹ƒ‰‡Ǧ„ƒ•‡† ‘””‡ –‹‘ȋ Ȍ‘ˆ’Š›•‹‘Ž‘‰‹ ƒŽ‘‹•‡ƒ”–‹ˆƒ –•‹Ǧˆ †ƒ–ƒȋ Ž‘˜‡”‡– 232 ƒŽǤǡ ʹͲͲͲȌǤ ƒ™ ’—Ž•‡ ƒ† ”‡•’‹”ƒ–‘”› †ƒ–ƒ ™‡”‡ ’”‘ ‡••‡† ‘ˆˆŽ‹‡ —•‹‰ ‹ǦŠ‘—•‡ •‘ˆ–™ƒ”‡ ˆ‘” 233 ‹–‡”ƒ –‹˜‡ ˜‹•—ƒŽ ƒ”–‹ˆƒ – ‘””‡ –‹‘ ƒ† ’‡ƒ †‡–‡ –‹‘ǡ ƒ† ™‡”‡ —•‡† –‘ •’‡ ‹ˆ› ˆ‹ˆ–ŠǦ‘”†‡” 234 ‘—”‹‡”‘†‡Ž•‘ˆ–Š‡ ƒ”†‹ƒ ƒ†”‡•’‹”ƒ–‘”›’Šƒ•‡Ǧ”‡Žƒ–‡†‘†—Žƒ–‹‘‘ˆ–Š‡•‹‰ƒŽȋƒ 235 ——”‡‡–ƒŽǤǡʹͲͲͻȌǡ›‹‡Ž†‹‰ͳͲ—‹•ƒ ‡”‡‰”‡••‘”•ˆ‘” ƒ”†‹ƒ ‘‹•‡ƒ†ͳͲˆ‘””‡•’‹”ƒ–‘”›‘‹•‡Ǥ 236 ††‹–‹‘ƒŽ ”‡‰”‡••‘”• ™‡”‡ ƒŽ —Žƒ–‡† ˆ‘” Š‡ƒ”– ”ƒ–‡ ˆ”‡“—‡ ›ǡ Š‡ƒ”– ”ƒ–‡ ˜ƒ”‹ƒ„‹Ž‹–›ǡ ȋ”ƒ™Ȍ 237 ƒ„†‘‹ƒŽ ‹” —ˆ‡”‡ ‡ǡ ”‡•’‹”ƒ–‘”› ˆ”‡“—‡ ›ǡ ”‡•’‹”ƒ–‘”› ƒ’Ž‹–—†‡ǡ ƒ† ”‡•’‹”ƒ–‹‘ ˜‘Ž—‡ 238 ’‡”—‹––‹‡ȋ‹”‡–ƒŽǤǡʹͲͲ͸Ȍǡ›‹‡Ž†‹‰ƒ–‘–ƒŽ‘ˆʹ͸ ”‡‰”‡••‘”•Ǥ 239  240 MRI data acquisition and multi-echo weighting 241   • ƒ• ™‡”‡ ƒ “—‹”‡† —•‹‰ ƒ ‹‡‡• ȋ”Žƒ‰‡ǡ ‡”ƒ›Ȍ   ˜ƒ–‘ ͳǤͷ  242 • ƒ‡”ǤʹȗǦ™‡‹‰Š–‡†„Ž‘‘†‘š›‰‡ƒ–‹‘Ž‡˜‡ŽǦ†‡’‡†‡–ȋȌ‹ƒ‰‡•™‡”‡”‡ ‘”†‡†—•‹‰ƒ 243 —•–‘‹œ‡†—Ž–‹Ǧ‡ Š‘ •‡“—‡ ‡™‹–Šƒ• ‡†‹‰•Ž‹ ‡ƒ “—‹•‹–‹‘ȋ͵ͷƒš‹ƒŽ•Ž‹ ‡•ǢǡʹǤʹ•Ǣǡ 244 ͻǤͶ•ǡʹͳ•ǡ͵͵•ǡͶͶ•ǡƒ†ͷ͸•Ǣ ‡‡”ƒŽ‹œ‡†—–‘ ƒŽ‹„”ƒ–‹‰ƒ”–‹ƒŽŽ›ƒ”ƒŽŽ‡Ž “—‹•‹–‹‘• 245 ȋ Ǣ ”‹•™‘Ž†‡–ƒŽǤǡʹͲͲʹȌƒ ‡Ž‡”ƒ–‹‘ˆƒ –‘”͵ǢˆŽ‹’ƒ‰Ž‡ǡͻͲιǢ•Ž‹ ‡ƒ–”‹š•‹œ‡ǡ͸Ͷš͸ͶǢ•Ž‹ ‡ 246 –Š‹ ‡••ǡ͵ǤͲǢ•Ž‹ ‡‰ƒ’ǡͲǤͷͳǢ ǡʹͳʹšʹͳʹǢ„ƒ†™‹†–Šǣʹ͸ͲͶ œȀ’šǢ‡ Š‘•’ƒ ‹‰ǣ 247 ͲǤͶͻ •ȌǤ ‘ ƒ ‘—– ˆ‘” ”‡‰‹‘ƒŽ ˜ƒ”‹ƒ–‹‘ ‹ •—• ‡’–‹„‹Ž‹–›Ǧ‹†— ‡† •‹‰ƒŽ †”‘’‘—–ǡ ˜‘š‡ŽǦ™‹•‡ 248 ™‡‹‰Š–‡† •—• ‘ˆ ƒŽŽ ‡ Š‘‡• ™‡”‡ ƒŽ —Žƒ–‡† „ƒ•‡† ‘ Ž‘ ƒŽ ‘–”ƒ•–Ǧ–‘Ǧ‘‹•‡ ”ƒ–‹‘ ȋ‘•‡” ‡– ƒŽǤǡ 249 ʹͲͲ͸ȌǤ•–”— –—”ƒŽ‹ƒ‰‡ȋͳ‹•‘–”‘’‹ Ȍ™ƒ•ƒ “—‹”‡†—•‹‰ƒͳǦ™‡‹‰Š–‡†͵ƒ‰‡–‹œƒ–‹‘Ǧ 250 ’”‡’ƒ”‡†”ƒ’‹†‰”ƒ†‹‡–Ǧ‡ Š‘•‡“—‡ ‡ȋǦ ǢǡʹǤ͹͵•ǢǡʹǤͻͷ•ǢˆŽ‹’ƒ‰Ž‡ǡ͹ιǢ ǡʹͷ͸š 251 ʹͷ͸šͳ͹͸ȌǤ 252  253 MRI data preprocessing in standard stereotactic space and statistical analyses 254  †ƒ–ƒ™‡”‡’”‡Ǧ’”‘ ‡••‡†‹•–ƒ†ƒ”†•–‡”‡‘–ƒ –‹ ȋ ͳͷʹȌ•’ƒ ‡ˆ‘”–Š‡’—”’‘•‡‘ˆ™Š‘Ž‡Ǧ 255 „”ƒ‹‰”‘—’ƒƒŽ›•‡•Ǥ—–—ƒŽ‹ˆ‘”ƒ–‹‘ƒš‹‹œƒ–‹‘Ǧ„ƒ•‡†”‹‰‹†Ǧ„‘†›”‡‰‹•–”ƒ–‹‘™ƒ•—•‡†–‘ 256 ”‡‰‹•–‡” •–”— –—”ƒŽ ƒ† ˆ— –‹‘ƒŽ ‹ƒ‰‡•Ǥ — –‹‘ƒŽ ‹ƒ‰‡• ™‡”‡ ‘–‹‘ ‘””‡ –‡† —•‹‰ ”‹‰‹†Ǧ

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257 „‘†›–”ƒ•ˆ‘”ƒ–‹‘•Ǥ–”— –—”ƒŽ‹ƒ‰‡•™‡”‡•‡‰‡–‡†‹–‘‰”ƒ›ƒ––‡”ǡ™Š‹–‡ƒ––‡”ǡƒ†  258 ‹ƒ‰‡• —•‹‰ ƒ —‹ˆ‹‡† ’”‘„ƒ„‹Ž‹•–‹  –‡’Žƒ–‡ ”‡‰‹•–”ƒ–‹‘ ƒ† –‹••—‡ Žƒ••‹ˆ‹ ƒ–‹‘ ‡–Š‘† 259 ȋ•Š„—”‡” ƒ† ”‹•–‘ǡ ʹͲͲͷȌǤ ‹••—‡ ‹ƒ‰‡• ™‡”‡ –Š‡ ”‡‰‹•–‡”‡† ™‹–Š •‹–‡Ǧ•’‡ ‹ˆ‹  –‹••—‡ 260 –‡’Žƒ–‡•—•‹‰ȋ•Š„—”‡”ǡʹͲͲ͹Ȍǡƒ†”‡‰‹•–‡”‡†ȋ—•‹‰ƒƒˆˆ‹‡–”ƒ•ˆ‘”ƒ–‹‘Ȍ™‹–Š 261 –Š‡  ͳͷʹ –‡’Žƒ–‡ ‹ Ž—†‡† ‹ ͺ ȋ ǣ̴ͲͲ͹Ͳ͵͹ȌǤ †‡–‹ ƒŽ –”ƒ•ˆ‘”ƒ–‹‘• ™‡”‡ 262 ƒ’’Ž‹‡†–‘ƒŽŽˆ— –‹‘ƒŽ‹ƒ‰‡•ǡ™Š‹ Š™‡”‡”‡•Ž‹ ‡†‹–‘ʹ‹•‘–”‘’‹ ˜‘š‡Ž•ƒ†•‘‘–Š‡†™‹–Š 263 ƒ͸   ƒ—••‹ƒ‡”‡ŽǤ 264 ‡ ”‡ƒ–‡† ˆ‘—” ˆ‹”•–ǦŽ‡˜‡Ž ‘†‡Ž• ˆ‘” ƒŽŽ •–ƒ‰‡• ‘ˆ –Š‡ ‡š’‡”‹‡– ȋ‹Ǥ‡Ǥǡ  “—‹•‹–‹‘ǡ 265 š–‹ –‹‘ǡ‡Ǧ‡ –‹ –‹‘ͳƒ†‡Ǧ‡š–‹ –‹‘ʹȌǤ‡•’‘•‡•–‘–Š‡•™‡”‡‘†‡Ž‡†—•‹‰„‘šǦ ƒ” 266 ”‡‰”‡••‘”• ȋ†—”ƒ–‹‘ ‘ˆ ͷ•ȌǤ —”‹‰ –Š‡ ƒ “—‹•‹–‹‘ ’Šƒ•‡ǡ ƒ††‹–‹‘ƒŽ ”‡‰”‡••‘”• ‹ Ž—†‡† –Š‡ 267 ”‡‹ˆ‘” ‡†Ϊ•ȋ†—”ƒ–‹‘‘ˆͷ•Ȍƒ†–Š‡•Š‘ ™Š‹ Š™ƒ•‘†‡Ž‡†ƒ•ƒ•–‹ ˆ— –‹‘Ǥ—”‹‰–Š‡ 268 ‡š–‹ –‹‘’Šƒ•‡ǡ”‡•’‘•‡•–‘–Š‡‡›‡Ǧ‘˜‡‡–„Ž‘ •™‡”‡ ‘†‡Ž‡† —•‹‰ „‘šǦ ƒ””‡‰”‡••‘”• 269 ™‹–Š ƒ †—”ƒ–‹‘ ‘ˆ ͳͲ•Ǥ ‡‰”‡••‘”• ™‡”‡ –‡’‘”ƒŽŽ› ‘˜‘Ž˜‡† ™‹–Š –Š‡ ƒ‘‹ ƒŽ Š‡‘†›ƒ‹  270 ”‡•’‘•‡ ˆ— –‹‘ ȋ  Ȍ ‹ Ž—†‡† ‹ ͺ ȋ ǣ̴ͲͲ͹Ͳ͵͹ȌǤ ††‹–‹‘ƒŽŽ›ǡ •‹š ‘˜‡‡– 271 ’ƒ”ƒ‡–‡””‡‰”‡••‘”•ȋ͵–”ƒ•Žƒ–‹‘•ǡ͵”‘–ƒ–‹‘•Ȍ†‡”‹˜‡†ˆ”‘”‹‰‹†Ǧ„‘†›‘–‹‘ ‘””‡ –‹‘ǡŠ‹‰ŠǦ 272 ’ƒ•• ˆ‹Ž–‡”‹‰ ȋͳȀͳʹͺ œ —–Ǧ‘ˆˆȌǡ ƒ† ȋͳȌ •‡”‹ƒŽ ‘””‡Žƒ–‹‘ ‘””‡ –‹‘• ™‡”‡ ‹ Ž—†‡† ‹ –Š‡ 273 ‘†‡ŽǤ‹‰Ž‡Ǧ•—„Œ‡ – ‘–”ƒ•–ƒ’•™‡”‡‡–‡”‡†‹–‘•‡ ‘†ǦŽ‡˜‡Ž‘‡Ǧ•ƒ’Ž‡–Ǧ–‡•–•Ǥ 274 Ž–Š‘—‰Š ƒ   ƒƒŽ›•‹• ‹ ‡š’‡”‹‡–• ™‹–Š ‡˜‡–Ǧ”‡Žƒ–‡† †‡•‹‰• ‹• ‘”‡ †‹ˆˆ‹ —Ž– –‘ 275 ‹–‡”’”‡–ȋǯ‡‹ŽŽ›‡–ƒŽǤǡʹͲͳʹȌǡ™‡†‹†’‡”ˆ‘”ƒ ƒƒŽ›•‹•ǡ•‹‹Žƒ”–‘š’‡”‹‡–ͳǡ™‹–Š–Š‡ 276 ƒ›‰†ƒŽƒ ƒ• ƒ •‡‡† ”‡‰‹‘ ‹ š’‡”‹‡– ʹ ƒ• ™‡ŽŽǤ ‡ †‹† ‘– ˆ‹† ƒ› •–ƒ–‹•–‹ ƒŽŽ› •‹‰‹ˆ‹ ƒ– 277 ‘‡ –‹˜‹–› Šƒ‰‡•Ǥ•ƒ‡š–”ƒ Š‡ ǡ™‡—•‡†–Š‡  ƒ•ƒ•‡‡†ƒ•™‡ŽŽǡ„—–ƒŽ•‘Š‡”‡™‡†‹†‘– 278 ‘„•‡”˜‡ ”‡Ž‹ƒ„Ž‡ ‘‡ –‹˜‹–› †‹ˆˆ‡”‡ ‡•Ǥ ††‹–‹‘ƒŽŽ›ǡ ™‡ ƒ’’Ž‹‡† ƒ „‡–ƒ •‡”‹‡• ‘””‡Žƒ–‹‘ 279 ȋ‹••ƒ‡–ƒŽǤǡʹͲͲͶȌ™‹–Š–Š‡ƒ›‰†ƒŽƒƒ•ƒ•‡‡†”‡‰‹‘Ǥ ‘™‡˜‡”ǡ™‡†‹†‘–ˆ‹†ƒ›•–ƒ–‹•–‹ ƒŽŽ› 280 •‹‰‹ˆ‹ ƒ– ‘‡ –‹˜‹–› Šƒ‰‡•—•‹‰–Š‹•‡–Š‘†‡‹–Š‡”ǤŠ‡”‡ƒ”‡–™‘’‘••‹„Ž‡‡š’Žƒƒ–‹‘•ˆ‘” 281 –Š‹•—ŽŽˆ‹†‹‰Ǥ ‹”•–ǡ–Š‡•Šƒ’‡ƒ†ƒ••—’–‹‘•”‡‰ƒ”†‹‰–Š‡  ƒ”‡‘”‡‹’‘”–ƒ–‹‡˜‡–Ǧ 282 ”‡Žƒ–‡††‡•‹‰•–Šƒ‹„Ž‘ ‡††‡•‹‰• ȋ ‹–‡Žƒ‡–ƒŽǤǡʹͲͲ͵ȌǤ‡ ‘†ǡ‹–Š‡’‡”‹‘†™Š‡”‡‹–‹• 283 ’‘••‹„Ž‡ –‘ Ž‘‘ ƒ– –ƒ•Ǧ†”‹˜‡ ‘‡ –‹˜‹–› Šƒ‰‡• ‹ ‡˜‡–Ǧ”‡Žƒ–‡† †‡•‹‰• ƒ”‡ •Š‘”–‡” –Šƒ ‹ 284 „Ž‘ ‡††‡•‹‰•ǤŠ—•ǡ ƒƒŽ›•‡•Šƒ˜‡Ž‡••’‘™‡”–Šƒ–ƒ•ƒ –‹˜‹–›ƒƒŽ›•‡•ƒ†–‡†–‘Šƒ˜‡ 285 •ƒŽŽ‡”‡ˆˆ‡ –•‹œ‡•–Šƒ„Ž‘ ‡††‡•‹‰•Ǥ 286 ƒ•‡†‘‘—”’”‹‘”‹Š›’‘–Š‡•‡•ǡ”‡•—Ž–•ˆ‘”–Š‡ƒ›‰†ƒŽƒƒ††‘”•ƒŽƒ–‡”‹‘” ‹‰—Žƒ–‡ 287 ‘”–‡šȋ†Ȍ™‡”‡ ‘””‡ –‡†ˆ‘””‡†— ‡†•‡ƒ” Š˜‘Ž—‡•—•‹‰•ƒŽŽ˜‘Ž—‡ ‘””‡ –‹‘•ȋȌ 288 ƒ†™‡”‡ˆƒ‹Ž›Ǧ™‹•‡‡””‘”ȋ Ȍ ‘””‡ –‡†—•‹‰˜‘š‡ŽǦŽ‡˜‡Ž•–ƒ–‹•–‹ •Ǥ‘ˆ–Š‡ƒ›‰†ƒŽƒ™ƒ• 289 „ƒ•‡†‘ƒ‰”‘—’ƒ•–Šƒ–™ƒ• ”‡ƒ–‡†„›ƒ˜‡”ƒ‰‹‰‹†‹˜‹†—ƒŽƒ›‰†ƒŽƒ•‡‰‡–ƒ–‹‘•ȋαʹͶȌ‘ˆ

