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Review | Development

Development and organization of the evolutionary conserved three-layered olfactory cortex

Olfactory cortex development

Esther Klingler Department of Basic , University of Geneva, Geneva, Switzerland.

DOI: 10.1523/ENEURO.0193-16.2016

Received: 6 July 2016

Revised: 11 November 2016

Accepted: 8 December 2016

Published: 20 January 2017

Author Contributions: EK designed and wrote the paper.

Conflict of Interest: Authors report no conflict of interest.

Correspondence should be addressed to Esther Klingler, Jabaudon Lab, Department of Basic Neuroscience, University of Geneva, CMU, 1, rue Michel Servet, 1206 Geneva – Switzerland, E-mail: [email protected]

Cite as: eNeuro 2017; 10.1523/ENEURO.0193-16.2016

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Accepted manuscripts are peer-reviewed but have not been through the copyediting, formatting, or proofreading process. This is an open-access article distributed under the terms of the Creative Commons Attribution 4.0 International (http://creativecommons.org/licenses/by/4.0), which permits unrestricted use, distribution and reproduction in any medium provided that the original work is properly attributed.

Copyright © 2017 the authors 

ϭ Manuscript Title Page Ϯ  ϯ  ϰ 1. Manuscript Title ϱ ‡˜‡Ž‘’‡–ƒ†‘”‰ƒ‹œƒ–‹‘‘ˆ–Š‡‡˜‘Ž—–‹‘ƒ”› ‘•‡”˜‡†–Š”‡‡ǦŽƒ›‡”‡†‘Žˆƒ –‘”› ‘”–‡šǤ ϲ  ϳ 2. Abbreviated Title ϴ Žˆƒ –‘”› ‘”–‡š†‡˜‡Ž‘’‡– ϵ  ϭϬ 3. List of all Author Names and Affiliations ϭϭ •–Š‡”Ž‹‰Ž‡”ǣ‹˜‡”•‹–›‘ˆ ‡‡˜ƒǡ‡’ƒ”–‡–‘ˆƒ•‹ ‡—”‘• ‹‡ ‡ǡ ‡‡˜ƒǡ™‹–œ‡”Žƒ†Ǥ ϭϮ  ϭϯ 4. Author Contributions ϭϰ †‡•‹‰‡†ƒ†™”‘–‡–Š‡’ƒ’‡”Ǥ ϭϱ  ϭϲ 5. Correspondence should be addressed to ϭϳ •–Š‡”Ž‹‰Ž‡”ǡ‡•–Š‡”ǤŽ‹‰Ž‡”̷—‹‰‡Ǥ Š ϭϴ ΪͶͳʹʹ͵͹ͻͶͳ͸͹ ϭϵ ƒ„ƒ—†‘ƒ„ ϮϬ ‡’ƒ”–‡–‘ˆƒ•‹ ‡—”‘• ‹‡ ‡ǡ‹˜‡”•‹–›‘ˆ ‡‡˜ƒǡ Ϯϭ ͳǡ”—‡‹ Š‡Ž‡”˜‡– ϮϮ ͳʹͲ͸ ‡‡˜ƒǦ™‹–œ‡”Žƒ† Ϯϯ  Ϯϰ 6. Number of Figures:͵ Ϯϱ 7. Number of Tables: Ͳ Ϯϲ 8. Number of Multimedia:Ͳ Ϯϳ  Ϯϴ 9. Number of words for Abstract: ͳͷͲ Ϯϵ 10. Number of words for Significance Statement: ͳͲͷ ϯϬ 11. Number of words for Introduction: ͷͲͷ ϯϭ 12. Number of words for Discussion: ͶͺͲ ϯϮ  ϯϯ 13. Acknowledgments ϯϰ –ŠƒǤ ƒ›ǡ Ǥ‹‘‹ƒ†Ǥ‘ƒ•‡ŽŽ‘ˆ‘” ‘•–”— –‹˜‡ ‘‡–•ƒ† ‘””‡ –‹‘•‘–Š‡ ϯϱ ƒ—• ”‹’–Ǥ –ŠƒǤ ‘—–‡„”‘œ‡ƒ†Ǥ ƒ„ƒ—†‘ˆ‘”–Š‡‹”•—’’‘”–Ǥ ϯϲ  ϯϳ 14. Conflict of Interest ϯϴ ‘ǣ—–Š‘””‡’‘”–•‘ ‘ˆŽ‹ –‘ˆ‹–‡”‡•– ϯϵ  ϰϬ 15. Funding sources ϰϭ Š‹• ™‘” ™ƒ• •—’’‘”–‡† „› •‡”ǡ ‹˜‡”•‹–› ‹‡””‡ ‡– ƒ”‹‡ —”‹‡ȋƒ”‹•ǡ ”ƒ ‡Ȍǡƒ† ϰϮ ‹˜‡”•‹–›‘ˆ ‡‡˜ƒȋ ‡‡˜ƒǡ™‹–œ‡”Žƒ†ȌǤ ϰϯ  ϰϰ 

ϭ  

ϰϱ Title: Development and organization of the evolutionary conserved three-layered

ϰϲ olfactory cortex.

ϰϳ Short title: Žˆƒ –‘”› ‘”–‡š†‡˜‡Ž‘’‡–

ϰϴ Abstract: Š‡ ‘Žˆƒ –‘”› ‘”–‡š ‹• ’ƒ”– ‘ˆ –Š‡ ƒƒŽ‹ƒ ‡”‡„”ƒŽ ‘”–‡š –‘‰‡–Š‡” ™‹–Š –Š‡

ϰϵ ‡‘ ‘”–‡š ƒ† –Š‡ Š‹’’‘ ƒ’—•Ǥ – ”‡ ‡‹˜‡• †‹”‡ – ‹’—– ˆ”‘ –Š‡ ‘Žˆƒ –‘”› „—Ž„• ƒ†

ϱϬ ’ƒ”–‹ ‹’ƒ–‡• ‹‘†‘” †‹• ”‹‹ƒ–‹‘ǡ ƒ••‘ ‹ƒ–‹‘ ƒ†Ž‡ƒ”‹‰ȋ‡‡”•ƒ† —œ—‹ǡ ʹͲͳ͵ȌǤ – ‹•

ϱϭ –Š‘—‰Š– –‘ „‡ ƒ ‡˜‘Ž—–‹‘ƒ”›Ǧ ‘•‡”˜‡† ’ƒŽ‡‘ ‘”–‡šǡ ™Š‹ Š •Šƒ”‡• ‘‘ Šƒ”ƒ –‡”‹•–‹ •

ϱϮ ™‹–Š–Š‡–Š”‡‡ǦŽƒ›‡”‡†‰‡‡”ƒŽ ‘”–‡š‘ˆ”‡’–‹Ž‡•ȋ„‘‹–‹œ‡–ƒŽǤǡʹͲͲʹȌǤŠ‡‘Žˆƒ –‘”› ‘”–‡šŠƒ•

ϱϯ „‡‡•–—†‹‡†ƒ•ƒDz•‹’Ž‡‘†‡Ždz–‘ƒ††”‡•••‡•‘”›’”‘ ‡••‹‰ǡ–Š‘—‰ŠŽ‹––Ž‡‹•‘™ƒ„‘—–‹–•

ϱϰ ’”‡ ‹•‡ ‡ŽŽ‘”‹‰‹ǡ†‹˜‡”•‹–›ƒ†‹†‡–‹–›ǤŠ‹Ž‡–Š‡†‡˜‡Ž‘’‡–ƒ†–Š‡ ‡ŽŽ—Žƒ”†‹˜‡”•‹–›‘ˆ–Š‡

ϱϱ •‹šǦŽƒ›‡”‡† ‡‘ ‘”–‡š ƒ”‡ ‹ ”‡ƒ•‹‰Ž› —†‡”•–‘‘†ǡ –Š‡ ‘Žˆƒ –‘”› ‘”–‡š ”‡ƒ‹• ’‘‘”Ž›

ϱϲ †‘ —‡–‡†ˆ‘”–Š‡•‡ƒ•’‡ –•Ǥ ‡”‡‹•ƒ”‡˜‹‡™‘ˆ —””‡–‘™Ž‡†‰‡‘–Š‡†‡˜‡Ž‘’‡–ƒ†

ϱϳ ‘”‰ƒ‹œƒ–‹‘ ‘ˆ –Š‡ ‘Žˆƒ –‘”› ‘”–‡šǡ ‡‡’‹‰ –Š‡ ƒƒŽ‘‰› ™‹–Š –Š‘•‡ ‘ˆ –Š‡ ‡‘ ‘”–‡šǤ Š‡

ϱϴ ‘’ƒ”‹•‘‘ˆ‘Žˆƒ –‘”› ‘”–‡šƒ†‡‘ ‘”–‡š™‹ŽŽƒŽŽ‘™–Š‡‘’‡‹‰ ‘ˆ ‡˜‘Ž—–‹‘ƒ”›

ϱϵ ’‡”•’‡ –‹˜‡•‘ ‘”–‹ ƒŽ†‡˜‡Ž‘’‡–Ǥ

ϲϬ

ϲϭ Significance statement: Š‡ ‘Žˆƒ –‘”› ‘”–‡š ‹• ƒ ‡˜‘Ž—–‹‘ƒ”›Ǧ ‘•‡”˜‡† ’ƒŽ‡‘ ‘”–‡š

ϲϮ ‹’Ž‹ ƒ–‡†‹‘†‘”’”‘ ‡••‹‰ǤŠ‹Ž‡•‡˜‡”ƒŽ•–—†‹‡•ƒ††”‡••‡†Š‘™–Š‡‘Žˆƒ –‘”› ‘”–‡š‡ ‘†‡•

ϲϯ ƒ†’”‘ ‡••‡•‘†‘”ƒ–‹ˆ‘”ƒ–‹‘ǡŽ‹––Ž‡‹•‘™ƒ„‘—–‹–•’”‡ ‹•‡ ‡ŽŽ—Žƒ”‘”‹‰‹ǡ†‹˜‡”•‹–›ƒ†

ϲϰ ‹†‡–‹–›Ǥ  ”ƒ˜‡Ž‹‰ ™Š‡”‡ –Š‡ ‡ŽŽ• ƒ”‡ „‘” ƒ† Š‘™ –Š‡› ‹‰”ƒ–‡ –‘™ƒ”†• ƒ† •‡––Ž‡ –Š‡

ϲϱ ‘Žˆƒ –‘”› ‘”–‡š †—”‹‰ †‡˜‡Ž‘’‡– ‹• ‘ˆ ƒ‹ ‹’‘”–ƒ ‡ –‘ —†‡”•–ƒ†‹–• ‹” —‹–

ϲϲ ‘”‰ƒ‹œƒ–‹‘ƒ†ˆ— –‹‘Ǥ ƒ††‹–‹‘ǡ ‘’ƒ”‹‰–Š‡†‡˜‡Ž‘’‡–•‘ˆ–Š‡‘Žˆƒ –‘”› ‘”–‡š™‹–Š

ϲϳ –Š‡‡‘ ‘”–‡š™‹ŽŽŠ‡Ž’‹†‡–‹ˆ›‹‰ ‘‘‡˜‘Ž—–‹‘ƒ”› ‘•‡”˜‡††‡˜‡Ž‘’‡–ƒŽ‡ Šƒ‹••

ϲϴ ƒ•™‡ŽŽƒ•‡™‡ Šƒ‹•••’‡ ‹ˆ‹ –‘–Š‡‡‘ ‘”–‡šǡ–Šƒ–ƒ’’‡ƒ”‡†Žƒ–‡”†—”‹‰‡˜‘Ž—–‹‘ƒ†

ϲϵ ’ƒ”–‹ ‹’ƒ–‡‹„—‹Ž†‹‰–Š‡ƒƒŽ‹ƒ ‘”–‡šǤ

ϭ 

ϳϬ Introduction

ϳϭ ƒƒŽ‹ƒ ‡”‡„”ƒŽ ‘”–‡š ‘’”‹•‡•–Š‡‡‘ ‘”–‡šǡ–Š‡Š‹’’‘ ƒ’—•ƒ†–Š‡‘Žˆƒ –‘”›

ϳϮ ‘”–‡šǤ Š‡ ‘Žˆƒ –‘”› ‘”–‡š ‹• ƒ ‡˜‘Ž—–‹‘ƒ”›Ǧ ‘•‡”˜‡† ’ƒŽ‡‘ ‘”–‡š Ž‘ ƒ–‡† ‹ –Š‡

ϳϯ ˜‡–”‘Žƒ–‡”ƒŽ ’ƒ”– ‘ˆ –Š‡ –‡Ž‡ ‡’ŠƒŽ‘ǡ ƒ† •Šƒ”‡• ‘‘ Šƒ”ƒ –‡”‹•–‹ • ™‹–Š –Š‡ ‰‡‡”ƒŽ

ϳϰ ‘”–‡š‘ˆ”‡’–‹Ž‡• ‘’‘•‡†‘ˆ–Š”‡‡–Š‹Žƒ›‡”•ȋ„‘‹–‹œ‡–ƒŽǤǡʹͲͲʹǢŠ‡’Š‡”†ǡʹͲͳͳǢ ‘—”‹‡”‡–

ϳϱ ƒŽǤǡ ʹͲͳͷȌǤ Š‹• –Š”‡‡ǦŽƒ›‡”‡† ‘”‰ƒ‹œƒ–‹‘ǡ ƒŽ•‘ ƒŽŽ‡† ƒŽŽ‘ ‘”–‡šǡ ‹• ‘•‡”˜‡† ‹ „‘–Š –Š‡

ϳϲ Š‹’’‘ ƒ’—•ȋ‡†‹ƒŽ’ƒŽŽ‹—†‡”‹˜ƒ–‹˜‡Ȍƒ†–Š‡‘Žˆƒ –‘”› ‘”–‡š ȋŽƒ–‡”ƒŽ ’ƒŽŽ‹— †‡”‹˜ƒ–‹˜‡Ȍ

ϳϳ ȋ„‘‹–‹œ‡–ƒŽǤǡʹͲͲʹȌǤŠ‡ƒƒŽ‹ƒ‘Žˆƒ –‘”› ‘”–‡š‹••—„†‹˜‹†‡†‹•‡˜‡”ƒŽ•–”— –—”‡•ǡ ‹–‡†

ϳϴ Š‡”‡ ƒŽ‘‰ ƒ–‡”‘Ǧ’‘•–‡”‹‘” ƒš‹•ǣ –Š‡ ƒ–‡”‹‘” ‘Žˆƒ –‘”› — Ž‡—•ǡ–Š‡‘Žˆƒ –‘”›–—„‡” Ž‡ǡ–Š‡

ϳϵ ’‹”‹ˆ‘” ‘”–‡šǡ–Š‡‘Žˆƒ –‘”›ƒ›‰†ƒŽƒȋ ‘”–‹ ƒŽƒ›‰†ƒŽƒƒ†— Ž‡—•‘ˆ–Š‡Ž‘–Ȍƒ†–Š‡Žƒ–‡”ƒŽ

ϴϬ ‡–‘”Š‹ƒŽ ‘”–‡šȋŽȌȋ ƒ„‡”Ž›ǡʹͲͲͳǢ”—Œ‡•‡–ƒŽǤǡʹͲͲͷǢƒ Š‡œǦ†”ƒ†‡ƒ†‡†”‹ ǡ

ϴϭ ʹͲͲͻȌǤ  ‘–”ƒ•––‘–Š‡ Žƒ••‹ ƒŽ•‡•‘”›’ƒ–Š™ƒ›™Š‡”‡•‡•‘”›‹ˆ‘”ƒ–‹‘‹•ˆ‹”•–”‡Žƒ›‡†‹

ϴϮ –Š‡–ŠƒŽƒ—•„‡ˆ‘”‡”‡ƒ Š‹‰–Š‡‡‘ ‘”–‡šǡ–Š‡‘Žˆƒ –‘”› ‘”–‡š”‡ ‡‹˜‡•†‹”‡ –•‡•‘”›‹’—–•

ϴϯ ˆ”‘–Š‡‹–”ƒŽƒ†–—ˆ–‡† ‡ŽŽ•‘ˆ‘Žˆƒ –‘”›„—Ž„ȋȌ˜‹ƒ–Š‡Žƒ–‡”ƒŽ‘Žˆƒ –‘”›–”ƒ –ȋȌǡƒ†‹•

ϴϰ –Š‘—‰Š––‘’Žƒ›ƒ‡›”‘Ž‡‹‘Žˆƒ –‹‘ȋ‹Ž•‘ƒ†ƒ‹‡ǡʹͲͲ͸Ǣ •ƒƒ •‘ǡʹͲͳͲǢ‡‹™ƒ†ƒ†

ϴϱ ŠƒŽƒ•ƒ‹ǡʹͲͳͳǢ‡‡”•ƒ†—œ—‹ǡʹͲͳ͵ȌǤŠ‡’‹”‹ˆ‘” ‘”–‡š‹•–Š‡‘•–•–—†‹‡†•–”— –—”‡‘ˆ

ϴϲ –Š‡ ‘Žˆƒ –‘”› ‘”–‡šǤ – ”‡ ‡‹˜‡• ‹’—–• ˆ”‘ –Š‡ ǡ ƒ• ™‡ŽŽ ƒ• ˆ”‘ –Š‡ ‘–Š‡” ”‡‰‹‘• ‘ˆ –Š‡

ϴϳ ‘Žˆƒ –‘”› ‘”–‡š ‹–‡† ƒ„‘˜‡ǡ ƒ† •‡†• ’”‘Œ‡ –‹‘• –‘ –Š‡ ƒ–‡”‹‘” ‘Žˆƒ –‘”› — Ž‡—•ǡ –Š‡

ϴϴ ‘Žˆƒ –‘”›–—„‡” Ž‡ǡ–Š‡ ‘”–‹ ƒŽƒ›‰†ƒŽƒȋ‘Ȍƒ†–Š‡Ž™‹–Š‹–Š‡‘Žˆƒ –‘”› ‘”–‡šǡƒ•™‡ŽŽ

ϴϵ ƒ• –‘ –Š‡ ‡†‹‘†‘”•ƒŽ — Ž‡—• ‘ˆ –Š‡ –ŠƒŽƒ—• ƒ† –‘ •‡˜‡”ƒŽ •—„†‹˜‹•‹‘• ‘ˆ –Š‡ ’”‡ˆ”‘–ƒŽ

ϵϬ ‘”–‡š ȋ‹ Ž—†‹‰ –Š‡ ‹ˆ”ƒŽ‹„‹ ǡ ‘”„‹–‘ˆ”‘–ƒŽ ƒ† ƒ‰”ƒ—ƒŽ ‹•—Žƒ” ‘”–‹ ‡•Ȍ ȋ ‘Š•‘ ‡– ƒŽǤǡ

ϵϭ ʹͲͲͲǢ •–”ƒ† ‡– ƒŽǤǡ ʹͲͲͳǢ ‡›‡” ‡– ƒŽǤǡ ʹͲͲ͸Ǣ ‡”” ‡– ƒŽǤǡ ʹͲͲ͹ȌǤ ‹ƒŽŽ›ǡ –Š‡ ’‹”‹ˆ‘” ‘”–‡š

ϵϮ •‡†• ‘•‡“—‡–ˆ‡‡†„ƒ ’”‘Œ‡ –‹‘•–‘–Š‡ȋ ƒ„‡”Ž›ƒ†”‹ ‡ǡͳͻ͹ͺǢŠ‹’Ž‡›ƒ††ƒ‡ǡ

ϵϯ ͳͻͺͶȌǤŠ‡’‹”‹ˆ‘” ‘”–‡šŠƒ•„‡‡•Š‘™–‘’ƒ”–‹ ‹’ƒ–‡‹‘†‘” †‹• ”‹‹ƒ–‹‘ǡ ƒ••‘ ‹ƒ–‹‘

ϵϰ ƒ† Ž‡ƒ”‹‰ ȋ‡‡”• ƒ† —œ—‹ǡ ʹͲͳ͵ȌǢ ‹– ƒŽŽ‘™• ‘„Œ‡ – ”‡ ‘‰‹–‹‘ ‹ ƒ •‡•‘”› Žƒ†• ƒ’‡ǡ

ϵϱ ™Š‘•‡”‡Ž‡˜ƒ–’‡” ‡’–—ƒŽ†‹‡•‹‘•ƒ”‡†›ƒ‹ ƒŽŽ›•Šƒ’‡†„›•‡•‘”›‡š’‡”‹‡ ‡ȋ ‘—”‹‡”

ϵϲ ‡–ƒŽǤǡʹͲͳͷȌǤ

Ϯ 

ϵϳ Š‹Ž‡–Š‡†‡˜‡Ž‘’‡–‘ˆ–Š‡•‹šǦŽƒ›‡”‡†‡‘ ‘”–‡šŠƒ•„‡‡Žƒ”‰‡Ž› †‡• ”‹„‡†ǡ –Š‡

ϵϴ †‡˜‡Ž‘’‡–‘ˆ–Š‡‘Žˆƒ –‘”› ‘”–‡š”‡ƒ‹•’‘‘”Ž›—†‡”•–‘‘†Ǥ‡—”‘•‘ˆ–Š‡‘Žˆƒ –‘”› ‘”–‡š

ϵϵ •‡‡ –‘ Šƒ˜‡ —Ž–‹’Ž‡ ‘”‹‰‹• „‘–Š ‹ –Š‡ ’ƒŽŽ‹ƒŽ ƒ† –Š‡ •—„’ƒŽŽ‹ƒŽ˜‡–”‹ —Žƒ”œ‘‡•ȋȌ

ϭϬϬ ȋƒ”‡› ‡– ƒŽǤǡ ʹͲͲ͸Ǣ ƒ” ‹ƒǦ‘”‡‘ ‡– ƒŽǤǡ ʹͲͲͺȌǤ ‘”‡‘˜‡” –Š‡•‡‡—”‘•‹‰”ƒ–‡‘˜‡”Ž‘‰

ϭϬϭ †‹•–ƒ ‡•ƒ††‹•’Žƒ› ‘’Ž‡š‹‰”ƒ–‘”›’”‘’‡”–‹‡•„‡ˆ‘”‡”‡ƒ Š‹‰–Š‡‹”ˆ‹ƒŽ†‡•–‹ƒ–‹‘‹–Š‡

ϭϬϮ ‘Žˆƒ –‘”› ‘”–‡šƒ†‘”‰ƒ‹œ‹‰‹–Š”‡‡Žƒ›‡”•Ǥ –Š‹•”‡˜‹‡™ǡ™‡†‹• —•• —””‡–‘™Ž‡†‰‡‘

ϭϬϯ –Š‡†‡˜‡Ž‘’‡–ƒ†‘”‰ƒ‹œƒ–‹‘‘ˆ–Š‡‘Žˆƒ –‘”› ‘”–‡šǣŠ‘™‹•–Š‡‘Žˆƒ –‘”› ‘”–‡š•’‡ ‹ˆ‹‡†

ϭϬϰ †—”‹‰†‡˜‡Ž‘’‡–ǫŠ‡”‡ƒ”‡–Š‡‘Žˆƒ –‘”› ‘”–‡š‡—”‘•„‘”ǫ ‘™†‘–Š‡›‹‰”ƒ–‡–‘–Š‡

ϭϬϱ ‘Žˆƒ –‘”› ‘”–‡š ƒ† ˆ‘”–Š‡ –Š”‡‡ Žƒ›‡”•ǫ Šƒ–‹•‘™ ƒ„‘—– ‡—”‘ †‹˜‡”•‹–› ™‹–Š‹ –Š‡

ϭϬϲ ‘Žˆƒ –‘”› ‘”–‡šƒ†Š‘™†‘–Š‡›‡ ‘†‡‘Žˆƒ –‘”›‹ˆ‘”ƒ–‹‘ǫŠ‡•‡ “—‡•–‹‘• ™‹ŽŽ „‡

ϭϬϳ ƒ††”‡••‡†‡‡’‹‰–Š‡ƒƒŽ‘‰›™‹–Š–Š‡‡‘ ‘”–‡š†‡˜‡Ž‘’‡–ǡ‹‘”†‡”–‘‘’‡‡˜‘Ž—–‹‘ƒ”›

ϭϬϴ ’‡”•’‡ –‹˜‡•‘ ‘”–‹ ƒŽ†‡˜‡Ž‘’‡–Ǥ

ϭϬϵ Arealization and neurogenesis

ϭϭϬ —”‹‰–Š‡–‡ˆ‹”•–†ƒ›•‘ˆ‘—•‡‡„”›‘‹ †‡˜‡Ž‘’‡–ǡ–Š‡’ƒŽŽ‹ƒŽ ‡—”‘‡’‹–Š‡Ž‹—

ϭϭϭ ’”‘Ž‹ˆ‡”ƒ–‡•ǡ ƒŽŽ‘™‹‰ –Š‡ ‡š’‘‡–‹ƒŽ ‰‡‡”ƒ–‹‘ ‘ˆ ’”‘‰‡‹‡• –Š”‘—‰Š •›‡–”‹  †‹˜‹•‹‘• ‹

ϭϭϮ –Š‡ȋ‘ –‘”‡–ƒŽǤǡʹͲͲͶȌǤ ”‘‡„”›‘‹ †ƒ›ͳͳȋͳͳȌǡ–Š‡ˆ‹”•–’‘•–Ǧ‹–‘–‹ ‡—”‘•ƒ”‡

ϭϭϯ ’”‘†— ‡† „› ƒ•›‡–”‹  †‹˜‹•‹‘•ǡ ‰‡‡”ƒ–‹‰ „‘–Š ’”‘‰‡‹‡•ǡ ™Š‹ Š •–ƒ› ‹ –Š‡ ǡ ƒ†

ϭϭϰ ‡—”‘„Žƒ•–•ǡ™Š‹ Š‹‰”ƒ–‡‘—–‘ˆ–Š‡–‘™ƒ”†–Š‡ ‘”–‹ ƒŽ•—”ˆƒ ‡Ǥ

ϭϭϱ Arealization of the cortex by signaling centers

ϭϭϲ ƒ•‡†‘‰‡‡‡š’”‡••‹‘ǡ–Š‡’ƒŽŽ‹—‹••—„†‹˜‹†‡†‹–‘ˆ‘—”ƒŒ‘”•—„Ǧ”‡‰‹‘•ǡƒ‡Ž›

ϭϭϳ –Š‡ ‡†‹ƒŽǡ †‘”•ƒŽǡ Žƒ–‡”ƒŽ ƒ† ˜‡–”ƒŽ ’ƒŽŽ‹—Ǣ ‡ƒ Š ’ƒŽŽ‹ƒŽ ”‡‰‹‘ ‹• –Š‘—‰Š– –‘ ‰‹˜‡ ”‹•‡ –‘

ϭϭϴ •’‡ ‹ˆ‹  ‘”–‹ ƒŽ•–”— –—”‡•ǡ•— Šƒ•–Š‡Š‹’’‘ ƒ’—•ǡ–Š‡‡‘ ‘”–‡šǡ–Š‡‘Žˆƒ –‘”› ‘”–‡šƒ†–Š‡

ϭϭϵ ƒ›‰†ƒŽƒȀ‡†‘’‹”‹ˆ‘” ‘”–‡š — Ž‡‹ ”‡•’‡ –‹˜‡Ž› ȋ—‡ŽŽ‡• ‡– ƒŽǤǡ ʹͲͲͲǢ — ‡– ƒŽǤǡ ʹͲͲͳȌǤ

ϭϮϬ –”‹•‹  ‡ Šƒ‹•• „ƒ•‡† ‘ ‘”’Š‘‰‡• ƒ† •‹‰ƒŽ‹‰ ‘Ž‡ —Ž‡• •‡ ”‡–‡† „› ’ƒ––‡”‹‰

ϭϮϭ ‡–‡”• ƒŽŽ‘™ –Š‡ ‰”ƒ†‡† ‡š’”‡••‹‘ ‘ˆ –”ƒ• ”‹’–‹‘ ˆƒ –‘”• „› ‘”–‹ ƒŽ ’”‘‰‡‹–‘”• ƒ† –Š‡

ϭϮϮ †‡˜‡Ž‘’‡–‘ˆ–Š‡†‹ˆˆ‡”‡– ‘”–‹ ƒŽˆ‹‡Ž†•ȋ̵‡ƒ”›‡–ƒŽǤǡʹͲͲ͹Ǣ”ƒ‹ƒ†‹‡”ƒ‹ǡʹͲͳͶȌǤŠ”‡‡

ϯ 

ϭϮϯ ’ƒ––‡”‹‰ ‡–‡”•Ž‹‡ƒ––Š‡„‘”†‡”•‘ˆ–Š‡–‡Ž‡ ‡’ŠƒŽ‘ƒ†’ƒ”–‹ ‹’ƒ–‡‹–Š‡ƒ”‡ƒŽ‹œƒ–‹‘‘ˆ

ϭϮϰ –Š‡ ‘”–‡šǣ–Š‡ ‘”–‹ ƒŽŠ‡ȋ„‡–™‡‡ ‘”–‹ ƒŽƒ† Š‘”‘‹†ƒŽˆ‹‡Ž†•Ȍǡ–Š‡ ‘‹••—”ƒŽ’Žƒ–‡ȋƒ–

ϭϮϱ –Š‡”‘•–”‘‡†‹ƒŽ’‘Ž‡‘ˆ–Š‡–‡Ž‡ ‡’ŠƒŽ‘Ȍǡƒ†–Š‡ ‘”–‹ ƒŽƒ–‹Š‡ȋƒ––Š‡’ƒŽŽ‹ƒŽǦ•—„’ƒŽŽ‹ƒŽ

ϭϮϲ „‘—†ƒ”›ǡ Ȍ ȋƒŽŽƒƒ ‹ ƒ† –‘›‘˜ƒǡ ʹͲͲ͸ȌǤ Š‡   Š‘‡‘„‘š’”‘–‡‹ʹȋ ʹȌǡ

ϭϮϳ ‡š’”‡••‡†‹ƒ”‘•–”‘ǦŽƒ–‡”ƒŽ –‘ ƒ—†‘Ǧ‡†‹ƒŽ‰”ƒ†‹‡–ǡ•—’’”‡••‡•Š‡ƒ†ƒ–‹Š‡ˆƒ–‡•ǡ

ϭϮϴ „‘–Š•–”— –—”‡•„‡‹‰‡š’ƒ†‡†‹Lhx2—–ƒ–‹ ‡ȋ—Ž Šƒ†‡–ƒŽǤǡʹͲͲͳǢ‘—‹‡–ƒŽǤǡʹͲͲͳǢ

ϭϮϵ ƒƒ‰ƒ™ƒƒ†̵‡ƒ”›ǡʹͲͲͳǢƒ‰ƒŽ‡‡–ƒŽǤǡʹͲͲͺȌǤŠ‡–”ƒ• ”‹’–‹‘ˆƒ –‘”Foxg1ǡ‡š’”‡••‡†‹

ϭϯϬ ƒ”‘•–”‘ǦŽƒ–‡”ƒŽ –‘ ƒ—†‘Ǧ‡†‹ƒŽ‰”ƒ†‹‡–ǡ•—’’”‡••‡•Š‡ˆƒ–‡ƒ†‹•”‡“—‹”‡†ˆ‘”Žƒ–‡”ƒŽ

ϭϯϭ ˆƒ–‡•ǡ‹ Ž—†‹‰–Šƒ–‘ˆ–Š‡ƒ–‹Š‡ȋ‘—‡–ƒŽǤǡͳͻͻͻǢ—œ‹‘ƒ†ƒŽŽƒƒ ‹ǡʹͲͲͷǢ ƒƒ•Š‹ƒ‡–

ϭϯϮ ƒŽǤǡʹͲͲ͹ǢŠ‹„ƒ–ƒ‡–ƒŽǤǡʹͲͲͺȌǤŠ‡ ‘”–‹ ƒŽƒ–‹Š‡‹•‘ˆƒŒ‘”‹’‘”–ƒ ‡ˆ‘”–Š‡•’‡ ‹ˆ‹ ƒ–‹‘

ϭϯϯ ‘ˆ–Š‡‘Žˆƒ –‘”› ‘”–‡šǤ –Š‡ˆ‘ŽŽ‘™‹‰•‡ –‹‘ǡ™‡™‹ŽŽ†‹• —••–Š‡ƒ–‹Š‡ •’‡ ‹ˆ‹ ƒ–‹‘ƒ†

ϭϯϰ ˆ— –‹‘•†—”‹‰†‡˜‡Ž‘’‡–ȋ•‡‡ȋ̵‡ƒ”›‡–ƒŽǤǡʹͲͲ͹Ǣ—„”ƒƒ‹ƒ‡–ƒŽǤǡʹͲͲͻǢ‘–‹‡Žƒ†

ϭϯϱ „‘‹–‹œǡʹͲͳͷȌˆ‘”ˆ—”–Š‡”‹ˆ‘”ƒ–‹‘ƒ„‘—–‘”‰ƒ‹œ‹‰ ‡–‡”•ƒ†ƒ”‡ƒŽ‹œƒ–‹‘‘ˆ–Š‡ ‡”‡„”ƒŽ

ϭϯϲ ‘”–‡šȌǤ

ϭϯϳ The antihem, major signaling center for the determination of the olfactory cortex