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290 ͳǦ™‡‹‰Š–‡†‹ƒ‰‡•ȋ—•‹‰  ǡ ǣ̴ͲͲʹͺʹ͵ǡ•‡‡ 291 Š––’•ǣȀȀˆ•ŽǤˆ”‹„Ǥ‘šǤƒ Ǥ—Ȁˆ•ŽȀˆ•Ž™‹‹Ȁ Ȍǡ™Š‹ Š™‡”‡™ƒ”’‡†‹–‘ •’ƒ ‡—•‹‰Ǥ 292 Š‡†™ƒ•†‡ˆ‹‡†„ƒ•‡†‘ƒˆ— –‹‘ƒŽ ƒ–Žƒ•ȋŠ‹”‡”‡–ƒŽǤǡʹͲͳʹȌǤ 293 294 MRI data preprocessing of the extinction session in native space and statistical analyses 295 ‘”–Š‡’”‹ƒ”›ˆ ƒƒŽ›•‹•ȋƒ›‰†ƒŽƒ”‡•’‘•‡†—”‹‰‡›‡‘˜‡‡–•Ȍǡ™‡’”‡’”‘ ‡••‡†  296 †ƒ–ƒ †—”‹‰ ‡š–‹ –‹‘ ‹ ƒ–‹˜‡ •’ƒ ‡ ȋi.e.ǡ ™‹–Š‘—– •–‡”‡‘–ƒ –‹  ‘”ƒŽ‹œƒ–‹‘Ȍ —•‹‰ ͺ 297 ȋ ǣ̴ͲͲ͹Ͳ͵͹ǡ Š––’ǣȀȀ™™™Ǥˆ‹ŽǤ‹‘Ǥ— ŽǤƒ Ǥ—Ȁ•’Ǣ ‡ŽŽ ‘‡ ‡’ƒ”–‡– ‘ˆ ƒ‰‹‰ 298 ‡—”‘• ‹‡ ‡ǡ‘†‘ǡȌǤ‹ ‡–Š‡”‡•—Ž–•ˆ”‘š’‡”‹‡–ͳ•Š‘™‡†–Š‡•—’’”‡••‹‘™ƒ•‘– 299 •’‡ ‹ˆ‹ –‘–Š‡ƒ›‰†ƒŽƒǡ™‡‘’–‡†ˆ‘”–Š‹•‘”‡•’‡ ‹ˆ‹ ƒƒŽ›•‹•–‘ƒ‡•—”‡–Šƒ––Š‡ƒ›‰†ƒŽƒ 300 ‡ˆˆ‡ –• ƒ”‡ ‘–ǡ ˆ‘” ‡šƒ’Ž‡ǡ †—‡ –‘ •‹‰ƒŽ „Ž—””‹‰ ˆ”‘ –Š‡ Š‹’’‘ ƒ’—• ‹–‘ –Š‡ ƒ›‰†ƒŽƒǤ 301 ††‹–‹‘ƒŽŽ›ǡ–Š‹•ƒƒŽ›•‹•ƒŽŽ‘™‡†—•–‘‡š–”ƒ –ƒƒ˜‡”ƒ‰‡†–‹‡ ‘—”•‡‘ˆ–Š‡ƒ›‰†ƒŽƒ•‹‰ƒŽǤ 302 ŽŽ ˆ— –‹‘ƒŽ • ƒ• ™‡”‡ ‘Ǧ”‡‰‹•–‡”‡† ™‹–Š •–”— –—”ƒŽ • ƒ• —•‹‰ —–—ƒŽ ‹ˆ‘”ƒ–‹‘ 303 ƒš‹‹œƒ–‹‘ǤŠ‡ƒ›‰†ƒŽƒ™ƒ•‹†‹˜‹†—ƒŽŽ›†‡ˆ‹‡†‹ƒ–‹˜‡•’ƒ ‡—•‹‰ƒ—–‘ƒ–‡†ƒƒ–‘‹ ƒŽ 304 •‡‰‡–ƒ–‹‘ ‘ˆ ͳǦ™‡‹‰Š–‡† ‹ƒ‰‡• —•‹‰   ȋ ǣ̴ͲͲʹͺʹ͵ǡ •‡‡ 305 Š––’•ǣȀȀˆ•ŽǤˆ”‹„Ǥ‘šǤƒ Ǥ—Ȁˆ•ŽȀˆ•Ž™‹‹Ȁ ȌǤ Š‡ ƒ›‰†ƒŽƒ •‡‰‡–ƒ–‹‘• —†‡”™‡– ˜‹•—ƒŽ 306 ‹•’‡ –‹‘Ǥ ‡ ‘„•‡”˜‡† –Šƒ– •‘‡ ƒ›‰†ƒŽƒ ˜‘š‡Ž• ˆ‘” •‘‡ ’ƒ”–‹ ‹’ƒ–• ™‡”‡ ‘– ’ƒ”– ‘ˆ –Š‡ 307 ƒ•ǡ ‹†‹ ƒ–‹‰ –Šƒ– –Š‡   •‡‰‡–ƒ–‹‘ ™ƒ• ”‡Žƒ–‹˜‡Ž› ‘•‡”˜ƒ–‹˜‡ ȋe.g., ‹ ƒ–‡”‹‘” 308 ’ƒ”–•ǡ•‡‡‘‘”‡‡–ƒŽǤǡʹͲͳͶȌǤ ‘”‡ƒ Š’ƒ”–‹ ‹’ƒ–ǡ™‡ƒ†‡•—”‡–Šƒ––Š‡ƒ›‰†ƒŽƒƒ•†‹†‘– 309 ‘–ƒ‹˜‘š‡Ž•–Šƒ–™‡”‡’ƒ”–‘ˆǡ‘”‡š–‡†‡†‹–‘ǡ–Š‡Š‹’’‘ ƒ’—•Ǥ‡–Š‡”‡ˆ‘”‡ ‘ Ž—†‡–Šƒ– 310 ‘—”ƒ›‰†ƒŽƒƒ••ƒ”‡”‡’”‡•‡–ƒ–‹˜‡‘ˆƒ›‰†ƒŽƒ˜‘Ž—‡ǡƒ†–Šƒ––Š‡•—’’”‡••‹‘‡ˆˆ‡ – ƒ„‡ 311 ƒ••‹‰‡† –‘ –Š‡ ƒ›‰†ƒŽƒǤ Š‡ ˜‡–”ƒŽ ‡†‹ƒŽ ’”‡ˆ”‘–ƒŽ ‘”–‡š ȋ˜ Ȍ ™ƒ• †‡ˆ‹‡† ƒ• ͳͲ  312 •’Š‡”‡ƒ”‘—†–Š‡ͲͶͲǦ͵ ‘‘”†‹ƒ–‡„ƒ•‡†‘ƒ’”‡˜‹‘—••–—†›ȋ Š‹ŽŽ‡”ƒ†‡Ž‰ƒ†‘ǡʹͲͳͲȌǤŠ‡ 313 ”‘–ƒŽ›‡ ‹‡Ž†•ȋ  Ȍ™‡”‡†‡ˆ‹‡†„ƒ•‡†‘ƒͷ•’Š‡”‡ƒ”‘—†–Š‡ ’‡ƒ ‘‘”†‹ƒ–‡• 314 ”‡’‘”–‡†‹ƒ‡–ƒǦƒƒŽ›•‹•ȋ ƒƒ†ƒ”‡–ƒŽǤǡʹͲͳ͵ȌǤ—„•‡“—‡–Ž›ǡ–Š‡  ƒ••™‡”‡–”ƒ•ˆ‡””‡† 315 „ƒ ‹–‘ƒ–‹˜‡•’ƒ ‡ˆ‘”‡ƒ Š‹†‹˜‹†—ƒŽ—•‹‰–Š‡”‡˜‡”•‡†•’ƒ–‹ƒŽ‘”ƒŽ‹œƒ–‹‘’ƒ”ƒ‡–‡”•Ǥ 316 ‘”•–ƒ–‹•–‹ ƒŽƒƒŽ›•‡•ǡ”‡•’‘•‡•–‘–Š‡‡›‡Ǧ‘˜‡‡–ƒ†‘Ǧ‡›‡‘˜‡‡–„Ž‘ •™‡”‡ 317 ‡•–‹ƒ–‡† —•‹‰ ƒ ˆ‹‹–‡ ‹’—Ž•‡ ”‡•’‘•‡ ȋ Ȍ ‘†‡Ž ™Š‹ Š ‹ Ž—†‡† ͻ –‹‡ „‹• ȋ α ʹǤʹ •Ȍ 318 •–ƒ”–‹‰ ‘‡ –‹‡Ǧ„‹ „‡ˆ‘”‡ –Š‡ ‘•‡– ‘ˆ –Š‡  ȋǦʹǤʹ •Ȍ ƒ† ‡†‹‰ ‘‡ –‹‡ „‹ ƒˆ–‡” –Š‡ ‡›‡Ǧ 319 ‘˜‡‡–„Ž‘ •ȋͳ͹Ǥ͸•ȌǤŠ‡”‡ˆ‘”‡ǡ„‹—„‡”•ͷ—–‹Žͺȋ͸Ǥ͸•ȂͳͷǤͶ•ȌƒŽ™ƒ›•ˆ‡ŽŽ™‹–Š‹–Š‡ 320 ‡›‡Ǧ‘˜‡‡–„Ž‘ •Ǥ ‘”–Š‡‘Ǧ‡›‡‘˜‡‡–„Ž‘ ǡ–Š‡•ƒ‡–‹‡ˆ”ƒ‡™ƒ•—•‡†ǤŠ‹•ˆ‹”•–Ǧ 321 Ž‡˜‡Ž ‘†‡Ž ƒ‡• ‘ ƒ••—’–‹‘• ”‡‰ƒ”†‹‰ –Š‡   •Šƒ’‡ ƒ† ›‹‡Ž†• ‹†‡’‡†‡– ”‡•’‘•‡ 322 ‡•–‹ƒ–‡• ˆ‘” ƒŽŽ ͻ –‹‡ „‹•ǡ ™Š‹ Š ƒ‡• ‹– ’‘••‹„Ž‡ –‘ ‹˜‡•–‹‰ƒ–‡ –Š‡ –‹‡ ‘—”•‡ ‘ˆ –Š‡

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323 ”‡•’‘•‡•Ǥ Š‡ ˆ‹”•–ǦŽ‡˜‡Ž ‘†‡Ž• ƒ††‹–‹‘ƒŽŽ› ‹ Ž—†‡† •‹š ‘˜‡‡– ’ƒ”ƒ‡–‡” ”‡‰”‡••‘”• ȋ͵ 324 –”ƒ•Žƒ–‹‘•ǡ͵”‘–ƒ–‹‘•Ȍ†‡”‹˜‡†ˆ”‘”‹‰‹†Ǧ„‘†›‘–‹‘ ‘””‡ –‹‘ǡʹ͸ ’Š›•‹‘Ž‘‰‹ ƒŽ 325 ‘‹•‡”‡‰”‡••‘”•ȋ•‡‡ƒ„‘˜‡ȌǡŠ‹‰ŠǦ’ƒ••ˆ‹Ž–‡”‹‰ȋͳȀͳʹͺ œ —–Ǧ‘ˆˆȌǡƒ†ȋͳȌ•‡”‹ƒŽ ‘””‡Žƒ–‹‘• 326 ‘””‡ –‹‘Ǥ‡‡š–”ƒ –‡†–Š‡ƒ˜‡”ƒ‰‡„‡–ƒ™‡‹‰Š–•™‹–Š‹–Š‡ƒ›‰†ƒŽƒƒ†  ˆ‘”‡ƒ Š–‹‡„‹ 327 ƒ† ‡ƒ Š Ǥ  ”‡’‡ƒ–‡†Ǧ‡ƒ•—”‡•  ™ƒ• ‘†— –‡† ˆ‘” ‡ƒ Š ”‡‰‹‘ ™‹–Š  ȋΪǡ ǦȌǡ 328 š–‹ –‹‘ƒ‹’—Žƒ–‹‘ȋ‡›‡ǡ‘Ǧ‡›‡Ȍƒ†‹‡„‹ȋͷǦͺȌƒ•™‹–Š‹Ǧ•—„Œ‡ –ˆƒ –‘”•Ǥ‡‡ ‹‰—”‡ͶǤ 329 330 Statistical analysis 331 ƒ’Ž‡ •‹œ‡ ‘ˆ ʹͶ ™ƒ• †‡–‡”‹‡† „ƒ•‡† ƒ ’‘‘Ž‡† ‡ˆˆ‡ – •‹œ‡ ˆ”‘ ˆ‘—” •–—†‹‡• –Šƒ– Šƒ˜‡ —•‡† 332 •‹‹Žƒ”’ƒ”ƒ†‹‰•ȋ Š‹ŽŽ‡”‡–ƒŽǤǡʹͲͳͲǡʹͲͳ͵Ǣ‰”‡‡–ƒŽǤǡʹͲͳʹǢ‹†–ƒ†‘‡–‡”ǡʹͲͳ͵ȌǡƒŽ’ŠƒαǤͲͷǡ ଶ 333 ƒ† ͳǦ„‡–ƒαǤͺͲǤ ƒ”–‹ƒŽ ‡–ƒǦ•“—ƒ”‡† ȋߟ௣Ȍ ‘” ‘Š‡ǯ• † ‡ˆˆ‡ – •‹œ‡ ‡•–‹ƒ–‡• ƒ”‡ ”‡’‘”–‡† ˆ‘” ƒŽŽ 334 ”‡Ž‡˜ƒ––‡•–•ǤŽ’Šƒ™ƒ••‡–ƒ–ǤͲͷ–Š”‘—‰Š‘—–ƒ†–™‘Ǧ–ƒ‹Ž‡†–Ǧ–‡•–•™‡”‡ ‘†— –‡†—Ž‡•••–ƒ–‡† 335 ‘–Š‡”™‹•‡Ǥ –ƒ–‹•–‹ ƒŽ ƒƒŽ›•‡• ™‡”‡ ’‡”ˆ‘”‡† —•‹‰  ͳͻ ȋ ǣ̴ͲͲʹͺ͸ͷǡ  ‘”’ǡ 336 ”‘ǡ‡™‘”Ȍǡƒ–Žƒ„͹ǤͳͶȋƒ–Š‘”•ǡ ǣ̴ͲͲͳ͸ʹʹȌǡƒ†ͺȋ ǣ̴ͲͲ͹Ͳ͵͹ǡ 337 Š––’ǣȀȀ™™™Ǥˆ‹ŽǤ‹‘Ǥ— ŽǤƒ Ǥ—Ȁ•’Ǣ ‡ŽŽ ‘‡ ‡’ƒ”–‡– ‘ˆ ƒ‰‹‰ ‡—”‘• ‹‡ ‡ǡ ‘†‘ǡ ȌǤ 338 ‘›‹œ‡††ƒ–ƒ™‹ŽŽ„‡ƒ†‡ƒ˜ƒ‹Žƒ„Ž‡–‘‘–Š‡”•—’‘ˆ‘”ƒŽ”‡“—‡•–Ǥ 339 340 Results 341 342  ͳ 343 344 A working memory task and goal-directed eye movements suppress amygdala activity 345 ƒ„Ž‘ †‡•‹‰ǡ’ƒ”–‹ ‹’ƒ–•’‡”ˆ‘”‡†ƒ–™‘Ǧ„ƒ ™‘”‹‰‡‘”›–ƒ•ƒ†‰‘ƒŽǦ†‹”‡ –‡†‡›‡ 346 ‘˜‡‡–• ™Š‹Ž‡ —†‡”‰‘‹‰ ˆ— –‹‘ƒŽ  Ǥ ‡ ˆ‘—† –›’‹ ƒŽ ƒ –‹˜ƒ–‹‘ ’ƒ––‡”• ™‹–Š‹ –Š‡ 347 †‘”•ƒŽˆ”‘–‘Ǧ’ƒ”‹‡–ƒŽ‡–™‘”ȋ‘”„‡––ƒƒ†Š—ŽƒǡʹͲͲʹǢ‹‡–ƒŽǤǡʹͲͲͻȌ†—”‹‰–Š‡–™‘Ǧ„ƒ  348 ȏe.gǤǡ–Š‡Ž‡ˆ–ƒ†”‹‰Š–†‘”•‘Žƒ–‡”ƒŽ ǣpδǤͲͲͳ’‡ƒǦ˜‘š‡Ž Ǧǡƒ†Ž‡ˆ–ƒ†”‹‰Š–’‘•–‡”‹‘” 349 ’ƒ”‹‡–ƒŽ ‘”–‡šǣ pδǤͲͲͳ ’‡ƒǦ˜‘š‡Ž ǦȐ ƒ† ‡›‡Ǧ‘˜‡‡– „Ž‘ • ȏe.gǤǡ –Š‡ Ž‡ˆ– ƒ† ”‹‰Š– 350 ˆ”‘–ƒŽ‡›‡ˆ‹‡Ž†•ǣpδǤͲͲͳ’‡ƒǦ˜‘š‡Ž Ǧǡƒ†Ž‡ˆ–ƒ†”‹‰Š–’‘•–‡”‹‘”’ƒ”‹‡–ƒŽ ‘”–‡šǣpδǤͲͲͳ 351 ’‡ƒǦ˜‘š‡Ž ǦȐǤ —”–Š‡”ǡ ƒ† ƒ• ‡š’‡ –‡†ǡ „‘–Š –Š‡ –™‘Ǧ„ƒ  „Ž‘ • ȏŽ‡ˆ–ǣ pαǤͲͲͳǡ ”‹‰Š–ǣ 352 pαǤͲͲ͵Ǣ ’‡ƒǦ˜‘š‡Ž ǦȐ ƒ† ‡›‡Ǧ‘˜‡‡– „Ž‘ • ȏŽ‡ˆ–ǣ pαǤͲ͵ͷǢ ”‹‰Š–ǣ pαǤͲͺǡ ’‡ƒǦ˜‘š‡Ž 353 ǦȐ Ž‡† –‘ †‡ƒ –‹˜ƒ–‹‘• ‹ –Š‡ ƒ›‰†ƒŽƒ ‘’ƒ”‡† –‘ ˆ‹šƒ–‹‘Ǥ Š‡ ƒ›‰†ƒŽƒ •—’’”‡••‹‘ 354 †—”‹‰–Š‡‡›‡Ǧ‘˜‡‡–„Ž‘ •™ƒ•‘–ƒ••–”‘‰ȋi.e.ǡ•—’’”‡••‹‘™ƒ•‘Ž›•‹‰‹ˆ‹ ƒ–‹–Š‡Ž‡ˆ– 355 ƒ›‰†ƒŽƒȌǡŠ‘™‡˜‡”ƒ†‹”‡ – ‘’ƒ”‹•‘”‡˜‡ƒŽ‡†‘†‹ˆˆ‡”‡ ‡‹ƒ›‰†ƒŽƒ†‡ƒ –‹˜ƒ–‹‘„‡–™‡‡