ϭϯϴ Š‡ ƒ–‹Š‡ ‹• Ž‘ ƒ–‡† ƒ– –Š‡ ǡ „‡–™‡‡ –Š‡ ˜‡–”ƒŽ ’ƒŽŽ‹— ƒ† –Š‡ †‘”•ƒŽ Žƒ–‡”ƒŽ

ϭϯϵ ‰ƒ‰Ž‹‘‹ ‡‹‡ ‡ȋ† Ȍȋ—‡–ƒŽǤǡʹͲͲͳȌǤŠ‡ƒ–‹Š‡‹•†‡Ž‹‡ƒ–‡†ˆ”‘ƒ†Œƒ ‡–”‡‰‹‘•

ϭϰϬ –Š”‘—‰Š–Š‡‡š Ž—•‹˜‡‡š’”‡••‹‘‘ˆ–Š‡–”ƒ• ”‹’–‹‘ˆƒ –‘”Dbx1‹–Š‡˜‡–”ƒŽ’ƒŽŽ‹—ǡƒ•

ϭϰϭ ™‡ŽŽƒ•–Š‡‡”‹ Š‡†‡š’”‡••‹‘‘ˆ–Š‡•‡ ”‡–‡† ”‹œœŽ‡†”‡Žƒ–‡†‰‡‡sFrp2ȋ‹‡–ƒŽǤǡʹͲͲͳǢ—

ϭϰϮ ‡–ƒŽǤǡʹͲͲͳǢ••‹ƒ ‘’‘—Ž‘•‡–ƒŽǤǡʹͲͲ͵Ǣ‡†‹ƒ‡–ƒŽǤǡʹͲͲͶȌǤ Š‡ ˜‡–”ƒŽ ’ƒŽŽ‹— ƒ† –Š‡

ϭϰϯ ƒ†Œƒ ‡–† „‘–Š†‹•’Žƒ›ƒ‡”‹ Š‡†‡š’”‡••‹‘‘ˆPax6–”ƒ• ”‹’–‹‘ˆƒ –‘”ǡ™Š‹ Š‹•

ϭϰϰ ”‡“—‹”‡† ˆ‘” –Š‡ †‡˜‡Ž‘’‡– ‘ˆ Žƒ–‡”ƒŽ ƒ† ˜‡–”ƒŽ ’ƒŽŽ‹ƒŽ ‹†‡–‹–‹‡• ȋ‘Žˆƒ –‘”› ‘”–‡š ƒ†

ϭϰϱ ƒ›†‰ƒŽ‘‹† ‘’Ž‡šȌȋ–‘›‘˜ƒ‡–ƒŽǤǡͳͻͻ͸Ǣ‘”‡••‘‡–ƒŽǤǡʹͲͲͲǢ‹‡–ƒŽǤǡʹͲͲͳǢ—‡–ƒŽǤǡ

ϭϰϲ ʹͲͲͳǢ ‹”ƒ–ƒ‡–ƒŽǤǡʹͲͲʹǢ‘Ž‡ǡʹͲͲͷǢ‹‘‡–ƒŽǤǡʹͲͲͺǢ‘ ƒ•‡–ƒŽǤǡʹͲͳͳȌǤ

ϭϰϳ Š‡’‘•‹–‹‘ƒ†–Š‡•’‡ ‹ˆ‹ ƒ–‹‘‘ˆ–Š‡ƒ–‹Š‡”‡Ž›‘–Š‡‡š’”‡••‹‘‘ˆ–”ƒ• ”‹’–‹‘

ϭϰϴ ˆƒ –‘”•Pax6ǡTlxƒ†Gsh2Ǥ Pax6—–ƒ–‹ ‡ǡ–Š‡˜‡–”ƒŽƒ†–Š‡Žƒ–‡”ƒŽ’ƒŽŽ‹—‡ –‘’‹ ƒŽŽ›

ϰ 

ϭϰϵ ‡š’”‡•••—„’ƒŽŽ‹ƒŽƒ”‡”•ǡ•— Šƒ•Mash1ǡGsh2ǡƒ†Dlx2ȋ–‘›‘˜ƒ‡–ƒŽǤǡͳͻͻ͸Ǣͳͻͻ͹ǢʹͲͲͲǢ

ϭϱϬ ‘”‡••‘ ‡– ƒŽǤǡ ʹͲͲͲǢ ‹ ‡– ƒŽǤǡ ʹͲͲͳǢ — ‡– ƒŽǤǡ ʹͲͲͳȌǤ Tlx —–ƒ–• †‹•’Žƒ› •‹‹Žƒ” „—– Ž‡••

ϭϱϭ †”ƒƒ–‹ ’Š‡‘–›’‡ȋ–‡ƒǡʹͲͲ͵ȌǤ Ž‹‡™‹–ŠGsh2 ‡š’”‡••‹‘ ‹ –Š‡ •—„’ƒŽŽ‹—ǡ Gsh2

ϭϱϮ —–ƒ–•†‹•’Žƒ›’ƒŽŽ‹ƒŽ‰‡‡‡š’”‡••‹‘‹•—„’ƒŽŽ‹ƒŽ†‘ƒ‹•ǡ•— Šƒ•–Š‡† ȋ‘”‡••‘‡–ƒŽǤǡ

ϭϱϯ ʹͲͲͲǢ—‡–ƒŽǤǡʹͲͲͳȌǤ͸‹•–Š‡”‡ˆ‘”‡”‡“—‹”‡†ˆ‘”–Š‡‡š’”‡••‹‘‘ˆ˜‡–”ƒŽ’ƒŽŽ‹—•’‡ ‹ˆ‹ 

ϭϱϰ ‰‡‡•ǡ™Š‹Ž‡  ʹ•—’’”‡••‡•–Š‡‹”‡š’”‡••‹‘‹–Š‡•—„’ƒŽŽ‹—ȋƒ”‡›‡–ƒŽǤǡʹͲͲͻȌǤ

ϭϱϱ Š‡‘”‰ƒ‹œ‡”ˆ— –‹‘‘ˆ–Š‡ƒ–‹Š‡‹•’‘‘”Ž›—†‡”•–‘‘†ƒ• ‘’ƒ”‡†–‘–Šƒ–‘ˆ–Š‡

ϭϱϲ ‘”–‹ ƒŽŠ‡‹Š‹’’‘ ƒ’—•†‡˜‡Ž‘’‡–ȋ—„”ƒƒ‹ƒ‡–ƒŽǤǡʹͲͲͻȌǤ’”‘‹‡–’ƒŽ‹•ƒ†‡‘ˆ

ϭϱϳ ”ƒ†‹ƒŽ‰Ž‹ƒŽˆ‹„‡”•†‡Ž‹‡ƒ–‡•–Š‡ǤŠ‡•‡ˆ‹„‡”•‘”‹‰‹ƒ–‡‹–Š‡ ‘”–‹ ‘Ǧ•–”‹ƒ–ƒŽŒ— –‹‘‘ˆ–Š‡

ϭϱϴ ƒ†‡š–‡†—’–‘–Š‡’‹ƒŽ•—”ˆƒ ‡‹–Š‡’‹”‹ˆ‘” ‘”–‡šȋ‘Žž”ƒ†—–Ž‡”ǡʹͲͲʹȌǤ ƒ„•‡ ‡

ϭϱϵ ‘ˆ–Š‡ƒ–‹Š‡‘”‹„‘–ŠPax6ƒ†Tlx—–ƒ–•ǡ–Š‡”ƒ†‹ƒŽ‰Ž‹ƒŽ’ƒŽ‹•ƒ†‡‹••‡˜‡”‡Ž›ƒˆˆ‡ –‡†ƒ–

ϭϲϬ –Š‡ Ǥ ‘”‡‘˜‡”ǡ Pax6 —–ƒ–• •Š‘™ Š‹‰Š‡” —„‡” ‘ˆ •—„’ƒŽŽ‹— †‡”‹˜‡†Ǧ‹–‡”‡—”‘• ‹

ϭϲϭ –Š‡ ‘”–‡šǡ •—‰‰‡•–‹‰ ƒ ”‘Ž‡ ‘ˆ –Š‡ ”ƒ†‹ƒŽ ‰Ž‹ƒŽ ’ƒŽ‹•ƒ†‡ ‹ ”‡•–”‹ –‹‰ –ƒ‰‡–‹ƒŽ ‹‰”ƒ–‹‘ ‘ˆ

ϭϲϮ ‹–‡”‡—”‘• †—”‹‰ †‡˜‡Ž‘’‡– ȋŠƒ’‘—–‘ ‡– ƒŽǤǡ ͳͻͻͻȌǤ  ƒ††‹–‹‘ǡ ™Š‹Ž‡ –Š‡ ƒ–‹Š‡

ϭϲϯ ‡š’”‡••‡•–•‹‰ƒŽ‹‰‹Š‹„‹–‘”sFrp2ǡWnt7b‹•‡š’”‡••‡†‹–Š‡† ƒ†Œƒ ‡––‘–Š‡ƒ–‹Š‡

ϭϲϰ ȋ‹ ‡– ƒŽǤǡ ʹͲͲͳǢ ••‹ƒ ‘’‘—Ž‘• ‡– ƒŽǤǡ ʹͲͲ͵ȌǤ Š‹• ”‡•–”‹ –‹‘ ‘ˆ –Š‡ – •‹‰ƒŽ‹‰ –‘ –Š‡

ϭϲϱ •—„’ƒŽŽ‹ƒŽ•‹†‡‘ˆ–Š‡‹•–”— –•–Š‡’‘•‹–‹‘‘ˆ–Š‡”ƒ†‹ƒŽ‰Ž‹ƒŽ’ƒŽ‹•ƒ†‡ǤŠ‡ƒ–‹Š‡ˆ—”–Š‡”

ϭϲϲ ‡š’”‡••‡• •’‡ ‹ˆ‹  ‘Ž‡ —Ž‡•ǡ Ž‹‡ –Š‡ ‡’‹†‡”ƒŽ ‰”‘™–Š ˆƒ –‘” ȋ Ȍ ˆƒ‹Ž› ‡„‡”•  Ƚǡ

ϭϲϳ ‡—”‡‰—Ž‹•ͳƒ†͵ȋ ͳȀ ͵Ȍǡƒ†ˆ‹„”‘„Žƒ•–‰”‘™–Šˆƒ –‘” ͹ ȋ‹ ‡– ƒŽǤǡ ʹͲͲͳǢ

ϭϲϴ ••‹ƒ ‘’‘—Ž‘• ‡– ƒŽǤǡ ʹͲͲ͵ȌǤ  ͳ Šƒ• „‡‡ •Š‘™ –‘ „‡ ‡••‡–‹ƒŽ ‹ –Š‡ ˆ‘”ƒ–‹‘ ƒ†

ϭϲϵ ƒ‹–‡ƒ ‡‘ˆ–Š‡”ƒ†‹ƒŽ‰Ž‹ƒŽ ‡ŽŽ•ȋ–‘‡–ƒŽǤǡͳͻͻ͹Ǣ Š‹†‡–ƒŽǤǡʹͲͲ͵ȌǤ

ϭϳϬ Signaling centers and production of different lineages of pioneer Cajal-Retzius cells

ϭϳϭ Š‡–Š”‡‡‘”‰ƒ‹œ‹‰ ‡–‡”•ƒ––Š‡’ƒŽŽ‹ƒŽ„‘”†‡”•ƒ”‡‘™–‘„‡–Š‡ƒ‹‘”‹‰‹•‘ˆ

ϭϳϮ ƒŒƒŽǦ‡–œ‹—•ȋȌ ‡ŽŽ•ȋƒ‹‰— Š‹Ǧ ƒ›ƒ•Š‹ǡʹͲͲͶǢ‹‡ŽŽ‡‡–ƒŽǤǡʹͲͲͷǢ‘•Š‹†ƒǡʹͲͲ͸ǢŠƒ‘‡–ƒŽǤǡ

ϭϳϯ ʹͲͲ͸ȌǤŠ‡•‡‡ƒ”Ž›Ǧ„‘”’ƒŽŽ‹ƒŽ ‡ŽŽ•†‹ˆˆ‡”‡–‹ƒ–‡„‡–™‡‡ͳͲǤͷƒ†ͳʹǤͷ‹‹ ‡ƒ†’‘’—Žƒ–‡

ϭϳϰ –Š‡ƒ”‰‹ƒŽœ‘‡‘ˆƒŽŽ ‘”–‹ ƒŽƒ”‡ƒ•ȋƒ”–ƒ†ƒ”–ǡͳͻ͹͹Ǣ‘‘†‡–ƒŽǤǡͳͻͻʹǢƒ”ÀǦƒ†‹ŽŽƒǡ

ϱ 

ϭϳϱ ͳͻͻͺȌǤƒ›  ‡ŽŽ•‡š’”‡•• Emx1ƒ†Tbr1’ƒŽŽ‹ƒŽ–”ƒ• ”‹’–‹‘ˆƒ –‘”•ȋ ‘”•‹‡–ƒŽǤǡʹͲͲʹǢ

ϭϳϲ ‡˜‡”‡–ƒŽǤǡʹͲͲ͵Ȍǡƒ†‡‡Ž‹ǡƒ•‡ ”‡–‡†‰Ž› ‘’”‘–‡‹–Šƒ–‰—‹†‡• ”ƒ†‹ƒŽ ‹‰”ƒ–‹‘ ‘ˆ

ϭϳϳ ‡‘ ‘”–‹ ƒŽ ‡—”‘• ȋƒ˜‹‡••ǡ ͳͻͺʹǢ ‘™‡ŽŽ ‡– ƒŽǤǡ ͳͻͻ͹Ǣ Ž ƒ–ƒ”ƒ‡–ƒŽǤǡͳͻͻͺȌǤŽŽ ‡ŽŽ•

ϭϳϴ ‡š’”‡••Reelinǡ„—–‘Ž›Š‡Ž‹‡ƒ‰‡†‡”‹˜‡†Ǧ ‡ŽŽ•‡š’”‡••–Š‡–—‘”’”‘–‡‹͹͵ȋp73Ȍǡ™Š‹Ž‡

ϭϳϵ ƒ–‹Š‡‘”•‡’–—†‡”‹˜‡†Ǧ ‡ŽŽ••’‡ ‹ˆ‹ ƒŽŽ›‡š’”‡••Dbx1ǤŠ‹•†‹˜‡”•‹–›‘ˆ ‡ŽŽŽ‹‡ƒ‰‡•

ϭϴϬ ƒ›’Žƒ›ƒ”‘Ž‡‹–Š‡†‡˜‡Ž‘’‡–‘ˆ ›–‘ƒ” Š‹–‡ –‘‹ †‹ˆˆ‡”‡ ‡•„‡–™‡‡–Š‡‡‘ ‘”–‡šǡ–Š‡

ϭϴϭ ‘Žˆƒ –‘”› ‘”–‡šƒ†–Š‡Š‹’’‘ ƒ’—•ȋ‹‡ŽŽ‡‡–ƒŽǤǡʹͲͲͷȌǤ

ϭϴϮ Migration and layer formation

ϭϴϯ Migration and layer formation in the

ϭϴϰ ‡—”‘‹‰”ƒ–‹‘‹–Š‡‡‘ ‘”–‡šŠƒ•„‡‡™‡ŽŽ•–—†‹‡†Ǥ ”‘ͳͳǡ–Š‡ˆ‹”•–‡—”‘„Žƒ•–•

ϭϴϱ ƒ ——Žƒ–‡ •—’‡”ˆ‹ ‹ƒŽŽ›‹–Š‡‡‘ ‘”–‹ ƒŽ‡—”‘‡’‹–Š‡Ž‹—ǡˆ‘”‹‰ –Š‡’”‡’Žƒ–‡„‡‡ƒ–Š–Š‡

ϭϴϲ  ‡ŽŽ Žƒ›‡” ȋƒ”‹Ǧƒ†‹ŽŽƒǡ ͳͻ͹ͺǢ –‡™ƒ”– ƒ† ‡ƒ”Žƒǡ ͳͻͻͲǢ ƒ”„‘—šǦ²‡ ƒ† ƒ•’ƒ”ǡ

ϭϴϳ ʹͲͲͺǢ ‹–ƒŽ‹• ƒ† ‘••‹‡”ǡ ʹͲͳͲȌǤ Š‡ ’”‡’Žƒ–‡ ‹• ƒ †‡˜‡Ž‘’‡–ƒŽ –”ƒ•‹–‘”› •–”— –—”‡ ‹ –Š‡

ϭϴϴ ‡‘ ‘”–‡š‘ˆ‘—•‡‡„”›‘•Ǥš‘•‘ˆ’”‡’Žƒ–‡‡—”‘•ƒ”‡–Š‘—‰Š––‘’Žƒ›ƒ”‘Ž‡‘ˆ’‹‘‡‡”•

ϭϴϵ ˆ‘” –Š‡ †‡˜‡Ž‘’‡– ‘ˆ •‡˜‡”ƒŽ ˆ‹„‡” –”ƒ –• ȋ—’°” ‡– ƒŽǤǡ ͳͻͻͺȌǤ ”‘ ͳ͵ǡ ‡™ ‡š ‹–ƒ–‘”›

ϭϵϬ ‡—”‘••‡––Ž‡–Š‡’”‡’Žƒ–‡ǡ†‹˜‹†‹‰‹–‹–™‘Žƒ›‡”•ǡ–Š‡ƒ”‰‹ƒŽœ‘‡ƒ––Š‡•—”ˆƒ ‡ƒ†–Š‡

ϭϵϭ •—„’Žƒ–‡ ‡ƒ” –Š‡ Ǥ ‹—Ž–ƒ‡‘—•Ž›ǡ ”ƒ†‹ƒŽ ‰Ž‹ƒ †‡˜‡Ž‘’• ƒ† ‹–• ’”‘ ‡••‡• ”‡ƒ Š –Š‡ ’‹ƒŽ

ϭϵϮ •—”ˆƒ ‡ȋƒ‹ ǡʹͲͲ͵Ǣ‘””‡ŽŽƒ† ږœǡʹͲͳͶȌǤŠ‡•‡”ƒ†‹ƒŽ‰Ž‹ƒŽ’”‘ ‡••‡•ƒŽŽ‘™–Š‡•—„•‡“—‡–

ϭϵϯ ‹‰”ƒ–‹‘‘ˆ‡—”‘„Žƒ•–•ǡ™Š‹ Š‹˜ƒ†‡–Š‡ ‘”–‹ ƒŽ’Žƒ–‡‹„‡–™‡‡–Š‡ƒ”‰‹ƒŽœ‘‡ƒ†–Š‡

ϭϵϰ •—„’Žƒ–‡ˆ”‘ͳ͵–‘ͳ͸ȋƒ”‹Ǧƒ†‹ŽŽƒǡͳͻ͹ͺǢ†‡ƒ”Ž‘•ƒ†‡ƒ”›ǡʹͲͲ͵ȌǤš ‹–ƒ–‘”›‡—”‘•

ϭϵϱ ‘ˆ ‘”–‹ ƒŽ Žƒ›‡”• ƒ”‡ –Š‡ ’”‘†— ‡† ‹ ƒ ‹•‹†‡Ǧ‘—– ƒ‡” ˆ”‘–Š‡†‡‡’–‘–Š‡•—’‡”ˆ‹ ‹ƒŽ

ϭϵϲ Žƒ›‡”•ǡ •— Š –Šƒ– Žƒ–‡Ǧ„‘” ‡—”‘• ‹‰”ƒ–‡ –Š”‘—‰Š‘—– ‡ƒ”Ž›Ǧ„‘” ‡—”‘• ȋ‘Ž›‡ƒ—š ‡– ƒŽǤǡ

ϭϵϳ ʹͲͲ͹Ǣƒ‹ ‡–ƒŽǤǡʹͲͲͻȌǤ –‡”‡—”‘•‘ˆ–Š‡‡‘ ‘”–‡šƒ”‡’”‘†— ‡† ‹ –Š‡ ‰ƒ‰Ž‹‘‹ 

ϭϵϴ ‡‹‡ ‡• ‘ˆ –Š‡ •—„’ƒŽŽ‹— ȋ‡†‹ƒŽǡ  Ǣ Žƒ–‡”ƒŽǡ  Ǣ ƒ† ƒ—†ƒŽǡ  Ȍ ȋ†‡”•‘ ‡– ƒŽǤǡ

ϭϵϵ ͳͻͻ͹Ȍǡƒ†‹‰”ƒ–‡ˆ‹”•––ƒ‰‡–‹ƒŽŽ›ƒ†–Š‡”ƒ†‹ƒŽŽ›–‘•‡––Ž‡–Š‡ ‘”–‹ ƒŽŽƒ›‡”•ǡ•–ƒ”–‹‰ˆ”‘

ϮϬϬ ͳ͵ ȋ ‹‰—”‡ ʹȌ ȋƒ”‹ ƒ† —„‡•–‡‹ǡ ʹͲͲͳǢ ±–‹ ‡– ƒŽǤǡ ʹͲͲ͸Ǣ ‡Žƒ ‡– ƒŽǤǡ ʹͲͳͳȌǤ  Ǧ

ϲ 

ϮϬϭ †‡”‹˜‡†‹–‡”‡—”‘•„‘”ƒ–†‹ˆˆ‡”‡–•–ƒ‰‡••‡––Ž‡†‹•–‹ –Žƒ›‡”•‘ˆ–Š‡‡‘ ‘”–‡šȋ‹›‘•Š‹ƒ†

ϮϬϮ ‹•Š‡ŽŽǡʹͲͳͳȌǤ

ϮϬϯ Early migration to the olfactory cortex

ϮϬϰ  ‘–”ƒ•––‘–Š‡‡‘ ‘”–‡šǡ–Š‡‘Žˆƒ –‘”› ‘”–‡š‹•Ž‘ ƒ–‡†˜‡–”‘ǦŽƒ–‡”ƒŽŽ›ƒ†–Š‡”‡ˆ‘”‡

ϮϬϱ †‘‡•‘–Ž‹‡–Š‡’ƒŽŽ‹ƒŽǤ‘’Ž‡š™ƒ˜‡•‘ˆ ‡ŽŽ‹‰”ƒ–‹‘ˆ”‘ ˜ƒ”‹‘—• ”‡‰‹‘• ‘ˆ –Š‡ 

ϮϬϲ –‘™ƒ”†–Š‡‘Žˆƒ –‘”› ‘”–‡šŠƒ˜‡„‡‡†‡• ”‹„‡†‹‡ƒ”Ž›‘—•‡†‡˜‡Ž‘’‡–ǡˆ”‘ͻǤ͹ͷ–‘ͳʹ

ϮϬϳ ȋ‘—”ƒ‡–ƒŽǤǡʹͲͲ͸Ǣ ƒ” ‹ƒǦ‘”‡‘‡–ƒŽǤǡʹͲͲͺȌǤŽˆƒ –‘”›‡—”‘•‰‡‡”ƒ–‡†‹–Š‡†‘”•ƒŽ’ƒ”–

ϮϬϴ ‘ˆ–Š‡–‡Ž‡ ‡’ŠƒŽ‘ƒ–ͻǤ͹ͷ‹‰”ƒ–‡˜‡–”ƒŽŽ›ƒ†ƒŽ‹‰–Š‡–Š”‘—‰Š’Š”‹Ǧͷ”‡’—Ž•‹˜‡

ϮϬϵ ƒ –‹˜‹–›ǡ™Š‹ Š‡š’”‡••‹‘‹•”‡‰—Žƒ–‡†„›Pax6–”ƒ• ”‹’–‹‘ˆƒ –‘”ǤŠ‡•‡˜‡–”ƒŽŽ›‹‰”ƒ–‹‰

ϮϭϬ ‡—”‘•†‹ˆˆ‡”‡–‹ƒ–‡„‘–Š‹Ž‘– ‡ŽŽ•ƒ†‘Žˆƒ –‘”› ‘”–‡š‡—”‘•ȋ‘—”ƒ‡–ƒŽǤǡʹͲͲ͸ȌǤ

Ϯϭϭ ‘– ‡ŽŽ•ƒ”‡ƒ•’‡ ‹ˆ‹ •—„’‘’—Žƒ–‹‘‘ˆͻǤͷ–‘ͳͳǤͷǦ„‘”‡‡Ž‹ΪȀ’͹͵Ϊ ‡ŽŽ•ǡƒ‡†

ϮϭϮ ƒˆ–‡”–Š‡‹”•’‡ ‹ˆ‹ ‡š’”‡••‹‘‘ˆlot1 ȋmGluR1ȾȌȋ‘–‡–Šƒ–Ž‘– ‡ŽŽ•ƒ”‡†‹•–‹‰—‹•Š‡†ˆ”‘

Ϯϭϯ ƒš‘•„›•ƒŽŽƒ† ƒ’‹–ƒŽŽ‡––‡”•ǡ”‡•’‡ –‹˜‡Ž›Ȍȋƒ–‘‡–ƒŽǤǡͳͻͻͺǢ ‹±‡œ‡–ƒŽǤǡʹͲͲʹǢ‹š‹–‡–ƒŽǤǡ

Ϯϭϰ ʹͲͳͶȌǤ ”‘‡—”ƒŽ ‰‡‡• Neurogenin1ƒ†Neurogenin2 ƒ”‡ ‘‡š’”‡••‡† ‹ †‘”•ƒŽ ƒ† ˜‡–”ƒŽ

Ϯϭϱ ’ƒŽŽ‹ƒŽ’”‘‰‡‹–‘”•ƒ†”‡“—‹”‡†ˆ‘”–Š‡†‹ˆˆ‡”‡–‹ƒ–‹‘‘ˆŽ‘– ‡ŽŽ•ȋ‹š‹–‡–ƒŽǤǡʹͲͳͶȌǤŠ‡•‡ ‡ŽŽ•

Ϯϭϲ ƒ”‡’”‘†— ‡†‹–Š‡™Š‘Ž‡‘ˆ–Š‡‡‘ ‘”–‡šˆ”‘ͻǤͷ–‘ͳͳǤͷǡƒ†‡š–‹‰”ƒ–‡–ƒ‰‡–‹ƒŽŽ›

Ϯϭϳ ƒ†˜‡–”ƒŽŽ›–‘™ƒ”†–Š‡ ‘”–‡š•—”ˆƒ ‡–‘‘”‰ƒ‹œ‡–Š‡•‡Ž˜‡•ƒ”‘—†–Š‡’”‡•—’–‹˜‡

Ϯϭϴ –‡””‹–‘”›‘ˆ–Š‡ƒ–ͳʹǤͷȋ‘‹‘ƒ‡–ƒŽǤǡʹͲͲͲȌǤŠ‡‹‰”ƒ–‹‘ƒ†’‘•‹–‹‘‘ˆŽ‘– ‡ŽŽ•”‡Ž›

Ϯϭϵ ‘ –Š‡ ‡š’”‡••‹‘ ‘ˆ ‰—‹†ƒ ‡ —‡•Ǥ Š‡ ‡‘ ‘”–‡š †‹•’Žƒ›• †‘”•‘Ǧ˜‡–”ƒŽ ‰”ƒ†‹‡–• ‘ˆ —‡•ǡ

ϮϮϬ ™Š‹ Š ’‘•‹–‹‘ –Š‡ ˆ‹”•– •–‡’• ‘ˆ Ž‘– ‡ŽŽ ‹‰”ƒ–‹‘Ǥ Š‡ •—„’ƒŽŽ‹— ‡š’”‡••‡• ”‡’—Ž•‹˜‡ —‡•ǡ

ϮϮϭ ™Š‹ Š’”‡˜‡–Ž‘– ‡ŽŽ•ˆ”‘‹˜ƒ†‹‰˜‡–”ƒŽ–‡””‹–‘”‹‡•ƒ†ƒ‹–ƒ‹–Š‡•—’‡”ˆ‹ ‹ƒŽŽ›ƒ––Š‡

ϮϮϮ Ǥ‘‰–Š‡•‡ —‡•ǡ‡–”‹ǦͳŠƒ•„‡‡•Š‘™–‘Ž‘ ƒŽŽ›ƒ––”ƒ –Ž‘– ‡ŽŽ•ƒ”‘—†–Š‡ǡ™Š‹Ž‡

ϮϮϯ ‡ƒ’Š‘”‹͵ ‹••‡ ”‡–‡†„›†‡‡’”‡‰‹‘•‘ˆ–Š‡˜‡–”ƒŽ–‡Ž‡ ‡’ŠƒŽ‘ƒ†‡‡’•Ž‘– ‡ŽŽ•ƒ––Š‡

ϮϮϰ ‡”‡„”ƒŽ •—”ˆƒ ‡ „› ‹–• ”‡’—Ž•‹˜‡ ƒ –‹‘ ȋƒ™ƒ•ƒ‹ǡ ʹͲͲ͸Ǣ –‘ ‡–ƒŽǤǡʹͲͲͺȌǤ‘– ‡ŽŽ•ƒ –ƒ•

ϮϮϱ ‰—‹†‡’‘•–•ˆ‘”–Š‡‡•–ƒ„Ž‹•Š‡–ƒ†’‘•‹–‹‘‹‰‘ˆƒš‘•ǡ™Š‹ Š‘ —”ˆ”‘ͳʹǤͷ–‘ͳ͵Ǥͷ

ϮϮϲ ‹‹ ‡ȋ“—ƒ”œ‘‹ǡʹͲͳͷȌǤ––Š‡•‡‡ƒ”Ž›•–ƒ‰‡•ǡŽ‘– ‡ŽŽ•†‹•’Žƒ›Ž‘‰’”‘ ‡••‡•ǡƒ†”‡•’‘†–‘

ϳ 

ϮϮϳ ‡Ž‡ –”‹ •–‹—Žƒ–‹‘‘ˆ–Š‡ȋƒ–‘‡–ƒŽǤǡͳͻͻͺǢ ‹”ƒ–ƒ‡–ƒŽǤǡʹͲͳʹȌǤš ‡’–ˆ‘”Ž‘– ‡ŽŽ•ǡƒ‰‡‡”ƒŽ

ϮϮϴ ”‘Ž‡‘ˆ ‡ŽŽ•ƒ•‰—‹†‡’‘•–•ˆ‘”ƒš‘†‡˜‡Ž‘’‡–•–‹ŽŽ”‡ƒ‹•–‘„‡‡Ž— ‹†ƒ–‡†ǡ„—– ‡ŽŽ•

ϮϮϵ Šƒ˜‡„‡‡‹˜‘Ž˜‡†‹–Š‡‰—‹†ƒ ‡‘ˆ‡–‘”Š‹ƒŽƒ†–ŠƒŽƒ‘ ‘”–‹ ƒŽƒš‘•ȋ‡ŽÀ‘‡–ƒŽǤǡʹͲͲʹǢ

ϮϯϬ ‡”ƒ‹‡–ƒŽǤǡʹͲͲ͸Ǣƒ”„‡”‡–ƒŽǤǡʹͲͳͷȌǤ‡ ‡–Ž›ǡ†‡ ”—–‘•ƒ† ‘ŽŽ‡ƒ‰—‡•†‹• ‘˜‡”‡†ƒŽƒ–‡”

Ϯϯϭ ƒ†—‡š’‡ –‡†”‘Ž‡ˆ‘”Ž‘– ‡ŽŽ•†—”‹‰†‡˜‡Ž‘’‡–ǣ„‡–™‡‡ͳ͵Ǥͷƒ†ͳͷǤͷǡŽ‘– ‡ŽŽ•”‡–”ƒ –

ϮϯϮ –Š‡‹”’”‘ ‡••‡•ǡƒ†‘’–ƒ”‘—†‡†Ǧ—’‘”’Š‘Ž‘‰›™‹–Š•ƒŽŽˆ‹Ž‘’‘†‹ƒƒ†‹‹–‹ƒ–‡ƒ‘‡„‘‹†ǦŽ‹‡

Ϯϯϯ ‹‰”ƒ–‹‘ƒ™ƒ›ˆ”‘–Š‡–‡””‹–‘”›–‘†‘”•ƒŽ‡‘ ‘”–‹ ƒŽ”‡‰‹‘•ȋ†‡ ”—–‘•‡–ƒŽǤǡʹͲͳ͸ȌǤŠ‹•

Ϯϯϰ ”‡•—Ž–• ‹ ƒ †‘—„Ž‡† †‡•‹–› ‘ˆ  ‡ŽŽ• ‹ –Š‡ ‡‘ ‘”–‡š ƒ† ƒ͸ͲΨ”‡†— –‹‘‘ˆŽ‘– ‡ŽŽ

Ϯϯϱ †‡•‹–›‹–Š‡–‡””‹–‘”›ǤŠ‹•”‡ƒŽŽ‘ ƒ–‹‘‘ˆŽ‘– ‡ŽŽ•‹•”‡‰—Žƒ–‡†„›‰Ž—–ƒƒ–‡

Ϯϯϲ ”‡ ‡’–‘”ǡ ƒ† ”‡“—‹”‡† ˆ‘” –Š‡ ’”‘’‡” ‘”‰ƒ‹œƒ–‹‘ ‘ˆ ‡‘ ‘”–‡šŽƒ›‡”ͳƒ†ˆ‘”Žƒ›‡”ʹȀ͵

Ϯϯϳ ’›”ƒ‹†ƒŽ ‡ŽŽƒ’‹ ƒŽ†‡†”‹–‡‘”’Š‘Ž‘‰›ƒ†•’‹‡†‡•‹–›ȋ†‡ ”—–‘•‡–ƒŽǤǡʹͲͳ͸ȌǤ‹ ‡–Š‡

Ϯϯϴ ‘Žˆƒ –‘”› ‘”–‡š’”‡†ƒ–‡•–Š‡‡‡”‰‡ ‡‘ˆ–Š‡‡‘ ‘”–‡š‹‡˜‘Ž—–‹‘ǡ–Š‡Ž‘– ‡ŽŽ”‡ƒŽŽ‘ ƒ–‹‘