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356 –Š‡–™‘Ǧ„ƒ ƒ†‡›‡Ǧ‘˜‡‡–•„Ž‘ •ǤŠ‡—•‹‰–Š‡–™‘Ǧ„ƒ „Ž‘ •ƒ•ƒˆ— –‹‘ƒŽŽ‘ ƒŽ‹œ‡” 357 ˆ‘” –Š‡ ƒ›‰†ƒŽƒǡ –Š‡ •—’’”‡••‹‘ ™ƒ• •‹‰‹ˆ‹ ƒ– ƒ• ™‡ŽŽ ȏŽ‡ˆ–ǣ pαǤͲͳͷǢ ”‹‰Š–ǣ pαǤͲͶͶǡ ’‡ƒǦ˜‘š‡Ž 358 ǦȐǡ‹†‹ ƒ–‹‰–Š‡•—’’”‡••‹‘‹•‹ƒ•‹‹Žƒ”Ž‘ ƒ–‹‘ˆ‘”„‘–Š–ƒ••Ǥ‘‰‘–Š‡””‡‰‹‘•ǡ 359 †‡ƒ –‹˜ƒ–‹‘™ƒ•ƒŽ•‘ˆ‘—†‹–Š‡˜‡–”‘‡†‹ƒŽ ȋpδǤͲͲͳǡ ǦȌ‹–Š‡–™‘Ǧ„ƒ „Ž‘ •Ǥ ‘” 360 ƒŽŽ™Š‘Ž‡Ǧ„”ƒ‹ƒ –‹˜ƒ–‹‘ƒ††‡ƒ –‹˜ƒ–‹‘”‡•—Ž–••‡‡ ‹‰—”‡ͳƒ†ƒ„Ž‡ͳǤ‡‡ ‹‰—”‡ͳˆ‘”ƒ 361 ‹ŽŽ—•–”ƒ–‹‘‘ˆ–Š‡Ž‘ ƒ–‹‘‘ˆ–Š‡•—’’”‡••‹‘‹–Š‡ƒ›‰†ƒŽƒǤ 362 ‹ƒŽŽ›ǡ™‡ ‘†— –‡†ƒ’•› Š‘’Š›•‹‘Ž‘‰‹ ƒŽ‹–‡”ƒ –‹‘ȋ ȌƒƒŽ›•‹•ƒ†ˆ‘—†‡Šƒ ‡† 363 ‘—’Ž‹‰„‡–™‡‡–Š‡ƒ›‰†ƒŽƒƒ†–Š‡†‘”•ƒŽˆ”‘–‘Ǧ’ƒ”‹‡–ƒŽ‡–™‘”†—”‹‰„‘–Š–Š‡–™‘Ǧ„ƒ  364 ȏ‡Ǥ‰Ǥǡ –Š‡ Ž‡ˆ– ƒ† ”‹‰Š– †‘”•‘Žƒ–‡”ƒŽ  ǣ pδǤͲͲͳ ’‡ƒǦ˜‘š‡Ž ǦȐ ƒ† ‡›‡Ǧ‘˜‡‡– „Ž‘ • 365 ȏ‡Ǥ‰Ǥǡ–Š‡Ž‡ˆ–ƒ†”‹‰Š–†‘”•‘Žƒ–‡”ƒŽ ǣpδǤͲͲͳ’‡ƒǦ˜‘š‡Ž Ǧǡƒ†Ž‡ˆ–ƒ†”‹‰Š–ˆ”‘–ƒŽ‡›‡ 366 ˆ‹‡Ž†•ǣ’δǤͲͲͳ’‡ƒǦ˜‘š‡Ž ǦȐǤ‡ƒŽ•‘ˆ‘—†‡Šƒ ‡† ‘—’Ž‹‰„‡–™‡‡–Š‡ƒ›‰†ƒŽƒƒ† 367 ˜‡–”‘‡†‹ƒŽ  ǡ †—”‹‰ „‘–Š –Š‡ –™‘Ǧ„ƒ  ȏpδǤͲͲͳ ’‡ƒǦ˜‘š‡Ž ǦȐ ƒ† ‡›‡Ǧ‘˜‡‡– 368 „Ž‘ •ȏpαǤͲͲͳ’‡ƒǦ˜‘š‡Ž ǦȐǤ ‘”ƒŽŽ ‘‡ –‹˜‹–›”‡•—Ž–••‡‡ ‹‰—”‡ͳƒ†ƒ„Ž‡ʹǤ 369  ‘ Ž—•‹‘ǡ‰‘ƒŽǦ†‹”‡ –‡†‡›‡‘˜‡‡–•ǡ•‹‹Žƒ”–‘ƒ™‘”‹‰‡‘”›–ƒ•ȋ‹‡–ƒŽǤǡ 370 ʹͲͲͻȌǡ •—’’”‡•• ƒ›‰†ƒŽƒ ƒ –‹˜‹–› ƒ† ‹†— ‡ ‡Šƒ ‡† ‘—’Ž‹‰ „‡–™‡‡ –Š‡ ƒ›‰†ƒŽƒ ƒ† 371 ”‡‰‹‘•‹˜‘Ž˜‡†‹ ‘‰‹–‹˜‡”‡‰—Žƒ–‹‘‘ˆ‡‘–‹‘ȋ Š•‡”ƒ† ”‘••ǡʹͲͲͷȌǤ 372  373  ʹ 374 375 Goal-directed eye movements suppress amygdala activity 376 ‹”•–ǡ ™‡ ‹˜‡•–‹‰ƒ–‡† ƒ›‰†ƒŽƒ ƒ –‹˜‹–› ‹ ”‡•’‘•‡ –‘ –Š‡ ‰‘ƒŽǦ†‹”‡ –‡† ‡›‡ ‘˜‡‡–• †—”‹‰ 377 ‡š–‹ –‹‘ Ž‡ƒ”‹‰Ǥ ‡ ƒƒŽ›œ‡† –Š‡ †ƒ–ƒ ‹ ƒ–‹˜‡ •’ƒ ‡ –‘ ƒ‡ •—”‡ –Šƒ– –Š‡ ‡ˆˆ‡ –• ‹ –Š‡ 378 ƒ›‰†ƒŽƒ ƒ”‡ ‘– †—‡ –‘ •‹‰ƒŽ „Ž—””‹‰ ˆ”‘ –Š‡ Š‹’’‘ ƒ’—• ‹–‘ –Š‡ ƒ›‰†ƒŽƒǤ ‡’Ž‹ ƒ–‹‰ 379 š’‡”‹‡– ͳǡ ‰‘ƒŽǦ†‹”‡ –‡† ‡›‡ ‘˜‡‡–• ‹ ”‡ƒ•‡† ˆ”‘–ƒŽ ‡›‡ ˆ‹‡Ž† ƒ –‹˜‹–› ȏFȋͳǡʹ͵Ȍαͳ͵Ǥͳͳǡ ଶ 380 pαǤͲͲͳǡߟ௣αǤ͵͸Ȑ ‘’ƒ”‡† –‘ ˆ‹šƒ–‹‘Ǥ ›‰†ƒŽƒ ƒ –‹˜‹–› ™ƒ• †‡ ”‡ƒ•‡† ȏFȋͳǡʹ͵ȌαͶǤͷ͹͸ǡ pαǤͲͶǡ ଶ 381 ߟ௣αǤͳ͹Ȑ ‘’ƒ”‡†–‘ˆ‹šƒ–‹‘ƒ†–Š‡”‡™ƒ•‘‹–‡”ƒ –‹‘™‹–ŠȋΪǡǦȌȏFȋͳǡʹ͵ȌαͳǤʹͻ͸ǡpαǤʹ͹ǡ ଶ 382 ߟ௣αǤͲͷȐǤ Š—•ǡ ‹ –™‘ ‹†‡’‡†‡– •–—†‹‡• ™‡ ˆ‘—† –Šƒ– ‰‘ƒŽǦ†‹”‡ –‡† ‡›‡ ‘˜‡‡–• •—’’”‡•• 383 ƒ›‰†ƒŽƒƒ –‹˜‹–›Ǥ‡‡ ‹‰—”‡ͶǤ 384 385 Skin conductance responses during fear acquisition and extinction 386 ‡ƒ•—”‡•†—”‹‰ƒ “—‹•‹–‹‘ȋ ‹‰—”‡͵Ȍ”‡˜‡ƒŽ‡†ƒ”‘„—•–†‹ˆˆ‡”‡–‹ƒŽ ‘†‹–‹‘‹‰‡ˆˆ‡ –ȋΪ ଶ 387 ˜‡”•—•ǦȌƒ ”‘••ƒŽŽ–”‹ƒŽ•ȏFȋͳǡʹʹȌαͳͺǤͷͶǡpαʹǤͺ͸ǦͶǡߟ௣αǤͶ͸Ȑƒ†–Š‡”‡™ƒ•‘‹–‡”ƒ –‹‘™‹–Š ଶ 388 Žƒ–‡”š–‹ –‹‘ƒ‹’—Žƒ–‹‘ȋ‡›‡ǡ‘Ǧ‡›‡ȌȏFȋͳǡʹʹȌαͳǤͻͶͷǡpαǤͳͺǡߟ௣αǤͲͺȐǤ—”‹‰‡ƒ”Ž›‡š–‹ –‹‘ǡ

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ଶ 389 –Š‡”‡™ƒ•ƒ†‹ˆˆ‡”‡–‹ƒŽ ‘†‹–‹‘‹‰‡ˆˆ‡ –ȏFȋͳǡʹ͵ȌαͶͻǤ͹͹ǡpα͵ǤͶ͸Ǧ͹ǡߟ௣αǤ͸ͺȐ™Š‹ Š„‡ ƒ‡‘Ǧ ଶ 390 •‹‰‹ˆ‹ ƒ– †—”‹‰ Žƒ–‡ ‡š–‹ –‹‘ ȏFȋͳǡʹ͵ȌαǤͺͻ͸ǡ pαǤ͵ͷǡߟ௣ αǤͲͶȐ ȋ ‹‰—”‡ ͵ȌǤ Š‡”‡ ™ƒ• ˆ—ŽŽ ଶ 391 ‡š–‹ –‹‘ ‘ –Š‡ Žƒ•– –”‹ƒŽ ȏFȋͳǡʹ͵ȌαǤʹ͸Ͳǡ pαǤ͸ͳǡߟ௣αǤͲͳȐ ƒ† ‘ ‹–‡”ƒ –‹‘ ™‹–Š š–‹ –‹‘ ଶ 392 ƒ‹’—Žƒ–‹‘ȋ‡›‡ǡ‘Ǧ‡›‡ȌȏFȋͳǡʹ͵ȌαǤͻͻͳǡpαǤ͵͵ǡߟ௣αǤͲͶȐǤŠ—•ǡ‡ƒ•—”‡•”‡˜‡ƒŽ‡†–Š‡”‡™ƒ• 393 •— ‡••ˆ—Žƒ “—‹•‹–‹‘ƒ†‡š–‹ –‹‘ǡ™Š‹ Š†‹†‘–†‹ˆˆ‡”•‹‰‹ˆ‹ ƒ–Ž›„‡–™‡‡–Š‡‡›‡‘˜‡‡– 394 ƒ†‘Ǧ‡›‡‘˜‡‡–ƒ‹’—Žƒ–‹‘Ǥ 395  396 Eye movements during extinction block spontaneous recovery of fear the following day 397 ”— ‹ƒŽŽ›ǡ ƒ† ƒ• ’”‡†‹ –‡†ǡ ƒ ”‡’‡ƒ–‡†Ǧ‡ƒ•—”‡•  ƒ ”‘•• ƒŽŽ ”‡Ǧ‡š–‹ –‹‘ͳ –”‹ƒŽ• ‘ †ƒ› 398 –Š”‡‡”‡˜‡ƒŽ‡†ƒ‹–‡”ƒ –‹‘„‡–™‡‡š–‹ –‹‘ƒ‹’—Žƒ–‹‘ȋ‡›‡ǡ‘Ǧ‡›‡Ȍƒ†‹‡ȋˆ‹”•–˜‡”•—• ଶ 399 •‡ ‘†ŠƒŽˆ‘ˆ”‡Ǧ‡š–‹ –‹‘ͳȌȏFȋͳǡʹʹȌα͸Ǥ͹ʹ͵ǡpαǤͲʹǡߟ௣αǤʹ͵ȐǤ ‘ŽŽ‘™Ǧ—’–‡•–•‘–Š‡•’‘–ƒ‡‘—• 400 ”‡ ‘˜‡”›‹†‡šǡ‹†‹ ƒ–‡†„›ƒ †‹ˆˆ‡”‡–‹ƒŽ”‡•’‘†‹‰ˆ”‘–Š‡Žƒ•––”‹ƒŽ‘ˆ‡š–‹ –‹‘ –‘–Š‡ˆ‹”•– 401 –”‹ƒŽ‘ˆ”‡Ǧ‡š–‹ –‹‘ͳȋ Š‹ŽŽ‡”‡–ƒŽǤǡʹͲͳͲȌǡ”‡˜‡ƒŽ‡†–Šƒ–•’‘–ƒ‡‘—•”‡ ‘˜‡”›†‹ˆˆ‡”‡†„‡–™‡‡ ଶ 402 ‡š–‹ –‹‘ ƒ‹’—Žƒ–‹‘• ȏFȋͳǡʹʹȌαͷǤͻ͹͸ǡ pαǤͲʹǡߟ௣αǤʹͳȐǤ • ‡š’‡ –‡†ǡ –Š‡”‡ ™ƒ• •’‘–ƒ‡‘—• 403 ”‡ ‘˜‡”›ˆ‘”‡š–‹ –‹‘™‹–Š‘—–ȏtȋʹʹȌα͵Ǥ͸ͲǡpαǤͲͲʹȐǡ„—–‘–™‹–ŠȏtȋʹʹȌαǤ͸ͻͶǡpαǤͷͲǢ ‹‰—”‡͵Ȑ 404 ‡›‡‘˜‡‡–•Ǥ 405 ‘ƒ‡•—”‡–Šƒ––Š‹•†‹ˆˆ‡”‡ ‡‹•’‘–ƒ‡‘—•”‡ ‘˜‡”›™ƒ•‘–†”‹˜‡„›”‡•’‘•‡•–‘–Š‡Ǧ• 406 ”ƒ–Š‡”–Šƒ„›”‡•’‘•‡•–‘–Š‡Ϊ•ǡ™‡ƒ††‹–‹‘ƒŽŽ› ‘’ƒ”‡†”‡•’‘•‡•–‘–Š‡ˆ‹”•–Ǧ•ƒ†Ϊ• ଶ 407 †—”‹‰ ”‡Ǧ‡š–‹ –‹‘ͳǤ ‡•’‘•‡• –‘ „‘–Š Ϊ• ȏFȋͳǡʹʹȌαͶǤ͵Ͳͷǡ pαǤͲͶͻͻǡߟ௣ αǤͳ͸Ȑ ƒ† Ǧ• ଶ 408 ȏFȋͳǡʹʹȌα͸Ǥ͹Ͳͷǡ pαǤͲʹǡߟ௣αǤʹ͵Ȑ †‹ˆˆ‡”‡† „‡–™‡‡ ‡›‡Ǧ‘˜‡‡– ‘†‹–‹‘•ǡ •Š‘™‹‰ –Šƒ– –Š‡ 409 ”‡†— –‹‘‹•’‘–ƒ‡‘—•”‡ ‘˜‡”›™ƒ•‘–†”‹˜‡•‘Ž‡Ž›„›”‡•’‘•‡•–‘–Š‡Ǧ•Ǥ 410 •—ǡ‡›‡‘˜‡‡–•†—”‹‰‡š–‹ –‹‘Ž‡ƒ”‹‰‹†‡‡†„Ž‘ ‡†•’‘–ƒ‡‘—•”‡ ‘˜‡”›Ǥ 411 412 Eye movement-induced amygdala suppression during extinction predicts a reduction in fear recovery 413 after reinstatement 414 ƒŽ›•‡•‘–Š‡”‡‹•–ƒ–‡‡–”‡ ‘˜‡”›‹†‡šǡ‹†‹ ƒ–‡†„›ƒ†‹ˆˆ‡”‡–‹ƒŽ”‡•’‘†‹‰ˆ”‘–Š‡Žƒ•– 415 –”‹ƒŽ ‘ˆ ”‡Ǧ‡š–‹ –‹‘ͳ –‘ –Š‡ ˆ‹”•– –”‹ƒŽ ‘ˆ ”‡Ǧ‡š–‹ –‹‘ʹ ȋ Š‹ŽŽ‡” ‡– ƒŽǤǡ ʹͲͳͲȌǡ •Š‘™‡† –Šƒ– ଶ 416 †‹ˆˆ‡”‡–‹ƒŽ”‡•’‘•‡•”‡–—”‡†‘ƒ˜‡”ƒ‰‡ȏFȋͳǡʹʹȌαʹ͵ǤͶͺ͸ǡpα͹Ǥ͸ͷǦͷǡߟ௣αǤͷʹȐǡƒ†–Šƒ––Š‡”‡™ƒ• ଶ 417 ‘ ‹–‡”ƒ –‹‘ ™‹–Š š–‹ –‹‘ ƒ‹’—Žƒ–‹‘ ȏFȋͳǡʹʹȌαǤͲͲͷǡ pαǤͻͶǡߟ௣δǤͲͲͳȐǤ ‘–ƒ„Ž›ǡ ‹ Ž—†‹‰ 418 •–”‡‰–Š‘ˆƒ›‰†ƒŽƒ†‡ƒ –‹˜ƒ–‹‘ƒ•ƒ ‘˜ƒ”‹ƒ–‡”‡˜‡ƒŽ‡†ƒ‹–‡”ƒ –‹‘„‡–™‡‡–Š‹•†‡ƒ –‹˜ƒ–‹‘ ଶ 419 ƒ† š–‹ –‹‘ ƒ‹’—Žƒ–‹‘ ȏFȋͳǡʹͳȌα͹Ǥʹͷʹǡ pαǤͲͳǡߟ௣αǤʹ͸ȐǤ ‘ŽŽ‘™Ǧ—’ –‡•–• •Š‘™‡† ƒ ’‘•‹–‹˜‡ 420 ‘””‡Žƒ–‹‘ „‡–™‡‡ ƒ›‰†ƒŽƒ †‡ƒ –‹˜ƒ–‹‘ ƒ† ”‡ ‘˜‡”› ˆ‘ŽŽ‘™‹‰ ”‡‹•–ƒ–‡‡– ˆ‘” –Š‡ ‡›‡Ǧ

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421 ‘˜‡‡–ƒ‹’—Žƒ–‹‘ȏrȋʹͳȌαǤ͵ͻpαǤͲʹͺǡ‘‡Ǧ–ƒ‹Ž‡†Ǣ ‹‰—”‡͵ȐǤ›‰†ƒŽƒ”‡•’‘•‡•†—”‹‰–Š‡ 422 ‘Ǧ‡›‡‘˜‡‡–„Ž‘ •†‹†‘–’”‡†‹ –”‡ ‘˜‡”›ˆ‘ŽŽ‘™‹‰”‡‹•–ƒ–‡‡–ȏFȋͳǡʹͳȌαͳǤ͵ͻʹǡpαǤʹͷǡ ଶ 423 ߟ௣αǤͲ͸ȐǤ††‹–‹‘ƒŽŽ›ǡƒ–‡‹‰‡”œǦ–‡•–ȋ–‡‹‰‡”ǡͳͻͺͲȌ ‘’ƒ”‹‰–Š‡ ‘””‡Žƒ–‹‘ ‘‡ˆˆ‹ ‹‡–•‹„‘–Š 424 ƒ‹’—Žƒ–‹‘ –›’‡• ”‡˜‡ƒŽ‡† –Š‡ ’‘•‹–‹˜‡ ‘””‡Žƒ–‹‘ ™ƒ• •–”‘‰‡” ˆ‘” –Š‡ ‡›‡Ǧ‘˜‡‡– 425 ƒ‹’—Žƒ–‹‘ ‘’ƒ”‡† –‘ –Š‡ ‘Ǧ‡›‡ ‘˜‡‡– ƒ‹’—Žƒ–‹‘ ȏzȋʹʹȌαʹǤͷͻͺǡ pαǤͲʹͶȐǤ ‡ 426 ˆ—”–Š‡”‘”‡ †‹† ‘– ˆ‹† ƒ ‹–‡”ƒ –‹‘ „‡–™‡‡ š–‹ –‹‘ ƒ‹’—Žƒ–‹‘ ƒ† ƒ›‰†ƒŽƒ ଶ 427 †‡ƒ –‹˜ƒ–‹‘ ˆ‘” –Š‡ •’‘–ƒ‡‘—• ”‡ ‘˜‡”› ‹†‡š ȏFȋͳǡʹͳȌαͳǤͷͻʹǡ pαǤʹʹǡߟ௣ αǤͲ͹ȐǤ ›‰†ƒŽƒ 428 †‡ƒ –‹˜ƒ–‹‘ †‹†ǡ Š‘™‡˜‡”ǡ ’”‡†‹ – –Š‡ †‹ˆˆ‡”‡ ‡ „‡–™‡‡ –Š‡ •’‘–ƒ‡‘—• ”‡ ‘˜‡”› ‹†‡š ƒ† 429 ”‡‹•–ƒ–‡‡– ”‡ ‘˜‡”› ‹†‡š ȏrȋʹͲȌαǤ͸ʹ pαǤͲͲʹȐǤ ‡•’‘•‡• ‹ –Š‡ ˆ”‘–ƒŽ ‡›‡ ˆ‹‡Ž†• ™‡”‡ ‘– ଶ 430 ƒ••‘ ‹ƒ–‡† ™‹–Š ”‡ ‘˜‡”› ˆ‘ŽŽ‘™‹‰ ”‡‹•–ƒ–‡‡– ȏFȋͳǡʹͳȌαǤͶͶǡ pαǤͷͳǡߟ௣αǤͲʹȐǤ ‘ ˆ—”–Š‡” ƒ‡ 431 •—”‡ –Š‹• ‡ˆˆ‡ – ‹• •’‡ ‹ˆ‹  ˆ‘” –Š‡ ƒ›‰†ƒŽƒǡ ™‡ ƒ††‡† –Š‡ ”‡‹•–ƒ–‡‡– ”‡ ‘˜‡”› ‹†‡š ƒ• ƒ 432 ‘˜ƒ”‹ƒ–‡ –‘ ƒ ‘˜‡–‹‘ƒŽ ‰”‘—’ ƒƒŽ›•‹• ‹ •–ƒ†ƒ”† •–‡”‡‘–ƒ –‹  ȋ ͳͷʹȌ •’ƒ ‡Ǥ ‹–Š –Š‹• 433 ƒƒŽ›•‹•ǡ™‡†‹†‘–ˆ‹†ƒ›‘–Š‡””‡‰‹‘ȋ‹ Ž—†‹‰–Š‡—•‡‘ˆˆ‘”–Š‡˜ ǡ  ƒ††Ȍ 434 –‘ „‡ ‘””‡Žƒ–‡† ™‹–Š –Š‡ ”‡‹•–ƒ–‡‡– ”‡ ‘˜‡”› ‹†‡š ‘–Š‡” –Šƒ –Š‡ ”‹‰Š– ƒ›‰†ƒŽƒ ȋpαǤͲʹͶǡ 435 ’‡ƒǦ˜‘š‡Ž ǦȌǤ 436  ‘ Ž—•‹‘ǡ †‹ˆˆ‡”‡–‹ƒŽ ˆ‡ƒ” ”‡•’‘•‡• ‘ ƒ˜‡”ƒ‰‡ ”‡ ‘˜‡”‡† ƒˆ–‡” ”‡‹•–ƒ–‡‡–ǡ 437 Š‘™‡˜‡”ǡ”‡ ‘˜‡”›ˆ‘”–Š‡‡›‡Ǧ‘˜‡‡– ‘†‹–‹‘™ƒ•ƒ––‡—ƒ–‡†™Š‡’ƒ”–‹ ‹’ƒ–•Šƒ†•–”‘‰‡” 438 ƒ›‰†ƒŽƒ†‡ƒ –‹˜ƒ–‹‘•†—”‹‰‡›‡‘˜‡‡–•Ǥ 439  440 Whole-brain group analysis in standard stereotactic space 441 ‡ˆ‹”•–˜‡”‹ˆ‹‡†™Š‡–Š‡”–Š‡ƒ “—‹•‹–‹‘’Šƒ•‡‡šŠ‹„‹–‡†–Š‡‡š’‡ –‡†–ƒ•Ǧ”‡Žƒ–‡†ƒ –‹˜ƒ–‹‘ƒ† 442 †‡ƒ –‹˜ƒ–‹‘ †—”‹‰  ’”‡•‡–ƒ–‹‘ —•‹‰ ‘˜‡–‹‘ƒŽ ‰”‘—’ ƒƒŽ›•‡• ‹ •–ƒ†ƒ”† •–‡”‡‘–ƒ –‹  443 ȋ ͳͷʹȌ •’ƒ ‡Ǥ ‡ ‘„•‡”˜‡† ”‘„—•– †‹ˆˆ‡”‡–‹ƒŽ  ”‡•’‘•‡• ‹ –Š‡ ƒ–‡”‹‘” ‹•—Žƒ ȏŽ‡ˆ–ǣ 444 pδǤͲͲͳ ”‹‰Š–ǣ pδǤͲͲͳ ’‡ƒǦ˜‘š‡Ž Ǧ ‘””‡ –‡†Ȑ ƒ† †‘”•ƒŽ ƒ–‡”‹‘” ‹‰—Žƒ–‡ ‘”–‡š ȋ†Ȍ 445 ȏpδǤͲͲͳ’‡ƒǦ˜‘š‡Ž Ǧ ‘””‡ –‡†Ȑƒ‘‰‘–Š‡”•Ǥ††‹–‹‘ƒŽŽ›ǡ™‡‘„•‡”˜‡†”‘„—•–†‡ƒ –‹˜ƒ–‹‘• 446 ‹–Š‡˜ ȏpαǤͲͲ͹’‡ƒǦ˜‘š‡Ž Ǧ ‘””‡ –‡†Ȑƒ‘‰‘–Š‡”•Ǥ‡‡ƒ„Ž‡͵Ǥ 447 ‡š–ǡ™‡‹˜‡•–‹‰ƒ–‡†”‡•’‘•‡’ƒ––‡”•–‘–Š‡†—”‹‰‡š–‹ –‹‘Ž‡ƒ”‹‰Ǥ ‹”•–ǡ™‡ˆ‘—† 448 ƒ –‹˜ƒ–‹‘’ƒ––‡”•‹–Š‡ƒ–‡”‹‘”‹•—ŽƒȏŽ‡ˆ–ǣpαǤͲͳ͸”‹‰Š–ǣpαǤͲͶ͵’‡ƒǦ˜‘š‡Ž Ǧ ‘””‡ –‡†Ȑƒ† 449 †ȏpαǤͲʹ’‡ƒǦ˜‘š‡Ž ǦȐƒ•ƒƒ‹‡ˆˆ‡ –‘ˆȋΪ˜‡”•—•ǦȌǤ‡†‹†‘–‘„•‡”˜‡ƒ› 450 †‡ƒ –‹˜ƒ–‹‘•Ǥ –‡”‡•–‹‰Ž›ǡ•‹‹Žƒ”–‘™Šƒ–‘—”ƒ–‹˜‡•’ƒ ‡ƒƒŽ›•‹•‘ˆ–Š‡ˆ”‘–ƒŽ‡›‡ˆ‹‡Ž†•ȋ  Ȍ 451 ƒ –‹˜ƒ–‹‘ƒŽ”‡ƒ†›•Š‘™‡†ȋ•‡‡ ‹‰—”‡ͶȌǡ™‡ˆ‘—†ƒ –‹˜ƒ–‹‘‹–Š‡  ‹”‡•’‘•‡–‘–Š‡–Šƒ– 452 ™ƒ• ‘—’Ž‡† –‘ –Š‡ ‡›‡ ‘˜‡‡–• ȏŽ‡ˆ–ǣ pαǤͲ͵Ͷ ’‡ƒǦ˜‘š‡Ž Ǧ ‘””‡ –‡†Ǣ ”‹‰Š–ǣ pαǤͲͲͳ ’‡ƒǦ 453 ˜‘š‡Ž ǦȐ ƒ• ƒ ƒ‹ ‡ˆˆ‡ – ‘ˆ ‡›‡ ‘˜‡‡–•Ǥ Š‹• ™ƒ• „‡ˆ‘”‡ –Š‡ ‡š‡ —–‹‘ ‘ˆ –Š‡ ‡›‡