Ϯϯϵ ‹–‘–Š‡‡‘ ‘”–‡š ‘—Ž†”‡ˆŽ‡ –ƒ‡˜‘Ž—–‹‘ƒ”› ‘Ǧ‘’–‹‘‘ˆ–Š‡•‡ƒ ‡•–”ƒŽ‰—‹†‡’‘•–•Ǥ –Š‡

ϮϰϬ ‡‘ ‘”–‡šǡ ‡ŽŽ•ƒ”‡’”‘‰”‡••‹˜‡Ž›‡Ž‹‹ƒ–‡†„›ƒ’‘’–‘•‹•ƒ††‹•ƒ’’‡ƒ”„›–Š‡‡†‘ˆ–Š‡

Ϯϰϭ •‡ ‘†’‘•–ƒ–ƒŽ™‡‡ȋ†‡Ž¿‘‡–ƒŽǤǡͳͻͻ͸ȌǤ –‹•›‡–— Ž‡ƒ”™Š‡–Š‡”Ž‘– ‡ŽŽ•”‡ƒ‹‹‰‹–Š‡

ϮϰϮ ‘Žˆƒ –‘”› ‘”–‡šƒ”‡‹–‡‰”ƒ–‡†‹–Š‡‘Žˆƒ –‘”›‡–™‘”‘”™Š‡–Š‡” –Š‡› ƒ”‡ ƒ –”ƒ•‹‡–

Ϯϰϯ ’‘’—Žƒ–‹‘ƒ•™‡ŽŽǤ

Ϯϰϰ ƒ” ‹ƒǦ‘”‡‘ƒ† ‘ŽŽ‡ƒ‰—‡•†‡• ”‹„‡†‘–Š‡”‹‰”ƒ–‹‘ƒ’•ƒ††‡•–‹ƒ–‹‘ƒ”‡ƒ•‘ˆ

Ϯϰϱ ‘Žˆƒ –‘”› ‘”–‡š ‡ŽŽ•„›‹ ‹Œ‡ –‹‘•ƒ–ͳͲǦͳʹˆ‘ŽŽ‘™‡†„›in toto‡„”›‘ —Ž–—”‡ˆ‘”ͳ†ƒ›

Ϯϰϲ ȋ ƒ” ‹ƒǦ‘”‡‘‡–ƒŽǤǡʹͲͲͺȌȋ ‹‰—”‡ͳȌǤ‡—”‘„Žƒ•–•ƒ”‡’”‘†— ‡†‹–Š‡’ƒŽŽ‹ƒŽǡ‹–Š‡†‘”•ƒŽ

Ϯϰϳ –‡Ž‡ ‡’ŠƒŽ‘ǡƒ† –Š‡ ”‘•–”‘Ǧ‡†‹ƒŽ–‡Ž‡ ‡’ŠƒŽ‹ ™ƒŽŽǡ ƒ• ™‡ŽŽƒ•‹ –Š‡ •—„’ƒŽŽ‹ƒŽǡ ‹–Š‡

Ϯϰϴ Žƒ–‡”ƒŽ ‰ƒ‰Ž‹‘‹  ‡‹‡ ‡ ƒ† –Š‡ •‡’–‘Ǧ‡‹‡–‹ƒŽ •—Ž —•Ǥ Š‡•‡ ‡—”‘„Žƒ•–• †‹•’Žƒ›

Ϯϰϵ –ƒ‰‡–‹ƒŽ‘””ƒ†‹ƒŽ‹‰”ƒ–‹‘ǡ•‘‡ ‡ŽŽ•†‹•’Žƒ›‹‰„‘–Š–›’‡•‘ˆ‹‰”ƒ–‹‘•‡“—‡–‹ƒŽŽ›ǤŠ‡

ϮϱϬ ƒ—–Š‘”••Š‘™‡†–Šƒ––Š‡ ‡ŽŽ†‡•–‹ƒ–‹‘ƒ†‹†‡–‹–›‹–Š‡‘Žˆƒ –‘”› ‘”–‡š†‹ˆˆ‡”†‡’‡†‹‰‘

Ϯϱϭ –Š‡‹”‘”‹‰‹‹–Š‡ȋ ‹‰—”‡ͳȌǤ—”’”‹•‹‰Ž›ǡ•‘‡‡—”‘•ˆ”‘ –Š‡ ’ƒŽŽ‹ƒŽ ”‘•–”‘‡†‹ƒŽ

ϮϱϮ –‡Ž‡ ‡’ŠƒŽ‹  ™ƒŽŽ  †‹ˆˆ‡”‡–‹ƒ–‡ ‹ ‹–‡”‡—”‘•ǡ ™Š‹Ž‡ ‘–Š‡”• ‰‡‡”ƒ–‡† ‹ –Š‡ †‘”•ƒŽ

Ϯϱϯ –‡Ž‡ ‡’ŠƒŽ‘†‹ˆˆ‡”‡–‹ƒ–‡‹‡š ‹–ƒ–‘”›‡—”‘•Ǥ ƒ††‹–‹‘ǡ‘Žˆƒ –‘”› ‘”–‡š‡—”‘•ˆ”‘–Š‡

ϴ 

Ϯϱϰ •—„’ƒŽŽ‹ƒŽ†‘”•ƒŽŽƒ–‡”ƒŽ‰ƒ‰Ž‹‘‹ ‡‹‡ ‡†‹ˆˆ‡”‡–‹ƒ–‡‹‡š ‹–ƒ–‘”›‡—”‘•ǡ™Š‹Ž‡‘–Š‡”•

Ϯϱϱ ‰‡‡”ƒ–‡†‹–Š‡•‡’–‘Ǧ‡‹‡–‹ƒŽ•—Ž —•ƒ‹Ž›†‹ˆˆ‡”‡–‹ƒ–‡‹‹–‡”‡—”‘•ȋ ƒ” ‹ƒǦ‘”‡‘‡–

Ϯϱϲ ƒŽǤǡʹͲͲͺȌȋ ‹‰—”‡ͳȌǤ ™Š‹ Š‘Žˆƒ –‘”› ‘”–‡šŽƒ›‡”–Š‡•‡‡ƒ”Ž› ‡ŽŽ•†‘•‡––Ž‡ƒ†™Šƒ–ƒ”‡–Š‡‹”

Ϯϱϳ •’‡ ‹ˆ‹  ˆ— –‹‘• ‹ –Š‡ ‹” —‹– ƒ”‡ ‘– ›‡– ‡Ž— ‹†ƒ–‡†Ǥ —”–Š‡” ‹˜‡•–‹‰ƒ–‹‘• ƒ”‡ ‡‡†‡† –‘

Ϯϱϴ †‡ ‹’Š‡”™Š‡–Š‡” ‡ŽŽ• ‘‹‰ˆ”‘†‹ˆˆ‡”‡–”‡‰‹‘•‹–Š‡‡š’”‡•••’‡ ‹ˆ‹ Ž‹‡ƒ‰‡ƒ”‡”•ǡ

Ϯϱϵ ™Š‹ Š ‘—Ž† Š‡Ž’ –‘ ‹†‡–‹ˆ› ‡ŽŽ ’‘’—Žƒ–‹‘• ƒ† –Š‡‹” •’‡ ‹ˆ‹  ˆ— –‹‘• ™‹–Š‹ –Š‡ ‘Žˆƒ –‘”›

ϮϲϬ ‘”–‡šǤ

Ϯϲϭ Late migration routes to the olfactory cortex

ϮϲϮ Š‡‘Žˆƒ –‘”› ‘”–‡š‡—”‘•„‘”‹–Š‡Žƒ–‡”ƒŽƒ†˜‡–”ƒŽ’ƒŽŽ‹—ƒ–‡„”›‘‹ •–ƒ‰‡•

Ϯϲϯ ͳʹ–‘ͳͷ‹”ƒ–‡„”›‘•ȋ ‘””‡•’‘†‹‰–‘ͳͳ–‘ͳͶ‹‹ ‡ȌŠƒ˜‡„‡‡‹‹–‹ƒŽŽ›†‡• ”‹„‡†–‘

Ϯϲϰ ‹‰”ƒ–‡”ƒ†‹ƒŽŽ›–‘™ƒ”†•–Š‡•—”ˆƒ ‡‘ˆ–Š‡˜‡–”‘Žƒ–‡”ƒŽ–‡Ž‡ ‡’ŠƒŽ‘ȋƒ›‡”ǡͳͻͺ͸ǢƒŽ˜‡”†‡

Ϯϲϱ ƒ†ƒ–ƒ ƒƒǡͳͻͻͶǢ†‡ƒ”Ž‘•‡–ƒŽǤǡͳͻͻ͸ȌǤŠ‡‘•–†‡• ”‹„‡†‹‰”ƒ–‘”›”‘—–‡ˆ‘”‘Žˆƒ –‘”›

Ϯϲϲ ‘”–‡š‡—”‘•‹•–Š‡Žƒ–‡”ƒŽ ‘”–‹ ƒŽ•–”‡ƒǤŠ‹•‹‰”ƒ–‘”›”‘—–‡‹•ˆ‘”‡†„› ‡ŽŽ•‹‰”ƒ–‹‰–‘

Ϯϲϳ –Š‡’‹”‹ˆ‘” ‘”–‡šǡ–Š‡‘Žˆƒ –‘”›–—„‡” Ž‡ǡƒ†–‘–Š‡‘Žˆƒ –‘”›ƒ›‰†ƒŽƒǤ‡—”‘„Žƒ•–•–ƒ‹‰–Š‹•

Ϯϲϴ ”‘—–‡ƒ”‡„‘”‹–Š‡ƒ––Š‡ƒ†‹‰”ƒ–‡–ƒ‰‡–‹ƒŽŽ›–‘–Š‡„”ƒ‹•—”ˆƒ ‡–Š”‘—‰Š˜‡–”ƒŽ

Ϯϲϵ ”‡‰‹‘•‘ˆ–Š‡–‡Ž‡ ‡’ŠƒŽ‘ǡƒ†–Š‡”ƒ†‹ƒŽŽ›–‘•‡––Ž‡–Š‡’‹”‹ˆ‘” ‘”–‡šȋ ‹‰—”‡ʹȌǤ Pax6

ϮϳϬ —–ƒ–‹ ‡ȋ™Š‹ Š†‹•’Žƒ›ƒ‹’ƒ‹”‡†ˆ‘”ƒ–‹‘Ȍǡ–Š‡”‘—–‹‰‘ˆ–Š‡Žƒ–‡”ƒŽ ‘”–‹ ƒŽ

Ϯϳϭ •–”‡ƒ–‘™ƒ”†•–Š‡ƒ›‰†ƒŽƒƒ†–Š‡‘Žˆƒ –‘”› ‘”–‡š‹•ƒŽ–‡”‡†ȋŠƒ’‘—–‘‡–ƒŽǤǡͳͻͻͻǢ‘Ž‡ǡ

ϮϳϮ ʹͲͲͷȌǤ  ‘–”ƒ•––‘‘–Š‡”‘™–ƒ‰‡–‹ƒŽ‹‰”ƒ–‘”›•–”‡ƒ•†‡• ”‹„‡† ˆ‘” ‹–‡”‡—”‘

Ϯϳϯ ‹‰”ƒ–‹‘ǡ –Š‡ ‡—”‘„Žƒ•–• ‘ˆ –Š‡ Žƒ–‡”ƒŽ ‘”–‹ ƒŽ •–”‡ƒ ƒ‹Ž› †‹ˆˆ‡”‡–‹ƒ–‡ ‹–‘ ‡š ‹–ƒ–‘”›

Ϯϳϰ ‡—”‘•‹–Š‡’‹”‹ˆ‘” ‘”–‡šȋ‘”„‹‡–ƒŽǤǡʹͲͲͳȌǤƒ‹ƒ† ‘ŽŽ‡ƒ‰—‡••–—†‹‡†–Š‹•‹‰”ƒ–‘”›

Ϯϳϱ •–”‡ƒ‹”ƒ–‡„”›‘•„›in utero‡Ž‡ –”‘’‘”ƒ–‹‘‘ˆƒ’Žƒ•‹†ƒŽŽ‘™‹‰ ‡š’”‡••‹‘„›–Š‡

Ϯϳϲ ‡ŽŽ•„‘”ƒ–ͳ͵‹–Š‡ƒ––Š‡Ž‡˜‡Ž‘ˆ–Š‡ȋƒ‹‡–ƒŽǤǡʹͲͲͺȌǤˆ–‡”͵†ƒ›•ǡ–Š‡ ‡ŽŽ•‹‰”ƒ–‡

Ϯϳϳ –ƒ‰‡–‹ƒŽŽ› –‘™ƒ”† ˜‡–”ƒŽ ”‡‰‹‘• ƒŽ‘‰ –Š‡ Žƒ–‡”ƒŽ ‘”–‹ ƒŽ •–”‡ƒǤ Š‡•‡ ‡ŽŽ• •‡––Ž‡ –Š‡

Ϯϳϴ ’‹”‹ˆ‘” ‘”–‡šƒˆ–‡”͸†ƒ›•ǡƒ†ƒ‹Ž›†‹ˆˆ‡”‡–‹ƒ–‡‹–‘‡—”‘ƒŽ–”ƒ• ”‹’–‹‘ˆƒ –‘”ͳΪ

Ϯϳϵ ‡ŽŽ•ȋ ‡˜‡”‡–ƒŽǤǡʹͲͲͳǢƒ‹‡–ƒŽǤǡʹͲͲͺȌǤ ‘™‡˜‡”ƒ”‡›ƒ† ‘ŽŽ‡ƒ‰—‡• •Š‘™‡† –Šƒ– –Š‡

ϵ 

ϮϴϬ ‹†‡–‹–›‘ˆ ‡ŽŽ•‹‰”ƒ–‹‰ƒŽ‘‰–Š‡Žƒ–‡”ƒŽ ‘”–‹ ƒŽ•–”‡ƒ‹• ‘– Š‘‘‰‡‘—• ȋƒ”‡› ‡– ƒŽǤǡ

Ϯϴϭ ʹͲͲ͸ȌǤŠ‡ƒ—–Š‘”•†‡• ”‹„‡†–™‘’‘’—Žƒ–‹‘•‘ˆ’”‘‰‡‹‡•‹‰”ƒ–‹‰ƒŽ‘‰–Š‡Žƒ–‡”ƒŽ ‘”–‹ ƒŽ

ϮϴϮ •–”‡ƒ ‹ ‘—•‡ ‡„”›‘•ǣ ͸Ϊ’”‘‰‡‹‡•ǡ™Š‹ Š†‡”‹˜‡ˆ”‘–Š‡’ƒŽŽ‹—ǡƒ†ʹΪ

Ϯϴϯ ’”‘‰‡‹‡•ǡ™Š‹ Š†‡”‹˜‡ˆ”‘–Š‡•—„’ƒŽŽ‹—ǤŠ‡•‡–™‘’‘’—Žƒ–‹‘•ƒ”‡‰‡‡”ƒ–‡†ƒ–’”‡ ‹•‡

Ϯϴϰ ƒ††‹•–‹ ––‡’‘”ƒŽ™‹†‘™•†—”‹‰†‡˜‡Ž‘’‡–Ǥ ”‘ͳͳǤͷǡPax6‹•Š‹‰ŠŽ›‡š’”‡••‡†‹–Š‡

Ϯϴϱ ’ƒŽŽ‹—ƒ†ƒ––Š‡‘ˆ–Š‡Ǥ––Š‹••–ƒ‰‡ǡ•‘‡͸Ϊ ‡ŽŽ•ƒ”‡˜‹•‹„Ž‡ƒŽ‘‰–Š‡Žƒ–‡”ƒŽ

Ϯϴϲ ‘”–‹ ƒŽ•–”‡ƒƒ†‹–Š‡’‹”‹ˆ‘” ‘”–‡šǤ–ͳͷǤͷǡ͸Ϊ ‡ŽŽ••‡––Ž‡–Š‡’‹”‹ˆ‘” ‘”–‡šƒ†–Š‡

Ϯϴϳ ƒ›‰†ƒŽ‘‹† ‘’Ž‡šǤʹΪ ‡ŽŽ•ƒ”‘•‡ˆ”‘–Š‡ƒ„‘—––™‘†ƒ›•ƒˆ–‡”͸Ϊ ‡ŽŽ•ǡƒ–ͳ͵ǤͷǤ

Ϯϴϴ ʹΪ ‡ŽŽ•ƒ”‡ƒ –‹˜‡Ž›‹‰”ƒ–‹‰ˆ”‘ͳͷǤͷǡƒ†„›ͳͺǤͷ–Š‡›ƒ ——Žƒ–‡ ‹ –Š‡ ’‹”‹ˆ‘”

Ϯϴϵ ‘”–‡šƒ†–Š‡‘Žˆƒ –‘”›ƒ›‰†ƒŽƒǤ͸Ϊ ‡ŽŽ•†‹•’Žƒ›ƒ‹‰”ƒ–‹‘ƒŽ‘‰ Ϊ ”ƒ†‹ƒŽ ‰Ž‹ƒŽ

ϮϵϬ ’”‘ ‡••‡•ˆ‘—†„‡–™‡‡–Š‡‘ˆ–Š‡ƒ†–Š‡’‹ƒŽ•—”ˆƒ ‡‘ˆ –Š‡ ˜‡–”‘ǦŽƒ–‡”ƒŽ

Ϯϵϭ –‡Ž‡ ‡’ŠƒŽ‘ƒ–ͳ͵Ǥͷƒ†ͳͷǤͷȋƒ”‡›‡–ƒŽǤǡʹͲͲ͸Ȍȋ ‹‰—”‡ʹȌǤ ƒ”ƒ†‘š‹ ƒŽŽ›ǡ –Š‡•‡ ‡ŽŽ•

ϮϵϮ †‹•’Žƒ›ƒ–ƒ‰‡–‹ƒŽ‹‰”ƒ–‹‘ƒŽ‘‰”ƒ†‹ƒŽ‰Ž‹ƒŽ’”‘ ‡••‡•Ǥ‘‡ʹΪ ‡ŽŽ•†‹•’Žƒ›‹‰”ƒ–‹‘

Ϯϵϯ ƒŽ‘‰ ”ƒ†‹ƒŽ ‰Ž‹ƒŽ ’”‘ ‡••‡• ƒ• ™‡ŽŽǡ „—– Dz‹ Šƒ‹dz ‹‰”ƒ–‹‘ Šƒ• ƒŽ•‘ „‡‡ ‘„•‡”˜‡† ‹ –Š‹•

Ϯϵϰ ’‘’—Žƒ–‹‘ȋƒ”‡›‡–ƒŽǤǡʹͲͲ͸Ȍȋ ‹‰—”‡ʹȌǤŠ‡ƒ—–Š‘”•Š›’‘–Š‡•‹œ‡ –Šƒ– ͸Ϊ ‡ŽŽ•

Ϯϵϱ †‹ˆˆ‡”‡–‹ƒ–‡‹‡š ‹–ƒ–‘”›‡—”‘•ǡ™Š‹Ž‡ʹΪ ‡ŽŽ•†‹ˆˆ‡”‡–‹ƒ–‡‹‹Š‹„‹–‘”›‡—”‘•ǡ•‹ ‡

Ϯϵϲ ʹΪ ‡ŽŽ•‡š’”‡••–Š‡‹–‡”‡—”‘ƒ”‡”ƒŽ„‹†‹ȋ Šƒ‡‡–ƒŽǤǡʹͲͲ͹ȌǤ

Ϯϵϳ Šƒ‘ƒ† ‘ŽŽ‡ƒ‰—‡•—”ƒ˜‡Ž‡†‘˜‡Ž†‹‡ ‡’ŠƒŽ‘Ǧ–‘Ǧ–‡Ž‡ ‡’ŠƒŽ‘‹‰”ƒ–‹‘•‹–‘–Š‡

Ϯϵϴ •‡’–—ǡ„—–ƒŽ•‘–Š‡’‹”‹ˆ‘” ‘”–‡šƒ†–Š‡ƒ›‰†ƒŽƒȋŠƒ‘‡–ƒŽǤǡʹͲͲͺȌǤ›‰‡‡–‹ ƒŽŽ›Žƒ„‡Ž‹‰

Ϯϵϵ –Š‡ Foxb1†‹‡ ‡’ŠƒŽ‹ Ž‹‡ƒ‰‡ǡ–Š‡ƒ—–Š‘”•‹†‡–‹ˆ‹‡†Žƒ„‡Ž‡† ‡ŽŽ•ˆ”‘ –Š‡ ƒ—†ƒŽ

ϯϬϬ Š›’‘–ŠƒŽƒ—•ǡ ™Š‹ Š ‹‰”ƒ–‡ ‹–‘ ˜‡–”ƒŽ Ž‡˜‡Ž• ‘ˆ –Š‡ –‡Ž‡ ‡’ŠƒŽ‘ –Š”‘—‰Š –Š‡ ‘–‹—‹–›

ϯϬϭ „‡–™‡‡ –Š‡ ˜‡–”ƒŽ †‹‡ ‡’ŠƒŽ‘ ƒ† –Š‡ –‡Ž‡ ‡’ŠƒŽ‘Ǥ Š‹• ‹‰”ƒ–‹‘ ‘ —”• ƒˆ–‡” –Š‡

ϯϬϮ ’”‡˜‹‘—•Ž› †‡• ”‹„‡† ‹‰”ƒ–‘”› •–”‡ƒ•ǣ ‹– •–ƒ”–• ˆ”‘ ͳͷǤͷ ƒ† ‹• •—„•–ƒ–‹ƒŽ ƒ– ͳͺǤͷ ‹

ϯϬϯ ‘—•‡ ‡„”›‘• ȋŠƒ‘ ‡– ƒŽǤǡ ʹͲͲͺȌǤ – „‹”–Šǡ ƒ„—†ƒ– Foxb1ǦŽ‹‡ƒ‰‡ ‡ŽŽ•‹‰”ƒ–‡†ˆ”‘–Š‡

ϯϬϰ –ŠƒŽƒ‹ ”‡‰‹‘‹–‘–Š‡‰Ž‘„—•’ƒŽŽ‹†—•ǡ–Š‡ƒ›‰†ƒŽƒǡƒ†–Š‡ ’‹”‹ˆ‘” ‘”–‡šǤ Š‡•‡ ‡ŽŽ•

ϯϬϱ ‡š’”‡•• ‹–‡”‡—”‘ ƒ”‡”•ǣ ‘•– ‘ˆ Foxb1ǦŽ‹‡ƒ‰‡ ‡ŽŽ•‹–Š‡ ‘”–‡šƒ†‹–Š‡ƒ›‰†ƒŽƒ

ϯϬϲ ‡š’”‡••–Š‡ Ž—–ƒƒ–‡‡ ƒ”„‘š›Žƒ•‡ȋ ȌȋŠƒ‘‡–ƒŽǤǡʹͲͲͺȌǤ‘‡Foxb1ǦŽ‹‡ƒ‰‡ ‡ŽŽ•‹–Š‡

ϭϬ 

ϯϬϳ ‘”–‡š™‡”‡ƒŽ”‡–‹‹Ϊǡ„—–‘‡‘ˆ–Š‡‡š’”‡••‡†ƒŽ„‹†‹ȋŠƒ‘‡–ƒŽǤǡʹͲͲͺȌǤ ”‘†‡–•ǡƒ•

ϯϬϴ ƒ”—Ž‡ǡ ‘”–‹ ƒŽ‹–‡”‡—”‘•ƒ”‡‰‡‡”ƒ–‡†‹–Š‡‰ƒ‰Ž‹‘‹ ‡‹‡ ‡•ȋƒ”Àƒ†—„‡•–‡‹ǡ

ϯϬϵ ʹͲͲ͵ȌǤŠ‡‹‰”ƒ–‘”›•–”‡ƒ— ‘˜‡”‡†„›Šƒ‘ƒ† ‘ŽŽ‡ƒ‰—‡•™‘—Ž†–Š‡”‡ˆ‘”‡ƒŽŽ‘™ƒ•’‡ ‹ˆ‹ 

ϯϭϬ ’‘‘Ž‘ˆ‹–‡”‡—”‘•–‘•‡––Ž‡–Š‡‘Žˆƒ –‘”› ‘”–‡š†—”‹‰Žƒ–‡‡„”›‘‹ •–ƒ‰‡•ǤŠ‡–Š‡”–Š‡•‡

ϯϭϭ ‹–‡”‡—”‘••‡––Ž‡ƒ•’‡ ‹ˆ‹ Žƒ›‡”‹–Š‡‘Žˆƒ –‘”› ‘”–‡š‘”™Š‡–Š‡”–Š‡›ƒ”‡Š‘‘‰‡‡‘—•Ž›

ϯϭϮ †‹•–”‹„—–‡†”‡ƒ‹‘’‡“—‡•–‹‘•ǡƒ•™‡ŽŽƒ•–Š‡‹”•’‡ ‹ˆ‹ ˆ— –‹‘‹–Š‡‡—”‘ƒŽ‡–™‘”Ǥ

ϯϭϯ Layer formation in the olfactory cortex

ϯϭϰ ‡‹‹• ‡–‘ˆ–Š‡‡‘ ‘”–‡šŽƒ›‡”‹•‹†‡Ǧ‘—–†‡˜‡Ž‘’‡–ǡ‹Œ‡ –‹‘•‘ˆȏ͵ Ȑ–Š›‹†‹‡

ϯϭϱ ‹ ͳͶǦʹʹ –‹‡†Ǧ’”‡‰ƒ– ”ƒ– ˆ‡ƒŽ‡• •Š‘™‡† •‹‰‹ˆ‹ ƒ–Ž› ‘”‡ ‡ƒ”Ž›Ǧ„‘” ‡ŽŽ• •‡––Ž‡† ‹

ϯϭϲ ’‹”‹ˆ‘” ‘”–‡š†‡‡’Žƒ›‡”͵ ‘’ƒ”‡†–‘Žƒ›‡”ʹȋƒ›‡”ǡͳͻͺ͸ǢƒŽ˜‡”†‡ƒ†ƒ–ƒ ƒƒǡͳͻͻͶȌǤ

ϯϭϳ Š‡’‹”‹ˆ‘” ‘”–‡šŽƒ›‡” ʹ ‹•‡ƒ•‹Ž›†‡Ž‹‡ƒ„Ž‡™‹–Š‹–•Š‹‰Š †‡•‹–›‘ˆ’›”ƒ‹†ƒŽ ‡ŽŽ„‘†‹‡•Ǥ

ϯϭϴ Š‹•Žƒ›‡”•–ƒ”–•–‘‘”‰ƒ‹œ‡ˆ”‘ͳ͸ƒ†‹•™‡ŽŽ†‡ˆ‹ƒ„Ž‡„›ͳͺ‹‘—•‡‡„”›‘•ȋŽ‹‰Ž‡”

ϯϭϵ ‡–ƒŽǤǡʹͲͳͷȌȋ ‹‰—”‡ʹȌǤ”†‹Œ‡ –‹‘•ƒ–†‹ˆˆ‡”‡–‘—•‡†‡˜‡Ž‘’‡–•–ƒ‰‡•ƒ†ƒƒŽ›•‡•‘ˆ

ϯϮϬ ‡ŽŽ ‹†‡–‹–› ƒ– ’‘•–ƒ–ƒŽ †ƒ› ͹ •Š‘™‡† –Šƒ– ‡ŽŽ „‹”–Š†ƒ–‡ •‹‰‹ˆ‹ ƒ–Ž›ƒˆˆ‡ –•‘–‘Ž›–Š‡

ϯϮϭ Žƒ‹ƒ”’‘•‹–‹‘‘ˆ ‡ŽŽ•„—–ƒŽ•‘–Š‡‹” ‡ŽŽ—Žƒ”ˆƒ–‡ȋƒ”ƒ‡–ƒŽǤǡʹͲͳͳȌǤ ‹ ‡ǡ‘•–Žƒ›‡”ʹ

ϯϮϮ ’›”ƒ‹†ƒŽ ‡ŽŽ•ƒ”‡„‘”ƒ–ͳʹǤͳͶ„‘”Ǧ ‡ŽŽ•ƒ”‡Ž‡••—‡”‘—•„—–•–‹ŽŽ†‹ˆˆ‡”‡–‹ƒ–‡ƒ‹Ž›‹

ϯϮϯ ’›”ƒ‹†ƒŽ ‡ŽŽ•Ǥ ‘™‡˜‡”ǡͳ͸„‘”Ǧ ‡ŽŽ•†‹•’Žƒ›‘”‡Š‡–‡”‘‰‡‡‘—• ‹†‡–‹–‹‡•ǣ Ͷ͵Ψ

ϯϮϰ †‹ˆˆ‡”‡–‹ƒ–‡‹’›”ƒ‹†ƒŽ‡—”‘•ǡʹͶΨ†‹ˆˆ‡”‡–‹ƒ–‡‹‘Ǧ’›”ƒ‹†ƒŽ‡—”‘•ȋ‹–‡”‡—”‘•Ȍǡ

ϯϮϱ ͵͵Ψ†‹ˆˆ‡”‡–‹ƒ–‡‹‘Ǧ‡—”‘ƒŽ ‡ŽŽ•ȋƒ•–”‘ ›–‡•Ȍȋƒ”ƒ‡–ƒŽǤǡʹͲͳͳȌǤŠ‡•‡”‡•—Ž–••Š‘™–Š‡

ϯϮϲ ‘•‡”˜ƒ–‹‘‘ˆƒˆ—†ƒ‡–ƒŽ†‡˜‡Ž‘’‡–ƒŽ Š”‘‘Ž‘‰›‹„‘–Š’ƒŽ‡‘Ǧƒ†‡‘Ǧ ‘”–‹ ‡•ǡ™‹–Š

ϯϮϳ –Š‡•‡“—‡–‹ƒŽ‰‡‡”ƒ–‹‘‘ˆ’›”ƒ‹†ƒŽ ‡ŽŽ•ǡ‹–‡”‡—”‘•ƒ†‰Ž‹ƒǤƒ„‡Ž‹‰‘ˆ ‡ŽŽ•‰‡‡”ƒ–‡†ƒ–

ϯϮϴ ’”‡ ‹•‡ –‹‡ ’‘‹–• †—”‹‰ †‡˜‡Ž‘’‡– ‘—Ž† „‡ Š‡Ž’ˆ—Ž –‘ ‹†‡–‹ˆ›‰‡‡–‹ ƒ”‡”•‘ˆ ‡ŽŽ

ϯϮϵ ’‘’—Žƒ–‹‘•‹–Š‡‘Žˆƒ –‘”› ‘”–‡šǤ‘ˆƒ”‘–Š‹‰‹•‘™ƒ„‘—–†‡˜‡Ž‘’‡–ƒŽ†›ƒ‹ •‘ˆ

ϯϯϬ Žƒ›‡”͵‡—”‘•Ǥ

ϯϯϭ š‘•ˆ”‘‹–”ƒŽƒ†–—ˆ–‡† ‡ŽŽ•ˆƒ• ‹ —Žƒ–‡–‘ˆ‘”–Š‡ƒ††‡˜‡Ž‘’ ‘ŽŽƒ–‡”ƒŽ•ǡ

ϯϯϮ ™Š‹ Š‹˜ƒ†‡–Š‡‹’•‹Žƒ–‡”ƒŽ‘Žˆƒ –‘”› ‘”–‡šŽƒ›‡”ͳˆ”‘ͳͷǤͷ–‘ͳ͹Ǥͷ‹ƒ ƒ—†ƒŽ–‘”‘•–”ƒŽ

ϭϭ 

ϯϯϯ •‡“—‡ ‡ ȋ ‹‰—”‡ ͵Ȍ ȋ ƒ„‡”Ž›ǡ ͳͻͺ͵Ǣ ƒ„‡”Ž› ƒ† ‡‹‰ǡ ͳͻͺ͵Ǣ ‹”ƒ–ƒ ƒ† —Œ‹•ƒ™ƒǡ ͳͻͻͻǢ

ϯϯϰ ƒ„‡”Ž›ǡʹͲͲͳȌǤƒ›‡”ͳ ƒ„‡†‹˜‹†‡†‹–™‘•—„ǦŽƒ›‡”•ǣ–Š‡•—’‡”ˆ‹ ‹ƒŽŽƒ›‡”ͳƒˆ‘”‡†„›–Š‡

ϯϯϱ –‡”‹ƒŽ• ‘ˆ  ƒš‘•ǡ ™Š‹ Š ƒ‡ •›ƒ’–‹  ‘–ƒ –• ™‹–Š –Š‡ ƒ’‹ ƒŽ †‡†”‹–‡• ‘ˆ Žƒ›‡” ʹȀ͵

ϯϯϲ ’›”ƒ‹†ƒŽ ‡ŽŽ•ǡ ƒ† –Š‡ Žƒ›‡” ͳ„ ˆ‘”‡† „› –Š‡ ƒ’‹ ƒŽ †‡†”‹–‡• ‘ˆ Žƒ›‡” ʹȀ͵ ’›”ƒ‹†ƒŽ ‡ŽŽ•

ϯϯϳ ȋ Š™‘„ƒ†”‹ ‡ǡͳͻͺͶǢ—œ—‹ƒ†‡‡”•ǡʹͲͳͳȌȋ ‹‰—”‡͵ȌǤ –Š‡ƒ†—Ž–‘—•‡„”ƒ‹ǡ–Š‡

ϯϯϴ Žƒ›‡”•ͳƒƒ†ͳ„ƒ”‡†‡Ž‹‡ƒ„Ž‡„›Ǧʹƒ†ƒŽ”‡–‹‹‹—‘•–ƒ‹‹‰•ǡ™Š‹ ŠŽƒ„‡ŽŽƒ›‡”ʹȀ͵

ϯϯϵ ’›”ƒ‹†ƒŽ ‡ŽŽ†‡†”‹–‡•ƒ†ƒš‘•”‡•’‡ –‹˜‡Ž›ȋƒ”ƒ‡–ƒŽǤǡʹͲͳͳȌǤ–„‹”–ŠǡŽƒ›‡”ͳ‹•‘–

ϯϰϬ ›‡– •—„†‹˜‹†‡†Ǥ ƒ›‡” ͳƒ •–ƒ”–• –‘ „‡ †‹•–‹‰—‹•Šƒ„Ž‡ ˆ”‘ ’‘•–ƒ–ƒŽ †ƒ› ͹ ™‹–Š ˜‹•‹„Ž‡ ‘Ǧ

ϯϰϭ •–ƒ‹‹‰ ‘ˆ Ǧʹ ƒ† ƒŽ”‡–‹‹ ȋƒ”ƒ ‡– ƒŽǤǡ ʹͲͳͳȌǤ Š‡•‡ ‘„•‡”˜ƒ–‹‘• •—‰‰‡•– –Šƒ– –Š‡

ϯϰϮ •›ƒ’–‹  ‘–ƒ –•„‡–™‡‡–‡”‹ƒŽ•ƒ†Žƒ›‡”ʹȀ͵’›”ƒ‹†ƒŽ ‡ŽŽƒ’‹ ƒŽ†‡†”‹–‡•†‡˜‡Ž‘’

ϯϰϯ †—”‹‰–Š‡ˆ‹”•–’‘•–ƒ–ƒŽ™‡‡Ǥ—”‹‰‘—•‡‡„”›‘‹ †‡˜‡Ž‘’‡–ǡ  ƒš‘• †‡˜‡Ž‘’

ϯϰϰ ‘ŽŽƒ–‡”ƒŽ•ǡ™Š‹ Šˆ‹”•–‹˜ƒ†‡–Š‡‘ƒ–ͳͷǤͷǡ–Š‡–Š‡’‹”‹ˆ‘” ‘”–‡š‘‡†ƒ›Žƒ–‡”ǡƒ†–Š‡

ϯϰϱ ‘”‡”‘•–”ƒŽŽ›Ž‘ ƒ–‡†”‡‰‹‘•‘ˆ–Š‡‘Žˆƒ –‘”› ‘”–‡šȋ ‹”ƒ–ƒƒ† —Œ‹•ƒ™ƒǡ ͳͻͻͻȌǤ Š‡•‡

ϯϰϲ ‘ŽŽƒ–‡”ƒŽ• ƒŽŽ‘™ ‘‡  ƒš‘ –‘ ƒ‡ •›ƒ’–‹  ‘–ƒ –• ™‹–Š Žƒ›‡”ʹȀ͵’›”ƒ‹†ƒŽ ‡ŽŽ•‘ˆ

ϯϰϳ †‹ˆˆ‡”‡–”‡‰‹‘•‘ˆ–Š‡‘Žˆƒ –‘”› ‘”–‡šȋŒ‹ƒ‡–ƒŽǤǡͳͻͺͶȌǤ‹ ‡ƒš‘•†‡˜‡Ž‘’ƒˆ–‡”„‹”–Šǡ

ϯϰϴ ‹–‹•”ƒ–‹‘ƒŽ–‘ƒ••—‡–Šƒ––Š‡ƒ–—”ƒ–‹‘‘ˆ–Š‡ ‹” —‹–‹•‡š’‡”‹‡ ‡Ǧ†‡’‡†‡–ǡƒ•‹–Šƒ•„‡‡

ϯϰϵ ™‡ŽŽ†‡• ”‹„‡†ˆ‘”–Š‡’”‹ƒ”›•‘ƒ–‘•‡•‘”›„ƒ””‡Ž ‘”–‡šȋˆ‘””‡˜‹‡™•‡‡‹–ƒŽ‹ƒ† ƒ„ƒ—†‘ǡ

ϯϱϬ ʹͲͳͶȌǤ – ™‘—Ž† „‡ ‘ˆ ‹–‡”‡•– –‘ ‹˜‡•–‹‰ƒ–‡ –Š‡ ‘•‡“—‡ ‡• ‘ˆ’‘•–ƒ–ƒŽ‘†‘”ƒ–•–‹—Ž—•

ϯϱϭ †‡’Ž‡–‹‘‹–Š‡‘”‰ƒ‹œƒ–‹‘ƒ†ƒ–—”ƒ–‹‘‘ˆ‘Žˆƒ –‘”› ‘”–‡šŽƒ›‡”ͳ ‘‡ –‹˜‹–›Ǥ

ϯϱϮ Cell types and organization of the circuit implicated in odor processing

ϯϱϯ Cell types and circuit organization within the

ϯϱϰ  –Š‡ ’‹”‹ˆ‘” ‘”–‡šǡ Žƒ›‡” ʹ ’›”ƒ‹†ƒŽ ‡ŽŽ• Šƒ˜‡ „‡‡ ‹‹–‹ƒŽŽ› Žƒ••‹ˆ‹‡† ‹ –™‘

ϯϱϱ ’‘’—Žƒ–‹‘• „ƒ•‡† ‘ –Š‡‹” †‹•–‹ – ˆ— –‹‘• ‹ ‘†‘” ’”‘ ‡••‹‰ǣ –Š‡ •‡‹Ž—ƒ” ƒ† –Š‡

ϯϱϲ •—’‡”ˆ‹ ‹ƒŽ’›”ƒ‹†ƒŽ ‡ŽŽ•ȋ ‹‰—”‡͵ȌǤŠ‡•‡‹Ž—ƒ” ‡ŽŽ•ƒ”‡Ž‘ ƒ–‡†‹–Š‡—’’‡”’ƒ”–‘ˆŽƒ›‡”ʹ

ϯϱϳ ȋʹƒȌƒ††‹•’Žƒ›ƒŠ‹‰Š•’‹‡†‡•‹–›‹–Š‡†‹•–ƒŽ”‡‰‹‘•‘ˆ–Š‡‹”ƒ’‹ ƒŽ†‡†”‹–‡•ǤŠ‡•‡ ‡ŽŽ•

ϯϱϴ ”‡ ‡‹˜‡’”‡†‘‹ƒ–Ž›ƒˆˆ‡”‡–‡š ‹–ƒ–‹‘ˆ”‘ƒš‘•ǡ–Š‡‹”ƒ –‹˜‹–›„‡‹‰Š‹‰ŠŽ› ‘””‡Žƒ–‡†

ϭϮ 

ϯϱϵ ™‹–Š  •–‹—Žƒ–‹‘ ȋ—œ—‹ ƒ† ‡‡”•ǡ ʹͲͳͳȌǤ Š‡ •—’‡”ˆ‹ ‹ƒŽ’›”ƒ‹†ƒŽ ‡ŽŽ•ƒ”‡Ž‘ ƒ–‡†

ϯϲϬ †‡‡’‡”‹Žƒ›‡”ʹȋʹ„Ȍƒ††‹•’Žƒ›„ƒ•ƒŽ†‡†”‹–‡•‡š–‡†‹‰‹–‘Žƒ›‡”͵ȋ ‹‰—”‡͵ȌǤŠ‡›”‡ ‡‹˜‡

ϯϲϭ ™‡ƒ‡” ƒˆˆ‡”‡– ‹’—–• ˆ”‘ –Š‡  ƒ† •–”‘‰‡” ‹–”ƒ ‘”–‹ ƒŽ ‡š ‹–ƒ–‘”› †”‹˜‡ –Š”‘—‰Š

ϯϲϮ ƒ••‘ ‹ƒ–‹˜‡ˆ‹„‡”•ǡ–Š‡‹”ƒ –‹˜‹–›–Š‡”‡ˆ‘”‡†‡’‡†‹‰‘Ž‘ ƒŽˆ‡‡†„ƒ Ž‘‘’•ȋ ƒ„‡”Ž›ƒ† ‡‹‰ǡ

ϯϲϯ ͳͻͺ͵Ǣ —œ—‹ ƒ† ‡‡”•ǡ ʹͲͳͳǢ ‹‡‰ƒ† ‡– ƒŽǤǡ ʹͲͳͳǢ ƒ‰‹™ƒ”ƒ ‡–ƒŽǤǡ ʹͲͳʹȌ ȋ ‹‰—”‡͵ȌǤ Š‡

ϯϲϰ Žƒ›‡”͵‹•Ž‡••†‡•‡‹ ‡ŽŽ„‘†‹‡•ƒ• ‘’ƒ”‡†–‘Žƒ›‡”ʹƒ† ‘’‘•‡†‘ˆ†‡‡’’›”ƒ‹†ƒŽ ‡ŽŽ•ǡ

ϯϲϱ ™Š‹ Š”‡ ‡‹˜‡‹‹ƒŽƒˆˆ‡”‡–‹’—–•ˆ”‘ƒš‘•ǡ„—–•—„•–ƒ–‹ƒŽ‹–”ƒ ‘”–‹ ƒŽ‡š ‹–ƒ–‹‘

ϯϲϲ ȋ ‹‰—”‡͵ȌǤ

ϯϲϳ ‘†ƒ–‡ǡ‘Ž‡ —Žƒ”‹†‡–‹–›‘ˆ–Š‡†‹•–‹ –’›”ƒ‹†ƒŽ‡—”‘™‹–Š‹–Š‡’‹”‹ˆ‘” ‘”–‡š

ϯϲϴ Šƒ•„‡‡—†‡”Ǧ‹˜‡•–‹‰ƒ–‡†Ǥ‘•–ƒ”‡”•†‡ˆ‹‹‰Žƒ›‡”•‹–Š‡‡‘ ‘”–‡šƒ”‡‡š’”‡••‡†‹–Š‡

ϯϲϵ ’‹”‹ˆ‘” ‘”–‡šƒ•™‡ŽŽȋ‘Ž›‡ƒ—š‡–ƒŽǤǡʹͲͲ͹Ȍȋ ‹‰—”‡͵ȌǤ’’‡”Žƒ›‡”ƒ”‡”•ͳƒ†ʹ

ϯϳϬ ȋ—„‡Ž‘•‡–ƒŽǤǡʹͲͳͲȌƒ”‡‡”‹ Š‡†‹’‹”‹ˆ‘” ‘”–‡šŽƒ›‡”ʹ„Ǥ‘‡ʹΪ ‡ŽŽ•ƒ”‡ƒŽ•‘’”‡•‡–

ϯϳϭ ‹Žƒ›‡”͵ǤƒŽŽ‘•ƒŽ’”‘Œ‡ –‹‘‡—”‘ƒ”‡”ʹȋŽ ƒ‘‡–ƒŽǤǡʹͲͲͺǢ‡‘‡‡–ƒŽǤǡʹͲͳͷȌ‹•

ϯϳϮ ‡š’”‡••‡†‹’‹”‹ˆ‘” ‘”–‡š Žƒ›‡”• ʹƒ† ͵Ǥ – ™‘—Ž† „‡ ‘ˆ‹–‡”‡•––‘‘™™Š‡–Š‡”ʹΪ

ϯϳϯ ’‹”‹ˆ‘” ‘”–‡š‡—”‘•’”‘Œ‡ ––‘ ‘–”ƒŽƒ–‡”ƒŽŠ‡‹•’Š‡”‡–Š”‘—‰Š–Š‡ƒ–‡”‹‘” ‘‹••—”‡ǡ

ϯϳϰ ƒ†ƒ”‡–Š‡”‡ˆ‘”‡–Š‡ ‘—–‡”’ƒ”–•‘ˆʹڐ‡‘ ‘”–‹ ƒŽ ƒŽŽ‘•ƒŽ’”‘Œ‡ –‹‘‡—”‘•ǤŠƒŽƒ‹ 

ϯϳϱ ”‡ ‹’‹‡–Žƒ›‡”Ͷ‡—”‘ƒ”‡”βȋ ƒ„ƒ—†‘‡–ƒŽǤǡʹͲͳʹȌ•Š‘™•˜‡”›™‡ƒ‡š’”‡••‹‘‹

ϯϳϲ –Š‡’‹”‹ˆ‘” ‘”–‡šǤŠ‹• ‘—Ž†„‡ƒ••‘ ‹ƒ–‡†™‹–Š–Š‡ˆƒ ––Šƒ–’‹”‹ˆ‘” ‘”–‡š‡—”‘•†‘‘–

ϯϳϳ ”‡ ‡‹˜‡•‡•‘”›‹’—–ˆ”‘–Š‡–ŠƒŽƒ—•„—–†‹”‡ –Ž›ˆ”‘–Š‡•ǤFezf2ǡ‘™–‘„‡‡”‹ Š‡†

ϯϳϴ ‹‡‘ ‘”–‡šŽƒ›‡”ͷȋ‘—ƒ—šƒ†”Ž‘––ƒǡʹͲͳͲǢŽƒ‘••ƒ‡–ƒŽǤǡʹͲͳ͵Ȍǡ‹•‡”‹ Š‡†‹’‹”‹ˆ‘”

ϯϳϵ ‘”–‡šŽƒ›‡”ʹƒȋ‹‘†ƒ–‘‡–ƒŽǤǡʹͲͳ͸ȌǤƒ›‡”ͷȀ͸ƒ”‡” ʹ‹•‡š’”‡••‡†‹’‹”‹ˆ‘” ‘”–‡š

ϯϴϬ Žƒ›‡”ʹȋ ‹‰—”‡͵ȌǤ ‡‡‡š’”‡••‹‘ ‘’ƒ”‹•‘„‡–™‡‡–Š‡‡‘ ‘”–‡šƒ†–Š‡’‹”‹ˆ‘” ‘”–‡š

ϯϴϭ Šƒ• „‡‡ ‡š–‡•‹˜‡Ž› ƒƒŽ›œ‡† ‹ –Š‡ ™‘” ‘ˆ —œœƒ–‹ ȋ—œœƒ–‹ǡ ʹͲͳͷȌǤ ‡ •Š‘™‡† –Šƒ– –Š‡

ϯϴϮ ‡‘ ‘”–‡šŽƒ›‡”•ʹȀ͵•Šƒ”‡ͶʹΨ‘ˆ‡”‹ Š‡†‰‡‡•™‹–Š–Š‡’‹”‹ˆ‘” ‘”–‡š™Š‹Ž‡–Š‡†‡‡’

ϯϴϯ Žƒ›‡”•Ͷƒ†͸ƒ”‡Ž‡••”‡Žƒ–‡†–‘–Š‡’‹”‹ˆ‘” ‘”–‡š™‹–Š‘Ž› ʹͻΨ ‘ˆ ‘Ǧ‡š’”‡••‡† ‰‡‡•

ϯϴϰ ȋ—œœƒ–‹ǡʹͲͳͷȌǤ‹”‹ˆ‘” ‘”–‡š‡—”‘••‡‡–Š‡”‡ˆ‘”‡–‘‡š’”‡•• ‘‘‘Ž‡ —Žƒ”ƒ”‡”•

ϯϴϱ ‡”‹ Š‡†‹‡‘ ‘”–‹ ƒŽ‡—”‘•ǡ„—––Š‡›†‘‘–†‹•’Žƒ›–Š‡•ƒ‡ ‘”‰ƒ‹œƒ–‹‘ ™‹–Š‹ Žƒ›‡”•Ǥ

ϭϯ 

ϯϴϲ ƒ•†‡ƒ† ‘ŽŽ‡ƒ‰—‡•†‡˜‡Ž‘’‡†ƒ‡™ ‘’—–ƒ–‹‘ƒŽ’‹’‡Ž‹‡ˆ‘”Š‹‰ŠǦ–Š”‘—‰Š’—–ƒƒŽ›•‹•

ϯϴϳ ƒ† ‘’ƒ”‹•‘‘ˆŽŽ‡”ƒ‹–Žƒ•in situŠ›„”‹†‹œƒ–‹‘•ƒ–Žƒ‹ƒ””‡•‘Ž—–‹‘–‘•–—†›‰‡‡

ϯϴϴ ‡š’”‡••‹‘‹–Š‡‡†‹ƒŽ‡–‘”Š‹ƒŽ ‘”–‡šǤ•‘„•‡”˜‡†ˆ‘”–Š‡’‹”‹ˆ‘” ‘”–‡šǡ˜‡”›ˆ‡™‰‡‡•

ϯϴϵ ƒ”‡—‹“—‡Ž›‡š’”‡••‡†‹–Š‡‡†‹ƒŽ‡–‘”Š‹ƒŽ ‘”–‡šǤ ƒ††‹–‹‘ƒ† ‘–”ƒ”›–‘–Š‡’‹”‹ˆ‘”

ϯϵϬ ‘”–‡šǡ †‡‡’ Žƒ›‡”• ‘ˆ –Š‡ ‡†‹ƒŽ ‡–‘”Š‹ƒŽ ‘”–‡š ƒ”‡ ”‡Žƒ–‹˜‡Ž› •‹‹Žƒ” –‘ –Š‘•‡ ‘ˆ –Š‡

ϯϵϭ ‡‘ ‘”–‡šǡ ™Š‹Ž‡ •—’‡”ˆ‹ ‹ƒŽ Žƒ›‡”• ƒ”‡ •—„•–ƒ–‹ƒŽŽ› ‘”‡ †‹˜‡”‰‡– ƒ– ƒ ‘Ž‡ —Žƒ” Ž‡˜‡Ž

ϯϵϮ ȋƒ•†‡ ‡– ƒŽǤǡ ʹͲͳͷȌǤ Š‡ ƒ—–Š‘”• ’”‘’‘•‡† –Šƒ– –Š‡ ‡†‹ƒŽ ‡–‘”Š‹ƒŽ ‘”–‡š‹•ƒ–›’‡‘ˆ

ϯϵϯ ’‡”‹ƒ” Š‹ ‘”–‡šȋ’ƒŽ‡‘ ‘”–‡šȌǡƒ–”ƒ•‹–‹‘ƒŽ•–”— –—”‡„‡–™‡‡–Š‡͸ǦŽƒ›‡”‡†‡‘ ‘”–‡šƒ†–Š‡

ϯϵϰ ͵ǦŽƒ›‡”‡†ƒ” Š‹ ‘”–‡šȋƒ•†‡‡–ƒŽǤǡʹͲͳͷȌǤ

ϯϵϱ ”‡ ‡–•–—†›ˆ”‘‹‘†ƒ–‘ƒ† ‘ŽŽ‡ƒ‰—‡•ƒ††”‡••‡†ˆ‘”–Š‡ˆ‹”•––‹‡–Š‡‘Ž‡ —Žƒ”

ϯϵϲ ‹†‡–‹–‹‡•‘ˆ’‹”‹ˆ‘” ‘”–‡š’”‘Œ‡ –‹‘‡—”‘•—•‹‰Žƒ•‡”Ǧ ƒ’–—”‡‹ ”‘†‹••‡ –‹‘‘ˆ’‹”‹ˆ‘”

ϯϵϳ ‘”–‡š Žƒ›‡”• ƒ†  †‡‡’ •‡“—‡ ‹‰ –‘ ‹†‡–‹ˆ› ‰‡‡• †‹ˆˆ‡”‡–‹ƒŽŽ› ‡š’”‡••‡† ™‹–Š‹

ϯϵϴ ’‹”‹ˆ‘” ‘”–‡š Žƒ›‡”• ‹ ‘„‹ƒ–‹‘ ™‹–Š ”‡–”‘‰”ƒ†‡ Žƒ„‡Ž‹‰ ˆ”‘ ’‹”‹ˆ‘” ‘”–‡š –ƒ”‰‡–•

ϯϵϵ ȋ‹‘†ƒ–‘‡–ƒŽǤǡʹͲͳ͸ȌǤ‹–Š–Š‡•‡ƒ’’”‘ƒ Š‡•ǡ–Š‡›•Š‘™‡†–Šƒ–Žƒ›‡”ʹƒ•‡‹Ž—ƒ” ‡ŽŽ•’”‘Œ‡ –

ϰϬϬ –‘‘ƒ†Žǡƒ†‡š’”‡••Reelinƒ†Fezf2ȋ ‹‰—”‡͵ȌǤŠ‡‡š’”‡••‹‘‘ˆ‡‡Ž‹„›‡š ‹–ƒ–‘”›

ϰϬϭ ’”‘Œ‡ –‹‘‡—”‘•‹ƒ†—Ž–„”ƒ‹‹••’‡ ‹ˆ‹ –‘–Š‡’‹”‹ˆ‘” ‘”–‡šǡ•‹ ‡‹–Š‡‡‘ ‘”–‡šǡ‡‡Ž‹

ϰϬϮ ‹•‘•–Ž›‡š’”‡••‡†„›‹–‡”‡—”‘•ȋŽ ƒ–ƒ”ƒ‡–ƒŽǤǡͳͻͻͺǢ‡•‘Ž†‡–ƒŽǤǡͳͻͻͺǢƒ‘•Ǧ‘”‡‘

ϰϬϯ ‡– ƒŽǤǡ ʹͲͲ͸ȌǤ Š‡•‡ ‡‡Ž‹Ϊ’‹”‹ˆ‘” ‘”–‡š•‡‹Ž—ƒ” ‡ŽŽ•”‡•‡„Ž‡–Š‡ ‡ŽŽ•ˆ”‘–Š‡

ϰϬϰ †‡˜‡Ž‘’‹‰„”ƒ‹ȋƒ” ‡ŽŽ‡”‡–ƒŽǤǡʹͲͳ͸ȌǤƒ” ‡ŽŽ‡”ƒ† ‘ŽŽ‡ƒ‰—‡•’”‘’‘•‡†–Šƒ––Š‡•‡ ”‡–‹‘‘ˆ

ϰϬϱ ‡‡Ž‹ „› —’’‡” ’‹”‹ˆ‘” ‘”–‡š Žƒ›‡” ʹ ‘—Ž† ’ƒ”–‹ ‹’ƒ–‡ ‹ –Š‡ ƒ–—”ƒ–‹‘ ‘ˆ ƒ ‹ƒ–—”‡

ϰϬϲ ‡—”‘•—„’‘’—Žƒ–‹‘‡š’”‡••‹‰Ǧƒ†Ž‘ ƒ–‡††‡‡’‡”‹Žƒ›‡”ʹȋƒ Š‡”‡–ƒŽǤǡ

ϰϬϳ ʹͲͲͳǢ—œœƒ–‹‡–ƒŽǤǡʹͲͲͻǢŽ‡’‹‡–ƒŽǤǡʹͲͳͳȌǤ –‡”‡•–‹‰Ž›ǡ‡‡Ž‹Ϊ•–‡ŽŽƒ–‡’”‹ ‹’ƒŽ ‡ŽŽ•Šƒ˜‡

ϰϬϴ „‡‡†‡• ”‹„‡†‹–Š‡ƒ†—Ž–‡†‹ƒŽ‡–‘”Š‹ƒŽ ‘”–‡šȋȌŽƒ›‡”ʹȋƒ”‰ƒ‡–ƒŽǤǡʹͲͳͲȌǤŠ‡

ϰϬϵ Žƒ›‡”ʹƒŽ•‘ ‘’”‹•‡•’›”ƒ‹†ƒŽ ‡ŽŽ•ǡ•‘‡‘ˆ–Š‡‡š’”‡••‹‰‘—„Ž‡ ‘”–‹Žƒ–‡‹’‘•–ƒ–ƒŽ

ϰϭϬ †‡˜‡Ž‘’‡– ȋƒ› ƒ† ”‡ Š–ǡ ʹͲͳ͸ȌǤ  ”‘Ž‡ ˆ‘” ‡‡Ž‹ ‹ –Š‡ Žƒ–‡ƒ–—”ƒ–‹‘‘ˆ‹ƒ–—”‡

ϰϭϭ ‡—”‘•—„’‘’—Žƒ–‹‘• ƒ–Š‡”‡ˆ‘”‡ƒ•™‡ŽŽ„‡ ‘•‹†‡”‡†‹–Š‡ ƒ•‡‘ˆ–Š‡Ǥ

ϭϰ 

ϰϭϮ ‡•‹†‡•ǡ’‹”‹ˆ‘” ‘”–‡š’›”ƒ‹†ƒŽ‡—”‘••‡†‹‰’”‘Œ‡ –‹‘•„ƒ –‘ƒ”‡Ž‘ ƒ–‡†‹

ϰϭϯ Žƒ›‡”ʹ„ƒ†Žƒ›‡”͵ǡ†‹•’Žƒ›‘”’Š‘Ž‘‰› Šƒ”ƒ –‡”‹•–‹ •‘ˆ•—’‡”ˆ‹ ‹ƒŽƒ††‡‡’’›”ƒ‹†ƒŽ ‡ŽŽ•

ϰϭϰ ƒ† ‡š’”‡•• Cux1 ȋ ‹‰—”‡ ͵ȌǤ –‡”‡•–‹‰Ž›ǡ ͵Ǧ”‡ ‘•–”— –‹‘ ‘ˆ ˆ‡‡†„ƒ  ’”‘Œ‡ –‹‰ ‡—”‘•

ϰϭϱ ƒˆ–‡””ƒ„‹‡•˜‹”—•‹Œ‡ –‹‘‹–Š‡‰”ƒ—Ž‡ ‡ŽŽŽƒ›‡”‘ˆ–Š‡ƒ‹•Š‘™‡†–Šƒ––Š‡•‡ ‡ŽŽ•ƒ”‡

ϰϭϲ ‘”‡—‡”‘—•‹–Š‡ƒ–‡”‹‘”–Šƒ‹–Š‡’‘•–‡”‹‘”’‹”‹ˆ‘” ‘”–‡šƒ††‹•’Žƒ›ƒ‘”ƒ†‘

ϰϭϳ ‘”‰ƒ‹œƒ–‹‘ǣ’‹”‹ˆ‘” ‘”–‡š‡—”‘•‡š––‘‘‡ƒ‘–Š‡”’”‘Œ‡ ––‘•‹‹Žƒ””‡‰‹‘•‘ˆ–Š‡

ϰϭϴ ‰”ƒ—Ž‡ ‡ŽŽŽƒ›‡”ȋƒ†ƒƒ„Šƒ‡–ƒŽǤǡʹͲͳ͸ȌǤ‡—”‘•’”‘Œ‡ –‹‰–‘‹ˆ”ƒŽ‹„‹ •—„†‹˜‹•‹‘‘ˆ

ϰϭϵ –Š‡‡†‹ƒŽ’”‡ˆ”‘–ƒŽ ‘”–‡šȋ Ǧ Ȍƒ”‡ƒ‹Ž›Ž‘ ƒ–‡†‹Žƒ›‡” ʹ„ ƒ† ‡š’”‡•• Ctip2Ǥ

ϰϮϬ ‘”‡‘˜‡”ƒˆ”ƒ –‹‘‘ˆŽƒ›‡”ʹ„‡—”‘•‡š’”‡••„‘–ŠCux1ƒ†Ctip2ƒ†’”‘Œ‡ ––‘„‘–Šƒ†

ϰϮϭ Ǧ ȋ ‹‰—”‡͵Ȍȋ‹‘†ƒ–‘‡–ƒŽǤǡʹͲͳ͸ȌǤŠ‹••–—†›’‘‹–•‘—–‹–‡”‡•–‹‰†‹ˆˆ‡”‡ ‡•ƒ„‘—––Š‡

ϰϮϮ ‘Ž‡ —Žƒ”‘”‰ƒ‹œƒ–‹‘•‘ˆ–Š‡’‹”‹ˆ‘” ‘”–‡šƒ†–Š‡‡‘ ‘”–‡šǣ™Š‹Ž‡‹–Š‡‡‘ ‘”–‡šǡCux1‹•

ϰϮϯ ‡š’”‡••‡† „› •—’‡”ˆ‹ ‹ƒŽ Žƒ›‡” ‡—”‘• ƒ† Fezf2„›†‡‡’Žƒ›‡”ͷȀ͸‡—”‘•‡š Ž—•‹˜‡Ž›ǡ–Š‹•

ϰϮϰ ‘”‰ƒ‹œƒ–‹‘‹•”‡˜‡”•‡†‹–Š‡’‹”‹ˆ‘” ‘”–‡šǡ™Š‡”‡Fezf2‹•‡š’”‡••‡†‹Žƒ›‡”ʹƒƒ†Cux1‹

ϰϮϱ Žƒ›‡”•ʹ„ƒ†͵Ǥ‘”‡‘˜‡”ǡCux1ƒ†Ctip2ƒ”‡‡š’”‡••‡†‹—–—ƒŽŽ›‡š Ž—•‹˜‡’‘’—Žƒ–‹‘•‹–Š‡

ϰϮϲ ‡‘ ‘”–‡šǡ™Š‹Ž‡ƒˆ”ƒ –‹‘‘ˆŽƒ›‡”ʹ„‡—”‘•‡š’”‡••„‘–Šƒ”‡”•‹–Š‡’‹”‹ˆ‘” ‘”–‡šǤ

ϰϮϳ –‡”‡•–‹‰Ž›ǡ–Š‡ƒ—–Š‘”•ˆ—”–Š‡”•Š‘™‡†–Šƒ–‹ReelinǦ†‡ˆ‹ ‹‡–‹ ‡ǡ™Š‡”‡ ‡ŽŽ•‘ˆ†‹•–‹ –

ϰϮϴ Žƒ›‡”•ƒ”‡‹–‡”‹‰Ž‡†ƒ ”‘••–Š‡’‹”‹ˆ‘” ‘”–‡šǡ–Š‡‘Ž‡ —Žƒ”‹†‡–‹–‹‡•‘ˆ‡—”‘•’”‘Œ‡ –‹‰

ϰϮϵ –‘–Š‡ȋͳΪȀ  ʹǦȌ ‘’ƒ”‡† –‘ ‡—”‘• ’”‘Œ‡ –‹‰ –‘ –Š‡ ‘ ȋͳǦȀ  ʹΪȌ ƒ”‡

ϰϯϬ ‘•‡”˜‡†ǡ †‡•’‹–‡ –Š‡‹” ’‘•‹–‹‘ †‡ˆ‡ –ǡ ƒ• †‡• ”‹„‡† ‹ –Š‡ ‡‘ ‘”–‡š ȋ‰ƒ™ƒ ‡– ƒŽǤǡ ͳͻͻͷǢ

ϰϯϭ ‡˜‡”‡–ƒŽǤǡʹͲͲ͵Ǣƒ‰‡‡”‡–ƒŽǤǡʹͲͳͲǢ‘›Ž‡‡–ƒŽǤǡʹͲͳͳǢ‹‘†ƒ–‘‡–ƒŽǤǡʹͲͳ͸ȌǤ

ϰϯϮ Odor processing in the piriform cortex

ϰϯϯ  ‹ ‡ǡ ‡ƒ Š ‘Žˆƒ –‘”› •‡•‘”› ‡—”‘ ‹ –Š‡ ‘Žˆƒ –‘”› ‡’‹–Š‡Ž‹— ‡š’”‡••‡• ‘Ž› ‘‡

ϰϯϰ ‘Žˆƒ –‘”›”‡ ‡’–‘”‰‡‡ǡƒ†‘Žˆƒ –‘”›•‡•‘”›‡—”‘•‡š’”‡••‹‰ƒ ‘‘‘Žˆƒ –‘”›”‡ ‡’–‘”

ϰϯϱ •‡† ‘˜‡”‰‡–’”‘Œ‡ –‹‘•–‘ʹ‰Ž‘‡”—Ž‹‹–Š‡ƒ‹ȋ— ƒ†š‡ŽǡͳͻͻͳǢ‘„ƒ‡”–•‡–

ϰϯϲ ƒŽǤǡͳͻͻ͸ǢƒŽ‹ ‡–ƒŽǤǡͳͻͻͻǢʹͲͳͲȌǤ‡•’‹–‡–Š‡’”‡ ‹•‡‘†‘–‘’‹ ‘”‰ƒ‹œƒ–‹‘ƒ––Š‡•‡Ž‡˜‡Ž•‘ˆ

ϰϯϳ –Š‡‘Žˆƒ –‘”› ‹” —‹–ǡ‹†‹˜‹†—ƒŽ‘†‘”ƒ–•‡˜‘‡ƒ”‡•’‘•‡‹•’ƒ”•‡Ž›ƒ†”ƒ†‘Ž›†‹•–”‹„—–‡†

ϭϱ 

ϰϯϴ •‡–• ‘ˆ ‡—”‘• ™‹–Š‹ –Š‡ ’‹”‹ˆ‘” ‘”–‡š ȋ‡ƒ‡” ‡– ƒŽǤǡ ʹͲͲ͹Ǣ ‘‘ ƒ† •ƒƒ •‘ǡ ʹͲͲͻǢ

ϰϯϵ –‡––Ž‡”ƒ†š‡ŽǡʹͲͲͻǢ Š‘•Š‡–ƒŽǤǡʹͲͳͳǢ‹›ƒ‹ Š‹‡–ƒŽǤǡʹͲͳͳǢ‘•—Ž•‹‡–ƒŽǤǡʹͲͳͳȌǤ•‹‰

ϰϰϬ •‹‰Ž‡Ǧ ‡ŽŽ”‡ ‘”†‹‰•ˆ”‘Š‡ƒ†Ǧ”‡•–”ƒ‹‡†ƒ™ƒ‡‹ ‡ǡŠƒƒ†—‘ƒƒŽ›œ‡†‘†‘””‡•’‘•‡

ϰϰϭ ’”‘ˆ‹Ž‡•‘ˆ‹†‹˜‹†—ƒŽ‡—”‘•‹–Š‡ƒ–‡”‹‘”’‹”‹ˆ‘” ‘”–‡šǤ’‘‘†‘”ƒ–’”‡•‡–ƒ–‹‘ǡʹͷΨ

ϰϰϮ ‘ˆ’”‘Œ‡ –‹‘‡—”‘•™‡”‡„”‘ƒ†Ž›‡š ‹–‡†ƒ†”ƒ”‡Ž›‹Š‹„‹–‡†ǡʹͷΨ•Š‘™‡†‘‡š ‹–ƒ–‹‘ƒ†

ϰϰϯ ƒ Ž‡ƒ” ‹Š‹„‹–‹‘ǡ ƒ† ͷͲΨ †‹•’Žƒ›‡† ˜‡”› •‡Ž‡ –‹˜‡ ”‡•’‘•‡• ‹–‡”•‘ˆ‡š ‹–ƒ–‹‘ƒ†

ϰϰϰ ‹Š‹„‹–‹‘ȋŠƒƒ†—‘ǡʹͲͳͲȌǤŠ‡ƒ’’ƒ”‡–”ƒ†‘ ‘‡ –‹‘• ˆ”‘ –Š‡  ‹–”ƒŽ ƒ†

ϰϰϱ –—ˆ–‡† ‡ŽŽƒš‘•–‘–Š‡’‹”‹ˆ‘” ‘”–‡š•—‰‰‡•––Šƒ––Š‡”‡’”‡•‡–ƒ–‹‘•‘ˆ‘†‘”•‹–Š‡‘Žˆƒ –‘”›

ϰϰϲ ‘”–‡šƒ”‡Ž‡ƒ”‡†„›‡š’‡”‹‡ ‡ǤŠ‡’‹”‹ˆ‘” ‘”–‡š™‘—Ž††‡–‡ –‘†‘”•„› ‘’ƒ”‹‰–Š‡–‘

ϰϰϳ ƒ’”‡˜‹‘—•Ž›ƒ “—‹”‡†Ž‹„”ƒ”›‘ˆ‘†‘”•ȋ—ŽŽ‹˜ƒƒ†‹Ž•‘ǡʹͲͲ͵Ȍǡƒ†„—‹Ž†‘†‘”—‹–•ˆ”‘–Š‡

ϰϰϴ Š‡‹ ƒŽ•‹†‡–‹ˆ‹‡†—’•–”‡ƒ‹–Š‡‘Žˆƒ –‘”› ‹” —‹–ȋ ‘Š•‘‡–ƒŽǤǡʹͲͲͲȌǤŠ‡’‹”‹ˆ‘” ‘”–‡š

ϰϰϵ ƒ –Š‡”‡ˆ‘”‡ „‡ •‡‡ ƒ• ƒ ‡‘”› –‘‘Ž ‘’–‹‹œ‡† ‹ –Š‡ •–‘”ƒ‰‡ ‘ˆ ‘†‘” •›ƒ’–‹ 

ϰϱϬ ”‡’”‡•‡–ƒ–‹‘•ȋƒ”ƒ‹‡–ƒŽǤǡͳͻͻͶȌǤ

ϰϱϭ Žˆƒ –‘”› ”‡’”‡•‡–ƒ–‹‘• ™‹–Š‹ –Š‡ ’‹”‹ˆ‘” ‘”–‡š ƒ”‡ •–”‘‰Ž› •Šƒ’‡† „› ”‡ —””‡–