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454 ‘˜‡‡–•ƒ†–Š—•‹†‹ ƒ–‡•ƒƒ–‹ ‹’ƒ–‹‘”‡•’‘•‡ǤŠ‡”‡™ƒ•‘ȋΪǡǦȌ„›š–‹ –‹‘ 455 ƒ‹’—Žƒ–‹‘ȋ‡›‡ǡ‘Ǧ‡›‡Ȍ‹–‡”ƒ –‹‘Ǥ 456 ”‹–‹ ƒŽŽ›ǡ ‹ ”‡•’‘•‡ –‘ –Š‡ ‡›‡ ‘˜‡‡– „Ž‘ • ȋ™Š‹ Š ˆ‘ŽŽ‘™‡† –Š‡  ’”‡•‡–ƒ–‹‘ 457 ƒˆ–‡”ƒ„”‹‡ˆ†‡Žƒ›Ȍ ™‡ ˆ‘—††‡ƒ –‹˜ƒ–‹‘ ‹–Š‡˜  ȏpαǤͲʹͷ’‡ƒǦ˜‘š‡Ž Ǧ ‘””‡ –‡†ȐǤ ‡‡ 458 ƒ„Ž‡Ͷˆ‘”–Š‡ˆ—ŽŽ™Š‘Ž‡Ǧ„”ƒ‹”‡•—Ž–•Ǥ††‹–‹‘ƒŽƒƒŽ›•‹•‘–Š‡˜  —•‹‰–Š‡ƒ–‹˜‡Ǧ•’ƒ ‡ ଶ 459 †ƒ–ƒ ‘ˆ‹”‡†˜ ™ƒ•†‡ƒ –‹˜ƒ–‡† ‘’ƒ”‡†–‘ˆ‹šƒ–‹‘ȏFȋͳǡʹ͵Ȍα͹Ǥʹ͸ͷǡpαǤͲͳ͵ǡߟ௣αǤʹͶȐǤŠ‹• ଶ 460 †‡ƒ –‹˜ƒ–‹‘ †‹† ‘– ’”‡†‹ – ”‡ ‘˜‡”› ˆ‘ŽŽ‘™‹‰ ”‡‹•–ƒ–‡‡– ȏFȋͳǡʹ͵ȌαǤͲ͸ͷǡ pαǤͺͲͳǡߟ௣αǤͲͲ͵ȐǤ 461 Š—•ǡ•‹‹Žƒ”–‘ƒ›‰†ƒŽƒ”‡•’‘•‡•ǡ˜ ”‡•’‘•‡•ƒ”‡•—’’”‡••‡††—”‹‰–Š‡‡›‡Ǧ‘˜‡‡– 462 „Ž‘ •Ǥ 463 ‹ƒŽŽ›ǡ†—”‹‰”‡Ǧ‡š–‹ –‹‘ͳǡ™‡ˆ‘—††‹ˆˆ‡”‡–‹ƒŽ”‡•’‘•‡•ȋΪ˜‡”•—•ǦȌ‹–Š‡ 464 ƒ–‡”‹‘” ‹•—Žƒ ȏpδǤͲͲͳ ’‡ƒǦ˜‘š‡Ž Ǧ ‘””‡ –‡†ȐǤ —”‹‰ ”‡Ǧ‡š–‹ –‹‘ʹǡ ™‡ ˆ‘—† †‹ˆˆ‡”‡–‹ƒŽ 465 ”‡•’‘•‡•‹–Š‡†ȏpδǤͲͲͳ’‡ƒǦ˜‘š‡Ž Ǧ ‘””‡ –‡†Ȑƒ†–Š‡”‡™ƒ•ƒ‹–‡”ƒ –‹‘™‹–Š 466 š–‹ –‹‘ƒ‹’—Žƒ–‹‘ȋ‡›‡ǡ‘Ǧ‡›‡ȌȏpδǤͲͲ͵’‡ƒǦ˜‘š‡Ž ǦȐǤ‡ˆ‘—††”‡•’‘•‡•ˆ‘” 467 –Š‡ ‘Ǧ‡›‡ ‘˜‡‡– ƒ‹’—Žƒ–‹‘ ȏpδǤͲͲ͹ ’‡ƒǦ˜‘š‡Ž Ǧ ‘””‡ –‡†Ȑǡ „—– ‘ †‹ˆˆ‡”‡–‹ƒŽ 468 ”‡•’‘•‡•ˆ‘”–Š‡‡›‡Ǧ‘˜‡‡–ƒ‹’—Žƒ–‹‘™ƒ•’”‡•‡–ǤŠ‹•‹•‹Ž‹‡™‹–Š’”‡˜‹‘—•ˆ‹†‹‰• 469 ȋ Š‹ŽŽ‡”‡–ƒŽǤǡʹͲͳ͵Ȍƒ† ‘—Ž†„‡„‡ ƒ—•‡–Š‹•ƒƒŽ›•‹•”‡ˆŽ‡ –•„‘–Š–Š‡’”‘ ‡••‘ˆ”‡ ‘˜‡”›ƒ•™‡ŽŽ 470 ƒ•–Š‡’”‘ ‡••‘ˆ”‡Ǧ‡š–‹ –‹‘Ǥ 471 472 Discussion 473 Š‹• •–—†› ƒ‹‡† –‘ –‡•– –Š‡ Š›’‘–Š‡•‡• –Šƒ– ‰‘ƒŽǦ†‹”‡ –‡† ‡›‡ ‘˜‡‡–•ǡ ƒ• —•‡† ‹ ›‡ 474 ‘˜‡‡– ‡•‡•‹–‹œƒ–‹‘ ƒ† ‡’”‘ ‡••‹‰ ȋȌ –Š‡”ƒ’›ǡ ƒ ‡Šƒ ‡ ‡š–‹ –‹‘ –Š”‘—‰Š 475 ƒ›‰†ƒŽƒ•—’’”‡••‹‘Ǥ ‹”•–ǡ™‡ˆ‘—†–Šƒ–‰‘ƒŽǦ†‹”‡ –‡†‡›‡‘˜‡‡–•ȋš’‡”‹‡–ͳƒ†ʹȌƒ• 476 ™‡ŽŽƒ•ƒ™‘”‹‰‡‘”›–ƒ•ȋš’‡”‹‡–ͳȌ†‡ƒ –‹˜ƒ–‡†–Š‡ƒ›‰†ƒŽƒǤ‡ ‘†ǡ™‡ˆ‘—†–Šƒ– 477 „‘–Š –ƒ•• ȋš’‡”‹‡– ͳȌ ƒŽ–‡”‡† ‘‡ –‹˜‹–› „‡–™‡‡ –Š‡ ƒ›‰†ƒŽƒ ƒ† –Š‡ †‘”•ƒŽ ˆ”‘–‘Ǧ 478 ’ƒ”‹‡–ƒŽ ‡–™‘” ƒ• ™‡ŽŽ ƒ• ‘‡ –‹˜‹–› „‡–™‡‡ –Š‡ ƒ›‰†ƒŽƒ ƒ† –Š‡ ˜‡–”‘‡†‹ƒŽ ’”‡ˆ”‘–ƒŽ 479 ‘”–‡šǤ Š‹”†ǡ ƒ ’”‡ ‹•‡Ž› –‹‡† ƒ’’Ž‹ ƒ–‹‘ ‘ˆ –Š‡ ‡›‡ ‘˜‡‡–• †—”‹‰ ‡š–‹ –‹‘ Ž‡ƒ”‹‰ 480 „Ž‘ ‡†•’‘–ƒ‡‘—•”‡ ‘˜‡”›ʹͶŠŽƒ–‡”ȋš’‡”‹‡–ʹȌǤ ‘—”–Šǡ™Š‹Ž‡ˆ‡ƒ””‡•’‘•‡•‘ƒ˜‡”ƒ‰‡ 481 ”‡ ‘˜‡”‡†ƒˆ–‡””‡‹•–ƒ–‡‡–ǡ”‡ ‘˜‡”›™ƒ•ƒ––‡—ƒ–‡†™Š‡’ƒ”–‹ ‹’ƒ–•Šƒ†•–”‘‰‡”ƒ›‰†ƒŽƒ 482 †‡ƒ –‹˜ƒ–‹‘• †—”‹‰ ‡›‡ ‘˜‡‡–• ȋš’‡”‹‡– ʹȌǤ ‹˜‡ –Šƒ– ™‡ ˆ‘—† •‹‹Žƒ” ƒ›‰†ƒŽƒ 483 •—’’”‡••‹‘‹ƒ‘–Š‡”–ƒ•–ƒš‹‰™‘”‹‰‡‘”›ȋš’‡”‹‡–ͳȌǡ–Š‡”‡’‘”–‡†‡ˆˆ‡ –•‘ˆ‡ƒ” 484 ”‡ ‘˜‡”›ƒ”‡Ž‹‡Ž›‘–•’‡ ‹ˆ‹ –‘‡›‡‘˜‡‡–•Ǥ 485  ’‘–‡–‹ƒŽ ‡š’Žƒƒ–‹‘ ˆ‘” why ƒ›‰†ƒŽƒ †‡ƒ –‹˜ƒ–‹‘ ‘ —”•ǡ ‹• –Šƒ– Žƒ”‰‡Ǧ• ƒŽ‡ „”ƒ‹ 486 ‡–™‘”•ƒ –”‡ ‹’”‘ ƒŽŽ›ȋ ‘š‡–ƒŽǤǡʹͲͲͷȌƒ† ‘’‡–‡ˆ‘””‡•‘—” ‡•ȋ ‡”ƒ•‡–ƒŽǤǡʹͲͳͶȌǤ —–‡

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487 •–”‡••‡‰ƒ‰‡•–Š‡ƒ›‰†ƒŽƒ„—–‹’ƒ‹”•†‘”•ƒŽˆ”‘–‘Ǧ’ƒ”‹‡–ƒŽ‡–™‘”ˆ— –‹‘‹‰ȋ ‡”ƒ•‡–ƒŽǤǡ 488 ʹͲͳͶȌǤ—”†ƒ–ƒ ‘ˆ‹”–Šƒ–‡‰ƒ‰‹‰–Š‡†‘”•ƒŽˆ”‘–‘Ǧ’ƒ”‹‡–ƒŽ‡–™‘”Šƒ•–Š‡‘’’‘•‹–‡‡ˆˆ‡ –‘ˆ 489 †‡ƒ –‹˜ƒ–‹‰–Š‡ƒ›‰†ƒŽƒǤ‡ˆ—”–Š‡”‘”‡ˆ‘—† ‘—’Ž‹‰„‡–™‡‡–Š‡ƒ›‰†ƒŽƒƒ††‘”•‘Žƒ–‡”ƒŽ 490 ƒ•™‡ŽŽƒ•˜‡–”‘‡†‹ƒŽ’”‡ˆ”‘–ƒŽ’ƒ–Š™ƒ›•ǤŠ‡˜ ’Žƒ›•ƒ ”— ‹ƒŽ”‘Ž‡‹‡š–‹ –‹‘Ž‡ƒ”‹‰ 491 ƒ† ‹• –Š‘—‰Š– –‘ ‡†‹ƒ–‡ †‘”•‘Žƒ–‡”ƒŽ  Ǧƒ›‰†ƒŽƒ ‹–‡”ƒ –‹‘• ȋ‡Ž‰ƒ†‘ ‡– ƒŽǤǡ ʹͲͲͺȌǤ —” 492 ˆ‹†‹‰ ƒŽ‹‰• Ž‘•‡Ž› ™‹–Š –Š‡ Ž‹–‡”ƒ–—”‡ ‘ ‘‰‹–‹˜‡ ”‡‰—Žƒ–‹‘ ‘ˆ ‡‘–‹‘ ȋ Š•‡” ƒ† ”‘••ǡ 493 ʹͲͲͷȌǡ™Š‹Ž‡”‡˜‡ƒŽ‹‰–Šƒ–‡”‡‡›‡‘˜‡‡–•ƒ”‡•—ˆˆ‹ ‹‡––‘‡‰ƒ‰‡–Š‡•‡’ƒ–Š™ƒ›•Ǥ 494  ‘•‡“—‡ ‡ ‘ˆ –Š‹• ”‡•‘—” ‡ ‘’‡–‹–‹‘ ‹‰Š– „‡ –Šƒ– ˆ‡ƒ” ‡š’”‡••‹‘ ‹• ƒ––‡—ƒ–‡†Ǥ 495 –ƒ”–Ž‡”‡•’‘•‡•ǡˆ‘”‹•–ƒ ‡ǡƒ”‡”‡†— ‡†™Š‡’‡”ˆ‘”‹‰ƒ™‘”‹‰‡‘”›–ƒ•ȋ›–ƒŽ‡–ƒŽǤǡ 496 ʹͲͳʹȌƒ†’ƒ–‹‡–•™‹–Šƒ›‰†ƒŽƒŽ‡•‹‘••Š‘™‡Šƒ ‡†™‘”‹‰‡‘”›’‡”ˆ‘”ƒ ‡ȋ‘”‰ƒ 497 ‡– ƒŽǤǡ ʹͲͳʹȌǤ ˆ –Š‹• ‡ Šƒ‹• —†‡”Ž‹‡• –Š‡ ”‘Ž‡ ‘ˆ ‡›‡ ‘˜‡‡–• ‹ ”‡†— ‹‰ –”ƒ—ƒ–‹  498 •›’–‘•ǡ–Š‡ƒ›–ƒ•–ƒš‹‰™‘”‹‰‡‘”›•Š‘—Ž†Šƒ˜‡•‹‹Žƒ”‡ˆˆ‡ –•Ǥ †‡‡†ǡ‡‘–‹‘ƒŽ‹–› 499 ƒ† ˜‹˜‹†‡•• ‘ˆ ƒ—–‘„‹‘‰”ƒ’Š‹ ƒŽ ‡‘”‹‡• ƒ• ™‡ŽŽ ƒ• ‹–”—•‹˜‡ ‡‘”‹‡• ƒ”‡ ”‡†— ‡† ™Š‡ 500 ‡‘”›”‡ƒ –‹˜ƒ–‹‘‹•’ƒ‹”‡†™‹–Š™‘”‹‰‡‘”›–ƒ••ȋ ‘Ž‡•‡–ƒŽǤǡʹͲͲͻǢ‰‡ŽŠƒ”†‡–ƒŽǤǡ 501 ʹͲͳͲǢ ƒ‡• ‡– ƒŽǤǡ ʹͲͳͷȌǤ ƒ•–Ž›ǡ ‘–Š‡” –›’‡• ‘ˆ ‘‰‹–‹˜‡ ‘–”‘Žǡ •— Š ƒ• ‡‘–‹‘ ”‡‰—Žƒ–‹‘ǡ 502 •—’’”‡•• ƒ›‰†ƒŽƒ ƒ –‹˜‹–› ƒ† ƒŽ–‡” ‡‘–‹‘ƒŽ‹–› †—”‹‰ ƒ—–‘„‹‘‰”ƒ’Š‹ ƒŽ ‡‘”› ”‡ ‘ŽŽ‡ –‹‘ 503 ȋ‡‘˜ƒ‡–ƒŽǤǡʹͲͳͷȌǤ—”†ƒ–ƒ–Š‡”‡ˆ‘”‡’”‘˜‹†‡ƒ‡š’Žƒƒ–‹‘ˆ‘”Š‘™„‘–Š‡›‡‘˜‡‡–•ƒ† 504 –ƒ••‹˜‘Ž˜‹‰ ‘‰‹–‹˜‡ ‘–”‘Ž ‘—Ž†ƒˆˆ‡ ––Š‡‡‘–‹‘ƒŽ‹–›‘ˆ‡‘”‹‡•Ǥ 505 ’‘–ƒ‡‘—•”‡ ‘˜‡”›™ƒ•†‹‹‹•Š‡†ƒˆ–‡”‡š–‹ –‹‘™‹–Š‡›‡‘˜‡‡–•ǤŠ‡†‘‹ƒ– 506 ˜‹‡™‘’‘•–Ǧ‡š–‹ –‹‘”‡ ‘˜‡”›ȋƒ”‡ǡʹͲͳͳǢ—•‘‘”‡–ƒŽǤǡʹͲͳͷ„ȌŠ‘Ž†•–Šƒ––Š‹• ƒ„‡†—‡ 507 –‘—’†ƒ–‹‰–Š‡‘”‹‰‹ƒŽǦƒ••‘ ‹ƒ–‹‘‘”–‘–Š‡ˆ‘”ƒ–‹‘‘ˆƒ•–”‘‰‡”‡™‡š–‹ –‹‘‡‘”›Ǥ 508 Ž‹‡™‹–Š–Š‡Žƒ––‡”ƒ ‘—–ǡ†‹ˆˆ‡”‡–‹ƒŽˆ‡ƒ””‡•’‘•‡•”‡ ‘˜‡”‡†ƒˆ–‡””‡‹•–ƒ–‡‡–ǡ‹†‹ ƒ–‹‰ 509 –Š‡Ǧƒ••‘ ‹ƒ–‹‘™ƒ•‘–ˆ—ŽŽ›‡Ž‹‹ƒ–‡†Ǥ•‹‹Žƒ””‡†— –‹‘‹•’‘–ƒ‡‘—•”‡ ‘˜‡”›™ƒ• 510 ‘„•‡”˜‡† ‹ ƒ •–—†› ‹ ™Š‹ Š –Š‡  ™ƒ• ”‡’Žƒ ‡† „› ƒ ‘Ǧƒ˜‡”•‹˜‡ –‘‡ †—”‹‰ ‡š–‹ –‹‘ 511 ȋ—•‘‘” ‡– ƒŽǤǡ ʹͲͳͷƒȌǤ ‡ ’‘••‹„‹Ž‹–›ǡ –Š‡”‡ˆ‘”‡ǡ ‹• –Šƒ– ‡›‡ ‘˜‡‡–• ˆ‘ŽŽ‘™‹‰ –Š‡  512 ’”‡•‡–ƒ–‹‘ǡ •‹‹Žƒ” –‘ ƒ –‘‡ǡ •–”‡‰–Š‡ ‡š–‹ –‹‘ „› ”‡†— ‹‰ –Š‡ ƒ„‹‰—‹–› ‘ˆ –Š‡  ‡‹–Š‡” 513 ’”‡†‹ –‹‰ –Š‡  ‘” ‘– ’”‡†‹ –‹‰ ƒ›–Š‹‰Ǥ Ž‹‡ ƒ –‘‡ǡ Š‘™‡˜‡”ǡ ‡›‡ ‘˜‡‡–• •—’’”‡•• 514 ƒ›‰†ƒŽƒ ƒ –‹˜‹–› ƒ† ’‘••‹„Ž› ƒ––‡—ƒ–‡ ˆ‡ƒ” ”‡•’‘•‡• ȋ›–ƒŽ ‡– ƒŽǤǡ ʹͲͳʹȌǤ Š‹• ƒ› ƒŽŽ‘™ ˆ‘” 515 ƒ††‹–‹‘ƒŽ Ž‡ƒ”‡† ‘–”‘ŽŽƒ„‹Ž‹–› ‘˜‡” ‘†‹–‹‘‡† ”‡•’‘•‡• ˜‹ƒ •—„•‡“—‡– •—’’”‡••‹‘Ǥ Š‹• 516 ‹–‡”’”‡–ƒ–‹‘ ƒŽ‹‰• ™‹–Š ˆ‹†‹‰• ‘ˆ ”‡†— ‡† •’‘–ƒ‡‘—• ”‡ ‘˜‡”› ‹ ”ƒ–• ™Š‡ –”ƒ‹‡† –‘ 517 ƒ –‹˜‡Ž›ƒ˜‘‹†–Š‡†—”‹‰‡š–‹ –‹‘Ž‡ƒ”‹‰ȋ‘• ƒ”‡ŽŽ‘ƒ†‡‘—šǡʹͲͳ͵ȌǤ 518 Š‡ƒ›‰†ƒŽƒ‹•ǡƒ††‹–‹‘ƒŽŽ›ǡ ”— ‹ƒŽŽ›‹˜‘Ž˜‡†‹‡ ‘†‹‰–Š‡Ǧƒ••‘ ‹ƒ–‹‘ȋƒ”‡ǡ 519 ʹͲͳͳȌǤ›‰†ƒŽƒ•—’’”‡••‹‘ˆ‘ŽŽ‘™‹‰”‡ƒ –‹˜ƒ–‹‘ ‘—Ž†–Š‡”‡ˆ‘”‡ƒŽ•‘ Šƒ˜‡Ž‡†–‘—’†ƒ–‹‰‘ˆ