ϰϱϮ ‡š ‹–ƒ–‘”›ƒ†‹Š‹„‹–‘”›‹–”ƒ ‘”–‹ ƒŽ ‘‡ –‹‘•ȋ ”ƒ•‡–ƒŽǤǡʹͲͳͳǢ‘‘ƒ† •ƒƒ •‘ǡʹͲͳͳȌǤ

ϰϱϯ Š‹Ž‡‡ƒ Š’‹”‹ˆ‘” ‘”–‡š’›”ƒ‹†ƒŽ ‡ŽŽ”‡ ‡‹˜‡•‘Ž›ʹͲͲ‹’—–•ˆ”‘‹–”ƒŽȀ–—ˆ–‡† ‡ŽŽ•ǡ

ϰϱϰ ‹– ”‡ ‡‹˜‡• ƒ– Ž‡ƒ•– ʹͲͲͲ ”‡ —””‡– ‡š ‹–ƒ–‘”› ‹’—–• ȋƒ˜‹•‘ ƒ† ŠŽ‡”•ǡ ʹͲͳͳǢ ”ƒ• ‡– ƒŽǤǡ

ϰϱϱ ʹͲͳͳȌǤ ‘–”ƒ”› –‘ –Š‡ ’”‹ƒ”› ˜‹•—ƒŽ ƒ† •‘ƒ–‘•‡•‘”› ‘”–‹ ‡•ǡ ”‡ —””‡– ’‹”‹ˆ‘” ‘”–‡š

ϰϱϲ ƒš‘•ˆ‘”•›ƒ’•‡•™‹–Š–Š‡•ƒ‡’”‘„ƒ„‹Ž‹–›™Šƒ–‡˜‡”–Š‡†‹•–ƒ ‡ˆ”‘–Š‡•‘ƒȋ ”ƒ•‡–

ϰϱϳ ƒŽǤǡ ʹͲͳͳȌǤ ‘ ƒŽ ‘”–‹ ƒŽ ƒ’’Ž‹ ƒ–‹‘ ‘ˆ „ƒ Ž‘ˆ‡ǡ ƒ „ ƒ–ƒ‰‘‹•–ǡ ƒ„‘Ž‹•Š‡• ‹–”ƒ ‘”–‹ ƒŽ

ϰϱϴ ƒ••‘ ‹ƒ–‹‘ƒŽ –”ƒ•‹••‹‘ ˆ”‘ ‡š ‹–ƒ–‘”› ‡—”‘• ™‹–Š‘—– ƒˆˆ‡ –‹‰ Ǧ‡˜‘‡† ‡š ‹–ƒ–‘”›

ϰϱϵ ”‡•’‘•‡• ȋƒ‰ ƒ† ƒ••‡Ž‘ǡ ͳͻͻͶǢ ”ƒ• ƒ† •ƒƒ •‘ǡ ʹͲͲͷȌǤ•‹‰–Š‹•’Šƒ”ƒ ‘Ž‘‰‹ 

ϰϲϬ ƒ’’”‘ƒ Šǡ‘‘ƒ† •ƒƒ •‘•Š‘™‡†–Šƒ––Š‡”‡ ”—‹–‡–‘ˆ‹–”ƒ ‘”–‹ ƒŽ‹’—–ǡ”ƒ–Š‡”–Šƒ

ϰϲϭ ‹’—–ǡ Žƒ”‰‡Ž› †‡–‡”‹‡• –Š‡ •–”‡‰–Š ‘ˆ ‘†‘”Ǧ‡˜‘‡† ‡š ‹–ƒ–‘”› •›ƒ’–‹  –”ƒ•‹••‹‘ ‹ –Š‡

ϰϲϮ ’‹”‹ˆ‘” ‘”–‡šȋ‘‘ƒ† •ƒƒ •‘ǡʹͲͳͳȌǤ•‹‰ƒ‘˜‡Ž–”ƒ•‰‡‹ ‘—•‡‘†‡ŽƒŽŽ‘™‹‰–Š‡

ϰϲϯ ‡š’”‡••‹‘ ‘ˆ Šƒ‡Ž ”Š‘†‘’•‹ ȋŠʹȌ ‹ ƒ •—„•‡– ‘ˆ •‡‹Ž—ƒ” ‡ŽŽ•ǡŠ‘›ƒ† ‘ŽŽ‡ƒ‰—‡•

ϰϲϰ •Š‘™‡†–Šƒ–ŠʹǦ‡š’”‡••‹‰•‡‹Ž—ƒ” ‡ŽŽ•ƒ˜‘‹†–ƒ”‰‡–‹‰ ‡‹‰Š„‘”‹‰ •‡‹Ž—ƒ” ‡ŽŽ•ǡ„—–

ϭϲ 

ϰϲϱ ’”‘˜‹†‡ •–”‘‰ ‘‘•›ƒ’–‹  ƒ••‘ ‹ƒ–‹‘ƒŽ ‡š ‹–ƒ–‹‘ ‘ˆ •—’‡”ˆ‹ ‹ƒŽ ƒ† †‡‡’ ’›”ƒ‹†ƒŽ ‡ŽŽ•

ϰϲϲ ȋŠ‘›‡–ƒŽǤǡʹͲͳͷȌǤ‹”‹ˆ‘” ‘”–‡š‹–‡”‡—”‘•ƒŽ•‘’Žƒ›ƒ‹’‘”–ƒ–”‘Ž‡‹‘†‘”’”‘ ‡••‹‰ǣ

ϰϲϳ –Š‡›ƒŽŽ‘™‹ ”‡ƒ•‹‰–Š‡†‹• ”‹‹ƒ–‹‘‘ˆƒ‘†‘”ƒ†’”‡˜‡–‘Ǧ•’‡ ‹ˆ‹ ‡š ‹–ƒ–‹‘•ǤŽ›

ϰϲϴ ͳͲΨ‘ˆ‡—”‘•™‹ŽŽ„‡ƒ –‹˜ƒ–‡†„›ƒ‰‹˜‡‘†‘”ȋ‡ƒ‡”‡–ƒŽǤǡʹͲͲ͹Ǣ‘‘ƒ† •ƒƒ •‘ǡʹͲͲͻǢ

ϰϲϵ Šƒ ƒ† —‘ǡ ʹͲͳͲȌǤ Š‹• •‹‰ƒŽ –”ƒ•ˆ‘”ƒ–‹‘ ƒ› ƒŽŽ‘™ –Š‡ ’‹”‹ˆ‘” ‘”–‡š –‘ ’‡” ‡‹˜‡

ϰϳϬ ‘’Ž‡š ‹š ‘ˆ ‘†‘”• ƒ• ‘†‘”ƒ– ‘„Œ‡ – •‡’ƒ”ƒ–‡† ˆ”‘ ‹–• ’—”‡ ‘’‘‡–•Ǥ ‡‡†ˆ‘”™ƒ”†

ϰϳϭ ‹Š‹„‹–‹‘‹•‡†‹ƒ–‡†„›Žƒ›‡”ͳŠ‘”‹œ‘–ƒŽƒ†‡—”‘‰Ž‹ƒˆ‘”‹–‡”‡—”‘•ǡ ™Š‹ Š ”‡ ‡‹˜‡

ϰϳϮ †‹”‡ –‹’—–•ˆ”‘–Š‡ǡ•›ƒ’•‡‘–‘ƒ’‹ ƒŽ†‡†”‹–‡•‘ˆ’›”ƒ‹†ƒŽ ‡ŽŽ•ǡƒ†ƒ”‡–Š‘—‰Š––‘

ϰϳϯ ’Žƒ›ƒ’”‘‹‡–”‘Ž‡‹†‡†”‹–‹ ‹–‡‰”ƒ–‹‘‘ˆƒˆˆ‡”‡–‹’—–ˆ”‘ƒš‘•‹ƒŽŽ’‹”‹ˆ‘”

ϰϳϰ ‘”–‡š ‡š ‹–ƒ–‘”› ‡ŽŽ•Ǥ ‡‡†„ƒ  ‹Š‹„‹–‹‘ ‹• ‡†‹ƒ–‡† „› Žƒ›‡” ʹȀ͵ „‹–—ˆ–‡† ”‡‰—Žƒ” •’‹‹‰

ϰϳϱ ‹–‡”‡—”‘•ǡ™Š‹ Š”‡ ‡‹˜‡Ž‹––Ž‡†‹”‡ –‹’—–ƒ†•›ƒ’•‡‘–‘’›”ƒ‹†ƒŽ ‡ŽŽ„‘†‹‡•ƒ†

ϰϳϲ „ƒ•ƒŽ†‡†”‹–‡•ǡ™‹–Š†‡‡’’›”ƒ‹†ƒŽ ‡ŽŽ•”‡ ‡‹˜‹‰–Š‡•–”‘‰‡•–”‡ —””‡–‹Š‹„‹–‹‘ȋ–‘‡•

ϰϳϳ ƒ† •ƒƒ •‘ǡ ʹͲͳͲǢ —œ—‹ ƒ† ‡‡”•ǡ ʹͲͳͲƒǢ ʹͲͳͲ„Ǣ ƒ‹‰— Š‹ǡ ʹͲͳͶǢ ƒ”‰‡ ‡– ƒŽǤǡ ʹͲͳ͸ȌǤ

ϰϳϴ ‡‹Ž—ƒ” ‡ŽŽ• Šƒ˜‡ „‡‡ •Š‘™ –‘ †‹”‡ –Ž› ƒ –‹˜ƒ–‡ Žƒ›‡” ͵ ‡”‰‹  ‹–‡”‡—”‘• ȋˆƒ•–

ϰϳϵ •’‹‹‰ǡ”‡‰—Žƒ”•’‹‹‰ƒ†‡—”‘‰Ž‹ƒˆ‘”Ȍǡ„—–‡‹–Š‡”‡—”‘‰Ž‹ƒˆ‘” ‘” Š‘”‹œ‘–ƒŽ

ϰϴϬ ‹–‡”‡—”‘•‘ˆŽƒ›‡”ͳƒȋŠ‘›‡–ƒŽǤǡʹͲͳͷȌǤŠ‡ˆ‡‡†„ƒ ‹Š‹„‹–‹‘ ™‘—Ž† †‘‹ƒ–‡ –Š‡

ϰϴϭ ˆ‡‡†ˆ‘”™ƒ”† ‹Š‹„‹–‹‘ ‹ ‘–”‘ŽŽ‹‰ –Š‡ ƒ –‹˜ƒ–‹‘ ‘ˆ ’‹”‹ˆ‘” ’›”ƒ‹†ƒŽ ‡ŽŽ• ȋ ”ƒ• ‡– ƒŽǤǡ

ϰϴϮ ʹͲͳͳȌǤ

ϰϴϯ Synaptic organization of the cortex: from 3 to 6 layers

ϰϴϰ Š‡ ‘”–‹ ƒŽ–›’‡‹ ”‘ ‹” —‹– ‘•‹•–•‘ˆƒ•—’‡”ˆ‹ ‹ƒŽ’Ž‡š‹ˆ‘”Žƒ›‡”ͳǡ™Š‡”‡‡š–”‹•‹ 

ϰϴϱ ƒ† ‹–”‹•‹  ’”‘Œ‡ –‹‘• ‡‡– –Š‡ ƒ’‹ ƒŽ †‡†”‹–‡• ‘ˆ ’›”ƒ‹†ƒŽ ‡—”‘• Ž‘ ƒ–‡† ‹ †‡‡’‡”

ϰϴϲ Žƒ›‡”•ǡ”‡•’‘•‹„Ž‡ˆ‘”–Š‡‰‡‡”ƒ–‹‘‘ˆ”‡ —””‡–‡š ‹–ƒ–‹‘ƒ†‹Š‹„‹–‹‘ȋ ƒ„‡”Ž›ǡͳͻͻͲȌǤŠ‡

ϰϴϳ ‡‘ ‘”–‡š •Šƒ”‡• –Š‡ „ƒ•‹  ‹ ”‘ ‹” —‹–• ™‹–Š –Š‡ ͵ Žƒ›‡”‡†ǦƒŽŽ‘ ‘”–‡šǡ„—–†‹•’Žƒ›•Š‹‰Š‡”

ϰϴϴ —„‡”‘ˆ‡—”‘•ƒ†Žƒ›‡”•ȋŠ‡’Š‡”†ǡʹͲͳͳǢ ‘—”‹‡”‡–ƒŽǤǡʹͲͳͷȌǤŠ‡‡‘ ‘”–‡š ‘—Ž†–Š‡

ϰϴϵ „‡†‡• ”‹„‡†ƒ•ƒ†‘—„Ž‡ǦƒŽŽ‘ ‘”–‡šǣ–™‘ ‘—’Ž‡•‘ˆ’›”ƒ‹†ƒŽ ‡ŽŽ’‘’—Žƒ–‹‘•ǡˆ‘”‡†„›—’’‡”

ϰϵϬ Žƒ›‡”•ʹǦ͵ƒ††‡‡’‡”Žƒ›‡”•ͷǦ͸ǡƒ”‡‡ƒ ŠŽ‘ ƒ–‡†„‡Ž‘™’Ž‡š‹ˆ‘”Žƒ›‡”•ͳƒ†Ͷ”‡•’‡ –‹˜‡Ž›ǡ

ϭϳ 

ϰϵϭ ƒ””›‹‰ ‡š–”‹•‹  ‹’—–• ȋŠ‡’Š‡”†ǡ ʹͲͳͳǢ —œœƒ–‹ǡ ʹͲͳͷȌǤ  ‘–”ƒ•– –‘ –Š‡ †‘”•ƒŽ ‘”–‡š ‘ˆ

ϰϵϮ ”‡’–‹Ž‡•‘”–‘–Š‡’‹”‹ˆ‘” ‘”–‡š‘ˆƒƒŽ•ǡ™Š‡”‡•‡•‘”›ƒˆˆ‡”‡–• ‘‡ˆ”‘–Š‡–‘’ǡ‹–Š‡

ϰϵϯ ‡‘ ‘”–‡š–Š‡–ŠƒŽƒ‹ ƒˆˆ‡”‡–•ƒ• ‡†ˆ”‘–Š‡†‡’–Šƒ†‡ˆˆ‡”‡–•†‡• ‡†–Š”‘—‰Š–Š‡•ƒ‡

ϰϵϰ Žƒ›‡”•Ǥ ‘–Š ‹’—–• ƒ† ‘—–’—–• Šƒ˜‡ –Š‡”‡ˆ‘”‡ ’‘–‡–‹ƒŽ ƒ ‡•• –‘ ƒŽŽ ‡ŽŽ• ‹ ‡˜‡”› Žƒ›‡”

ϰϵϱ ȋŠ‡’Š‡”†ǡ ʹͲͳͳȌǤ ‘”‡‘˜‡”ǡ –Š‡ ‰‡‡”ƒ–‹‘ ‘ˆ •—’‡”ˆ‹ ‹ƒŽ ƒ† †‡‡’ Žƒ›‡”• ‹ –Š‡ ‡‘ ‘”–‡š

ϰϵϲ ƒŽŽ‘™•‡š’ƒ•‹‘‘ˆ–Š‡ ‘„‹ƒ–‘”‹ƒŽ’‘••‹„‹Ž‹–‹‡•ˆ‘”‹–”ƒ ‘”–‹ ƒŽƒ†•—„ ‘”–‹ ƒŽ’”‘ ‡••‹‰

ϰϵϳ ȋŠ‡’Š‡”†ǡ ʹͲͳͳȌǤ‡‘ ‘”–‡š—’’‡” Žƒ›‡”•ƒ”‡ ‡˜‘Ž—–‹‘ƒ”› –Š‡‘•–”‡ ‡–Ǥ –‡”‡•–‹‰Ž›ƒ†

ϰϵϴ ‘—–‡”‹–—‹–‹˜‡Ž›ǡ’‹”‹ˆ‘” ‘”–‡š‡—”‘•‡š’”‡••‘”‡‘Ž‡ —Žƒ”ƒ”‡”• ‘‘–‘–Š‡•‡

ϰϵϵ ‡‘ ‘”–‡š —’’‡” Žƒ›‡”•Ǥ Š‡•‡ •‹‹Žƒ”‹–‹‡• Ž‡ƒ† –‘ –Š‡ Š›’‘–Š‡•‹• ‘ˆ ƒ ‘Žˆƒ –‘”› ‘”–‡šǦ„ƒ•‡†

ϱϬϬ †‡˜‡Ž‘’‡–ƒŽ’”‘‰”ƒˆ‘”–Š‡‡˜‘Ž—–‹‘‘ˆ–Š‡‡‘ ‘”–‹ ƒŽŽƒ›‡”•ʹȀ͵ȋ—œœƒ–‹ǡʹͲͳͷȌǤŠ‡‡ƒ”Ž›

ϱϬϭ ‡‘ ‘”–‹ ƒŽ ‘Ž— ™‘—Ž† –Š‡”‡ˆ‘”‡ „‡ –Š‡ ”‡•—Ž– ‘ˆ –Š‡ •—’‡”’‘•‹–‹‘ ‘ˆ –Š‡ Žƒ–‡”ƒŽ ‘”–‡š

ϱϬϮ ȋ‘Žˆƒ –‘”› ‘”–‡šȌƒ†–Š‡†‘”•ƒŽ ‘”–‡šȋ—œœƒ–‹ǡʹͲͳͷȌǤƒ”‡ˆ—Ž•–—†‹‡•‘ˆ’‹”‹ˆ‘” ‘”–‡š ‡ŽŽ

ϱϬϯ Ž‹‡ƒ‰‡• ƒ† ‡—”‘ ‘Ž‡ —Žƒ” ‹†‡–‹–‹‡• ƒ”‡ ‡‡†‡† –‘ —†‡”•–ƒ† ™Šƒ– ƒ”‡ –Š‡ •’‡ ‹ˆ‹  ‡ŽŽ

ϱϬϰ –›’‡•ˆ‘—†‹‡˜‘Ž—–‹‘ƒ”›Dz‘Ž†dz‘Žˆƒ –‘”› ‘”–‡šƒ†‡‘ ‘”–‡šŽƒ›‡”•Ǥ

ϱϬϱ Concluding remarks

ϱϬϲ Š‡‘Žˆƒ –‘”› ‘”–‡šǡ•‘Ǧ ƒŽŽ‡†Ǯ•‹’Ž‡ǯ ‘”–‡š‹–Š‡Ž‹–‡”ƒ–—”‡™Š‡ ‘’ƒ”‡†–‘–Š‡•‹šǦ

ϱϬϳ Žƒ›‡”‡† ‡‘ ‘”–‡šǡ ‹• ‘ˆ–‡ ’”‡•‡–‡† ƒ• ƒ ‰‡‡”ƒŽ ‘†‡Ž ˆ‘” ‘”–‹ ƒŽ•‡•‘”›’”‘ ‡••‹‰Ǥ

ϱϬϴ ‘™‡˜‡” –Š‡ ‘Ž‡ —Žƒ” ‹†‡–‹–‹‡• ƒ† –Š‡ •’‡ ‹ˆ‹  ˆ— –‹‘• ‘ˆ ‡ŽŽ• ‘’‘•‹‰ ‡ƒ Š ‘Žˆƒ –‘”›

ϱϬϵ ‘”–‡šŽƒ›‡””‡ƒ‹’‘‘”Ž›†‡• ”‹„‡†ǤŠ‡•‡“—‡–‹ƒŽ‰‡‡”ƒ–‹‘‘ˆ’›”ƒ‹†ƒŽ ‡ŽŽ•ǡ‹–‡”‡—”‘•

ϱϭϬ ƒ†‰Ž‹ƒǡƒ•™‡ŽŽƒ•–Š‡‹•‹†‡Ǧ‘—–‹‰”ƒ–‹‘‘ˆ’›”ƒ‹†ƒŽ ‡ŽŽ••‡‡–‘„‡ ‘•‡”˜‡†’”‘ ‡••‡•‹

ϱϭϭ „‘–Š‘Žˆƒ –‘”› ‘”–‡šƒ†‡‘ ‘”–‡š†—”‹‰†‡˜‡Ž‘’‡–Ǥ –Š‡‡‘ ‘”–‡šǡ‡š ‹–ƒ–‘”›‡—”‘•

ϱϭϮ †‡”‹˜‡ˆ”‘’ƒŽŽ‹ƒŽƒ†‹‰”ƒ–‡”ƒ†‹ƒŽŽ›ǡ™Š‹Ž‡‹Š‹„‹–‘”›‡—”‘•†‡”‹˜‡ˆ”‘•—„’ƒŽŽ‹ƒŽ

ϱϭϯ ƒ†‹‰”ƒ–‡–ƒ‰‡–‹ƒŽŽ›ǤŠ‹••‡‡•‘––‘„‡ƒ•‘„˜‹‘—•‹–Š‡‘Žˆƒ –‘”› ‘”–‡šǡ™Š‡”‡•‘‡

ϱϭϰ ‡—”‘• ˆ”‘ –Š‡ ’ƒŽŽ‹— †‹ˆˆ‡”‡–‹ƒ–‡ ‹ ‹Š‹„‹–‘”› ‡—”‘• ƒ† •‘‡ ‡—”‘• ˆ”‘ –Š‡

ϱϭϱ •—„’ƒŽŽ‹— ‹ ‡š ‹–ƒ–‘”› ‡—”‘• ȋ ƒ” ‹ƒǦ‘”‡‘ ‡– ƒŽǤǡ ʹͲͲͺȌ ȋ ‹‰—”‡ͳȌǤŠ‡”‘—–‡•‘ˆ

ϱϭϲ ‹‰”ƒ–‹‘–‘™ƒ”†–Š‡‘Žˆƒ –‘”› ‘”–‡šƒ”‡’ƒ”–‹ —Žƒ”Ž› ‘’Ž‹ ƒ–‡† •‹ ‡ ‘Žˆƒ –‘”› ‘”–‡š

ϭϴ 

ϱϭϳ ‡—”‘•Šƒ˜‡—Ž–‹’Ž‡‘”‹‰‹•‹–Š‡ƒ†•‘‡‘ˆ–Š‡‹‰”ƒ–‡‘˜‡”Ž‘‰†‹•–ƒ ‡–‘”‡ƒ Š

ϱϭϴ –Š‡‹” ˆ‹ƒŽ Ž‘ ƒ–‹‘Ǥ Š‡•‡ ”‘—–‡• ‹’Ž‹ ƒ–‡ –ƒ‰‡–‹ƒŽ ƒ• ™‡ŽŽ ƒ• ”ƒ†‹ƒŽ ‹‰”ƒ–‹‘•Ǥ Š‡ ‘•–

ϱϭϵ •–—†‹‡†‹‰”ƒ–‘”›•–”‡ƒ–‘–Š‡‘Žˆƒ –‘”› ‘”–‡š‹•–Š‡Žƒ–‡”ƒŽ ‘”–‹ ƒŽ•–”‡ƒ–Š‘—‰Š––‘„‡–Š‡

ϱϮϬ ƒ‹ •–”‡ƒ ˆ‘” ‘Žˆƒ –‘”› ‘”–‡š ‡š ‹–ƒ–‘”› ‡—”‘•Ǥ ‘‡ ”ƒ†‹ƒŽ ‰Ž‹ƒŽ ’”‘ ‡••‡• ȋ”ƒ†‹ƒŽ ‰Ž‹ƒŽ

ϱϮϭ ’ƒŽ‹•ƒ†‡ȌŠƒ˜‡„‡‡‹†‡–‹ˆ‹‡†ƒŽ‘‰–Š‹••–”‡ƒƒ†’‘••‹„Ž›ƒŽŽ‘™–Š‡–ƒ‰‡–‹ƒŽǦŽ‹‡‹‰”ƒ–‹‘

ϱϮϮ ‘ˆ‡š ‹–ƒ–‘”›‡—”‘•ƒŽ‘‰”ƒ†‹ƒŽ‰Ž‹ƒȋƒ”‡›‡–ƒŽǤǡʹͲͲ͸Ȍȋ ‹‰—”‡ʹȌǤ

ϱϮϯ  Š‡†‹˜‡”•‹–›‘ˆ–Š‡‹”‘”‹‰‹•‹’Ž‹‡•–Šƒ––Š‡ ‡ŽŽ• ‘’‘•‹‰ –Š‡ ‘Žˆƒ –‘”› ‘”–‡š

ϱϮϰ ‘‡ˆ”‘†‹ˆˆ‡”‡– ‡ŽŽŽ‹‡ƒ‰‡•ƒ†•—‰‰‡•–•–Šƒ–†‹ˆˆ‡”‡– ‡ŽŽ’‘’—Žƒ–‹‘••Š‘—Ž†„‡†‡ˆ‹ƒ„Ž‡

ϱϮϱ ƒ‘‰‘Žˆƒ –‘”› ‘”–‡šŽƒ›‡”•Ǥ‘ˆƒ”–™‘’›”ƒ‹†ƒŽ ‡ŽŽ’‘’—Žƒ–‹‘•Šƒ˜‡ƒ‹Ž›„‡‡†‡• ”‹„‡†

ϱϮϲ ‹–Š‡’‹”‹ˆ‘” ‘”–‡šŽƒ›‡”ʹ„ƒ•‡†‘–Š‡‹”‘”’Š‘Ž‘‰‹ ƒŽƒ†‡Ž‡ –”‘’Š›•‹‘Ž‘‰‹ ƒŽ’”‘’‡”–‹‡•

ϱϮϳ ȋ—œ—‹ƒ†‡‡”•ǡʹͲͳͳȌȋ ‹‰—”‡͵ȌǤ ‘™‡˜‡”ǡ”‡ ‡–•–—†‹‡••–ƒ”–‡††‡ ‘†‹‰–Š‡‘Ž‡ —Žƒ”

ϱϮϴ •‹‰ƒ–—”‡•‘ˆ’”‘Œ‡ –‹‘‡—”‘•‹”‡•’‡ ––‘–Š‡‹” ‘‡ –‹˜‹–›’”‘’‡”–‹‡•ȋ‹‘†ƒ–‘‡–ƒŽǤǡʹͲͳ͸Ǣ

ϱϮϵ ƒ†ƒƒ„Šƒ‡–ƒŽǤǡʹͲͳ͸Ȍȋ ‹‰—”‡͵ȌǤ —”–Š‡”‹˜‡•–‹‰ƒ–‹‘•™‹ŽŽƒŽŽ‘™—†‡”•–ƒ†‹‰™Š‡–Š‡”

ϱϯϬ –Š‡†‹ˆˆ‡”‡–‘”‹‰‹•ƒ†‹‰”ƒ–‘”›’”‘’‡”–‹‡•‘ˆ‘Žˆƒ –‘”› ‘”–‡š ‡—”‘• †—”‹‰

ϱϯϭ †‡˜‡Ž‘’‡– ƒ”‡ ‘””‡Žƒ–‡† ™‹–Š –Š‡‹” Š‡–‡”‘‰‡‡‘—• ‹†‡–‹–‹‡• ƒ† ˆ— –‹‘• ‹ ‘†‘”

ϱϯϮ ’”‘ ‡••‹‰Ǥ ††”‡••‹‰ ‘Žˆƒ –‘”› ‘”–‡š ‡—”‘ –”ƒ• ”‹’–‹‘ƒŽ ‹†‡–‹–‹‡• –Š”‘—‰Š‘—–

ϱϯϯ †‡˜‡Ž‘’‡– ™‹ŽŽ Š‡Ž’ –‘ ‘Ž‡ —Žƒ”Ž› Šƒ”ƒ –‡”‹œ‡ ‡ŽŽ ’‘’—Žƒ–‹‘• ™‹–Š •’‡ ‹ˆ‹  ’”‘’‡”–‹‡• ‹

ϱϯϰ –Š‡‘Žˆƒ –‘”› ‘”–‡šǡ‹‘”†‡”–‘ ‘’ƒ”‡–Š‡™‹–Š’‘’—Žƒ–‹‘•‹†‡–‹ˆ‹‡†‹–Š‡‡‘ ‘”–‡šǡ‹

ϱϯϱ –‡”• ‘ˆ Žƒ›‡” Ž‘ ƒŽ‹œƒ–‹‘ǡ ‘‡ –‹˜‹–› ȋ‹’—– ƒ† ‘—–’—–Ȍǡ ƒ† ˆ— –‹‘Ǥ Š‡ ’‹”‹ˆ‘” ‘”–‡š

ϱϯϲ ƒ†–Š‡†‘”•ƒŽ ‘”–‡š‘ˆ”‡’–‹Ž‡•ȋ ‘””‡•’‘†‹‰–‘˜‹•—ƒŽ ‘”–‡šȌ„‘–Š•‡‡–‘’”‘ ‡•••‡•‘”›

ϱϯϳ ‹’—–•ƒ•Š‹‰ŠǦ ‘”–‹ ƒŽƒ”‡ƒ•”ƒ–Š‡”–Šƒ’”‹ƒ”›•‡•‘”›‡‘ ‘”–‡šǤ‘’—–ƒ–‹‘•’‡”ˆ‘”‡†

ϱϯϴ „›Š‹‰ŠǦ‘”†‡” ‘”–‹ ƒŽƒ”‡ƒ••‡‡–Š‡”‡ˆ‘”‡–‘„‡ƒ ‡•–”ƒŽ™Š‹Ž‡ ‘’—–ƒ–‹‘• ’‡”ˆ‘”‡† ƒ–

ϱϯϵ ‹‹–‹ƒŽ •–ƒ‰‡• ‘ˆ ‡‘ ‘”–‹ ƒŽ ’”‘ ‡••‹‰ ƒ’’‡ƒ”‡† Žƒ–‡” ‹ ‡˜‘Ž—–‹‘ǡ ’‘••‹„Ž› Ž‹‡† –‘ –Š‡

ϱϰϬ ƒ††‹–‹‘•‘ˆ‡™Žƒ›‡”•ȋ ‘—”‹‡”‡–ƒŽǤǡʹͲͳͷȌǤ”ƒ˜‡Ž‹‰’”‡ ‹•‡ ‡—”‘ƒŽ ‘”‹‰‹• ƒ†

ϱϰϭ ‹†‡–‹–‹‡• ‹ „‘–Š –Š‡ ‡‘ ‘”–‡š ƒ† –Š‡ ‘Žˆƒ –‘”› ‘”–‡š ™‹ŽŽ ˆ—”–Š‡” ‡Ž— ‹†ƒ–‡ ‡˜‘Ž—–‹‘ƒ”›