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520 –Š‡Ǧƒ••‘ ‹ƒ–‹‘ȋe.g.,ƒ•Ž‡••ƒ˜‡”•‹˜‡Ȍ”ƒ–Š‡”–Šƒ‘Ž›ˆƒ ‹Ž‹–ƒ–‹‰‡™Ž‡ƒ”‹‰ǤŽ–Š‘—‰Š–Š‡ 521 ˆ‹†‹‰ –Šƒ– ˆ‡ƒ” ”‡ ‘˜‡”‡† ˆ‘ŽŽ‘™‹‰ ”‡‹•–ƒ–‡‡– ™‘—Ž† •’‡ƒ ƒ‰ƒ‹•– –Š‹• ‡š’Žƒƒ–‹‘ǡ –Š‹• 522 ˆ‹†‹‰ ‹• ‘– ‘ Ž—•‹˜‡ ‘ ™Š‡–Š‡” –Š‡ ‡›‡ ‘˜‡‡–• ‹  –Š‡”ƒ’› ‹†— ‡ —Ž‡ƒ”‹‰ ‘” 523 ‡™Ž‡ƒ”‹‰Ǥ‹•‡–ƒ‰Ž‹‰—Ž‡ƒ”‹‰ˆ”‘‡™Ž‡ƒ”‹‰‹ƒŽƒ„‘”ƒ–‘”›‡š’‡”‹‡–”‡Ž›‹‰‘ 524 „‡Šƒ˜‹‘”ƒŽ‘„•‡”˜ƒ–‹‘•‹Š—ƒ•‹•’”‘„Ž‡ƒ–‹ ǡ•‹ ‡ƒ›„‡Šƒ˜‹‘”ƒŽ‘—– ‘‡‹• ‘•‹•–‡–™‹–Š 525 —Ž–‹’Ž‡ ‡š’Žƒƒ–‹‘•Ǥ ‘”‡‘˜‡”ǡ ”‡ ‡– ˜‹‡™• ‘ •ƒˆ‡–› Ž‡ƒ”‹‰ ŠƒŽŽ‡‰‡ ƒ •–”‹ – †‹ Š‘–‘› 526 „‡–™‡‡ —Ž‡ƒ”‹‰ ƒ† ‡™ Ž‡ƒ”‹‰ ȋŽ‡ ƒ†  Š‹ŽŽ‡”ǡ ʹͲͳ͸ȌǤ —–—”‡ •–—†‹‡• —•‹‰ ƒ ƒ‹ƒŽ 527 ‘†‡Ž‘ˆǡ‹ ‘„‹ƒ–‹‘™‹–Š–‡ Š‹“—‡•™‹–Š™Š‹ Šƒ‡‘”›–”ƒ ‡‘†‹ˆ‹ ƒ–‹‘ ƒ„‡ 528 ‡ƒ•—”‡†ǡ ‘—Ž† •Š‡† ‘”‡ Ž‹‰Š– ‘ –Š‹• ƒ––‡”Ǥ ‘” ‡šƒ’Ž‡ǡ —•‹‰ ‡‰”ƒǦ„ƒ•‡† –ƒ‰‰‹‰ 529 –‡ Š‹“—‡•ǡƒ”‡ ‡–•–—†›•Š‘™‡†Š‘™•’‡ ‹ˆ‹ ‡—”‘•ƒ”‡ƒ –‹˜ƒ–‡††—”‹‰Ž‡ƒ”‹‰ƒ†”‡ ƒŽŽ‘ˆ 530 ˆ‡ƒ” ‡‘”‹‡•ǡ ƒ† Š‘™ –Š‡•‡ ƒ”‡ ”‡Žƒ–‡† –‘ ˆ‡ƒ” ƒ––‡—ƒ–‹‘ ȋŠƒŽƒˆ ‡– ƒŽǤǡ ʹͲͳͺȌǤ ‘‰‡–Š‡”ǡ ™‡ 531 ’”‘’‘•‡ –Šƒ– ‡›‡ ‘˜‡‡–• †—”‹‰ ‡š–‹ –‹‘ Ž‡ƒ”‹‰ ƒ› ƒˆˆ‡ – ˆ‡ƒ” ”‡ ‘˜‡”› „› ‡Šƒ ‹‰ 532 ‡š–‹ –‹‘˜‹ƒ‡™Ž›Ž‡ƒ”‡†‹•–”—‡–ƒŽ ‘–”‘Ž‘˜‡”Ǧ‡˜‘‡†ˆ‡ƒ””‡•’‘•‡•ˆ‘ŽŽ‘™‹‰‡‘”› 533 ”‡ƒ –‹˜ƒ–‹‘ȋ‘• ƒ”‡ŽŽ‘ƒ†‡‘—šǡʹͲͳ͵Ȍǡ’‘••‹„Ž›–Š”‘—‰Š’ƒ–Š™ƒ›•‡‰ƒ‰‡††—”‹‰ ‘‰‹–‹˜‡ 534 ”‡‰—Žƒ–‹‘‘ˆ‡‘–‹‘ȋ Š•‡”ƒ† ”‘••ǡʹͲͲͷȌǤ 535 Š‡ ‘„•‡”˜‡† ƒ›‰†ƒŽƒ •—’’”‡••‹‘ ™ƒ• Ž‘ ƒ–‡† –‘™ƒ”†• –Š‡ †‘”•ƒŽ ”ƒ–Š‡” –Šƒ ˜‡–”ƒŽ 536 ’ƒ”–Ǥ ‘™‡˜‡”ǡ™‡ƒ”‡Š‡•‹–ƒ–‹ƒ••‹‰‹‰–Š‹•†‡ƒ –‹˜ƒ–‹‘–‘ƒ•’‡ ‹ˆ‹ •—„”‡‰‹‘Ǥ ‘’ƒ”‹•‘ 537 „‡–™‡‡†‘”•ƒŽƒ†˜‡–”ƒŽ‹•†‹ˆˆ‹ —Ž–†—‡–‘‹Š‡”‡–’”‘„Ž‡•‘ˆ‰”ƒ†‹‡– •‡“—‡ ‡•ǡ•— Šƒ• 538 •‹‰ƒŽŽ‘••‘”†‹•–‘”–‹‘•ǡ™Š‹ Šƒ”‡‹ ”‡ƒ•‡†–‘™ƒ”†•–Š‡˜‡–”ƒŽ’ƒ”–‘ˆ–Š‡„”ƒ‹ȋ‡”„‘Ž†–‡–ƒŽǤǡ 539 ʹͲͲͳǢŽƒ†›‡–ƒŽǤǡʹͲͳ͵ȌǤŠ‡•‡‹Š‡”‡–’”‘„Ž‡•ȋi.e.ǡ†‹•–ƒ ‡„‡–™‡‡ƒ„”ƒ‹ƒ”‡ƒƒ†–Š‡ 540 Š‡ƒ† ‘‹Ž‘” ƒ˜‹–‹‡•Ȍ ƒ‘–„‡ˆ—ŽŽ›”‡•‘Ž˜‡†ȋ‡”„‘Ž†–‡–ƒŽǤǡʹͲͲͳǢŽƒ†›‡–ƒŽǤǡʹͲͳ͵ȌǤŠ‡–Š‡” 541 –Š‡‡ˆˆ‡ –™‡‘„•‡”˜‡† ƒ„‡ƒ––”‹„—–‡†–‘ƒ•’‡ ‹ˆ‹ •—„”‡‰‹‘‘ˆ–Š‡ƒ›‰†ƒŽƒ–Š‡”‡ˆ‘”‡”‡ƒ‹• 542 ƒ‘’‡“—‡•–‹‘Ǥ 543 ‡ˆ‘—†ƒ–‹ ‹’ƒ–‘”›  ƒ –‹˜ƒ–‹‘‹”‡•’‘•‡–‘–Š‡ ‘—’Ž‡†™‹–Š‡›‡‘˜‡‡–• 544 „‡ˆ‘”‡ ‡›‡ ‘˜‡‡–• –‘‘ ’Žƒ ‡Ǥ Š‹• ‹• ‹ Ž‹‡ ™‹–Š ƒ–‹ ‹’ƒ–‘”› ”‡•’‘•‡• ‘„•‡”˜‡† —•‹‰ 545 ‡Ž‡ –”‘’Š›•‹‘Ž‘‰‹ ƒŽ ”‡ ‘”†‹‰• ‹ ‘‡›• ȋe.g.ǡ Š‘— ƒ† Š‘’•‘ǡ ʹͲͲͻȌǤ  ‘—” •–—†›ǡ –Š‡

546 Ϊ‡›‡ƒ†Ǧ‡›‡ƒŽ™ƒ›•’”‡†‹ –‡†ƒ‡›‡Ǧ‘˜‡‡–„Ž‘ ǡ–Š‡”‡ˆ‘”‡ƒ–‹ ‹’ƒ–‘”›”‡•’‘•‡• ‘—Ž† 547 „‡‡š’‡ –‡†ǤŽ›ˆ‹˜‡‘—–‘ˆʹͶ’ƒ”–‹ ‹’ƒ–•”‡’‘”–‡†–Š‡ƒ••‘ ‹ƒ–‹‘„‡–™‡‡–Š‡ƒ†–Š‡‡›‡Ǧ 548 ‘˜‡‡– „Ž‘ •Ǥ š Ž—†‹‰ –Š‡•‡ ’ƒ”–‹ ‹’ƒ–• †‹† ‘– ƒˆˆ‡ – –Š‡ ”‡•—Ž–•Ǥ Š‡   ƒ –‹˜ƒ–‹‘ǡ 549 ‘”‡‘˜‡”ǡ †‹† ‘– ’”‡†‹ – –Š‡ ”‡†— –‹‘ ‹ •’‘–ƒ‡‘—• ”‡ ‘˜‡”›ǡ •—‰‰‡•–‹‰ –Š‡ ƒ–‹ ‹’ƒ–‘”› 550 ”‡•’‘•‡•‹–Š‡  †‹†‘–ƒˆˆ‡ –ˆ‡ƒ””‡ ‘˜‡”›Ǥ 551 ‡•’‹–‡  „‡‹‰ ƒ ‡˜‹†‡ ‡Ǧ„ƒ•‡† –Š‡”ƒ’› ȋ‹••‘ ‡– ƒŽǤǡ ʹͲͳ͵Ǣ ‡‡ ƒ† —‹Œ’‡”•ǡ 552 ʹͲͳ͵Ȍǡ ‹– Šƒ• ”‡ ‡‹˜‡† •—„•–ƒ–‹ƒŽ ”‹–‹ ‹• ȋ‡˜‹ŽŽ›ǡ ʹͲͲʹǢ ‘‰‡”• ƒ† ‹Ž˜‡”ǡ ʹͲͲʹȌǤ —” ”‡•—Ž–•

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553 •Š‡† ‡™ Ž‹‰Š– ‘ –Š‡ ™‘”‹‰ ‡ Šƒ‹•• ‘ˆ –Š‹• –”‡ƒ–‡–Ǥ ‡ ƒ ‘—– ‘ˆ  Š‘Ž†• –Šƒ–ǡ 554 —Ž‹‡ ‡š’‘•—”‡ –Š‡”ƒ’› ȋƒ”‡ǡ ʹͲͳͳǢ ‹••‘ ‡– ƒŽǤǡ ʹͲͳ͵Ȍǡ  ‹†— ‡• —Ž‡ƒ”‹‰ ȋŠƒ’‹”‘ǡ 555 ͳͻͺͻǢ‡˜‹ŽŽ›ǡʹͲͲʹǢ˜ƒ†‡ ‘—–ƒ†‰‡ŽŠƒ”†ǡʹͲͳʹȌǤ‡ˆ‘—†‡˜‹†‡ ‡•’‡ƒ‹‰ƒ‰ƒ‹•––Š‹• 556 Žƒ‹ǡ•‹ ‡ˆ‡ƒ””‡ ‘˜‡”‡†ˆ‘ŽŽ‘™‹‰”‡‹•–ƒ–‡‡–Ǥ ‘™‡˜‡”ǡƒ•‡š’Žƒ‹‡†ƒ„‘˜‡ǡ‹–‹•’‘••‹„Ž‡–Šƒ– 557 ‘—” †‡•‹‰ †‹† ‘– ƒ’–—”‡ ƒŽŽ ’”‘ ‡••‡• –Šƒ– ‘ —” †—”‹‰  –”‡ƒ–‡–ǡ –Š‡”‡ˆ‘”‡ǡ ˆ—”–Š‡” 558 ”‡•‡ƒ” Š‹•‡ ‡••ƒ”›–‘‹˜‡•–‹‰ƒ–‡™Š‡–Š‡”ƒ›‹˜‘Ž˜‡—Ž‡ƒ”‹‰Ǥ —–—”‡•–—†‹‡• ‘—Ž† 559 ˆ‘ —• ‘ ‹ ‘”’‘”ƒ–‹‰ ‡›‡ ‘˜‡‡–• ‹ ’ƒ”ƒ†‹‰• –Šƒ– ‘”‡ Ž‹‡Ž› ƒ’–—”‡ ”‡ ‘•‘Ž‹†ƒ–‹‘ 560 ȋ Š‹ŽŽ‡”‡–ƒŽǤǡʹͲͳ͵Ǣ ƒ‡•‡–ƒŽǤǡʹͲͳͷȌǤ 561 ‘–Š‡” ‘–”‘˜‡”•› ”‡‰ƒ”†‹‰  ‘ ‡”• –Š‡ ”‘Ž‡ ‘ˆ ‡›‡ ‘˜‡‡–•ǡ ™Š‹ Š •‘‡ 562 ”‡‰ƒ”†ƒ• ”— ‹ƒŽȋŠƒ’‹”‘ǡͳͻͺͻȌǡ™Š‹Ž‡‘–Š‡”•ƒ”‰—‡–Š‡›Šƒ˜‡‘ƒ††‡†˜ƒŽ—‡ȋ‘‰‡”•ƒ†‹Ž˜‡”ǡ 563 ʹͲͲʹȌ‘”‡”‡Ž›•‡”˜‡ƒ•ƒ†‹•–”ƒ –‘”ȋ‡˜‹ŽŽ›ǡʹͲͲʹȌǤ—”†ƒ–ƒ†‡‘•–”ƒ–‡–Šƒ–‡›‡‘˜‡‡–• 564 Šƒ˜‡ƒ††‡†˜ƒŽ—‡ƒ„‘˜‡•–ƒ†ƒ”†‡š–‹ –‹‘Ž‡ƒ”‹‰Ǥ ‘™‡˜‡”ǡ–Š‡†ƒ–ƒˆ”‘š’‡”‹‡–ͳ•—‰‰‡•– 565 –Šƒ–ƒ›–ƒ•–ƒš‹‰™‘”‹‰‡‘”›™‘—Ž†•—’’”‡••ƒ›‰†ƒŽƒƒ –‹˜‹–›ƒ†Šƒ˜‡•‹‹Žƒ”‡ˆˆ‡ –•Ǥ 566 †‡‡†ǡ –Š‡”‡ ‹• ƒ Žƒ”‰‡ „‘†› ‘ˆ ”‡•‡ƒ” Š ‹†‹ ƒ–‹‰ –Šƒ– ™‘”‹‰ ‡‘”› –ƒ•• ”‡†— ‡ –Š‡ 567 ‡‘–‹‘ƒŽ‹–› ‘ˆ ‡‘”‹‡• ȋ‰‡ŽŠƒ”† ‡– ƒŽǤǡ ʹͲͳͳȌ ƒ• ™‡ŽŽ ƒ• ‡‘”› ‹–”—•‹‘• ȋ ‘Ž‡• ‡– ƒŽǤǡ 568 ʹͲͲͻǢ ƒ‡•‡–ƒŽǤǡʹͲͳͷǢ ›ƒ†—”ƒ‹‡–ƒŽǤǡʹͲͳ͹ȌǤŠ‡•‡ƒ‹’—Žƒ–‹‘•Šƒ˜‡ƒŽ•‘„‡‡•Š‘™–‘„‡ 569 ‡ˆˆ‡ –‹˜‡‹ƒ Ž‹‹ ƒŽ•‡––‹‰ȋ ›ƒ†—”ƒ‹‡–ƒŽǤǡʹͲͳ͹ȌǤŽ›–ƒ••™‹–Šƒ™‘”‹‰‡‘”›Ž‘ƒ†ƒ’’‡ƒ” 570 –‘Šƒ˜‡•— Š‡ˆˆ‡ –•ǡ™Š‹Ž‡˜‹•—ƒŽ†‹•–”ƒ –‹‘„›‹–•‡Žˆ†‘‡•‘–ȋ†‡”†‘ƒ†˜ƒ†‡ ‘—–ǡʹͲͳ͸ȌǤ 571 —” †ƒ–ƒ ƒ”‡ ƒŽ•‘ ‹ Ž‹‡ ™‹–Š •–—†‹‡• —•‹‰ ‘–Š‡” –›’‡• ‘ˆ ˆ‘ —•‡† ƒ––‡–‹‘ǡ •— Š ƒ• ‡‘–‹‘ 572 ”‡‰—Žƒ–‹‘•–”ƒ–‡‰‹‡•ȋ‡‘˜ƒ‡–ƒŽǤǡʹͲͳͷȌǡ™Š‹ Šƒ”‡ƒ ‘’ƒ‹‡†„›ƒ›‰†ƒŽƒ•—’’”‡••‹‘ƒ• 573 ™‡ŽŽǤ •—ǡƒŽ–Š‘—‰Š‘–Š‡”ƒ‹’—Žƒ–‹‘•ƒ›Šƒ˜‡ƒ•‹‹Žƒ”‡ˆˆ‡ –ǡ‘—”†ƒ–ƒ•—’’‘”––Š‡˜‹‡™–Šƒ– 574 ‡›‡‘˜‡‡–•Šƒ˜‡ƒˆ— –‹‘ƒŽ”‘Ž‡‹Ǥ 575 ‡˜‡”ƒŽŽ‹‹–ƒ–‹‘•”‡‰ƒ”†‹‰‘—”•–—†›‡‡†–‘„‡‡–‹‘‡†Ǥ ‹”•–ǡ•‘‡‘ˆ‘—”ˆ‹†‹‰• 576 ƒ”‡ ‘Ž› Œ—•– •–ƒ–‹•–‹ ƒŽŽ› •‹‰‹ˆ‹ ƒ–ǡ ƒ† ™‡”‡ ‘„–ƒ‹‡† ‹ ƒ Ž‹‹–‡† •ƒ’Ž‡ ȋ α ʹͶ ‹ ‡ƒ Š 577 ‡š’‡”‹‡–ȌǤ Š‹Ž‡ ƒ›‰†ƒŽƒ •—’’”‡••‹‘ †—‡ –‘ ‡›‡ ‘˜‡‡–• ™ƒ• ”‡’Ž‹ ƒ–‡† ƒ ”‘•• 578 ‡š’‡”‹‡–•ǡ–Š‡‡ˆˆ‡ –‘‡š–‹ –‹‘Ž‡ƒ”‹‰™ƒ•‘Ž›–‡•–‡†‹š’‡”‹‡–ʹƒ†–Š‡”‡ˆ‘”‡ƒ™ƒ‹–• 579 ‹†‡’‡†‡–”‡’Ž‹ ƒ–‹‘Ǥ‡ ‘†ǡ–Š‡‡š’‡”‹‡–ƒŽ‘†‡Ž‘ˆ™‡†‡˜‡Ž‘’‡†‹–Š‹••–—†›Šƒ• 580 ‹Š‡”‡–Ž‹‹–ƒ–‹‘•„‡ ƒ—•‡‹–‹•‹’‘••‹„Ž‡–‘ ƒ’–—”‡‡˜‡”›ƒ•’‡ –‘ˆ–Š‹•–Š‡”ƒ’›ȋe.g.ǡ”‡‰ƒ”†‹‰ 581 –‹‹‰ ‘ˆ –”ƒ—ƒ ”‡ ƒŽŽ ƒ† ‡›‡ ‘˜‡‡–•ǡ ‘” ‡ˆˆ‡ –• ‘ˆ ”‡’‡ƒ–‡† •‡••‹‘•Ȍ ‹ ƒ ‘–”‘ŽŽ‡† 582 ‡š’‡”‹‡–Ǥ —–—”‡ •–—†‹‡• •Š‘—Ž† –Š‡”‡ˆ‘”‡ ˆ‘ —• ‘ ȋͳȌ ‡•–ƒ„Ž‹•Š‹‰ –Š‡ ”‡’”‘†— ‹„‹Ž‹–› ƒ† 583 ‰‡‡”ƒŽ‹œƒ„‹Ž‹–›‘ˆ‘—”ˆ‹†‹‰•ǡȋʹȌ‹˜‡•–‹‰ƒ–‹‰–Š‡•’‡ ‹ˆ‹ ‹–›‘ˆ–Š‡‘„•‡”˜‡†‡ˆˆ‡ –•‘‡š–‹ –‹‘ 584 Ž‡ƒ”‹‰ –‘ ‡›‡ ‘˜‡‡–•Ǥ † ȋ͵Ȍ ˆ—”–Š‡” ‹ŽŽ—‹ƒ–‹‰ –Š‡ ƒ—•ƒŽ Šƒ‹ ˆ”‘ –ƒš‹‰ ™‘”‹‰ 585 ‡‘”›–‘ƒ›‰†ƒŽƒ•—’’”‡••‹‘ƒ†‡Šƒ ‡†‡š–‹ –‹‘Ž‡ƒ”‹‰Ǥ