ϱϰϮ ‘•‡”˜‡†’”‘’‡”–‹‡•‘ˆ•‡•‘”› ‘”–‹ ‡•Ǥ

ϱϰϯ

ϭϵ 

ϱϰϰ References

ϱϰϱ „‘‹–‹œ ǡ‘–‹‡Ž ǡ‘”ƒŽ‡•ǡ‘ ŠƒȋʹͲͲʹȌ˜‘Ž—–‹‘ƒ”›†‹˜‡”‰‡ ‡‘ˆ–Š‡”‡’–‹Ž‹ƒƒ†–Š‡

ϱϰϲ ƒƒŽ‹ƒ „”ƒ‹•ǣ ‘•‹†‡”ƒ–‹‘• ‘ ‘‡ –‹˜‹–› ƒ† †‡˜‡Ž‘’‡–Ǥ ”ƒ‹ ‡•‡ƒ” Š ‡˜‹‡™•

ϱϰϳ ͵ͻǣͳͶͳȂͳͷ͵Ǥ

ϱϰϴ Ž ƒ‘ǡŠ‹”‹˜‡ŽŽƒǡƒ—–œ‡„‡”‰ǡ‘„”‡˜ƒ ǡ ƒ”‹Óƒ• ǡ ”‘•• Š‡†Ž ǡ  ‘‡ŽŽ 

ϱϰϵ ȋʹͲͲͺȌ ƒ–„ʹ ‡‰—Žƒ–‡• ƒŽŽ‘•ƒŽ ”‘Œ‡ –‹‘ ‡—”‘ †‡–‹–› ‹ –Š‡ ‡˜‡Ž‘’‹‰ ‡”‡„”ƒŽ

ϱϱϬ ‘”–‡šǤ‡—”‘ͷ͹ǣ͵͸ͶȂ͵͹͹Ǥ

ϱϱϭ Ž ƒ–ƒ”ƒ ǡ —‹œ ǡ ” ƒ‰‡Ž‘ ǡ œƒ ǡ †‡ ‡ ‡ƒ ǡ —””ƒ ǡ ‘–‡Ž‘ ǡ ‘”‹ƒ‘  ȋͳͻͻͺȌ

ϱϱϮ ‡‰‹‘ƒŽƒ†‡ŽŽ—Žƒ”ƒ––‡”•‘ˆreelinš’”‡••‹‘‹–Š‡ ‘”‡„”ƒ‹‘ˆ–Š‡‡˜‡Ž‘’‹‰

ϱϱϯ ƒ††—Ž–‘—•‡Ǥ ‘—”ƒŽ‘ˆ‡—”‘• ‹‡ ‡ͳͺǣ͹͹͹ͻȂ͹͹ͻͻǤ

ϱϱϰ †‡”•‘ǡ‹•‡•–ƒ–ǡŠ‹ǡ—„‡•–‡‹ ȋͳͻͻ͹Ȍ –‡”‡—”‘ ‹‰”ƒ–‹‘ ˆ”‘ „ƒ•ƒŽ

ϱϱϱ ˆ‘”‡„”ƒ‹–‘‡‘ ‘”–‡šǣ†‡’‡†‡ ‡‘Žš‰‡‡•Ǥ ‹‡ ‡ʹ͹ͺǣͶ͹ͶȂͶ͹͸Ǥ

ϱϱϲ –‘ǡƒ” Š‹‘‹ǡ‡‡ ǡƒ‹ ȋͳͻͻ͹Ȍ‘Ž‡‘ˆ Ȁ‡—”‡‰—Ž‹ •‹‰ƒŽ‹‰ ‹

ϱϱϳ ‹–‡”ƒ –‹‘•„‡–™‡‡‹‰”ƒ–‹‰‡—”‘•ƒ†”ƒ†‹ƒŽ‰Ž‹ƒ‹–Š‡†‡˜‡Ž‘’‹‰ ‡”‡„”ƒŽ ‘”–‡šǤ

ϱϱϴ ‡˜‡Ž‘’‡–ͳʹͶǣ͵ͷͲͳȂ͵ͷͳͲǤ

ϱϱϵ ”ƒ‹ǡ‹‡”ƒ‹ȋʹͲͳͶȌ‡˜‡Ž‘’‡–ƒ†‡˜‘Ž—–‹‘‘ˆ ‘”–‹ ƒŽˆ‹‡Ž†•Ǥ ‡—”‘• ‹‡ ‡ ‡•‡ƒ” Š

ϱϲϬ ͺ͸ǣ͸͸Ȃ͹͸Ǥ

ϱϲϭ ••‹ƒ ‘’‘—Ž‘•ǡ ”‘˜‡ǡƒ‰•†ƒŽ‡ȋʹͲͲ͵Ȍ †‡–‹ˆ‹ ƒ–‹‘‘ˆƒƒš͸Ǧ†‡’‡†‡–‡’‹†‡”ƒŽ

ϱϲϮ ‰”‘™–Šˆƒ –‘”ˆƒ‹Ž›•‹‰ƒŽ‹‰•‘—” ‡ƒ––Š‡Žƒ–‡”ƒŽ‡†‰‡‘ˆ–Š‡‡„”›‘‹  ‡”‡„”ƒŽ ‘”–‡šǤ 

ϱϲϯ ‡—”‘• ‹ʹ͵ǣ͸͵ͻͻȂ͸ͶͲ͵Ǥ

ϱϲϰ ƒ‹ ǡƒ‘•ǡƒ”ƒƒ•‹˜ƒǡ‹††‹“‹ ǡ ƒ ǡ‘—” ‘ ȋʹͲͲͺȌŠ‡‘Ž‡‘ˆƒ†

ϱϲϱ  ͳ‹‹‰”ƒ–‹‘–Š”‘—‰Š–Š‡ƒ–‡”ƒŽ‘”–‹ ƒŽ–”‡ƒ‘ˆ‡˜‡Ž‘’‹‰ ‘”‡„”ƒ‹Ǥ‡˜‡—”‘• ‹

ϱϲϲ ͵ͲǣͳͶͶȂͳͷ͸Ǥ

ϮϬ 

ϱϲϳ ƒ”„‡”ǡ”ƒ‹ǡ‘”‹•Š‹–ƒǡ‹‰‹‡”ǡƒ—•‡”‡– ǡ‘”‡ŽŽ‘ǡ‡†‘‡ ǡ‘’’‘Žƒǡ

ϱϲϴ ‘–”‡‘—Ž‹•ǡˆ”‹‡‰‡” ǡ‹••‹” ǡ ‘˜‹†ƒǡ ƒ„ƒ—†‘ǡ”‘—šǦ ‹ŽŽƒ”†‡ƒ—šǡ ƒŽŽ‹ǡ

ϱϲϵ ‹‡”ƒ‹ȋʹͲͳͷȌ‹‰”ƒ–‹‘’‡‡†‘ˆƒŒƒŽǦ‡–œ‹—•‡ŽŽ•‘†—Žƒ–‡†„›‡•‹ —Žƒ””ƒˆˆ‹ ‹‰

ϱϳϬ ‘–”‘Ž•–Š‡‹œ‡‘ˆ ‹‰Š‡”Ǧ”†‡”‘”–‹ ƒŽ”‡ƒ•Ǥ—””‡–‹‘Ž‘‰›ʹͷǣʹͶ͸͸ȂʹͶ͹ͺǤ

ϱϳϭ ƒ”ƒ‹ǡ‡”‰ƒǡ ‘”™‹–œ ǡ ƒ••‡Ž‘ȋͳͻͻͶȌ‘†—Žƒ–‹‘‘ˆ ƒ••‘ ‹ƒ–‹˜‡ ‡‘”›

ϱϳϮ ˆ— –‹‘ ‹ ƒ „‹‘’Š›•‹ ƒŽ •‹—Žƒ–‹‘ ‘ˆ ”ƒ– ’‹”‹ˆ‘” ‘”–‡šǤ ‘—”ƒŽ ‘ˆ ‡—”‘’Š›•‹‘Ž‘‰›

ϱϳϯ ͹ʹǣ͸ͷͻȂ͸͹͹Ǥ

ϱϳϰ ƒ›‡”ȋͳͻͺ͸Ȍ‡—”‘‰‡‡•‹•‹–Š‡”ƒ–’”‹ƒ”›‘Žˆƒ –‘”› ‘”–‡šǤ – ‡˜‡—”‘• ‹ͶǣʹͷͳȂʹ͹ͳǤ

ϱϳϱ ‡‡”• ǡ —œ—‹ ȋʹͲͳ͵Ȍ‡—”‘•ƒ† ‹” —‹–• ˆ‘”‘†‘” ’”‘ ‡••‹‰‹–Š‡’‹”‹ˆ‘” ‘”–‡šǤ

ϱϳϲ ”‡†•‹‡—”‘• ‹‡ ‡•͵͸ǣͶʹͻȂͶ͵ͺǤ

ϱϳϳ ‹‡ŽŽ‡ ǡ ”‹˜‡ƒ—ǡƒ”„‘—šǦ²‡ǡ‹‰‡ƒ—ǡ‹‰”‹•–ǡ”„‡”ǡƒ••‡ˆǡ‹‡”ƒ‹ȋʹͲͲͷȌ

ϱϳϴ —Ž–‹’Ž‡‘”‹‰‹•‘ˆƒŒƒŽǦ‡–œ‹—• ‡ŽŽ•ƒ––Š‡„‘”†‡”•‘ˆ–Š‡†‡˜‡Ž‘’‹‰’ƒŽŽ‹—Ǥƒ–‡—”‘• ‹

ϱϳϵ ͺǣͳͲͲʹȂͳͲͳʹǤ

ϱϴϬ ‘””‡ŽŽǡ ږœȋʹͲͳͶȌ‘Ž‡‘ˆ”ƒ†‹ƒŽ‰Ž‹ƒŽ ‡ŽŽ•‹ ‡”‡„”ƒŽ ‘”–‡šˆ‘Ž†‹‰Ǥ—””‡–’‹‹‘‹

ϱϴϭ ‡—”‘„‹‘Ž‘‰›ʹ͹ǣ͵ͻȂͶ͸Ǥ

ϱϴϮ ‘›Ž‡ǡ‡”ƒ”†ǡŠ‘’•‘ǡ‰ǡ‘‡ǡ‘”–”—†ǡ ƒ™”›Ž› œ ǡ ‘‡•ǡ ‡˜‡” ǡ

ϱϴϯ ‡‹ȋʹͲͳͳȌ‡ŽŽǦ–›’‡Ǧ•’‡ ‹ˆ‹  ‘•‡“—‡ ‡•‘ˆ”‡‡Ž‹†‡ˆ‹ ‹‡ ›‹–Š‡‘—•‡‡‘ ‘”–‡šǡ

ϱϴϰ Š‹’’‘ ƒ’—•ǡƒ†ƒ›‰†ƒŽƒǤ ‘’‡—”‘ŽͷͳͻǣʹͲ͸ͳȂʹͲͺͻǤ

ϱϴϱ ”—Œ‡•ǡ ŽŽ‹‰ǡ‡›‡”ȋʹͲͲͷȌ ˆ‹‡Ž†‰—‹†‡–‘–Š‡ƒ–‡”‹‘”‘Žˆƒ –‘”›— Ž‡—•ȋ ‘”–‡šȌǤ

ϱϴϲ ”ƒ‹‡•‡ƒ” Š‡˜‹‡™•ͷͲǣ͵ͲͷȂ͵͵ͷǤ

ϱϴϳ — ǡš‡ŽȋͳͻͻͳȌ‘˜‡Ž—Ž–‹‰‡‡ˆƒ‹Ž›ƒ›‡ ‘†‡‘†‘”ƒ–”‡ ‡’–‘”•ǣƒ‘Ž‡ —Žƒ”„ƒ•‹•

ϱϴϴ ˆ‘”‘†‘””‡ ‘‰‹–‹‘Ǥ‡ŽŽ͸ͷǣͳ͹ͷȂͳͺ͹Ǥ

ϱϴϵ —Ž Šƒ†ǡ ”‘˜‡ǡ‘”–‡” ǡ‘Ž‡ȋʹͲͲͳȌ ǦŠ‘‡‘†‘ƒ‹‰‡‡ Ššʹ ”‡‰—Žƒ–‡• –Š‡

Ϯϭ 

ϱϵϬ ˆ‘”ƒ–‹‘‘ˆ–Š‡ ‘”–‹ ƒŽŠ‡Ǥ‡ Šƒ‹••‘ˆ‡˜‡Ž‘’‡–ͳͲͲǣͳ͸ͷȂͳ͹ͷǤ

ϱϵϭ ƒ” ‡ŽŽ‡” ǡ‘˜‹”ƒǦ•–‡„ƒǡƒ Š‡” ǡƒ•–”±ǡ —‹”ƒ†‘ȋʹͲͳ͸Ȍ‡—”‘ Š‡‹ ƒŽŠ‡‘–›’‡

ϱϵϮ ‘ˆ‡‡Ž‹ —‘”‡ƒ –‹˜‡‡ŽŽ•‹–Š‡‹”‹ˆ‘”‘”–‡šƒ›‡” Ǥ ”‘–‡ŽŽ‡—”‘• ‹ͳͲǣ͸ͷǤ

ϱϵϯ ƒ”‡›ǡŽˆ‘•‘ǡ‘Š‡ǡƒ‹ ǡ‡”›ǡ–‘‹ ƒǡŽ‘–‹ ǡ”‡‰ƒǡ ‹•Š‡ŽŽ ǡ‘”„‹

ϱϵϰ  ȋʹͲͲ͸Ȍ ‡ŽŽ ‹‰”ƒ–‹‘ ƒŽ‘‰ –Š‡ ƒ–‡”ƒŽ ‘”–‹ ƒŽ –”‡ƒ –‘ –Š‡ ‡˜‡Ž‘’‹‰ ƒ•ƒŽ

ϱϵϱ ‡Ž‡ ‡’ŠƒŽ‹ ‹„‹ ›•–‡Ǥ ‘—”ƒŽ‘ˆ‡—”‘• ‹‡ ‡ʹ͸ǣͳͳͷ͸ʹȂͳͳͷ͹ͶǤ

ϱϵϲ ƒ”‡› ǡ ‘ ƒ• ǡ ‹”ƒ–ƒ ǡ ƒ•ˆ‹‡Ž† ǡ ‘”„‹  ȋʹͲͲͻȌ ‹ˆˆ‡”‡–‹ƒŽ ‡‰—Žƒ–‹‘ ‘ˆ

ϱϵϳ ‡Ž‡ ‡’ŠƒŽ‹  ƒŽŽ‹ƒŽǦ—„’ƒŽŽ‹ƒŽ ‘—†ƒ”› ƒ––‡”‹‰ „› ƒš͸ ƒ† •ŠʹǤ ‡”‡„”ƒŽ ‘”–‡š

ϱϵϴ ͳͻǣ͹ͶͷȂ͹ͷͻǤ

ϱϵϵ ƒ˜‹‡••ȋͳͻͺʹȌ‡‘ ‘”–‹ ƒŽŠ‹•–‘‰‡‡•‹•‹‘”ƒŽƒ†”‡‡Ž‡”‹ ‡ǣƒ†‡˜‡Ž‘’‡–ƒŽ•–—†›

ϲϬϬ „ƒ•‡†—’‘ȏ͵ Ȑ–Š›‹†‹‡ƒ—–‘”ƒ†‹‘‰”ƒ’Š›Ǥ”ƒ‹‡•‡ƒ” Šʹͷ͸ǣʹͻ͵Ȃ͵ͲʹǤ

ϲϬϭ ‡”ƒ‹ǡŠƒ‘ǡ ”‘–• Š‡”ȋʹͲͲ͸Ȍ‡˜‡Ž‘’‡–‘ˆ–Š‡–‘”Š‹‘Ǧ ‹’’‘ ƒ’ƒŽ”‘Œ‡ –‹‘ǣ

ϲϬϮ —‹†ƒ ‡„›ƒŒƒŽǦ‡–œ‹—•‡ŽŽš‘•ǤƒŽ•‘ˆ–Š‡‡™‘” ƒ†‡›‘ˆ ‹‡ ‡•ͻͳͳǣͶ͵Ȃ

ϲϬϯ ͷͶǤ

ϲϬϰ Šƒ’‘—–‘ǡ ¡”–‡”ǡ ږœȋͳͻͻͻȌŠ‡”‘Ž‡‘ˆƒš͸‹”‡•–”‹ –‹‰ ‡ŽŽ‹‰”ƒ–‹‘„‡–™‡‡

ϲϬϱ †‡˜‡Ž‘’‹‰ ‘”–‡šƒ†„ƒ•ƒŽ‰ƒ‰Ž‹ƒǤ‡˜‡Ž‘’‡–ǣͳȂͳͳǤ

ϲϬϲ Š‘› ǡ—œ—‹ǡŠ‹ƒǡ—†‹•ƒ–‘•‘ǡ‡Ž•‘ǡ‡‡”• ȋʹͲͳͷȌ’–‘‰‡‡–‹ ƒ’’‹‰

ϲϬϳ ‘ˆ –”ƒ ‘”–‹ ƒŽ ‹” —‹–• ”‹‰‹ƒ–‹‰ ˆ”‘ ‡‹Ž—ƒ” ‡ŽŽ• ‹ –Š‡ ‹”‹ˆ‘” ‘”–‡šǤ ‡”‡„”ƒŽ

ϲϬϴ ‘”–‡šǣ„Š˜ʹͷͺȂͳ͵Ǥ

ϲϬϵ ‘ ƒ•ǡ ‡‘”‰ƒŽƒǡƒ‰‹ ǡŽ‡‰‰ ǡ‡••ƒ”‹•ǡ ƒ›†ƒ” ǡ ƒŽŽ‘ǡ”‹ ‡ ǡ‘”„‹ 

ϲϭϬ ȋʹͲͳͳȌƒš͸ •‡“—‹”‡†ƒ––Š‡‡Ž‡ ‡’ŠƒŽ‹ ƒŽŽ‹ƒŽǦ—„’ƒŽŽ‹ƒŽ‘—†ƒ”›ˆ‘”–Š‡ ‡‡”ƒ–‹‘

ϲϭϭ ‘ˆ‡—”‘ƒŽ‹˜‡”•‹–›‹–Š‡‘•–ƒ–ƒŽ‹„‹ ›•–‡Ǥ ‘—”ƒŽ‘ˆ ‡—”‘• ‹‡ ‡ ͵ͳǣͷ͵ͳ͵Ȃ

ϲϭϮ ͷ͵ʹͶǤ

ϮϮ 

ϲϭϯ ‘”„‹ ǡ‡”›ǡ ‹•Š‡ŽŽ ȋʹͲͲͳȌ‡Ž‡ ‡’ŠƒŽ‹  ‡ŽŽ•–ƒ‡ƒ–ƒ‰‡–ǣ‘Ǧ”ƒ†‹ƒŽ‹‰”ƒ–‹‘‹–Š‡

ϲϭϰ ƒƒŽ‹ƒˆ‘”‡„”ƒ‹Ǥƒ–‡—”‘• ‹Ͷǣͳͳ͹͹ȂͳͳͺʹǤ

ϲϭϱ —„‡Ž‘•ǡ‡„ƒ•–‹žǦ‡””ƒ‘ǡ‡ ƒ”‹ǡƒŽ ƒ‰‘––‘ǡ‹•‡”‘•ǡ‹ǡ‘’ƒœ‘ǡŽ˜ƒ”‡œǦ

ϲϭϲ ‘Žƒ†‘ǡ‡†‘†‘ ǡ‘˜‘Ž‡–ƒǡƒŽ•Šǡ‹‡–‘ȋʹͲͳͲȌ—šͳ ƒ† —šʹ ‡‰—Žƒ–‡

ϲϭϳ ‡†”‹–‹  ”ƒ Š‹‰ǡ ’‹‡ ‘”’Š‘Ž‘‰›ǡ ƒ† ›ƒ’•‡• ‘ˆ –Š‡ ’’‡” ƒ›‡” ‡—”‘• ‘ˆ –Š‡

ϲϭϴ ‘”–‡šǤ‡—”‘͸͸ǣͷʹ͵Ȃͷ͵ͷǤ

ϲϭϵ ƒ˜‹•‘ ǡ ŠŽ‡”•  ȋʹͲͳͳȌ ‡—”ƒŽ ‹” —‹– ‡ Šƒ‹•• ˆ‘” ’ƒ––‡” †‡–‡ –‹‘ ƒ† ˆ‡ƒ–—”‡

ϲϮϬ ‘„‹ƒ–‹‘‹‘Žˆƒ –‘”› ‘”–‡šǤ‡—”‘͹ͲǣͺʹȂͻͶǤ

ϲϮϭ †‡ƒ”Ž‘• ǡ‘’‡œǦƒ• ƒ”ƒ“—‡ǡƒŽ˜‡”†‡ ȋͳͻͻ͸Ȍƒ”Ž›‘Žˆƒ –‘”›ˆ‹„‡”’”‘Œ‡ –‹‘•ƒ† ‡ŽŽ

ϲϮϮ ‹‰”ƒ–‹‘‹–‘–Š‡”ƒ––‡Ž‡ ‡’ŠƒŽ‘Ǥ –‡”ƒ–‹‘ƒŽ ‘—”ƒŽ‘ˆ‡˜‡Ž‘’‡–ƒŽ‡—”‘• ‹‡ ‡

ϲϮϯ ͳͶǣͺͷ͵Ȃͺ͸͸Ǥ

ϲϮϰ †‡ƒ”Ž‘• ǡ‡ƒ”›ȋʹͲͲ͵Ȍ ”‘™–Šƒ†ƒ”‰‡–‹‰‘ˆ—„’Žƒ–‡š‘•ƒ†•–ƒ„Ž‹•Š‡–‘ˆ

ϲϮϱ ƒŒ‘”‘”–‹ ƒŽƒ–Š™ƒ›•Ǥ ‡—”‘• ‹ǣͳȂͳͺǤ

ϲϮϲ †‡ ”—–‘•ǡ‘—˜‹‡” ǡ”ƒ‹ǡŠ‹‘ǡ‘ƒ‡ǡ‡‹–ƒǡ ƒ” ‹ƒǦ‘‹‰—‡œǡŠƒ”ƒ›ǡ

ϲϮϳ ‹”ƒ–ƒǡ‹‡–Šƒ Š‡”ǡ ”‘˜‡ǡ‹••‹” ǡƒ•ƒ†‘ǡ‹‡”ƒ‹ǡ ƒ”‡ŽȋʹͲͳ͸Ȍ‡ƒŽŽ‘ ƒ–‹‘

ϲϮϴ ‘ˆŽˆƒ –‘”›ƒŒƒŽǦ‡–œ‹—•‡ŽŽ•Šƒ’‡•‡‘ ‘”–‡š” Š‹–‡ –—”‡Ǥ‡—”‘ͻʹǣͶ͵ͷȂͶͶͺǤ

ϲϮϵ †‡Ž¿‘ ǡ ‡‹”‹ Šǡ—’°” ǡ‘””‡ŽŽǡ ”‘–• Š‡”ǡ‘”‹ƒ‘ȋͳͻͻ͸Ȍ‹ˆˆ‡”‡–‹ƒŽ•—”˜‹˜ƒŽ‘ˆ

ϲϯϬ ƒŒƒŽǦ‡–œ‹—• ‡ŽŽ• ‹ ‘”‰ƒ‘–›’‹  —Ž–—”‡• ‘ˆ Š‹’’‘ ƒ’—• ƒ† ‡‘ ‘”–‡šǤ ‘—”ƒŽ ‘ˆ

ϲϯϭ ‡—”‘• ‹‡ ‡ͳ͸ǣ͸ͺͻ͸Ȃ͸ͻͲ͹Ǥ

ϲϯϮ †‡ŽÀ‘ ǡ‘Ž±ǡ‘””‡ŽŽǡƒ”–À‡œǡ‘”‹ƒ‘ȋʹͲͲʹȌ ˜‘Ž˜‡‡–‘ˆƒŒƒŽǦ‡–œ‹—• ‡ŽŽ•‹

ϲϯϯ ”‘„—•–ƒ†Žƒ›‡”Ǧ•’‡ ‹ˆ‹ ”‡‰‡‡”ƒ–‹‘‘ˆ–Š‡‡–‘”Š‹‘ǦŠ‹’’‘ ƒ’ƒŽ ’ƒ–Š™ƒ›•Ǥ —”‘’‡ƒ

ϲϯϰ ‘—”ƒŽ‘ˆ‡—”‘• ‹‡ ‡ͳͷǣͳͺͺͳȂͳͺͻͲǤ

ϲϯϱ ‹‘†ƒ–‘ǡ—‹ƒ”–†‡”‹‘–ǡ‹ǡ‡”‹ƒǡ‡””‹ǡ‘— Š ǡŽƒ–œƒǡ ƒ”‡ŽǡŠ‘‹

Ϯϯ 

ϲϯϲ ǡ Ž‡‹• Šƒ  ȋʹͲͳ͸Ȍ ‘Ž‡ —Žƒ” •‹‰ƒ–—”‡• ‘ˆ ‡—”ƒŽ ‘‡ –‹˜‹–› ‹ –Š‡ ‘Žˆƒ –‘”›

ϲϯϳ ‘”–‡šǤƒ–—”‡‘—‹ ƒ–‹‘•͹ǣͳʹʹ͵ͺǤ

ϲϯϴ ‹š‹–ǡ‹Ž‹•‘ ǡƒ ‹‘ ǡŠƒ‡”ǡ†ƒ‹ǡ‹ǡ‡‹•ǡ—””ƒ• ŠǡŠƒ ǡŽ•‘

ϲϯϵ ǡƒ’Žƒǡ‹‡”ǡ Š——”ƒ•ȋʹͲͳͶȌ‡—”‘‰ͳƒ†‡—”‘‰ʹ‘–”‘Ž™‘ƒ˜‡•

ϲϰϬ ‘ˆ‡—”‘ƒŽ‹ˆˆ‡”‡–‹ƒ–‹‘‹–Š‡‹”‹ˆ‘”‘”–‡šǤ ‘—”ƒŽ‘ˆ‡—”‘• ‹‡ ‡͵Ͷǣͷ͵ͻȂͷͷ͵Ǥ

ϲϰϭ ‘—ǡ‹ǡƒ‹ȋͳͻͻͻȌ—ƒŽ‘Ž‡‘ˆ”ƒ‹ ƒ –‘”Ǧͳ‹‡‰—Žƒ–‹‰ ”‘™–Šƒ†ƒ––‡”‹‰‘ˆ–Š‡

ϲϰϮ ‡”‡„”ƒŽ ‡‹•’Š‡”‡•Ǥ‡”‡„”ƒŽ‘”–‡šͻǣͷͶ͵ȂͷͷͲǤ

ϲϰϯ •–”ƒ† ǡ‘”‘‡•‡ǡ ‘Š•‘ ǡ ‡‹‰ǡ‘†‡Žǡ‡Šƒǡ ƒ„‡”Ž›ȋʹͲͲͳȌ‡™

ϲϰϰ •—„†‹˜‹•‹‘‘ˆƒ–‡”‹‘”’‹”‹ˆ‘” ‘”–‡šƒ†ƒ••‘ ‹ƒ–‡††‡‡’— Ž‡—•™‹–Š‘˜‡Žˆ‡ƒ–—”‡•‘ˆ

ϲϰϱ ‹–‡”‡•–ˆ‘”‘Žˆƒ –‹‘ƒ†‡’‹Ž‡’•›Ǥ ‘’‡—”‘ŽͶ͵ͶǣʹͺͻȂ͵Ͳ͹Ǥ

ϲϰϲ ‘—”‹‡” ǡòŽŽ‡”ǡƒ—”‡– ȋʹͲͳͷȌ‘‘‹‰ˆ‘”–Š‡”‘‘–•‘ˆ ‘”–‹ ƒŽ•‡•‘”› ‘’—–ƒ–‹‘‹

ϲϰϳ –Š”‡‡ǦŽƒ›‡”‡† ‘”–‹ ‡•Ǥ—””‡–’‹‹‘‹‡—”‘„‹‘Ž‘‰›͵ͳǣͳͳͻȂͳʹ͸Ǥ

ϲϰϴ ”ƒ• ǡ •ƒƒ •‘  ȋʹͲͲͷȌ ›ƒ’•‡Ǧ•’‡ ‹ˆ‹  †‘™”‡‰—Žƒ–‹‘ ‘ˆ  ”‡ ‡’–‘”• „› ‡ƒ”Ž›

ϲϰϵ ‡š’‡”‹‡ ‡ǣ ƒ ”‹–‹ ƒŽ ’‡”‹‘† ˆ‘” ’Žƒ•–‹ ‹–› ‘ˆ •‡•‘”› ‹’—– –‘ ‘Žˆƒ –‘”› ‘”–‡šǤ ‡—”‘

ϲϱϬ Ͷ͹ǣͳͲͳȂͳͳͶǤ

ϲϱϭ ”ƒ•ǡ—••‘ ǡ‘•—Ž•‹ǡ—ŽŽ‹‰ƒǡ‹‡‰‡Ž„ƒ—ǡš‡Ž  ȋʹͲͳͳȌ ‡ —””‡–

ϲϱϮ ‹” —‹–”››ƒ‹ ƒŽŽ›Šƒ’‡•–Š‡ –‹˜ƒ–‹‘‘ˆ‹”‹ˆ‘”‘”–‡šǤ‡—”‘͹ʹǣͶͻȂͷ͸Ǥ

ϲϱϯ ƒ” ‹ƒǦ‘”‡‘ ǡ‘’‡œǦƒ• ƒ”ƒ“—‡ǡ†‡ƒ”Ž‘• ȋʹͲͲͺȌƒ”Ž›‡Ž‡ ‡’ŠƒŽ‹  ‹‰”ƒ–‹‘

ϲϱϰ ‘’‘‰”ƒ’Š‹ ƒŽŽ›‘˜‡”‰‹‰‹–Š‡Žˆƒ –‘”›‘”–‡šǤ‡”‡„”ƒŽ‘”–‡šͳͺǣͳʹ͵ͻȂͳʹͷʹǤ

ϲϱϱ ‡Žƒǡ ”‹˜‡ƒ—ǡ‡Š‘”–‡”ǡ‡‹••‹‡”ǡƒ”‡ŽƒǡŽƒǡ‹‡”ƒ‹ǡƒ”‹ȋʹͲͳͳȌ‹†‡

ϲϱϲ ‹˜‡”•‹–› ‘ˆ ‘”–‹ ƒŽ ‡”‰‹  –‡”‡—”‘• ‡”‹˜‡• ˆ”‘ –Š‡ „”›‘‹  ”‡‘’–‹  ”‡ƒǤ

ϲϱϳ ‘—”ƒŽ‘ˆ‡—”‘• ‹‡ ‡͵ͳǣͳ͸ͷ͹ͲȂͳ͸ͷͺͲǤ

ϲϱϴ Šƒ‡ǡ—ǡ‘‰ ǡ ƒ– Š ǡ—„‡•–‡‹ ǡ‡”ȋʹͲͲ͹Ȍ ‹•–‹ – ‹•Ǧ‡‰—Žƒ–‘”›

Ϯϰ 

ϲϱϵ Ž‡‡–• ˆ”‘ –Š‡ ŽšͳȀŽšʹ ‘ —• ƒ” ‹ˆˆ‡”‡– ”‘‰‡‹–‘” ‡ŽŽ ‘’—Žƒ–‹‘• ‹ –Š‡

ϲϲϬ ƒ‰Ž‹‘‹  ‹‡ ‡• ƒ† ‹ˆˆ‡”‡– —„–›’‡• ‘ˆ †—Ž– ‘”–‹ ƒŽ –‡”‡—”‘•Ǥ ‘—”ƒŽ ‘ˆ

ϲϲϭ ‡—”‘• ‹‡ ‡ʹ͹ǣͷͲͳʹȂͷͲʹʹǤ

ϲϲϮ Š‘•Šǡƒ”•‘ǡ ‡ˆœ‹ ǡƒ”‘›ǡ—–ˆ‘”–Šǡ‘ƒǡƒŽ†™‹ȋʹͲͳͳȌ‡•‘”›ƒ’•‹

ϲϲϯ –Š‡‘Žˆƒ –‘”› ‘”–‡š†‡ˆ‹‡†„›Ž‘‰Ǧ”ƒ‰‡˜‹”ƒŽ–”ƒ ‹‰‘ˆ•‹‰Ž‡‡—”‘•Ǥƒ–—”‡Ͷ͹ʹǣʹͳ͹Ȃ

ϲϲϰ ʹʹͲǤ

ϲϲϱ ‘”•‹ ǡƒŽŽ‡›ǡ‡‰•Š‡‰ǡ—‡ŽŽ‡•ǡǡ ‘‡•ȋʹͲͲʹȌ‘”–‹ ƒŽš ‹–ƒ–‘”›‡—”‘•

ϲϲϲ ƒ† Ž‹ƒǡ —– ‘– ‡”‰‹  ‡—”‘•ǡ ”‡ ”‘†— ‡† ‹ –Š‡ šͳǦš’”‡••‹‰ ‹‡ƒ‰‡Ǥ 

ϲϲϳ ‡—”‘• ‹ǣͳȂ͸Ǥ

ϲϲϴ ƒ„‡”Ž›  ȋͳͻͺ͵Ȍ –”— –—”‡ ‘ˆ –Š‡ ’‹”‹ˆ‘” ‘”–‡š ‘ˆ –Š‡ ‘’‘••—Ǥ Ǥ ‡• ”‹’–‹‘ ‘ˆ ‡—”‘

ϲϲϵ –›’‡•™‹–Š‰‘Ž‰‹‡–Š‘†•Ǥ ‘’‡—”‘Žʹͳ͵ǣͳ͸͵Ȃͳͺ͹Ǥ

ϲϳϬ ƒ„‡”Ž›ȋͳͻͻͲȌ‘’ƒ”ƒ–‹˜‡ƒ•’‡ –•‘ˆ‘Žˆƒ –‘”› ‘”–‡šǤ ǣ‡”‡„”ƒŽ‘”–‡šȋ ‘‡• ǡ‡–‡”•

ϲϳϭ ǡ‡†•ȌǤ‘•–‘ǡǣ’”‹‰‡”Ǥ

ϲϳϮ ƒ„‡”Ž›  ȋʹͲͲͳȌ ƒ”ƒŽŽ‡ŽǦ†‹•–”‹„—–‡† ”‘ ‡••‹‰ ‹ Žˆƒ –‘”› ‘”–‡šǣ ‡™ •‹‰Š–• ˆ”‘

ϲϳϯ ‘”’Š‘Ž‘‰‹ ƒŽ ƒ† Š›•‹‘Ž‘‰‹ ƒŽ ƒŽ›•‹• ‘ˆ ‡—”‘ƒŽ ‹” —‹–”›Ǥ Š‡‹ ƒŽ ‡•‡• ʹ͸ǣͷͷͳȂ

ϲϳϰ ͷ͹͸Ǥ

ϲϳϱ ƒ„‡”Ž›ǡ ‡‹‰ȋͳͻͺ͵Ȍ–”— –—”‡‘ˆ–Š‡’‹”‹ˆ‘” ‘”–‡š‘ˆ–Š‡‘’‘••—Ǥ Ǥ ‹‡•–”— –—”‡‘ˆ

ϲϳϲ ‡ŽŽ„‘†‹‡•ƒ†‡—”‘’‹ŽǤ ‘’‡—”‘Žʹͳ͸ǣ͸ͻȂͺͺǤ

ϲϳϳ ƒ„‡”Ž›ǡ”‹ ‡ ȋͳͻ͹ͺȌ••‘ ‹ƒ–‹‘ƒ† ‘‹••—”ƒŽˆ‹„‡”•›•–‡•‘ˆ–Š‡‘Žˆƒ –‘”› ‘”–‡š‘ˆ

ϲϳϴ –Š‡”ƒ–Ǥ ‘’‡—”‘Žͳ͹ͺǣ͹ͳͳȂ͹ͶͲǤ

ϲϳϵ ƒ‰‹™ƒ”ƒǡƒŽǡƒ–‘ ǡ‹‡‹• Šǡ—”–Š›ȋʹͲͳʹȌ’–‘’Š›•‹‘Ž‘‰‹ ƒŽ ƒƒŽ›•‹• ‘ˆ

ϲϴϬ ƒ••‘ ‹ƒ–‹‘ƒŽ ‹” —‹–•‹–Š‡‘Žˆƒ –‘”› ‘”–‡šǤ ”‘–‡—”ƒŽ‹” —‹–•͸ǣͳȂͳͻǤ

ϲϴϭ ƒƒ•Š‹ƒǡ ‡”ƒ†‡•ǡ ‡„‡”– ǡ ‹•Š‡ŽŽ ȋʹͲͲ͹ȌŠ‡‘Ž‡‘ˆ ‘š‰ͳƒ†‘”•ƒŽ‹†Ž‹‡

Ϯϱ 

ϲϴϮ ‹‰ƒŽ‹‰‹–Š‡ ‡‡”ƒ–‹‘‘ˆƒŒƒŽǦ‡–œ‹—•—„–›’‡•Ǥ ‘—”ƒŽ‘ˆ ‡—”‘• ‹‡ ‡ ʹ͹ǣͳͳͳͲ͵Ȃ

ϲϴϯ ͳͳͳͳͳǤ

ϲϴϰ ‡˜‡”  ǡ ‡‘‰‹ ǡ ‰Ž—† ǡ ƒœƒ ǡ ‹  ȋʹͲͲ͵Ȍ ƒŒƒŽȂ‡–œ‹—• ‡ŽŽ• ‹ –Š‡ ‘—•‡ǣ

ϲϴϱ –”ƒ• ”‹’–‹‘ ˆƒ –‘”•ǡ ‡—”‘–”ƒ•‹––‡”•ǡ ƒ† „‹”–Š†ƒ›• •—‰‰‡•–ƒ’ƒŽŽ‹ƒŽ‘”‹‰‹Ǥ

ϲϴϲ ‡˜‡Ž‘’‡–ƒŽ”ƒ‹‡•‡ƒ” ŠͳͶͳǣ͵ͻȂͷ͵Ǥ

ϲϴϳ ‡˜‡” ǡŠ‹ǡ —•–‹ ‡ǡ •—‡ŠǡŠ‡‰ǡ‹‰ƒǡ—Žˆ‘‡ǡ ‘ˆˆ‹‡–ǡƒ’ƒ‰‘‹ǡ

ϲϴϴ —„‡•–‡‹ ȋʹͲͲͳȌ„”ͳ”‡‰—Žƒ–‡•†‹ˆˆ‡”‡–‹ƒ–‹‘‘ˆ–Š‡’”‡’Žƒ–‡ƒ†Žƒ›‡”͸Ǥ

ϲϴϵ ‡—”‘ǣ͵ͷ͵Ȃ͵͸͸Ǥ

ϲϵϬ ‹”ƒ–ƒǡ —Œ‹•ƒ™ƒ ȋͳͻͻͻȌ˜‹”‘‡–ƒŽ ‘–”‘Ž‘ˆ ‘ŽŽƒ–‡”ƒŽ„”ƒ Š‹‰ƒ†–ƒ”‰‡–‹˜ƒ•‹‘‘ˆ

ϲϵϭ ‹–”ƒŽ ‡ŽŽƒš‘•†—”‹‰†‡˜‡Ž‘’‡–Ǥ ‡—”‘„‹‘Ž͵ͺǣͻ͵ȂͳͲͶǤ

ϲϵϮ ‹”ƒ–ƒǡ—ƒ†ƒǡƒ™ƒ•ƒ‹ǡ —”—ƒ™ƒǡ‹„ƒǡ‘“—‡– ǡƒ‰ƒ ǡ ——†ƒ  ȋʹͲͳʹȌ

ϲϵϯ —‹†‡’‘•– ‡—”‘• ˆ‘” –Š‡ Žƒ–‡”ƒŽ ‘Žˆƒ –‘”› –”ƒ –ǣ š’”‡••‹‘ ‘ˆ ‡–ƒ„‘–”‘’‹  ‰Ž—–ƒƒ–‡

ϲϵϰ ”‡ ‡’–‘”ͳƒ†‹‡”˜ƒ–‹‘„›‰Ž—–ƒƒ–‡”‰‹ ‘Žˆƒ –‘”›„—Ž„ƒš‘•Ǥ‡˜‡Ž‡—”‘„‹‘͹ʹǣͳͷͷͻȂ

ϲϵϱ ͳͷ͹͸Ǥ

ϲϵϲ ‹”ƒ–ƒ ǡ ‘—”ƒ ǡ ƒƒ‰‹ ǡ ƒ–‘ ǡ ‘‹‘ƒ ǡ —Œ‹•ƒ™ƒ ǡ •—‹  ȋʹͲͲʹȌ ‘•ƒ‹ 

ϲϵϳ †‡˜‡Ž‘’‡–‘ˆ–Š‡‘Žˆƒ –‘”› ‘”–‡š™‹–Šƒš͸Ǧ†‡’‡†‡–ƒ†Ǧ‹†‡’‡†‡– ‘’‘‡–•Ǥ

ϲϵϴ ‡˜‡Ž‘’‡–ƒŽ”ƒ‹‡•‡ƒ” ŠǣͳȂͳͲǤ

ϲϵϵ ‘™‡ŽŽǡ ƒ™‡•ǡ‘”‹ƒ‘ǡ‘‘’‡” ȋͳͻͻ͹Ȍ‡—”‘ƒŽ’‘•‹–‹‘‹–Š‡†‡˜‡Ž‘’‹‰„”ƒ‹‹•

ϳϬϬ ”‡‰—Žƒ–‡†„›‘—•‡†‹•ƒ„Ž‡†ǦͳǤƒ–—”‡ǣͳȂͷǤ

ϳϬϭ •ƒƒ •‘  ȋʹͲͳͲȌ †‘” ”‡’”‡•‡–ƒ–‹‘• ‹ ƒƒŽ‹ƒ ‘”–‹ ƒŽ ‹” —‹–•Ǥ —””‡– ’‹‹‘ ‹

ϳϬϮ ‡—”‘„‹‘Ž‘‰›ʹͲǣ͵ʹͺȂ͵͵ͳǤ

ϳϬϯ –‘ ǡ ƒ™ƒ•ƒ‹ ǡ ƒƒ•Š‹ƒ ǡ ƒ–•—†ƒ ǡ ‹„ƒ ǡ ‹”ƒ–ƒ  ȋʹͲͲͺȌ ‡ƒ’Š‘”‹ ͵  ‘ˆ‹‡•

ϳϬϰ ‡–”ƒŽ ƒ‰‡–‹ƒŽ ‹‰”ƒ–‹‘ ‘ˆ ƒ–‡”ƒŽ Žˆƒ –‘”› ”ƒ – ‡—”‘• ‘–‘ –Š‡ ‡Ž‡ ‡’ŠƒŽ‘

Ϯϲ 

ϳϬϱ —”ˆƒ ‡Ǥ ‘—”ƒŽ‘ˆ‡—”‘• ‹‡ ‡ʹͺǣͶͶͳͶȂͶͶʹʹǤ

ϳϬϲ ƒ„ƒ—†‘ǡŠ‹†‡” ǡ‹• Šˆ‹‡Ž† ǡ ƒŽƒœ‘ ǡƒ Ž‹• ȋʹͲͳʹȌ Ⱦ ‹†— ‡• „ƒ””‡ŽǦŽ‹‡

ϳϬϳ ‡—”‘ƒŽ Ž—•–‡”•‹–Š‡†‡˜‡Ž‘’‹‰‡‘ ‘”–‡šǤ‡”‡„‘”–‡šʹʹǣͻͻ͸ȂͳͲͲ͸Ǥ

ϳϬϴ ‹±‡œ ǡ ג‡œǦƒ• ƒ”ƒ“—‡ ǡ ƒŽ˜‡”†‡ ǡ ‡ ƒ”Ž‘•  ȋʹͲͲʹȌ ƒ‰‡–‹ƒŽ ‹‰”ƒ–‹‘ ‹

ϳϬϵ ‡‘ ‘”–‹ ƒŽ‡˜‡Ž‘’‡–Ǥ‡˜‡Ž‘’‡–ƒŽ‹‘Ž‘‰›ʹͶͶǣͳͷͷȂͳ͸ͻǤ

ϳϭϬ ‘Š•‘ ǡ ŽŽ‹‰ ǡ ‡Šƒ ǡ ƒ„‡”Ž›  ȋʹͲͲͲȌ ‡™ ˆ‡ƒ–—”‡• ‘ˆ ‘‡ –‹˜‹–› ‹ ’‹”‹ˆ‘”

ϳϭϭ ‘”–‡š ˜‹•—ƒŽ‹œ‡† „› ‹–”ƒ ‡ŽŽ—Žƒ” ‹Œ‡ –‹‘ ‘ˆ ’›”ƒ‹†ƒŽ ‡ŽŽ••—‰‰‡•––Šƒ–̶’”‹ƒ”›̶

ϳϭϮ ‘Žˆƒ –‘”› ‘”–‡š ˆ— –‹‘• Ž‹‡ Dzƒ••‘ ‹ƒ–‹‘dz ‘”–‡š ‹ ‘–Š‡” •‡•‘”› •›•–‡•Ǥ ‘—”ƒŽ ‘ˆ

ϳϭϯ ‡—”‘• ‹‡ ‡ʹͲǣ͸ͻ͹ͶȂ͸ͻͺʹǤ

ϳϭϰ ƒ™ƒ•ƒ‹ȋʹͲͲ͸Ȍ‡–”‹ͳ”‡‰—Žƒ–‡•˜‡–”ƒŽ–ƒ‰‡–‹ƒŽ‹‰”ƒ–‹‘‘ˆ‰—‹†‡’‘•–‡—”‘•‹–Š‡

ϳϭϱ Žƒ–‡”ƒŽ‘Žˆƒ –‘”›–”ƒ –Ǥ‡˜‡Ž‘’‡–ͳ͵͵ǣͺͶͷȂͺͷ͵Ǥ

ϳϭϲ ‡””ǡ‰•–‡”ǡ —”–ƒǡ—”™‡ŽŽȋʹͲͲ͹Ȍ — –‹‘ƒŽ‡—”‘ƒƒ–‘› ‘ˆ –Š‡

ϳϭϳ ’ƒ”ƒŠ‹’’‘ ƒ’ƒŽ”‡‰‹‘ǣŠ‡Žƒ–‡”ƒŽƒ†‡†‹ƒŽ‡–‘”Š‹ƒŽƒ”‡ƒ•Ǥ ‹’’‘ ƒ’—• ͳ͹ǣ͸ͻ͹Ȃ

ϳϭϴ ͹ͲͺǤ

ϳϭϵ ‹ ǡ †‡”•‘ ǡ —„‡•–‡‹ ǡ ‘™‡•–‡‹  ǡ Ž‡ƒ•—”‡   ȋʹͲͲͳȌ ƒšǦ͸ ‡‰—Žƒ–‡•

ϳϮϬ š’”‡••‹‘‘ˆ Ǧʹƒ†–Ǧ͹„‹–Š‡†‡˜‡Ž‘’‹‰Ǥ ‡—”‘• ‹ǣͳȂͷǤ

ϳϮϭ Ž‡’‹ ǡ ”‘‡„‡”‰ ǡ Š‡—‰ ǡ ‡––‡ƒ ǡ ‡’‡”ƒ  ȋʹͲͳͳȌ ”‘’‡”–‹‡• ‘ˆ

ϳϮϮ †‘—„Ž‡ ‘”–‹ǦȋȌǦ‡š’”‡••‹‰ ‡ŽŽ• ‹ –Š‡ ’‹”‹ˆ‘” ‘”–‡š ‘’ƒ”‡† –‘ –Š‡ ‡—”‘‰‡‹ 

ϳϮϯ †‡–ƒ–‡‰›”—•‘ˆƒ†—Ž–‹ ‡Ǥ‘͸ǣ‡ʹͷ͹͸ͲǤ

ϳϮϰ Ž‹‰Ž‡”ǡƒ”–‹ǡ ƒ” ‹ƒǡ‘”‡ƒ—Ǧ ƒ—˜ƒ”“—‡ǡ ƒŽ ǡŠƒ”‡›”‡ ǡ ‹‘˜ƒ‹‹ǡŠ‡†‘–ƒŽ

ϳϮϱ ǡ ‹”ƒ—Ž– ǡ ‘—–‡„”‘œ‡ȋʹͲͳͷȌŠ‡ ›–‘•‡Ž‡–‘Ǧƒ••‘ ‹ƒ–‡†’”‘–‡‹ ͳ‹•‹˜‘Ž˜‡†

ϳϮϲ ‹ƒš‘‰—‹†ƒ ‡ǡƒ†‹•”‡“—‹”‡†ˆ‘”’‹”‹ˆ‘” ‘”–‡šƒ†ƒ–‡”‹‘” ‘‹••—”‡†‡˜‡Ž‘’‡–Ǥ

ϳϮϳ ‡˜‡Ž‘’‡–ͳͶʹǣʹͲʹ͸ȂʹͲ͵͸Ǥ

Ϯϳ 

ϳϮϴ Žƒ‘••ƒ‡ǡ‡ŽŽ‘‡ǡ ‘Ž†‹‰ǡ‹–ƒŽ‹ ǡ‘•• ǡ‘‹ǡò• Š‡”ǡ ƒ„ƒ—†‘ȋʹͲͳ͵Ȍ ˜‹˜‘

ϳϮϵ ”‡’”‘‰”ƒ‹‰ ‘ˆ ‹” —‹– ‘‡ –‹˜‹–› ‹ ’‘•–‹–‘–‹  ‡‘ ‘”–‹ ƒŽ ‡—”‘•Ǥ ƒ– ‡—”‘• ‹

ϳϯϬ ͳ͸ǣͳͻ͵ȂʹͲͲǤ

ϳϯϭ ƒ”‰‡ǡ‘‰Ž‡”ǡ‹‡Ž‘ǡ•™ƒŽ†ǦȋʹͲͳ͸ȌƒŽƒ ‡†ˆ‡‡†ˆ‘”™ƒ”† ‹Š‹„‹–‹‘ ƒ†

ϳϯϮ †‘‹ƒ–”‡ —””‡–‹Š‹„‹–‹‘‹‘Žˆƒ –‘”› ‘”–‡šǤ”‘ ‡‡†‹‰•‘ˆ–Š‡ƒ–‹‘ƒŽ ƒ†‡›‘ˆ

ϳϯϯ  ‹‡ ‡•ͳͳ͵ǣʹʹ͹͸ȂʹʹͺͳǤ

ϳϯϰ ‡‹™ƒ†  ǡ ŠƒŽƒ•ƒ‹   ȋʹͲͳͳȌ Žˆƒ –‘”› ‡–™‘”•ǣ ˆ”‘ •‡•ƒ–‹‘ –‘ ’‡” ‡’–‹‘Ǥ —””‡–

ϳϯϱ ’‹‹‘‹ ‡‡–‹ •Ƭ‡˜‡Ž‘’‡–ʹͳǣͺͲ͸ȂͺͳͳǤ

ϳϯϲ ‡‘‡ ǡ ‡ƒ˜‡” ǡ ‡”‡ œ‹ ǡ ‘„”‡˜ƒ ǡ —‰—‡ƒ”† ǡ ”‘•• Š‡†Ž ǡ  ‘‡ŽŽ 

ϳϯϳ ȋʹͲͳͷȌƒ–„ʹ‡‰—Žƒ–‡•–Š‡‹ˆˆ‡”‡–‹ƒ–‹‘‘ˆ‘–ŠƒŽŽ‘•ƒŽƒ† —„ ‡”‡„”ƒŽ ”‘Œ‡ –‹‘

ϳϯϴ ‡—”‘•‹–Š‡‡˜‡Ž‘’‹‰‡”‡„”ƒŽ‘”–‡šǤ‡”‡„”ƒŽ‘”–‡šʹͷǣ͵ͶͲ͸Ȃ͵ͶͳͻǤ

ϳϯϵ —œœƒ–‹ ȋʹͲͳͷȌŠ›’‘–Š‡•‹•ˆ‘”–Š‡‡˜‘Ž—–‹‘‘ˆ–Š‡—’’‡”Žƒ›‡”•‘ˆ–Š‡‡‘ ‘”–‡š–Š”‘—‰Š ‘Ǧ

ϳϰϬ ‘’–‹‘‘ˆ–Š‡‘Žˆƒ –‘”› ‘”–‡š†‡˜‡Ž‘’‡–ƒŽ’”‘‰”ƒǤ ”‘–‡—”‘• ‹ͻǣͶʹͻȂͳʹǤ

ϳϰϭ —œœƒ–‹ ǡ‘ˆƒ–‹ǡ ƒ•‘Ž‘ǡ‡”‡––‘ȋʹͲͲͻȌƒ†Ǧ š’”‡••‹‘ †‡–‹ˆ‹‡• ƒ

ϳϰϮ ‘’—Žƒ–‹‘ ‘ˆ ‡—”‘• ”‡ˆ‡”‡–‹ƒŽŽ› ‹•–”‹„—–‡† ‹ ••‘ ‹ƒ–‹˜‡”‡ƒ•‘ˆ‹ˆˆ‡”‡–ƒŽŽ‹ƒŽ

ϳϰϯ ‡”‹˜ƒ–‹˜‡•ƒ†‡”–‡„”ƒ–‡’‡ ‹‡•Ǥ‡”‡„”ƒŽ‘”–‡šͳͻǣͳͲʹͺȂͳͲͶͳǤ

ϳϰϰ ƒŽŽƒƒ ‹ǡ–‘›‘˜ƒȋʹͲͲ͸Ȍ ‡‡‡–™‘”• ‘–”‘ŽŽ‹‰‡ƒ”Ž› ‡”‡„”ƒŽ ‘”–‡š ƒ”‡ƒŽ‹œƒ–‹‘Ǥ

ϳϰϱ —”‘’‡ƒ ‘—”ƒŽ‘ˆ‡—”‘• ‹‡ ‡ʹ͵ǣͺͶ͹Ȃͺͷ͸Ǥ

ϳϰϲ ƒŽ‹ ǡ ‘œƒŽ‡œǦ”‹•–‡ŽŽ‡”ǡ —–‹›ƒƒȋʹͲͳͲȌ†‘”ƒ–‡ ‡’–‘”•Ǥ ǣŠ‡‡—”‘„‹‘Ž‘‰›

ϳϰϳ ‘ˆŽˆƒ –‹‘Ǥ

ϳϰϴ ƒŽ‹ ǡ ‹”‘‘ ǡƒ–‘ǡ— ȋͳͻͻͻȌ‘„‹ƒ–‘”‹ƒŽ”‡ ‡’–‘” ‘†‡•ˆ‘”‘†‘”•Ǥ‡ŽŽͻ͸ǣ͹ͳ͵Ȃ

ϳϰϵ ͹ʹ͵Ǥ

ϳϱϬ ƒ‰ƒŽ‡ǡ ‹”‘ƒ™ƒǡƒ–›ƒ‹ǡ ‘—Ž Šƒ†”ƒǡŠ‹„‹”‡ǡ—„”ƒƒ‹ƒǡŠ‡––›ǡ

Ϯϴ 

ϳϱϭ ƒ”–›‘‰ƒǡƒ—Ž ǡƒ‹ǡ‹ǡ Žƒƒ‰ƒǡ‘Ž‡ǡ‘—‹ ȋʹͲͲͺȌ Ššʹ •‡Ž‡ –‘”

ϳϱϮ ƒ –‹˜‹–› •’‡ ‹ˆ‹‡• ‘”–‹ ƒŽ ‹†‡–‹–› ƒ† •—’’”‡••‡• Š‹’’‘ ƒ’ƒŽ‘”‰ƒ‹œ‡”ˆƒ–‡Ǥ ‹‡ ‡

ϳϱϯ ͵ͳͻǣ͵ͲͶȂ͵ͲͻǤ

ϳϱϰ ƒ”‹ ǡ —„‡•–‡‹  ȋʹͲͲͳȌ  Ž‘‰ǡ ”‡ƒ”ƒ„Ž‡ Œ‘—”‡›ǣ –ƒ‰‡–‹ƒŽ ‹‰”ƒ–‹‘ ‹ –Š‡

ϳϱϱ –‡Ž‡ ‡’ŠƒŽ‘Ǥƒ–‡˜‡—”‘• ‹ʹǣ͹ͺͲȂ͹ͻͲǤ

ϳϱϲ ƒ”‹Ǧƒ†‹ŽŽƒȋͳͻ͹ͺȌ—ƒŽ‘”‹‰‹‘ˆ–Š‡ƒƒŽ‹ƒ‡‘ ‘”–‡šƒ†‡˜‘Ž—–‹‘‘ˆ–Š‡ ‘”–‹ ƒŽ

ϳϱϳ ’Žƒ–‡Ǥƒ–„”›‘ŽͳͷʹǣͳͲͻȂͳʹ͸Ǥ

ϳϱϴ ƒ”Àǡ—„‡•–‡‹ ȋʹͲͲ͵Ȍ‡ŽŽ‹‰”ƒ–‹‘‹–Š‡ˆ‘”‡„”ƒ‹Ǥ—‡˜‡—”‘• ‹ʹ͸ǣͶͶͳȂͶͺ͵Ǥ

ϳϱϵ ƒ”ÀǦƒ†‹ŽŽƒ  ȋͳͻͻͺȌ ƒŒƒŽȂ‡–œ‹—• ‡ŽŽ• ƒ† –Š‡ †‡˜‡Ž‘’‡– ‘ˆ –Š‡ ‡‘ ‘”–‡šǤ ”‡†• ‹

ϳϲϬ ‡—”‘• ‹‡ ‡•ʹͳǣ͸ͶȂ͹ͳǤ

ϳϲϭ ‡†‹ƒǡ‡‰ƒœ ǡ ‘œžŽ‡œ ǡ‡ƒ•–”‘ ǡ—„‡•–‡‹ ǡ—‡ŽŽ‡•ȋʹͲͲͶȌš’”‡••‹‘‘ˆ„šͳǡ

ϳϲϮ ‡—”‘‰‡‹ʹǡ‡ƒ’Š‘”‹ͷǡƒ†Š‡”‹ͺǡƒ†šͳ†‹•–‹‰—‹•Š˜‡–”ƒŽƒ†Žƒ–‡”ƒŽ’ƒŽŽ‹ƒŽ

ϳϲϯ Š‹•–‘‰‡‡–‹ †‹˜‹•‹‘•‹–Š‡†‡˜‡Ž‘’‹‰‘—•‡ Žƒ—•–”‘ƒ›‰†ƒŽ‘‹† ‘’Ž‡šǤ ‘’‡—”‘Ž

ϳϲϰ Ͷ͹ͶǣͷͲͶȂͷʹ͵Ǥ

ϳϲϱ ‡›‡” ǡ ŽŽ‹‰ ǡ ”—Œ‡•  ȋʹͲͲ͸Ȍ ‹ˆˆ‡”‡ ‡• ‹ Š‡‘Ǧ ƒ† ›–‘ƒ” Š‹–‡ –—”ƒŽ ˆ‡ƒ–—”‡•

ϳϲϲ ™‹–Š‹’ƒ”•’”‹ ‹’ƒŽ‹•‘ˆ–Š‡”ƒ–ƒ–‡”‹‘”‘Žˆƒ –‘”›— Ž‡—••—‰‰‡•–ˆ— –‹‘ƒŽ•’‡ ‹ƒŽ‹œƒ–‹‘Ǥ

ϳϲϳ ‘’‡—”‘ŽͶͻͺǣ͹ͺ͸Ȃ͹ͻͷǤ

ϳϲϴ ±–‹ǡƒ—†‘‹ Ǧǡƒ‹©ǡƒ”ƒ˜‡Žƒ• ȋʹͲͲ͸Ȍ‡ŽŽƒ†‘Ž‡ —Žƒ”‡ Šƒ‹••‹˜‘Ž˜‡†‹

ϳϲϵ –Š‡‹‰”ƒ–‹‘‘ˆ ‘”–‹ ƒŽ‹–‡”‡—”‘•Ǥ—”‘’‡ƒ ‘—”ƒŽ‘ˆ‡—”‘• ‹‡ ‡ʹ͵ǣͺͻͶȂͻͲͲǤ

ϳϳϬ ‹›ƒ‹ Š‹ǡƒ– ǡ‘—••ƒ˜‹ ǡƒ‰ǡ‹ ‡”•Šƒ ǡƒŽŽǡƒ‹‰— Š‹ ǡƒ•‹ ǡ —ƒ‰

ϳϳϭ  ǡ ‡ǡƒŽŽƒ™ƒ›ǡ ‘”‘™‹–œǡ—‘ȋʹͲͳͳȌ‘”–‹ ƒŽ”‡’”‡•‡–ƒ–‹‘•‘ˆ‘Žˆƒ –‘”›‹’—–

ϳϳϮ „›–”ƒ•Ǧ•›ƒ’–‹ –”ƒ ‹‰Ǥƒ–—”‡Ͷ͹ʹǣͳͻͳȂͳͻ͸Ǥ

ϳϳϯ ‹›‘•Š‹ ǡ ‹•Š‡ŽŽ  ȋʹͲͳͳȌ ‡”‰‹  –‡”‡—”‘ ‹‡ƒ‰‡• ‡Ž‡ –‹˜‡Ž› ‘”– ‹–‘ ’‡ ‹ˆ‹ 

Ϯϵ 

ϳϳϰ ‘”–‹ ƒŽƒ›‡”•†—”‹‰ƒ”Ž›‘•–ƒ–ƒŽ‡˜‡Ž‘’‡–Ǥ‡”‡„”ƒŽ‘”–‡šʹͳǣͺͶͷȂͺͷʹǤ

ϳϳϱ ‘Žž” ǡ —–Ž‡”  ȋʹͲͲʹȌ Š‡ ‘”–‹ ‘•–”‹ƒ–ƒŽ Œ— –‹‘ǣ  ”— ‹ƒŽ ”‡‰‹‘ ˆ‘” ˆ‘”‡„”ƒ‹

ϳϳϲ †‡˜‡Ž‘’‡–ƒ†‡˜‘Ž—–‹‘Ǥ‹‘‡••ƒ›•ʹͶǣͷ͵ͲȂͷͶͳǤ

ϳϳϳ ‘Ž›‡ƒ—š ǡ”Ž‘––ƒǡ‡‡œ‡• ǡƒ Ž‹• ȋʹͲͲ͹Ȍ‡—”‘ƒŽ•—„–›’‡•’‡ ‹ˆ‹ ƒ–‹‘‹–Š‡

ϳϳϴ ‡”‡„”ƒŽ ‘”–‡šǤƒ–‡˜‡—”‘• ‹ͺǣͶʹ͹ȂͶ͵͹Ǥ

ϳϳϵ ‘„ƒ‡”–•ǡƒ‰ ǡ—Žƒ ǡŠƒ‘ǡ‡‡•ǡ‡†‡Ž•‘Šǡ†‘†•‘ ǡš‡Žȋͳͻͻ͸Ȍ

ϳϴϬ ‹•—ƒŽ‹œ‹‰ƒŽˆƒ –‘”›‡•‘”›ƒ’Ǥ‡ŽŽͺ͹ǣ͸͹ͷȂ͸ͺ͸Ǥ

ϳϴϭ ‘–‹‡Ž ǡ „‘‹–‹œ  ȋʹͲͳͷȌ ƒŽŽ‹ƒŽ ’ƒ––‡”‹‰ ƒ† –Š‡ ‘”‹‰‹ ‘ˆ–Š‡‹•‘ ‘”–‡šǤ ”‘–‡—”‘• ‹

ϳϴϮ ͻǣͳ͸͸ȂͳͲǤ

ϳϴϯ ‘—‹ǡ‘”–‡” ǡƒŽ•ŠȋʹͲͲͳȌƒ––‡”‹‰‘ˆ–Š‡‘”•ƒŽ‡Ž‡ ‡’ŠƒŽ‘ ƒ† ‡”‡„”ƒŽ

ϳϴϰ ‘”–‡š„›ƒ‘‘ˆŽƒ–‡ǦŠšʹƒ–Š™ƒ›Ǥ‡—”‘͵ʹǣͷͻͳȂ͸ͲͶǤ

ϳϴϱ —œ‹‘ǡƒŽŽƒƒ ‹ȋʹͲͲͷȌ ‘š‰ͳ ‘ˆ‹‡•ƒŒƒŽǦ‡–œ‹—•‡—”‘‘‰‡‡•‹• ƒ† Š‹’’‘ ƒ’ƒŽ

ϳϴϲ ‘”’Š‘‰‡‡•‹•–‘–Š‡†‘”•‘‡†‹ƒŽ’ƒŽŽ‹—Ǥ ‡—”‘• ‹ʹͷǣͶͶ͵ͷȂͶͶͶͳǤ

ϳϴϳ ƒ Š‡” ǡ”‡•’‘ǡ ™‡ȋʹͲͲͳȌ‘—„Ž‡ ‘”–‹‡š’”‡••‹‘‹–Š‡ƒ†—Ž–”ƒ––‡Ž‡ ‡’ŠƒŽ‘Ǥ

ϳϴϴ —” ‡—”‘• ‹ͳͶǣ͸ʹͻȂ͸ͶͶǤ

ϳϴϵ ƒƒ‰ƒ™ƒǡ̵‡ƒ”›ȋʹͲͲͳȌ‘„‹ƒ–‘”‹ƒŽ‡š’”‡••‹‘’ƒ––‡”•‘ˆ ǦŠ‘‡‘†‘ƒ‹ƒ†

ϳϵϬ ‘–Š‡””‡‰—Žƒ–‘”›‰‡‡•’ƒ” ‡ŽŽƒ–‡†‡˜‡Ž‘’‹‰–ŠƒŽƒ—•Ǥ ‡—”‘• ‹ʹͳǣʹ͹ͳͳȂʹ͹ʹͷǤ

ϳϵϭ ƒ”„‘—šǦ²‡ ǡ ƒ•’ƒ”  ȋʹͲͲͺȌ ±˜‡Ž‘’’‡‡– †— ‘”–‡š ±”±„”ƒŽ ǣ ƒ’’‘”–• ”± ‡–• †‡•

ϳϵϮ ±–—†‡• Š‡œŽƒ•‘—”‹•‡–Ž‡•’”‹ƒ–‡•Ǥ’‹Ž‡’•‹‡ǣͳȂͻǤ

ϳϵϯ ‘ –‘”ǡƒ”–À‡œǦ‡”†‡Ó‘ǡ ˜‹ ǡ”‹‡‰•–‡‹ȋʹͲͲͶȌ‘”–‹ ƒŽ‡—”‘•ƒ”‹•‡‹•›‡–”‹ 

ϳϵϰ ƒ† ƒ•›‡–”‹  †‹˜‹•‹‘ œ‘‡• ƒ† ‹‰”ƒ–‡ –Š”‘—‰Š •’‡ ‹ˆ‹  ’Šƒ•‡•Ǥ ƒ– ‡—”‘• ‹ ͹ǣͳ͵͸Ȃ

ϳϵϱ ͳͶͶǤ

ϯϬ 

ϳϵϲ ‘—”ƒǡ ‘Ž„‡”‰ ǡ ”‹•‡ ǡ•—‹ȋʹͲͲ͸Ȍƒš͸Ǧ†‡’‡†‡–„‘—†ƒ”›†‡ˆ‹‡•ƒŽ‹‰‡–‘ˆ

ϳϵϳ ‹‰”ƒ–‹‰‘Žˆƒ –‘”› ‘”–‡š‡—”‘•˜‹ƒ–Š‡”‡’—Ž•‹˜‡ƒ –‹˜‹–›‘ˆ ‡’Š”‹ ͷǤ ‡˜‡Ž‘’‡–

ϳϵϴ ͳ͵͵ǣͳ͵͵ͷȂͳ͵ͶͷǤ

ϳϵϵ ̵‡ƒ”› ǡ Š‘— Ǧ ǡ ƒŠƒ”ƒ  ȋʹͲͲ͹Ȍ ”‡ƒ ƒ––‡”‹‰ ‘ˆ –Š‡ ƒƒŽ‹ƒ ‘”–‡šǤ ‡—”‘

ϴϬϬ ͷ͸ǣʹͷʹȂʹ͸ͻǤ

ϴϬϭ ‰ƒ™ƒǡ‹›ƒ–ƒǡƒƒŒ‹ƒǡƒ‰›—ǡ‡‹‡ǡ ‡ƒƒǡƒƒ‘–‘ ǡ‹‘•Š‹„ƒȋͳͻͻͷȌ

ϴϬϮ Š‡ ”‡‡Ž‡” ‰‡‡Ǧƒ••‘ ‹ƒ–‡† ƒ–‹‰‡ ‘ ƒŒƒŽǦ‡–œ‹—• ‡—”‘• ‹• ƒ ”— ‹ƒŽ ‘Ž‡ —Ž‡ ˆ‘”

ϴϬϯ Žƒ‹ƒ”‘”‰ƒ‹œƒ–‹‘‘ˆ ‘”–‹ ƒŽ‡—”‘•Ǥ‡—”‘ͳͶǣͺͻͻȂͻͳʹǤ

ϴϬϰ Œ‹ƒ ǡ‘”‹ǡ‹•Š‹ȋͳͻͺͶȌŠ‡–”ƒŒ‡ –‘”›‘ˆ‹–”ƒŽ ‡ŽŽƒš‘•‹–Š‡”ƒ„„‹–‘Žˆƒ –‘”› ‘”–‡š

ϴϬϱ ”‡˜‡ƒŽ‡†„›‹–”ƒ ‡ŽŽ—Žƒ” ‹Œ‡ –‹‘Ǥ ‘’‡—”‘Žʹ͵Ͳǣ͹͹Ȃͺ͹Ǥ

ϴϬϲ ƒ†ƒƒ„Šƒǡ•ƒƒ†ƒ ǡƒ”ƒ„”‹ƒǡ‹œ‡”ǡƒŽŽƒ™ƒ›ǡ ƒ‰‡ ǡ‡Œ‘™•‹ ȋʹͲͳ͸Ȍ

ϴϬϳ ‹˜‡”•‡ ‡’”‡•‡–ƒ–‹‘• ‘ˆ Žˆƒ –‘”› ˆ‘”ƒ–‹‘ ‹ ‡–”‹ˆ—‰ƒŽ ‡‡†„ƒ  ”‘Œ‡ –‹‘•Ǥ

ϴϬϴ ‘—”ƒŽ‘ˆ‡—”‘• ‹‡ ‡͵͸ǣ͹ͷ͵ͷȂ͹ͷͶͷǤ

ϴϬϵ ‡•‘Ž†ǡ ’ƒ‰ƒ–‹‡ŽŽ‘ ǡ‹•— ǡœ—‘˜ǡ‘•–ƒǡ —‹†‘––‹ǡƒ”— Š‘ ȋͳͻͻͺȌ‡‡Ž‹‹•

ϴϭϬ ’”‡ˆ‡”‡–‹ƒŽŽ› ‡š’”‡••‡† ‹ ‡—”‘• •›–Š‡•‹œ‹‰ ‰ƒƒǦƒ‹‘„—–›”‹  ƒ ‹† ‹ ‘”–‡š ƒ†

ϴϭϭ Š‹’’‘ ƒ’—•‘ˆƒ†—Ž–”ƒ–•Ǥ”‘ ‡‡†‹‰•‘ˆ–Š‡ƒ–‹‘ƒŽ ƒ†‡›‘ˆ ‹‡ ‡•ͻͷǣ͵ʹʹͳȂ͵ʹʹ͸Ǥ

ϴϭϮ ‹‘ ǡ —‘  ǡ •Š‡”›Ǧƒ†ƒ ǡ ‘Žž” ǡ –‘›‘˜ƒ  ȋʹͲͲͺȌ Ž–‡”‡† ‘Ž‡ —Žƒ”