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586  ‘ Ž—•‹‘ǡ‘—”ˆ‹†‹‰••Š‘™‡›‡‘˜‡‡–•Šƒ˜‡ƒ††‡†˜ƒŽ—‡‹•ƒˆ‡–›Ž‡ƒ”‹‰ƒ„‘˜‡ 587 •–ƒ†ƒ”†‡š–‹ –‹‘ƒŽ‘‡ǤŠ‹•‡ˆˆ‡ –ǡ™Š‹Ž‡Ž‹‡Ž›‘–•’‡ ‹ˆ‹ –‘‡›‡‘˜‡‡–•ǡ‹•ƒ••‘ ‹ƒ–‡†™‹–Š 588 ƒ›‰†ƒŽƒ†‡ƒ –‹˜ƒ–‹‘ ’‘••‹„Ž› ƒ•ƒ ‘•‡“—‡ ‡‘ˆ”‡ ‹’”‘ ƒŽŽ› ‘—’Ž‡†ƒ –‹˜ƒ–‹‘‘ˆ–Š‡†‘”•ƒŽ 589 ˆ”‘–‘Ǧ’ƒ”‹‡–ƒŽ ‡–™‘”ǡ ˜‹ƒ ˜‡–”‘‡†‹ƒŽ ’”‡ˆ”‘–ƒŽ ’ƒ–Š™ƒ›• •‹‹Žƒ” –‘ –Š‘•‡ ‹˜‘Ž˜‡† ‹ 590 ‘‰‹–‹˜‡ ”‡‰—Žƒ–‹‘ ‘ˆ ‡‘–‹‘Ǥ  ‡› ƒ†˜ƒ–ƒ‰‡ ‘ˆ ƒ›‰†ƒŽƒ †‡ƒ –‹˜ƒ–‹‘ –Š”‘—‰Š „‡Šƒ˜‹‘”ƒŽ 591 ƒ‹’—Žƒ–‹‘•ǡ”ƒ–Š‡”–Šƒ˜‹ƒ’Šƒ”ƒ ‘Ž‘‰‹ ƒŽ–”‡ƒ–‡–•ǡ‹•–Šƒ––Š‡›ƒ”‡‘Ǧ‹˜ƒ•‹˜‡ǡ’”‡ ‹•‡‹ 592 –‹‡ƒ††—”ƒ–‹‘ǡƒ†•Š‘™–‘„‡ Ž‹‹ ƒŽŽ›‡ˆˆ‡ –‹˜‡ȋ‹••‘‡–ƒŽǤǡʹͲͳ͵ȌǤ—”ˆ‹†‹‰•’”‘˜‹†‡ƒ 593 ’ƒ”•‹‘‹‘—• ƒ ‘—– ˆ‘” Š‘™ ƒ ™‹†‡ ”ƒ‰‡ ‘ˆ „‡Šƒ˜‹‘”ƒŽ ƒ‹’—Žƒ–‹‘• ‹ Ž—†‹‰ ™‘”‹‰ 594 ‡‘”›–ƒ••ǡƒ‰ƒ‡‘ˆ‡–”‹•ƒ†‡›‡‘˜‡‡–• ƒƒŽ–‡””‡–‡–‹‘‘ˆ‡‘–‹‘ƒŽ‡‘”‹‡•Ǥ 595 596 Acknowledgments 597 ‡–Šƒƒ•Šƒ‘—ƒǡ ”‡†‡”‹‡‹Œ‡”•ǡƒ† ——Ž‹ŽŽ‡”‹‰ˆ‘”‡š’‡”–ƒ†˜‹ ‡‘ƒ†ƒ—”ƒ 598 ˜ƒ‡‡”†‡„—”‰ˆ‘”Š‡Ž’™‹–Š†ƒ–ƒ ‘ŽŽ‡ –‹‘ǤŠ‹•™‘”™ƒ••—’’‘”–‡†„›‰”ƒ–•ˆ”‘–Š‡ 599 ‡•‡ƒ” Š ‘—†ƒ–‹‘ ƒ† –Š‡ —”‘’‡ƒ ‡•‡ƒ” Š ‘— ‹Ž ȋǦʹͲͳͷǦ‘  ͸ͺʹͷͻͳȌǤ —†‹‰ 600 •‘—” ‡•Šƒ†‘”‘Ž‡‹•–—†›†‡•‹‰ǡ†ƒ–ƒƒƒŽ›•‹•ǡ‘”™”‹–‹‰‘ˆ–Š‡ƒ—• ”‹’–Ǥ  References 601 ‰”‡ǡ‰ƒ ǡ ”‹ ǡŒÚ”•–”ƒ† ǡƒ”••‘Ǧǡ —”ƒ”ǡ ”‡†”‹•‘ȋʹͲͳʹȌ‹•”—’–‹‘ 602 ‘ˆ ”‡ ‘•‘Ž‹†ƒ–‹‘ ‡”ƒ•‡• ƒ ˆ‡ƒ” ‡‘”› –”ƒ ‡ ‹ –Š‡ Š—ƒ ƒ›‰†ƒŽƒǤ  ‹‡ ‡ ͵͵͹ǣͳͷͷͲȂ 603 ͳͷͷʹǤ 604 •Š„—”‡” ȋʹͲͲ͹Ȍˆƒ•–†‹ˆˆ‡‘‘”’Š‹ ‹ƒ‰‡”‡‰‹•–”ƒ–‹‘ƒŽ‰‘”‹–ŠǤ‡—”‘‹ƒ‰‡͵ͺǣͻͷȂͳͳ͵Ǥ 605 •Š„—”‡” ǡ ”‹•–‘ ȋʹͲͲͷȌ‹ˆ‹‡†•‡‰‡–ƒ–‹‘Ǥ‡—”‘‹ƒ‰‡ʹ͸ǣͺ͵ͻȂͺͷͳǤ 606 ‡ ǡ–‡‡”ǡ”‘™ ȋͳͻͻ͸Ȍ‡ ‡’”‡••‹‘ ˜‡–‘”›Ǧ ǤŠ‡•› Š‘Ž‘‰‹ ƒŽ‘”’‘”ƒ–‹‘Ǥ 607 ‹” ǡ ‹ƒ‘† ǡ ‹–Š ǡ ƒ†‡––‹‹  ȋʹͲͲ͸Ȍ ‡’ƒ”ƒ–‹‰ ”‡•’‹”ƒ–‘”›Ǧ˜ƒ”‹ƒ–‹‘Ǧ”‡Žƒ–‡† 608 ˆŽ— –—ƒ–‹‘•ˆ”‘‡—”‘ƒŽǦƒ –‹˜‹–›Ǧ”‡Žƒ–‡†ˆŽ— –—ƒ–‹‘•‹ˆ Ǥ‡—”‘‹ƒ‰‡͵ͳǣͳͷ͵͸ȂͳͷͶͺǤ 609 ‹••‘ ǡ ‘„‡”–• ǡ †”‡™ ǡ ‘‘’‡” ǡ ‡™‹•  ȋʹͲͳ͵Ȍ •› Š‘Ž‘‰‹ ƒŽ –Š‡”ƒ’‹‡• ˆ‘” Š”‘‹  610 ’‘•–Ǧ–”ƒ—ƒ–‹ •–”‡••†‹•‘”†‡”ȋȌ‹ƒ†—Ž–•Ǥ‘ Š”ƒ‡ƒ–ƒ„ƒ•‡›•–‡˜ǣͶȂ͹Ǥ 611 ‘• ǡ‡ ‡”•ǡ‹†–ȋʹͲͳʹȌŠ‡‡ˆˆ‡ –•‘ˆ‘”ƒ†”‡‡”‰‹ „Ž‘ ƒ†‡‘‡š–‹ –‹‘‹Š—ƒ•Ǥ 612 ‹‘Ž•› Š‘ŽͺͻǣͷͻͺȂ͸ͲͷǤ 613 Ž‡ ǡ  Š‹ŽŽ‡”  ȋʹͲͳ͸Ȍ ‡™ ‡ƒ”‹‰ ƒ† Ž‡ƒ”‹‰ǣ –”ƒ‰‡”• ‘”  ‘’Ž‹ ‡• ‹ Š”‡ƒ– 614 ‡‘”›––‡—ƒ–‹‘ǫ”‡†•‡—”‘• ‹͵ͻǣ͵ͶͲȂ͵ͷͳǤ 615 ‘”„‡––ƒǡŠ—Žƒ ȋʹͲͲʹȌ‘–”‘Ž‘ˆ‰‘ƒŽǦ†‹”‡ –‡†ƒ†•–‹—Ž—•Ǧ†”‹˜‡ƒ––‡–‹‘‹–Š‡„”ƒ‹Ǥ 616 ƒ–‡˜‡—”‘• ‹͵ǣʹͳͷȂʹʹͻǤ 617 ᑣ„‹‡  ǡ ‡†‘—š  ȋʹͲͲͶȌ ‹•”—’–‹‘ ‘ˆ ”‡ ‘•‘Ž‹†ƒ–‹‘ „—– ‘– ‘•‘Ž‹†ƒ–‹‘ ‘ˆ ƒ—†‹–‘”› ˆ‡ƒ” 618 ‘†‹–‹‘‹‰„›‘”ƒ†”‡‡”‰‹ „Ž‘ ƒ†‡‹–Š‡ƒ›‰†ƒŽƒǤ‡—”‘• ‹‡ ‡ͳʹͻǣʹ͸͹Ȃʹ͹ʹǤ 619 ‡Ž‰ƒ†‘ǡ‡ƒ”‹‰ ǡ‡†‘—š ǡŠ‡Ž’•ȋʹͲͲͺȌ‡—”ƒŽ ‹” —‹–”›—†‡”Ž›‹‰–Š‡”‡‰—Žƒ–‹‘‘ˆ 620 ‘†‹–‹‘‡†ˆ‡ƒ”ƒ†‹–•”‡Žƒ–‹‘–‘‡š–‹ –‹‘Ǥ‡—”‘ͷͻǣͺʹͻȂͺ͵ͺǤ 621 ‡‘˜ƒ ǡ ‘Ž ‘• ǡ ‘Ž ‘•  ȋʹͲͳͷȌ ‡—”ƒŽ ‘””‡Žƒ–‡• ‘ˆ Dz†‹•–”ƒ –‹‰dz ˆ”‘ ‡‘–‹‘ †—”‹‰ 622 ƒ—–‘„‹‘‰”ƒ’Š‹ ƒŽ”‡ ‘ŽŽ‡ –‹‘Ǥ‘ ‘‰ˆˆ‡ –‡—”‘• ‹ͳͲǣʹͳͻȂʹ͵ͲǤ 623 ‡˜‹ŽŽ›  ȋʹͲͲʹȌ            Š”‘‘Ž‘‰› ‘ˆ –• 624 ‡˜‡Ž‘’‡–ƒ† ‹‡–‹ˆ‹ –ƒ†‹‰Ǥ ‹‡˜‡– ‡ƒŽƒ –‹ ‡ͳǣͳͳ͵Ȃͳ͵ͺǤ

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625 —•‘‘” ǡƒ’‡•‡ǡ‡ ‡Ž‹ǡ‡‘—š ǡŠ‡Ž’•ȋʹͲͳͷƒȌ‘˜‡Ž–›Ǧ ƒ ‹Ž‹–ƒ–‡†š–‹ –‹‘ǣ 626 ”‘˜‹†‹‰ƒ‘˜‡Ž—– ‘‡‹Žƒ ‡‘ˆƒš’‡ –‡†Š”‡ƒ–‹‹‹•Š‡•‡ ‘˜‡”›‘ˆ‡ˆ‡•‹˜‡ 627 ‡•’‘•‡•Ǥ‹‘Ž•› Š‹ƒ–”›͹ͺǣʹͲ͵ȂʹͲͻǤ 628 —•‘‘” ǡ‹˜ǡƒ™ǡŠ‡Ž’•ȋʹͲͳͷ„Ȍ‡–Š‹‹‰š–‹ –‹‘Ǥ‡—”‘ͺͺǣͶ͹Ȃ͸͵Ǥ 629 †‡Ž„‡”‰ȋͳͻ͹ʹȌŽ‡ –”‘†‡”ƒŽ‡ ‘˜‡”›ƒ–‡ǡ ‘ƒŽǦ”‹‡–ƒ–‹‘ǡƒ†˜‡”•‹‘Ǥ•› Š‘’Š›•‹‘Ž‘‰› 630 ͻǣͷͳʹȂͷʹͲǤ 631 ‰‡ŽŠƒ”† ǡ˜ƒ†‡ ‘—–ƒǡ‡‡–•ȋʹͲͳͳȌƒš‹‰™‘”‹‰‡‘”›”‡†— ‡•˜‹˜‹†‡•• 632 ƒ†‡‘–‹‘ƒŽ‹–‡•‹–›‘ˆ‹ƒ‰‡•ƒ„‘—––Š‡—‡‡ǯ•ƒ›–”ƒ‰‡†›Ǥ ‡Šƒ˜Š‡”š’•› Š‹ƒ–”› 633 Ͷʹǣ͵ʹȂ͵͹Ǥ 634 ‰‡ŽŠƒ”† ǡ ˜ƒ ‹Œ‡ ǡ ˜ƒ †‡ ‘—–  ȋʹͲͳͲȌ Š‡ ‹’ƒ – ‘ˆ –ƒš‹‰ ™‘”‹‰ ‡‘”› ‘ 635 ‡‰ƒ–‹˜‡ƒ†’‘•‹–‹˜‡‡‘”‹‡•Ǥ—” •› Š‘–”ƒ—ƒ–‘ŽͳǣͳȂͺǤ 636 ‘šǡ›†‡”ǡ‹ ‡– ǡ‘”„‡––ƒǡƒ••‡ǡƒ‹ ŠŽ‡ȋʹͲͲͷȌŠ‡Š—ƒ„”ƒ‹‹• 637 ‹–”‹•‹ ƒŽŽ›‘”‰ƒ‹œ‡†‹–‘†›ƒ‹ ǡƒ–‹ ‘””‡Žƒ–‡†ˆ— –‹‘ƒŽ‡–™‘”•Ǥ”‘ ƒ–Ž ƒ† ‹ 638 ͳͲʹǣͻ͸͹͵Ȃͻ͸͹ͺǤ 639 ”‹•–‘Ǥǡ—‡ Š‡Žǡ ‹ Ǥǡ‘””‹• ǡ‘ŽŽ•ǡ‘ŽƒǤȋͳͻͻ͹Ȍ•› Š‘’Š›•‹‘Ž‘‰‹ ƒŽƒ†‘†—Žƒ–‘”› 640 –‡”ƒ –‹‘•‹‡—”‘‹ƒ‰‹‰Ǥ‡—”‘‹ƒ‰‡͸ǣʹͳͺȂʹʹͻǤ 641 ‡”•Šƒ ǡ ƒ”–Ž‡›ȋʹͲͳͷȌ †‹˜‹†—ƒŽ†‹ˆˆ‡”‡ ‡•‹Ž‡ƒ”‹‰’”‡†‹ ––Š‡”‡–—”‘ˆˆ‡ƒ”Ǥ‡ƒ” 642 ‡Šƒ˜Ͷ͵ǣʹͶ͵ȂʹͷͲǤ 643 ‡”•Šƒ ǡ‹˜ȋʹͲͳʹȌš’Ž‘”‹‰ƒŽƒ–‡– ƒ—•‡–Š‡‘”›‘ˆ Žƒ••‹ ƒŽ ‘†‹–‹‘‹‰Ǥ‡ƒ”‡Šƒ˜ 644 ͶͲǣʹͷͷȂʹ͸ͺǤ 645 ‡”•Šƒ  ǡ ƒ†—Ž‡• — ǡ ‘”ƒ ǡ ‹˜  ȋʹͲͳͶȌ –ƒ–‹•–‹ ƒŽ ‘’—–ƒ–‹‘• †‡”Ž›‹‰ –Š‡ 646 ›ƒ‹ •‘ˆ‡‘”›’†ƒ–‹‰Ǥ‘‘’—–‹‘ŽͳͲǤ 647 ‹–‡Žƒǡ‡›ǡ•Š„—”‡” ǡ ”‹•–‘ ȋʹͲͲ͵Ȍ‘†‡Ž‹‰”‡‰‹‘ƒŽƒ†’•› Š‘’Š›•‹‘Ž‘‰‹  648 ‹–‡”ƒ –‹‘• ‹ ˆ ǣ Š‡ ‹’‘”–ƒ ‡ ‘ˆ Š‡‘†›ƒ‹  †‡ ‘˜‘Ž—–‹‘Ǥ ‡—”‘‹ƒ‰‡ ͳͻǣʹͲͲȂ 649 ʹͲ͹Ǥ 650 Ž‘˜‡” ǡ‹ǡ‡••ȋʹͲͲͲȌ ƒ‰‡Ǧ„ƒ•‡†‡–Š‘†ˆ‘””‡–”‘•’‡ –‹˜‡ ‘””‡ –‹‘‘ˆ’Š›•‹‘Ž‘‰‹ ƒŽ 651 ‘–‹‘‡ˆˆ‡ –•‹ˆ ǣ Ǥƒ‰‡•‘‡†ͶͶǣͳ͸ʹȂͳ͸͹Ǥ 652 ”‡‡ǡ”ƒ‰‡Žǡ ‡ –‡ƒ—ǡƒƒ”ȋʹͲͳͶȌ‡˜‡Ž‘’‡–ƒ†˜ƒŽ‹†ƒ–‹‘‘ˆƒ—•—’‡”˜‹•‡† 653 • ‘”‹‰ •›•–‡ ȋ—–‘‘ƒ–‡Ȍ ˆ‘” •‹ ‘†— –ƒ ‡ ”‡•’‘•‡ ƒƒŽ›•‹•Ǥ –  •› Š‘’Š›•‹‘Ž 654 ͻͳǣͳͺ͸Ȃͳͻ͵Ǥ 655 ”‹•™‘Ž† ǡ ƒ‘„ ǡ ‡‹†‡ƒ ǡ ‹––ƒ ǡ ‡ŽŽ—• ǡ ƒ‰ ǡ ‹‡ˆ‡” ǡ ƒƒ•‡  ȋʹͲͲʹȌ 656 ‡‡”ƒŽ‹œ‡† —–‘ ƒŽ‹„”ƒ–‹‰ ƒ”–‹ƒŽŽ› ƒ”ƒŽŽ‡Ž  “—‹•‹–‹‘• ȋ ȌǤ ƒ‰ ‡•‘ ‡† 657 Ͷ͹ǣͳʹͲʹȂͳʹͳͲǤ 658 ƒƒ‡” ǡ ‘Žƒ”ǡ ‡”ƒ•ǡ‘•†‘”ˆȋʹͲͳͶȌ”‡˜‹‡™‘Š—ƒ”‡‹•–ƒ–‡‡–•–—†‹‡•᩿ǣƒ 659 ‘˜‡”˜‹‡™ƒ†‡–Š‘†‘Ž‘‰‹ ƒŽ ŠƒŽŽ‡‰‡•Ǥ‡ƒ”‡Ǥ 660 ‡‡‰‡” ǡ‡••ȋʹͲͲʹȌŠƒ–†‘‡•ˆ –‡ŽŽ—•ƒ„‘—–‡—”‘ƒŽƒ –‹˜‹–›ǫƒ–‡˜‡—”‘• ‹͵ǣͳȂͳͲǤ 661 ‡”ƒ• ǡ ‡ ‡•  ǡ ‘´Ž•ǡ ‡”ž†‡œ ȋʹͲͳͶȌ›ƒ‹ ƒ†ƒ’–ƒ–‹‘‘ˆŽƒ”‰‡Ǧ• ƒŽ‡„”ƒ‹ 662 ‡–™‘”•‹”‡•’‘•‡–‘ƒ —–‡•–”‡••‘”•Ǥ”‡†•‡—”‘• ‹͵͹ǣ͵ͲͶȂ͵ͳͶǤ 663 ‡”ƒ• ǡ ‡ ‡•  ǡ‘‡Ž‘ˆ•ǡ ‡”ž†‡œ ȋʹͲͳ͵Ȍ ‡ƒ”„”ƒ†› ƒ”†‹ƒƒ†ƒ –‹˜ƒ–‹‘‘ˆ–Š‡ 664 Š—ƒ’‡”‹ƒ“—‡†— –ƒŽ‰”‡›Ǥ‡—”‘‹ƒ‰‡͸͸ǣʹ͹ͺȂʹͺ͹Ǥ 665 ‘Ž‡•ǡ ƒ‡•ǡ‘‘†‡Ǧƒ–‡ǡ‡‡’”‘•‡ȋʹͲͲͻȌƒŽƒ›‹‰–Š‡‘’—–‡” ƒ‡Dz‡–”‹•dz 666 ‡†— ‡–Š‡—‹Ž†Ǧ’‘ˆ Žƒ•Š„ƒ •ˆ‘””ƒ—ƒǫ”‘’‘•ƒŽˆ”‘‘‰‹–‹˜‡ ‹‡ ‡Ǥ‘‡ 667 Ͷǣ‡Ͷͳͷ͵Ǥ 668 —”Ž‡ƒǡƒŽ–‡” ǡ‡Š‡ƒǡ—‘ŽŒƒ ǡƒ–‘”‘ǡ Š‹†–ǡ Š‡ŽŽǡ—••Š‘ˆˆ ǡ‡›•‡”• 669 ǡƒ‹‡”ǡ‡†”‹ ǡ—”ƒȋʹͲͳͲȌ —ƒƒ›‰†ƒŽƒ”‡ƒ –‹˜‹–›‹•†‹‹‹•Š‡†„›–Š‡ȾǦ 670 ‘”ƒ†”‡‡”‰‹ ƒ–ƒ‰‘‹•–’”‘’”ƒ‘Ž‘ŽǤ•› Š‘Ž‡†ͶͲǣͳͺ͵ͻȂͳͺͶͺǤ 671 ›ƒ†—”ƒ‹ǡŽƒ ™‡ŽŽǡ‡‹•‡”Ǧ–‡†ƒǡƒ–•‘ǡ‘•ƒŽŽǡ ‡††‡• ǡ‘„”‡ǡ ‘Ž‡• 672  ȋʹͲͳ͹Ȍ ”‡˜‡–‹‰ ‹–”—•‹˜‡ ‡‘”‹‡• ƒˆ–‡” –”ƒ—ƒ ˜‹ƒ ƒ „”‹‡ˆ ‹–‡”˜‡–‹‘ ‹˜‘Ž˜‹‰ 673 ‡–”‹• ‘’—–‡” ‰ƒ‡ ’Žƒ› ‹ –Š‡ ‡‡”‰‡ › †‡’ƒ”–‡–ǣ ƒ ’”‘‘ˆǦ‘ˆǦ ‘ ‡’– ”ƒ†‘‹œ‡†

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674 ‘–”‘ŽŽ‡†–”‹ƒŽǤ‘Ž•› Š‹ƒ–”›Ǥ 675 ƒƒ†ƒ”ǡ ‹‡Ž†‹‰ ǡ‰ƒ ȋʹͲͳ͵Ȍ—ƒ–‹–ƒ–‹˜‡‡–ƒǦƒƒŽ›•‹•‘ˆˆ ƒ†•–—†‹‡•”‡˜‡ƒŽ• 676 ‘•‹•–‡– ƒ –‹˜ƒ–‹‘ ‹ ˆ”‘–‘Ǧ•–”‹ƒ–ƒŽǦ’ƒ”‹‡–ƒŽ ”‡‰‹‘• ƒ† ‡”‡„‡ŽŽ— †—”‹‰ ƒ–‹•ƒ ƒ†‡• 677 ƒ†’”‘•ƒ ƒ†‡•Ǥ ”‘–•› Š‘ŽͶǣͳȂͳͷǤ 678 ƒ‡•ǡ‘•ƒŽŽǡ ‘’’‹––ǡ—„”‹†‰‡ǡ ‡††‡• ǡ‹Ž–‘ǡ ‘Ž‡•ȋʹͲͳͷȌ‘’—–‡” 679 ‰ƒ‡ ’Žƒ› ”‡†— ‡• ‹–”—•‹˜‡ ‡‘”‹‡• ‘ˆ ‡š’‡”‹‡–ƒŽ –”ƒ—ƒ ˜‹ƒ ”‡ ‘•‘Ž‹†ƒ–‹‘Ǧ—’†ƒ–‡ 680 ‡ Šƒ‹••Ǥ•› Š‘Ž ‹ǣͳȂͳͷǤ 681 ‹†–ǡ‘‡–‡”ȋʹͲͳ͵Ȍ‡ ‘•‘Ž‹†ƒ–‹‘‹ƒŠ—ƒˆ‡ƒ” ‘†‹–‹‘‹‰•–—†›ǣƒ–‡•–‘ˆ‡š–‹ –‹‘ƒ• 682 —’†ƒ–‹‰‡ Šƒ‹•Ǥ‹‘Ž•› Š‘ŽͻʹǣͶ͵ȂͷͲǤ 683 ‹†– ǡ ‘‡–‡” ǡ ‡”˜Ž‹‡–  ȋʹͲͲͻȌ ‡›‘† ‡š–‹ –‹‘ǣ ‡”ƒ•‹‰ Š—ƒ ˆ‡ƒ” ”‡•’‘•‡• ƒ† 684 ’”‡˜‡–‹‰–Š‡”‡–—”‘ˆˆ‡ƒ”Ǥƒ–‡—”‘• ‹ͳʹǣʹͷ͸ȂʹͷͺǤ 685 ‡‡ ǡ —‹Œ’‡”•  ȋʹͲͳ͵Ȍ  ‡–ƒǦƒƒŽ›•‹• ‘ˆ –Š‡ ‘–”‹„—–‹‘ ‘ˆ ‡›‡ ‘˜‡‡–• ‹ ’”‘ ‡••‹‰ 686 ‡‘–‹‘ƒŽ‡‘”‹‡•Ǥ ‡Šƒ˜Š‡”š’•› Š‹ƒ–”›ͶͶǣʹ͵ͳȂʹ͵ͻǤ 687 ƒ”‡ȋʹͲͳͳȌ‡‡‹‰ƒ’‘–Ž‡••‹†ǣš–‹ –‹‘ǡ‡ ‘•‘Ž‹†ƒ–‹‘ǡƒ†”ƒ•—”‡‘ˆ ‡ƒ”‡‘”›Ǥ 688 ‡—”‘͹Ͳǣͺ͵ͲȂͺͶͷǤ 689 ‡”„‘Ž†– ǡ ”ƒ••‘ ǡ ”—Š ǡ ”ƒŠ  ȋʹͲͲͳȌ — –‹‘ƒŽ   ‘ˆ –Š‡ Š—ƒ ƒ›‰†ƒŽƒǫ 690 ‡—”‘‹ƒ‰‡ͳͶǣʹͷ͵Ȃʹͷ͹Ǥ 691 ‘‘”‡ǡ —ǡ‘‘ǡǯ ‡ƒ”ǡ ‘”†ƒǡ‘Ž ‘•ǡ‘Ž ‘• ȋʹͲͳͶȌ‘’”‡Š‡•‹˜‡”‘–‘ ‘Ž 692 ˆ‘”ƒ—ƒŽ‡‰‡–ƒ–‹‘‘ˆ–Š‡‡†‹ƒŽ‡’‘”ƒŽ‘„‡–”— –—”‡•Ǥ ‹•š’Ǥ 693 ‘”‰ƒ ǡ ‡”„—”‰ ǡ Š‘”–‘ ǡ –‡‹  ǡ ‘  ȋʹͲͳʹȌ ƒ”ƒ†‘š‹ ƒŽ ˆƒ ‹Ž‹–ƒ–‹‘ ‘ˆ ™‘”‹‰ 694 ‡‘”›ƒˆ–‡”„ƒ•‘Žƒ–‡”ƒŽƒ›‰†ƒŽƒ†ƒƒ‰‡Ǥ‘‡͹Ǥ 695 ‘• ƒ”‡ŽŽ‘ ǡ ‡‘—š  ȋʹͲͳ͵Ȍ  –‹˜‡ ˜‘‹†ƒ ‡ ‡ƒ”‹‰ ‡“—‹”‡• ”‡ˆ”‘–ƒŽ —’’”‡••‹‘ ‘ˆ 696 ›‰†ƒŽƒǦ‡†‹ƒ–‡†‡ˆ‡•‹˜‡‡ƒ –‹‘•Ǥ ‡—”‘• ‹͵͵ǣ͵ͺͳͷȂ͵ͺʹ͵Ǥ 697 ƒ†‡”ǡ Šƒˆ‡ ǡ‡‘—š ȋʹͲͲͲȌ ‡ƒ”‡‘”‹‡•”‡“—‹”‡’”‘–‡‹•›–Š‡•‹•‹–Š‡ƒ›‰†ƒŽƒˆ‘” 698 ”‡ ‘•‘Ž‹†ƒ–‹‘ƒˆ–‡””‡–”‹‡˜ƒŽǤƒ–—”‡ͶͲ͸ǣ͹ʹʹȂ͹ʹ͸Ǥ 699 ǯ‡‹ŽŽ› ǡ ‘‘Ž”‹ Š ǡ ‡Š”‡•  ǡ ‹–Š ǡ ‘Šƒ•‡Ǧ‡”‰  ȋʹͲͳʹȌ ‘‘Ž• ‘ˆ –Š‡ –”ƒ†‡ǣ 700 •› Š‘’Š›•‹‘Ž‘‰‹ ƒŽ‹–‡”ƒ –‹‘•ƒ†ˆ— –‹‘ƒŽ ‘‡ –‹˜‹–›Ǥ‘ ‘‰ˆˆ‡ –‡—”‘• ‹͹ǣ͸ͲͶȂ 701 ͸ͲͻǤ 702  Š•‡”ǡ ”‘•• ȋʹͲͲͷȌŠ‡ ‘‰‹–‹˜‡ ‘–”‘Ž‘ˆ‡‘–‹‘Ǥ”‡†•‘‰ ‹ͻǣʹͶʹȂʹͶͻǤ 703 †‡”†‘ǡ˜ƒ†‡ ‘—–ȋʹͲͳ͸Ȍ‘’ƒ”‹•‘•‘ˆ‡›‡‘˜‡‡–•ƒ†ƒ– Š‡† Šƒ‰‹‰ 704 ˜‹•—ƒŽ‹’—–Ǥ ‡Šƒ˜Š‡”š’•› Š‹ƒ–”›ͷ͵ǣ͵ͶȂͶͲǤ 705 ‘•‡”ǡ‡”•Ž—‹• ǡ ‘‘‰†—‹ ǡ‘””‹• ȋʹͲͲ͸Ȍ ‘–”ƒ•–•‡•‹–‹˜‹–›‡Šƒ ‡‡–ƒ† 706 ƒ”–‹ˆƒ – ”‡†— –‹‘ ™‹–Š —Ž–‹‡ Š‘  ǣ ƒ”ƒŽŽ‡ŽǦƒ “—‹”‡† ‹Š‘‘‰‡‡‹–›Ǧ†‡•‡•‹–‹œ‡† ˆ Ǥ 707 ƒ‰‡•‘‡†ͷͷǣͳʹʹ͹Ȃͳʹ͵ͷǤ 708 ‹ǡ ‡”ƒ• ǡ˜ƒƒ”Ž‡ ǡ—‘ ǡ ‡”ž†‡œ ȋʹͲͲͻȌ —–‡’•› Š‘Ž‘‰‹ ƒŽ•–”‡••”‡†— ‡• 709 ™‘”‹‰‡‘”›Ǧ”‡Žƒ–‡†ƒ –‹˜‹–›‹–Š‡†‘”•‘Žƒ–‡”ƒŽ’”‡ˆ”‘–ƒŽ ‘”–‡šǤ‹‘Ž•› Š‹ƒ–”›͸͸ǣʹͷȂ 710 ͵ʹǤ 711 ‹••ƒ ǡ ƒœœƒŽ‡› ǡ ǯ•’‘•‹–‘  ȋʹͲͲͶȌ ‡ƒ•—”‹‰ ˆ— –‹‘ƒŽ ‘‡ –‹˜‹–› †—”‹‰ †‹•–‹ – 712 •–ƒ‰‡•‘ˆƒ ‘‰‹–‹˜‡–ƒ•Ǥ‡—”‘‹ƒ‰‡ʹ͵ǣ͹ͷʹȂ͹͸͵Ǥ 713 ‘‰‡”• ǡ ‹Ž˜‡”  ȋʹͲͲʹȌ •  ƒ ‡š’‘•—”‡ –Š‡”ƒ’›ǫ  ”‡˜‹‡™ ‘ˆ –”ƒ—ƒ ’”‘–‘ ‘Ž•Ǥ  Ž‹ 714 •› Š‘ŽͷͺǣͶ͵ȂͷͻǤ 715  Š‹ŽŽ‡” ǡ ‡Ž‰ƒ†‘  ȋʹͲͳͲȌ ˜‡”Žƒ’’‹‰ ‡—”ƒŽ •›•–‡• ‡†‹ƒ–‹‰ ‡š–‹ –‹‘ǡ ”‡˜‡”•ƒŽ ƒ† 716 ”‡‰—Žƒ–‹‘‘ˆˆ‡ƒ”Ǥ”‡†•‘‰ ‹ͳͶǣʹ͸ͺȂʹ͹͸Ǥ 717  Š‹ŽŽ‡”ǡƒ‡ ǡ‡‘—š ǡ‘ˆ‹Ž•Ǧ ǡŠ‡Ž’•ȋʹͲͳ͵Ȍš–‹ –‹‘†—”‹‰”‡ ‘•‘Ž‹†ƒ–‹‘ 718 ‘ˆ –Š”‡ƒ– ‡‘”› †‹‹‹•Š‡• ’”‡ˆ”‘–ƒŽ ‘”–‡š ‹˜‘Ž˜‡‡–Ǥ ”‘  ƒ–Ž  ƒ†  ‹    719 ͳͳͲǣʹͲͲͶͲȂʹͲͲͶͷǤ 720  Š‹ŽŽ‡”ǡ‘ˆ‹Ž•Ǧ ǡƒ‹‘ǡ ‘Š•‘ǡ‡†‘—š ǡŠ‡Ž’•ȋʹͲͳͲȌ”‡˜‡–‹‰–Š‡”‡–—”‘ˆ 721 ˆ‡ƒ”‹Š—ƒ•—•‹‰”‡ ‘•‘Ž‹†ƒ–‹‘—’†ƒ–‡‡ Šƒ‹••Ǥƒ–—”‡Ͷ͸͵ǣͶͻȂͷ͵Ǥ 722  Š™ƒ”œ ȋͳͻ͹ͺȌ•–‹ƒ–‹‰–Š‡‹‡•‹‘‘ˆƒ‘†‡ŽǤ–ƒ–͸ǣͶ͸ͳȂͶ͸ͶǤ

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723 Šƒ’‹”‘  ȋͳͻͺͻȌ ›‡ ‘˜‡‡– †‡•‡•‹–‹œƒ–‹‘ǣ  ‡™ –”‡ƒ–‡– ˆ‘” ’‘•–Ǧ–”ƒ—ƒ–‹  •–”‡•• 724 †‹•‘”†‡”Ǥ ‡Šƒ˜Š‡”š’•› Š‹ƒ–”›ʹͲǣʹͳͳȂʹͳ͹Ǥ 725 Š‹”‡”ǡ›ƒŽ‹ǡ›ŠŽ‡˜•ƒ‹ƒǡ‡‘ǡ ”‡‹ ‹—•ȋʹͲͳʹȌ‡ ‘†‹‰•—„Œ‡ –Ǧ†”‹˜‡ ‘‰‹–‹˜‡ 726 •–ƒ–‡•™‹–Š™Š‘Ž‡Ǧ„”ƒ‹ ‘‡ –‹˜‹–›’ƒ––‡”•Ǥ‡”‡„‘”–‡šʹʹǣͳͷͺȂͳ͸ͷǤ 727 Žƒ†›ǡƒŽ†‹‰‡”ǡ”ƒœ ǡ”ǫǫ•–Ž ǡ ǫǫˆŽ‹ Šǡƒœ‡„‡”‰‡”ǡ‘•‡”ǡ‹†‹• Š„‡”‰‡” 728 ȋʹͲͳ͵Ȍ ‹‰ŠǦ”‡•‘Ž—–‹‘ˆ— –‹‘ƒŽ ‘ˆ–Š‡Š—ƒƒ›‰†ƒŽƒƒ–͹Ǥ—” ƒ†‹‘Žͺʹǣ͹ʹͺȂ͹͵͵Ǥ 729 –‡‹‰‡” ȋͳͻͺͲȌ‡•–•ˆ‘” ‘’ƒ”‹‰‡Ž‡‡–•‘ˆƒ ‘””‡Žƒ–‹‘ƒ–”‹šǤ•› Š‘Ž—ŽŽͺ͹ǣʹͶͷȂʹͷͳǤ 730 Š‡  ‡ƒǤ ȋʹͲͳ͹Ȍ  ȋ‡”•‹‘ ͲǤͺǤͳǤͳȌǤ ȏ‘’—–‡” •‘ˆ–™ƒ”‡Ȑ ˜ƒ‹Žƒ„Ž‡ ƒ–ǣ Š––’•ǣȀȀŒƒ•’Ǧ 731 •–ƒ–•Ǥ‘”‰Ȁ͓Ǥ 732 ƒ——”‡ǡ Žƒ†™‹ǡƒ†„‡Ž–ǡƒ‡ ‡—˜‡Žǡƒ•‡› ǡƒŠǡ‹ȋʹͲͲͻȌ 733 ƒ”†‹‘”‡•’‹”ƒ–‘”›‡ˆˆ‡ –•‘†‡ˆƒ—Ž–Ǧ‘†‡‡–™‘”ƒ –‹˜‹–›ƒ•‡ƒ•—”‡†™‹–Šˆ Ǥ —”ƒ‹ 734 ƒ’’͵Ͳǣ͵Ͳ͵ͳȂ͵ͲͶʹǤ 735 ˜ƒ †‡ ‘—– ǡ ƒ”–‡Ž•‹ ǡ ‰‡ŽŠƒ”†  ȋʹͲͳ͵Ȍ  ǣ ›‡ ‘˜‡‡–• †—”‹‰ ”‡–”‹‡˜ƒŽ 736 ”‡†— ‡ •—„Œ‡ –‹˜‡ ˜‹˜‹†‡•• ƒ† ‘„Œ‡ –‹˜‡ ‡‘”› ƒ ‡••‹„‹Ž‹–› †—”‹‰ ˆ—–—”‡ ”‡ ƒŽŽǤ ‘‰ 737 ‘–ʹ͹ǣͳ͹͹Ȃͳͺ͵Ǥ 738 ˜ƒ†‡ ‘—–ǡ‰‡ŽŠƒ”† ȋʹͲͳʹȌ ‘™†‘‡•™‘”ǫ š’•› Š‘’ƒ–Š‘Ž͵ǣ͹ʹͶȂ͹͵ͺǤ 739 ƒ †‡” Ž‘‡‰  ȋͳͻͺͲȌ ƒŽ‹†‹–› ‘ˆ –Š‡ ‡ŽˆǦ‡‘‘”†‡Ž‹‰•Ǧ”ƒ‰‡Ž‹Œ•– ȋ —– Š ˜‡”•‹‘ ‘ˆ –Š‡ 740 ’‹‡Ž„‡”‰‡”–ƒ–‡Ǧ”ƒ‹–š‹‡–› ˜‡–‘”›ȌǤ‡†‹Œ†• Š”˜‘‘”•› Š‘Ž‡Šƒƒ” ”‡•‰‡„‹‡†‡ 741 ͵ͷǣʹͶ͵ȂʹͶͻǤ 742 ›–ƒŽ ǡ ‘”™‡ŽŽ ǡ ”‹ ǡ ”‹ŽŽ‘  ȋʹͲͳʹȌ ‡• ”‹„‹‰ –Š‡ ‹–‡”’Žƒ› „‡–™‡‡ ƒš‹‡–› ƒ† 743 ‘‰‹–‹‘ǣˆ”‘‹’ƒ‹”‡†’‡”ˆ‘”ƒ ‡—†‡”Ž‘™ ‘‰‹–‹˜‡Ž‘ƒ†–‘”‡†— ‡†ƒš‹‡–›—†‡”Š‹‰Š 744 Ž‘ƒ†Ǥ•› Š‘’Š›•‹‘Ž‘‰›ͶͻǣͺͶʹȂͺͷʹǤ 745 Š‘— ǡ Š‘’•‘   ȋʹͲͲͻȌ ‘‰‹–‹˜‡Ž› †‹”‡ –‡† •’ƒ–‹ƒŽ •‡Ž‡ –‹‘ ‹ –Š‡ ˆ”‘–ƒŽ ‡›‡ ˆ‹‡Ž† ‹ 746 ƒ–‹ ‹’ƒ–‹‘‘ˆ˜‹•—ƒŽ•–‹—Ž‹–‘„‡†‹• ”‹‹ƒ–‡†Ǥ‹•‹‘‡•ͶͻǣͳʹͲͷȂͳʹͳͷǤ  747

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Figures

Figure 1Ǥ (A) Experimental design. (B) Activations and deactivations of the two-back task and eye movements compared to fixation. The threshold for significance for the amygdala (whole-brain threshold p<.005 uncorrected and peak voxel FWE-SVC p<.05) is applied to the whole brain to show the specificity of the effect. Whole-brain corrected inferential statistics are reported in Table 1. (C) PPI cluster with the left and right amygdala as a seed for the two-back task and eye movements compared to fixation. Images are thresholded at p<.05, whole-brain FWE corrected. (D) Deactivation during the two-back blocks and eye-movement blocks (in green) overlaid onto the average T1 scan from all participants. In red is the anatomical location of the amygdala. It can be seen that the deactivation (overlap is in yellow) is located towards the dorsal part in both tasks. EM: eye movements. 748

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Figure 2Ǥ Overview of the experimental design. (A) The experiment took place on three consecutive days and included an acquisition phase (day one), an extinction phase (day two), and a re-extinction (re-extinction1) phase and re-extinction after reinstatement (re-extinction2) phase (day three). (B) Time line of a single trial during extinction when participants made eye movements. CS: conditioned stimulus.

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749 

Figure 3.The time course of the skin conductance responses (SCR) measured during acquisition and extinction, re-extinction1, and re-extinction2. Error bars represent ± standard error of the mean. CS: Conditioned stimulus, Eye: eye movements, No-eye: no-eye movements.

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Figure 4. Differential skin conductance responses (SCR) measured during (A) acquisition and (B) early and late extinction. (C) Differential spontaneous recovery index (first trial of re-extinction1 minus last trial of extinction) and (D) differential reinstatement recovery index (first trial re- extinction2 minus last trial re-extinction1). (E) Correlation between amygdala deactivation during eye-movement blocks on day two and differential reinstatement recovery index for extinction learning with eye movements. Error bars represent ± standard error of the mean. *= p < .05. Eye: eye movements, No-eye: no-eye movements. 

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 Figure 5. Eye movements suppress amygdala activity. (A) Single-subject example of the automated amygdala segmentation, ventral medial prefrontal cortex (vmPFC) was defined by a 10 mm radius around the 0 40 -3 coordinate (Schiller and Delgado, 2010) and frontal eye field (FEF) as a bilateral 5 mm radius around the MNI peak coordinates reported in a meta-analysis of neuroimaging studies of eye movements (Jamadar et al., 2013). (B) Amygdala and vmPFC deactivation and FEF activation during eye-movement and no-eye movement blocks within the extinction phase. Error bars represent ± SEM. (C) Eye-movement recordings during eye-movement and no-eye movement blocks within the extinction phase. The black and white lines reflect the mean across all participants and the gray shaded area the standard error of the mean (SEM). Eye: eye movements, No-eye: no-eye movements

29

750 Tables 751 752 Table 1 - Peak voxel coordinates and statistics of activations and deactivations of the two-back task and eye movements compared to fixation (Experiment 1) Region Side x(mm) y(mm) z(mm) Peak T P-value

Two-Back blocks > fixation a Anterior insula R 34 26 0 14.70 P<.001 Dorsolateral prefrontal cortex (dlPFC) L -40 2 30 8.57 P<.001 Dorsolateral prefrontal cortex (dlPFC) R 44 38 24 7.16 P<.001 Posterior parietal cortex L -34 -56 50 11.63 P<.001 Posterior parietal cortex R 48 -36 46 11.88 P<.001 Two-Back blocks < fixation Superior frontal gyrus L -18 42 46 13.38 P<.001 Precuneus L -4 -52 14 12.72 P<.001 Cerebellum R 34 -78 -34 10.80 P<.001 Rolandic operculum R 48 -22 18 10.12 P<.001 Angular gyrus L -48 -64 26 10.08 P<.001 Middle temporal gyrus L -58 -8 -6 9.69 P<.001 Fusiform gyrus L -32 -40 -12 9.60 P<.001 Angular gyrus R 56 -66 36 9.25 P<.001 Fusiform gyrus R 30 -42 -10 7.83 P=.007 Inferior frontal gyrus R 52 32 6 7.80 P=.007 Inferior orbital frontal gyrus R 38 36 -10 7.44 P=.015 Cerebellum L -26 -84 -32 7.13 P=.025 Ventral medial prefrontal cortex (vmPFC) L/R -6 46 0 8.39 P<.001* Amygdala L -26 -10 -18 5.64 P=.001* Amygdala R 24 -8 -16 4.73 P=.003* Eye movements > fixation Calcarine R 6 -70 8 7.37 P<.001 Precentral gyrus (Frontal Eye Fields) L -40 -10 50 6.57 P<.001 Precentral gyrus (Frontal Eye Fields) R 32 -4 52 8.48 P=.002 Posterior parietal cortex R 28 -52 50 5.81 P=.001 Posterior parietal cortex L -26 -52 54 5.77 P=.001 Middle cingulate cortex L -12 -20 40 5.22 P=.018 Eye movements < fixation Amygdala L -24 -8 -14 3.17 P=.035* Amygdala R 28 -8 -16 3.13 P =.08* Inferior occipital gyrus R 34 -98 -8 5.38 P =.008 Inferior occipital gyrus L -26 -98 -10 5.31 P =.012 753 Notes: all coordinates are defined in MNI152 space. All statistics listed are significant at p<.05, whole-brain family-wise error corrected unless indicated otherwise. aTo reduce 754 the number of peak voxels, only the peak voxels in the predefined ROIs are reported. *Small volume corrected for region of interest. 755

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756 Table 2 - Peak voxel coordinates and statistics of PPI analyses (with the amygdala as a seed) of the two-back task and eye movements compared to fixation (Experiment 1) Region Side x(mm) y(mm) z(mm) Peak T P-value

Two-Back blocks > fixation Superior motor area (SMA) R 2 -4 60 11.10 p<.001 Dorsolateral prefrontal cortex (dlPFC) L -4 56 36 6.84 P<.001 Dorsolateral prefrontal cortex (dlPFC) R 62 6 24 6.72 P<.001 Posterior parietal cortex L -32 -40 56 8.63 P<.001 Posterior parietal cortex R 6 -36 48 7.36 P<.001 Ventromedial prefrontal cortex (vmPFC) L/R -2 38 -6 6.12 P<.001* Eye movements > fixation Middle cingulate cortex R 6 -38 34 10.21 P<.001 Dorsolateral prefrontal cortex (dlPFC) L -22 40 36 6.89 P<.001 Dorsolateral prefrontal cortex (dlPFC) R 4 66 10 6.52 P<.001 Posterior parietal cortex L -4 -62 30 7.47 P<.001 Posterior parietal cortex R 52 -24 14 7.24 P<.001 Precentral gyrus (frontal eye fields) L -44 -18 46 7.01 P<.001 Precentral gyrus (frontal eye fields) R 44 -14 56 7.04 P<.001 Ventromedial prefrontal cortex (vmPFC) L/R -4 42 -2 6.44 P<.001* 757 Notes: all coordinates are defined in MNI152 space. All statistics listed are significant at p<.05, whole-brain family-wise error corrected unless indicated otherwise. To reduce the 758 number of peak voxels, only the peak voxels in the predefined ROIs are reported. *Small volume corrected for region of interest 759

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760 Table 3 - Peak voxel coordinates and statistics during acquisition Region Side x(mm) y(mm) z(mm) Peak T P-value

CS+ > CS- Inferior frontal gyrus / anterior insula R 50 24 4 9.6797 P<.001 Inferior frontal gyrus / anterior insula L -36 24 4 8.90 P<.001 Supramarginal gyrus R 54 -40 28 8.6493 P<.001 Dorsal anterior cingulate cortex (dACC) R 8 26 28 7.8007 P<.001 Supramarginal gyrus L -54 -40 32 6.1253 P=.002 Cerrebellum R 2 -54 -36 5.9356 P=.004 Cerrebellum L -34 -54 -32 5.9167 P=.004

CS- > CS+ Middle occipital gyrus L -44 -74 30 5.8497 P=.006 Hippocampus L -24 -18 -18 5.8302 P=.006 Ventral medial prefrontal cortex (vmPFC) L -10 52 -6 5.7878 P=.007 Precuneus R 10 -54 14 5.7507 P=.008 Paracentral lobule R 8 -26 70 5.6498 P=.012 Precuneus L -8 -60 16 5.6123 P=.014 Fusiform gyrus / Para-hippocampal gyrus R 32 -44 -4 5.3385 P=.038 761 Notes: all coordinates are defined in MNI152 space. All statistics listed are significant at p<.05, whole-brain family-wise error corrected unless indicated otherwise. 762

763 764

32 de Voogd et al. – Eye movements enhance extinction – Manuscript - Page | 33

765 Table 4 - Peak voxel coordinates and statistics during extinction learning Region Side x(mm) y(mm) z(mm) Peak T P-value

CS presentation CS+ > CS- [main effect CS] Anterior insula R 42 18 -6 6.36 P=.001 Anterior insula L -32 22 -4 5.75 P=.009 Frontal middle orbital cortex R 20 60 -12 5.61 P=.02 Dorsal anterior cingulate cortex (dACC) R 4 28 30 5.31 P=.04

Eye movements > No-eye movements [main effect eye movements] Calcrine L 10 -68 18 10.82 P<.001 Supplemental Motor Area L -4 -6 62 7.94 P<.001 Posterior parietal cortex L -26 -52 52 7.60 P<.001 Precentral gyrus (frontal eye fields) L -44 -8 52 7.58 P<001 Precentral gyrus (frontal eye fields) R 44 -8 50 7.94 P<.001 Posterior parietal cortex R 26 -52 48 7.53 P<.001 Middle temporal gyrus R 44 -64 10 6.26 P=.001 Middle cingulate cortex L -14 -20 38 5.98 P=044 Eye movement blocks Eye movements > fixation Lingual gyrus L -10 -88 -2 15.13 p<.001 Precentral gyrus (frontal eye fields) R 48 -4 42 10.26 P<.001 Putamen L -22 -2 12 9.80 P<.001 Supplemental Motor Area L -4 -4 62 8.21 P=.003 Precentral gyrus (frontal eye fields) L -52 -4 38 8.02 P=.005 Putamen R 22 8 10 7.46 P=.016 Eye movements < fixation Posterior insula R 38 -18 12 11.22 P<.001 Parahippocampal gyrus R 18 -14 -24 9.16 P<.001 Posterior insula L -36 -20 8 8.99 P=.001 Inferior occipital gyrus R 32 -96 -4 7.80 P=.008 Superior parietal gyrus R 18 -44 68 7.44 P=.017 Inferior occipital gyrus L -28 -94 -6 7.34 P=.020 Ventral medial prefrontal cortex (vmPFC) L -8 24 -12 7.22 P=.025 Post-central gyrus R 44 -26 54 7.01 P=.036 Amygdala R 32 4 -20 4.43 P=.008* 766 Notes: all coordinates are defined in MNI152 space. All statistics listed are significant at p<.05, whole-brain family-wise error corrected unless indicated otherwise. 767 *Small volume corrected for region of interest. 768

33 de Voogd et al. – Eye movements enhance extinction – Manuscript - Page | 34

769 Table 5 - Peak voxel coordinates and statistics during re-extinction1 and re-extinction2 Region Side x(mm) y(mm) z(mm) Peak T P-value

Re-extinction1 Cs+ > CS- Anterior insula R 50 20 -2 6.3 P<.001 Re-extinction2 CS+>CS- Dorsal Anterior cingulate cortex R 8 26 50 5.52 P=.024 Thalamus R 6 -6 2 5.52 P=.024 Cs by Extinction manipulation interaction Dorsal Anterior cingulate cortex L -8 28 38 4.89 P=.003* CS+ > CS- [no-eye movements] Dorsal Anterior cingulate cortex R 8 36 40 6.24 P=.007

770 Notes: all coordinates are defined in MNI152 space. All statistics listed are significant at p<.05, whole-brain family-wise error corrected unless indicated otherwise. 771 *Small volume corrected for region of interest. 772 773

34 AB amygdala ... 2-back EMEM 2-back rest 2-back ... 2-back blocks ...eye-movement blocks 2-back blocks

2 amygdala

5 amygdala

2

3

eye-movement blocks amygdala 3 activation deactivation ...... 3.2 4.0 4.8 5.6 CD

dlPFC parietal cortex vmPFC

x=-4 x=-32 z=-6 x = -22 x = -23 x = -24 dlPFC FEF vmPFC

x=-32 x=-22 z=-2 eye-movement blocks 2-back blocks eye-movement blocks 2-back blocks x = -22 x = -23 x = -24 ppi amygdala seed 6 6.5 7 7.5 A Day 1 Day 2 Day 3 Day 3 Acquisition Extinction Re-extinction 1 Re-extinction 2

Reinstatement

CS+ CS+

CS+ CS+ CS+ CS+ CS+ CS+ eye noeye eye noeye eye noeye

CS- CS- CS- CS- CS- CS- CS- CS- eye noeye eye noeye eye noeye

B Eye movements during extinction

CS+ eye ......

4 seconds 0.5-1.5 seconds 10 seconds 6-8 seconds A 1 CS+ eye 0.9 CS+ no-eye CS- eye 0.8 CS- no-eye 0.7 0.6 0.5 0.4 SCR (√μS) 0.3 0.2 0.1 0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 1 2 3 4 5 6 7 8 9 10 11 12 B trials (acquisition) trials (extinction) 1 0.9 0.8 0.7 0.6 0.5 0.4 SCR (√μS) 0.3 0.2 0.1 0 123456 123456 trials (re-extinction1) trials (re-extinction2) AB 0.5 0.5 eye 0.45 0.45 no-eye 0.4 0.4 0.35 0.35 0.3 0.3 0.25 0.25 0.2 0.2 0.15 0.15 0.1 0.1 mean differential SCR (√μS) mean differential SCR (√μS) mean differential 0.05 0.05 0 0 acquisition early extinction late extinction C D E 1 1 0.9 0.9 1.5 0.8 * 0.8 0.7 0.7 1 0.6 0.6 0.5 * 0.5 0.5 0.4 0.4 0 0.3 0.3 -0.5 0.2 0.2 mean differential SCR (√μS) mean differential mean differential SCR (√μS) mean differential 0.1 0.1 -1 0 0 index recovery Δ reinstatement -0.04 -0.02 0 0.02 spontaneous recovery after amygdala activity recovery reinstatement during eye movements AB eye amygdala 0.01 0.01 no-eye

0 0

-0.01 -0.01 vmPFC Amygdala Arbitrary units (au) -0.02 cs+ eye/no eye -0.02 cs- eye/no eye -4 -2 0 2 4 6 8 10 12 14 16 -4 -2 0 2 4 6 8 10 12 14 16 seconds seconds

0.01 0.01 FEF

0 0

-0.01 -0.01 vmPFC C Arbitrary units (au) -0.02 cs+ eye/no eye -0.02 cs- eye/no eye 1.5 -4-20246810121416 -4-20246810121416 0 seconds seconds -1.5 1.5 0.4 0.4 0 -1.5 cs+ eye/no eye Arbitrary units (au) −2 0 2 4 6 8 10121416 0.2 0.2 seconds 1.5 0 0 0 FEF -1.5 -0.2 -0.2

1.5 Arbitrary units (au) 0 -0.4 cs+ eye/no eye -0.4 cs- eye/no eye -1.5 cs- eye/no eye

Arbitrary units (au) −2 0 2 4 6 8 10121416 -4 -2 0 2 4 6 8 10 12 14 16 -4-20 2 4 6 8 10121416 seconds seconds seconds