ϴϭϯ ‡‰‹‘ƒŽ‹œƒ–‹‘ƒ†‘”ƒŽŠƒŽƒ‘ ‘”–‹ ƒŽ‘‡ –‹‘•‹‘”–‡šǦ’‡ ‹ˆ‹ ƒš͸‘ Ǧ—–

ϴϭϰ ‹ ‡Ǥ ‘—”ƒŽ‘ˆ‡—”‘• ‹‡ ‡ʹͺǣͺ͹ʹͶȂͺ͹͵ͶǤ

ϴϭϱ ‘‘ǡ •ƒƒ •‘ ȋʹͲͲͻȌ†‘”‡’”‡•‡–ƒ–‹‘•‹Žˆƒ –‘”›‘”–‡šǣȈȈ’ƒ”•‡ȈȈ‘†‹‰ǡ Ž‘„ƒŽ

ϴϭϲ Š‹„‹–‹‘ǡƒ†• ‹ŽŽƒ–‹‘•Ǥ‡—”‘͸ʹǣͺͷͲȂͺ͸ͳǤ

ϴϭϳ ‘‘ǡ •ƒƒ •‘ ȋʹͲͳͳȌƒŒ‘”‘Ž‡ˆ‘” –”ƒ ‘”–‹ ƒŽ‹” —‹–•‹–Š‡–”‡‰–Šƒ†—‹‰‘ˆ

ϴϭϴ †‘”Ǧ˜‘‡†š ‹–ƒ–‹‘‹Žˆƒ –‘”›‘”–‡šǤ‡—”‘͹ʹǣͶͳȂͶͺǤ

ϯϭ 

ϴϭϵ —‡ŽŽ‡• ǡ —™ƒƒ ǡ —‡ŽŽ‡• ǡ —Žˆ‘‡ ǡ Š‹ƒ—”ƒ ǡ ‡Ž‡Š‡” ǡ ‹‰ƒ ǡ —„‡•–‡‹ 

ϴϮϬ ȋʹͲͲͲȌƒŽŽ‹ƒŽƒ†•—„’ƒŽŽ‹ƒŽ†‡”‹˜ƒ–‹˜‡•‹–Š‡‡„”›‘‹  Š‹ ƒ†‘—•‡–‡Ž‡ ‡’ŠƒŽ‘ǡ

ϴϮϭ –”ƒ ‡†„›–Š‡‡š’”‡••‹‘‘ˆ–Š‡‰‡‡•ŽšǦʹǡšǦͳǡšǦʹǤͳǡƒšǦ͸ǡƒ†„”ǦͳǤ ‘’‡—”‘Ž

ϴϮϮ ͶʹͶǣͶͲͻȂͶ͵ͺǤ

ϴϮϯ ƒ‹   ȋʹͲͲ͵Ȍ ‡˜‡Ž‘’‡–ƒŽ ƒ† ˜‘Ž—–‹‘ƒ”› †ƒ’–ƒ–‹‘• ‘ˆ ‘”–‹ ƒŽ ƒ†‹ƒŽ Ž‹ƒǤ ‡”‡„”ƒŽ

ϴϮϰ ‘”–‡šǣͳȂͻǤ

ϴϮϱ ƒ‹ ǡ›‘—„ǡ”‡—‹‰ ǡ‘‹‰—‡œ ȋʹͲͲͻȌ‡ ‹•‹‘„›†‹˜‹•‹‘ǣƒ‹‰ ‘”–‹ ƒŽƒ’•Ǥ

ϴϮϲ ”‡†•‹‡—”‘• ‹‡ ‡•͵ʹǣʹͻͳȂ͵ͲͳǤ

ϴϮϳ ƒ‘•Ǧ‘”‡‘ ǡ ƒŽƒœ‘  ǡ ‘””‡”‘ ǡ ƒ”–À‡œǦ‡”†‡Ó‘ ǡ Žƒ• ž  ȋʹͲͲ͸Ȍ š–”ƒ ‡ŽŽ—Žƒ”

ϴϮϴ ƒ–”‹š ‘Ž‡ —Ž‡• ƒ† •›ƒ’–‹  ’Žƒ•–‹ ‹–›ǣ ‹—‘ƒ’’‹‰ ‘ˆ ‹–”ƒ ‡ŽŽ—Žƒ” ƒ† •‡ ”‡–‡†

ϴϮϵ ‡‡Ž‹‹–Š‡ƒ†—Ž–”ƒ–„”ƒ‹Ǥ—” ‡—”‘• ‹ʹ͵ǣͶͲͳȂͶʹʹǤ

ϴϯϬ ƒ•†‡ ǡò”‡Ž‹ ǡ ‘ƒ‰Š ǡ‘Žƒ ȋʹͲͳͷȌƒ‹ƒ”ƒ†‘”•‘˜‡–”ƒŽ‘Ž‡ —Žƒ”

ϴϯϭ ”‰ƒ‹œƒ–‹‘‘ˆ–Š‡‡†‹ƒŽ–‘”Š‹ƒŽ‘”–‡š‡˜‡ƒŽ‡†„›ƒ”‰‡Ǧ• ƒŽ‡ƒ–‘‹ ƒŽƒŽ›•‹•

ϴϯϮ ‘ˆ ‡‡š’”‡••‹‘Š‡ Š‹ ǡ‡†Ǥ‘‘’—–‹‘Žͳͳǣ‡ͳͲͲͶͲ͵ʹȂ‡ͳͲͲͶͲ͵ͺǤ

ϴϯϯ ƒ› ǡ ”‡ Š–  ȋʹͲͳ͸Ȍ –”— –—”ƒŽ †‡˜‡Ž‘’‡– ƒ† †‘”•‘˜‡–”ƒŽ ƒ–—”ƒ–‹‘ ‘ˆ –Š‡ ‡†‹ƒŽ

ϴϯϰ ‡–‘”Š‹ƒŽ ‘”–‡šǤ‡‹ˆ‡ͷǣͳ͹ͷǤ

ϴϯϱ ‡ƒ‡”ǡŠ‡Ǧ ǡ—›Ž‡ǡŽ‘ƒǡ‹Ž•‘ȋʹͲͲ͹Ȍ’ƒ–‹ƒŽ ƒ† –‡’‘”ƒŽ

ϴϯϲ †‹•–”‹„—–‹‘‘ˆ‘†‘”ƒ–Ǧ‡˜‘‡†ƒ –‹˜‹–›‹–Š‡’‹”‹ˆ‘” ‘”–‡šǤ ‡—”‘• ‹ʹ͹ǣͳͷ͵ͶȂͳͷͶʹǤ

ϴϯϳ ‘—ƒ—šǡ”Ž‘––ƒȋʹͲͳͲȌ ‡œˆʹ†‹”‡ –•–Š‡†‹ˆˆ‡”‡–‹ƒ–‹‘‘ˆ ‘”–‹ ‘ˆ—‰ƒŽ‡—”‘•ˆ”‘•–”‹ƒ–ƒŽ

ϴϯϴ ’”‘‰‡‹–‘”•‹˜‹˜‘Ǥƒ–‡—”‘• ‹ͳ͵ǣͳ͵ͶͷȂͳ͵Ͷ͹Ǥ

ϴϯϵ ƒ Š‡œǦ†”ƒ†‡ ǡ‡†”‹ ȋʹͲͲͻȌŠ‡ƒ‹‘Žˆƒ –‘”›•›•–‡ ƒ† •‘ ‹ƒŽ Ž‡ƒ”‹‰ ‹

ϴϰϬ ƒƒŽ•Ǥ‡Šƒ˜‹‘—”ƒŽ”ƒ‹‡•‡ƒ” ŠʹͲͲǣ͵ʹ͵Ȃ͵͵ͷǤ

ϴϰϭ ƒ”ƒǡ‹ Šƒ”†ǡ ”‡‡”ȋʹͲͳͳȌ‡˜‡Ž‘’‡–ƒŽ›ƒ‹ •‘ˆ‹”‹ˆ‘”‘”–‡šǤ‡”‡„”ƒŽ

ϯϮ 

ϴϰϮ ‘”–‡šʹͳǣͳʹ͵ͳȂͳʹͶͷǤ

ϴϰϯ ƒ–‘ ǡ ‹”ƒ–ƒ ǡ ‰ƒ™ƒ ǡ —Œ‹•ƒ™ƒ  ȋͳͻͻͺȌ ‡“—‹”‡‡– ˆ‘” ƒ”Ž›Ǧ ‡‡”ƒ–‡† ‡—”‘•

ϴϰϰ ‡ ‘‰‹œ‡† „› ‘‘ Ž‘ƒŽ –‹„‘†› ‘–ͳ ‹ –Š‡ ‘”ƒ–‹‘ ‘ˆ ƒ–‡”ƒŽ Žˆƒ –‘”› ”ƒ –Ǥ 

ϴϰϱ ‡—”‘• ‹ǣͳȂͳͳǤ

ϴϰϲ  Š‹†ǡ ”ƒ–Šǡ‡”‡ Š‹†ǡ‘›Ž‡•ǡƒ” Š‹‘‹ǡ‡•–ƒ ǡ –‘  ȋʹͲͲ͵Ȍ

ϴϰϳ ‡—”‡‰—Ž‹ͳǦ‡”„ʹ•‹‰ƒŽ‹‰‹•”‡“—‹”‡†ˆ‘”–Š‡‡•–ƒ„Ž‹•Š‡–‘ˆ ”ƒ†‹ƒŽ ‰Ž‹ƒ ƒ† –Š‡‹”

ϴϰϴ –”ƒ•ˆ‘”ƒ–‹‘‹–‘ƒ•–”‘ ›–‡•‹ ‡”‡„”ƒŽ ‘”–‡šǤ”‘ ‡‡†‹‰•‘ˆ–Š‡ƒ–‹‘ƒŽ ƒ†‡›‘ˆ

ϴϰϵ  ‹‡ ‡•ͳͲͲǣͶʹͷͳȂͶʹͷ͸Ǥ

ϴϱϬ  Š™‘„ ǡ”‹ ‡ ȋͳͻͺͶȌŠ‡†‡˜‡Ž‘’‡–‘ˆŽƒ‹ƒ–‹‘‘ˆƒˆˆ‡”‡– ˆ‹„‡”• –‘ –Š‡ ‘Žˆƒ –‘”›

ϴϱϭ ‘”–‡š‹”ƒ–•ǡ™‹–Šƒ††‹–‹‘ƒŽ‘„•‡”˜ƒ–‹‘•‹–Š‡ƒ†—Ž–Ǥ ‘’‡—”‘Žʹʹ͵ǣʹͲ͵ȂʹʹʹǤ

ϴϱϮ Š‡’Š‡”† ȋʹͲͳͳȌŠ‡‹ ”‘ ‹” —‹–‘ ‡’–’’Ž‹‡†–‘‘”–‹ ƒŽ˜‘Ž—–‹‘ǣˆ”‘Š”‡‡Ǧƒ›‡”

ϴϱϯ –‘‹šǦƒ›‡”‘”–‡šǤ ”‘–‡—”‘ƒƒ–ͷǣͳȂͳͷǤ

ϴϱϰ Š‹„ƒ–ƒǡ—”‘ƒ™ƒǡƒƒ‘ ǡŠ—”ƒǡ‹œƒ™ƒȋʹͲͲͺȌ‹ ”‘Ǧͻ ‘†—Žƒ–‡• ƒŒƒŽǦ

ϴϱϱ ‡–œ‹—• ‡ŽŽ ‹ˆˆ‡”‡–‹ƒ–‹‘ „› —’’”‡••‹‰ ‘š‰ͳ š’”‡••‹‘ ‹ ‘—•‡ ‡†‹ƒŽ ƒŽŽ‹—Ǥ

ϴϱϲ ‘—”ƒŽ‘ˆ‡—”‘• ‹‡ ‡ʹͺǣͳͲͶͳͷȂͳͲͶʹͳǤ

ϴϱϳ Š‹’Ž‡›ǡ†ƒ‡ ȋͳͻͺͶȌ–Š‡ ‘‡ –‹‘•‘ˆ–Š‡‘—•‡‘Žˆƒ –‘”› „—Ž„ǣ  •–—†› —•‹‰

ϴϱϴ ‘”–Š‘‰”ƒ†‡ ƒ† ”‡–”‘‰”ƒ†‡ –”ƒ•’‘”– ‘ˆ ™Š‡ƒ– ‰‡” ƒ‰‰Ž—–‹‹ ‘Œ—‰ƒ–‡† –‘ Š‘”•‡”ƒ†‹•Š

ϴϱϵ ’‡”‘š‹†ƒ•‡Ǥ”ƒ‹‡•‡ƒ” Š—ŽŽ‡–‹ͳʹǣ͸͸ͻȂ͸ͺͺǤ

ϴϲϬ ƒ”– ǡƒ”–ȋͳͻ͹͹ȌŠ‡Ž‘ ƒ–‹‘‘ˆ— Ž‡‹‘ˆ†‹ˆˆ‡”‡–Žƒ„‡ŽŽ‹‰ ‹–‡•‹–‹‡• ‹

ϴϲϭ ƒ—–‘”ƒ†‹‘‰”ƒ’Š•‘ˆ–Š‡ƒ–‡”‹‘”ˆ‘”‡„”ƒ‹‘ˆ’‘•–ƒ–‹ƒŽ‹ ‡‹Œ‡ –‡†™‹–Šȏ͵ Ȑ–Š›‹†‹‡‘

ϴϲϮ –Š‡‡Ž‡˜‡–Šƒ†–™‡Žˆ–Š†ƒ›•’‘•–Ǧ ‘ ‡’–‹‘Ǥ ‘—”ƒŽ‘ˆƒ–‘›ͳʹ͵ǣͷͳʹȂͷʹͷǤ

ϴϲϯ ‘•—Ž•‹ǡ‹••‹–•›ƒŽ‘‘ǡ—–ˆ‘”–Šǡš‡Žǡƒ––ƒȋʹͲͳͳȌ‹•–‹ –”‡’”‡•‡–ƒ–‹‘•‘ˆ

ϴϲϰ ‘Žˆƒ –‘”›‹ˆ‘”ƒ–‹‘‹†‹ˆˆ‡”‡– ‘”–‹ ƒŽ ‡–”‡•Ǥƒ–—”‡Ͷ͹ʹǣʹͳ͵Ȃʹͳ͸Ǥ

ϯϯ 

ϴϲϱ “—ƒ”œ‘‹  ȋʹͲͳͷȌ ‡—”‘ƒŽ ƒ† ‹ ”‘‰Ž‹ƒŽ ”‡‰—Žƒ–‘”• ‘ˆ ‘”–‹ ƒŽ ™‹”‹‰ǣ —•—ƒŽ ƒ† ‘˜‡Ž

ϴϲϲ ‰—‹†‡’‘•–•Ǥ ”‘–‡—”‘• ‹ͻǣͳȂͳ͸Ǥ

ϴϲϳ –‡ƒ ȋʹͲͲ͵ȌŽšƒ†ƒš͸ ‘Ǧ‘’‡”ƒ–‡‰‡‡–‹ ƒŽŽ›–‘‡•–ƒ„Ž‹•Š–Š‡’ƒŽŽ‹‘Ǧ•—„’ƒŽŽ‹ƒŽ„‘—†ƒ”›

ϴϲϴ ‹–Š‡‡„”›‘‹ ‘—•‡–‡Ž‡ ‡’ŠƒŽ‘Ǥ‡˜‡Ž‘’‡–ͳ͵Ͳǣͳͳͳ͵ȂͳͳʹʹǤ

ϴϲϵ –‡––Ž‡”ǡš‡ŽȋʹͲͲͻȌ‡’”‡•‡–ƒ–‹‘•‘ˆ†‘”‹–Š‡‹”‹ˆ‘”‘”–‡šǤ‡—”‘͸͵ǣͺͷͶȂͺ͸ͶǤ

ϴϳϬ –‡™ƒ”– ǡ‡ƒ”ŽƒȋͳͻͻͲȌ ‹„”‘‡ –‹ǦŽ‹‡Ž—‘”‡ƒ –‹˜‹–›‹–Š‡‡˜‡Ž‘’‹‰‡”‡„”ƒŽ

ϴϳϭ ‘”–‡šǤ ‡—”‘• ‹ǣͳȂͻǤ

ϴϳϮ –‘‡• ǡ •ƒƒ •‘  ȋʹͲͳͲȌ ”‘ †‡†”‹–‡ –‘ •‘ƒǣ †›ƒ‹  ”‘—–‹‰‘ˆ‹Š‹„‹–‹‘„›

ϴϳϯ ‘’Ž‡‡–ƒ”›‹–‡”‡—”‘‹ ”‘ ‹” —‹–•‹‘Žˆƒ –‘”› ‘”–‡šǤ‡—”‘͸͹ǣͶͷʹȂͶ͸ͷǤ

ϴϳϰ –‘›‘˜ƒ ǡ ”‹–• Š ǡ ƒŽ–Š‡” ǡ ”—••  ȋͳͻͻ͸Ȍ ‘”‡„”ƒ‹ ’ƒ––‡”‹‰ †‡ˆ‡ –• ‹ ƒŽŽ ‡›‡

ϴϳϱ —–ƒ–‹ ‡Ǥ‡˜‡Ž‘’‡–ͳʹʹǣ͵Ͷͷ͵Ȃ͵Ͷ͸ͷǤ

ϴϳϲ –‘›‘˜ƒǡ ږœǡ ”—••ǡ”‹ ‡ ȋͳͻͻ͹Ȍƒš͸Ǧ†‡’‡†‡–”‡‰—Žƒ–‹‘‘ˆƒ†Š‡•‹˜‡’ƒ––‡”‹‰ǡǦ

ϴϳϳ ƒ†Š‡”‹ ‡š’”‡••‹‘ ƒ† „‘—†ƒ”› ˆ‘”ƒ–‹‘ ‹ †‡˜‡Ž‘’‹‰ ˆ‘”‡„”ƒ‹Ǥ ‡˜‡Ž‘’‡–

ϴϳϴ ͳʹͶǣ͵͹͸ͷȂ͵͹͹͹Ǥ

ϴϳϵ –‘›‘˜ƒǡ”‡‹ Š‡Žǡ ƒŽŽ‘‡–ǡ ”—••ȋʹͲͲͲȌƒš͸‘†—Žƒ–‡•–Š‡‘”•‘˜‡–”ƒŽƒ––‡”‹‰

ϴϴϬ ‘ˆ–Š‡ƒƒŽ‹ƒ‡Ž‡ ‡’ŠƒŽ‘Ǥ ‡—”‘• ‹ǣͳȂͻǤ

ϴϴϭ —„”ƒƒ‹ƒǡ‡‡†‹‘•ǡŠ‡––›ǡ‘Ž‡ȋʹͲͲͻȌ‹‰ƒŽ•ˆ”‘–Š‡‡†‰‡•ǣŠ‡ ‘”–‹ ƒŽŠ‡

ϴϴϮ ƒ† ƒ–‹Š‡ ‹ –‡Ž‡ ‡’ŠƒŽ‹  †‡˜‡Ž‘’‡–Ǥ ‡‹ƒ”• ‹ ‡ŽŽ ƒ† ‡˜‡Ž‘’‡–ƒŽ ‹‘Ž‘‰›

ϴϴϯ ʹͲǣ͹ͳʹȂ͹ͳͺǤ

ϴϴϰ —ŽŽ‹˜ƒǡ‹Ž•‘ȋʹͲͲ͵Ȍ‘Ž‡ —Žƒ”„‹‘Ž‘‰›‘ˆ‡ƒ”Ž›‘Žˆƒ –‘”›‡‘”›Ǥ‡ƒ”‡ͳͲǣͳȂͶǤ

ϴϴϱ —’°” ǡ‘”‹ƒ‘ǡ›Ž‹‰• ȋͳͻͻͺȌŠ‡ˆ— –‹‘•‘ˆ–Š‡’”‡’Žƒ–‡‹†‡˜‡Ž‘’‡–ƒ†‡˜‘Ž—–‹‘

ϴϴϲ ‘ˆ–Š‡‡‘ ‘”–‡šƒ†Š‹’’‘ ƒ’—•Ǥ”ƒ‹‡•‡ƒ” Š‡˜‹‡™•ǣͳȂʹͷǤ

ϯϰ 

ϴϴϳ —œ—‹ ǡ ‡‡”•  ȋʹͲͳͲƒȌ ‹•–‹ –‹˜‡ Žƒ••‡• ‘ˆ ‡”‰‹  ‹–‡”‡—”‘• ’”‘˜‹†‡ Žƒ›‡”Ǧ

ϴϴϴ •’‡ ‹ˆ‹ ’Šƒ•‹ ‹Š‹„‹–‹‘‹–Š‡ƒ–‡”‹‘”’‹”‹ˆ‘” ‘”–‡šǤ‡”‡„‘”–‡šʹͲǣʹͻ͹ͳȂʹͻͺͶǤ

ϴϴϵ —œ—‹ǡ‡‡”• ȋʹͲͳͲ„Ȍ Š‹„‹–‘”›‡—”‘•‹–Š‡ƒ–‡”‹‘”’‹”‹ˆ‘” ‘”–‡š‘ˆ–Š‡‘—•‡ǣ

ϴϵϬ Žƒ••‹ˆ‹ ƒ–‹‘—•‹‰‘Ž‡ —Žƒ”ƒ”‡”•Ǥ ‘’‡—”‘Žͷͳͺǣͳ͸͹ͲȂͳ͸ͺ͹Ǥ

ϴϵϭ —œ—‹ ǡ ‡‡”•  ȋʹͲͳͳȌ ™‘ ƒ›‡”• ‘ˆ ›ƒ’–‹  ”‘ ‡••‹‰ „› ”‹ ‹’ƒŽ ‡—”‘• ‹

ϴϵϮ ‹”‹ˆ‘”‘”–‡šǤ ‘—”ƒŽ‘ˆ‡—”‘• ‹‡ ‡͵ͳǣʹͳͷ͸Ȃʹͳ͸͸Ǥ

ϴϵϯ ƒ‹‰— Š‹Ǧ ƒ›ƒ•Š‹  ȋʹͲͲͶȌ ‡‡”ƒ–‹‘ ‘ˆ ‡‡Ž‹Ǧ‘•‹–‹˜‡ ƒ”‰‹ƒŽ ‘‡ ‡ŽŽ• ˆ”‘ –Š‡

ϴϵϰ ƒ—†‘‡†‹ƒŽƒŽŽ‘ˆ‡Ž‡ ‡’ŠƒŽ‹ ‡•‹ Ž‡•Ǥ ‘—”ƒŽ‘ˆ‡—”‘• ‹‡ ‡ʹͶǣʹʹͺ͸ȂʹʹͻͷǤ

ϴϵϱ ƒ‰ǡ ƒ••‡Ž‘ȋͳͻͻͶȌ‡Ž‡ –‹˜‡•—’’”‡••‹‘‘ˆ‹–”‹•‹ „—–‘–ƒˆˆ‡”‡–ˆ‹„‡”•›ƒ’–‹ 

ϴϵϲ –”ƒ•‹••‹‘„›„ƒ Ž‘ˆ‡‹–Š‡’‹”‹ˆ‘”ȋ‘Žˆƒ –‘”›Ȍ ‘”–‡šǤ”ƒ‹‡•‡ƒ” Š͸ͷͻǣ͹ͷȂͺͳǤ

ϴϵϳ ƒ‹‰— Š‹ ȋʹͲͳͶȌ ‡‡–‹ †‹••‡ –‹‘‘ˆ ‡”‰‹ ‡—”ƒŽ ‹” —‹–•‹‘—•‡‡‘ ‘”–‡šǤǣͳȂʹʹǤ

ϴϵϴ ‘Ž‡  ȋʹͲͲͷȌ ‡Ž‡ –‹˜‡ ‡“—‹”‡‡– ‘ˆ ƒš͸ǡ —– ‘– šʹǡ ‹ –Š‡ ’‡ ‹ˆ‹ ƒ–‹‘ ƒ†

ϴϵϵ ‡˜‡Ž‘’‡–‘ˆ‡˜‡”ƒŽ— Ž‡‹‘ˆ–Š‡›‰†ƒŽ‘‹†‘’Ž‡šǤ ‘—”ƒŽ ‘ˆ ‡—”‘• ‹‡ ‡

ϵϬϬ ʹͷǣʹ͹ͷ͵Ȃʹ͹͸ͲǤ

ϵϬϭ ‘‹‘ƒ ǡ •—‹ ǡ ƒ–‘ ǡ ‘—‡ ǡ ƒƒ—”ƒ ǡ —Œ‹•ƒ™ƒ ǡ ‹”ƒ–ƒȋʹͲͲͲȌ‡‘ ‘”–‹ ƒŽ

ϵϬϮ ”‹‰‹ ƒ† ƒ‰‡–‹ƒŽ ‹‰”ƒ–‹‘ ‘ˆ —‹†‡’‘•– ‡—”‘• ‹ –Š‡ ƒ–‡”ƒŽŽˆƒ –‘”›”ƒ –Ǥ 

ϵϬϯ ‡—”‘• ‹ǣͳȂͳͳǤ

ϵϬϰ ‘”‡••‘ ǡ‘––‡”ǡƒ’„‡ŽŽȋʹͲͲͲȌ ‡‡–‹  ‘–”‘Ž‘ˆ†‘”•ƒŽǦ˜‡–”ƒŽ ‹†‡–‹–› ‹ –Š‡

ϵϬϱ –‡Ž‡ ‡’ŠƒŽ‘ǣ‘’’‘•‹‰”‘Ž‡•ˆ‘”ƒš͸ƒ† •ŠʹǤ‡˜‡Ž‘’‡–ͳʹ͹ǣͶ͵͸ͳȂͶ͵͹ͳǤ

ϵϬϲ ƒŽ˜‡”†‡ ǡƒ–ƒ ƒƒȋͳͻͻͶȌ‡˜‡Ž‘’‡–ƒ†‡ƒ”Ž›’‘•–ƒ–ƒŽƒ–—”ƒ–‹‘‘ˆ–Š‡”ƒ–’”‹ƒ”›

ϵϬϳ ‘Žˆƒ –‘”› ‘”–‡šǤ‡˜‡Ž‘’‡–ƒŽ”ƒ‹‡•‡ƒ” ŠͺͲǣȋͳȂʹȌǣͻ͸ȂͳͳͶǤ

ϵϬϴ ƒ”‰ƒǡ‡‡ǡ‘Ž–‡•œ ȋʹͲͳͲȌƒ”‰‡–Ǧ•‡Ž‡ –‹˜‡ ‡”‰‹  ‘–”‘Ž‘ˆ‡–‘”Š‹ƒŽ ‘”–‡š‘—–’—–Ǥ

ϯϱ 

ϵϬϵ ƒ–—”‡—„Ž‹•Š‹‰ ”‘—’ͳ͵ǣͺʹʹȂͺʹͶǤ

ϵϭϬ ‹–ƒŽ‹ ǡ ƒ„ƒ—†‘ȋʹͲͳͶȌ›ƒ’–‹ „‹‘Ž‘‰›‘ˆ„ƒ””‡Ž ‘”–‡š ‹” —‹–ƒ••‡„Ž›Ǥ‡‹ƒ”•‹‡ŽŽ

ϵϭϭ ƒ†‡˜‡Ž‘’‡–ƒŽ‹‘Ž‘‰›͵ͷǣͳͷ͸Ȃͳ͸ͶǤ

ϵϭϮ ‹–ƒŽ‹•ǡ‘••‹‡” ȋʹͲͳͲȌ‡™‹•‹‰Š–•‹–‘ ‘”–‹ ƒŽ‹–‡”‡—”‘• †‡˜‡Ž‘’‡– ƒ†

ϵϭϯ Žƒ••‹ˆ‹ ƒ–‹‘ǣ ‘–”‹„—–‹‘ ‘ˆ †‡˜‡Ž‘’‡–ƒŽ •–—†‹‡•  ƒ‹  ǡ ‹•Š‡ŽŽ ǡ ‡†•Ǥ ‡˜‡Ž

ϵϭϰ ‡—”‘„‹‘͹ͳǣ͵ͶȂͶͶǤ

ϵϭϱ ƒ‰‡‡” ǡƒ˜‹†ǡŠƒ‘ǡ ƒƒ•ǡ–ƒ‹‰‡” ȋʹͲͳͲȌŠ‡‘ƒ–‘•‡•‘”› ‘”–‡š ‘ˆ ”‡‡Ž‡”

ϵϭϲ —–ƒ– ‹ ‡ Š‘™• „•‡– ƒ›‡”‹‰ —– –ƒ – ‘”ƒ–‹‘ ƒ† ‡Šƒ˜‹‘”ƒŽ  –‹˜ƒ–‹‘ ‘ˆ

ϵϭϳ ‘Ž—ƒ”‘ƒ–‘–‘’‹ ƒ’•Ǥ ‘—”ƒŽ‘ˆ‡—”‘• ‹‡ ‡͵Ͳǣͳͷ͹ͲͲȂͳͷ͹ͲͻǤ

ϵϭϴ ‹‡‰ƒ† ǡ‡‡†ǡ‡†‡Ž• ǡ‡‹„‘Ž†ǡ Š‹–œǡ ‘Š‡‹‰ ȋʹͲͳͳȌ‘’Ž‡‡–ƒ”›

ϵϭϵ ‡•‘”›ƒ†••‘ ‹ƒ–‹˜‡‹ ”‘ ‹” —‹–”›‹”‹ƒ”›Žˆƒ –‘”›‘”–‡šǤ ‘—”ƒŽ‘ˆ‡—”‘• ‹‡ ‡

ϵϮϬ ͵ͳǣͳʹͳͶͻȂͳʹͳͷͺǤ

ϵϮϭ ‹Ž•‘ ǡƒ‹‡ ȋʹͲͲ͸Ȍƒ”Ž›˜‡–•‹Žˆƒ –‘”›”‘ ‡••‹‰Ǥ—‡˜‡—”‘• ‹ǣͳȂͶͳǤ

ϵϮϮ ‘‘† ǡƒ”–‹ǡ”‹ ‡ ȋͳͻͻʹȌ˜‹†‡ ‡–Šƒ––Š‡‡ƒ”Ž‹‡•–‰‡‡”ƒ–‡† ‡ŽŽ• ‘ˆ –Š‡ —”‹‡

ϵϮϯ ‡”‡„”ƒŽ ‘”–‡š ˆ‘” ƒ –”ƒ•‹‡– ’‘’—Žƒ–‹‘ ‹ –Š‡ •—„’Žƒ–‡ ƒ† ƒ”‰‹ƒŽ œ‘‡Ǥ

ϵϮϰ ‡˜‡Ž‘’‡–ƒŽ”ƒ‹‡•‡ƒ” Š͸͸ǣͳ͵͹ȂͳͶͲǤ

ϵϮϱ ‘•Š‹†ƒ ȋʹͲͲ͸Ȍ ƒ••‹˜‡Ž‘••‘ˆƒŒƒŽǦ‡–œ‹—• ‡ŽŽ•†‘‡•‘– †‹•”—’–‡‘ ‘”–‹ ƒŽŽƒ›‡”‘”†‡”Ǥ

ϵϮϲ ‡˜‡Ž‘’‡–ͳ͵͵ǣͷ͵͹ȂͷͶͷǤ

ϵϮϳ —ǡ‘––‡”ǡ—„‡•–‡‹ ȋʹͲͲͳȌ •Šʹƒ†ƒš͸’Žƒ› ‘’Ž‡‡–ƒ”›”‘Ž‡•‹†‘”•‘˜‡–”ƒŽ

ϵϮϴ ’ƒ––‡”‹‰‘ˆ–Š‡ƒƒŽ‹ƒ–‡Ž‡ ‡’ŠƒŽ‘Ǥ‡˜‡Ž‘’‡–ͳʹͺǣͳͻ͵ȂʹͲͷǤ

ϵϮϵ Šƒ ǡ —‘  ȋʹͲͳͲȌ ‹˜‡”•‡ ƒ––‡”• ‘ˆ †‘” ‡’”‡•‡–ƒ–‹‘ „› ‡—”‘• ‹ –Š‡ –‡”‹‘”

ϵϯϬ ‹”‹ˆ‘”‘”–‡š‘ˆ™ƒ‡‹ ‡Ǥ ‘—”ƒŽ‘ˆ‡—”‘• ‹‡ ‡͵Ͳǣͳ͸͸͸ʹȂͳ͸͸͹ʹǤ

ϯϲ 

ϵϯϭ Šƒ‘ǡ —ƒǡŽ‡ƒ•—”‡ ȋʹͲͲ͸Ȍ–”ƒ•‰‡‹ ƒ”‡”‘—•‡Ž‹‡Žƒ„‡Ž•ƒŒƒŽȂ‡–œ‹—• ‡ŽŽ•

ϵϯϮ ˆ”‘–Š‡ ‘”–‹ ƒŽŠ‡ƒ†–ŠƒŽƒ‘ ‘”–‹ ƒŽƒš‘•Ǥ”ƒ‹‡•‡ƒ” ŠͳͲ͹͹ǣͶͺȂͷ͵Ǥ

ϵϯϯ Šƒ‘ǡœƒ„×ǡƒ ǡ—‘ǡŠ‘—ǡŽ˜ƒ”‡œǦ‘Žƒ†‘ ȋʹͲͲͺȌ ‡‡–‹ ƒ’’‹‰‘ˆ ‘š„ͳǦ ‡ŽŽ

ϵϯϰ Ž‹‡ƒ‰‡ •Š‘™• ‹‰”ƒ–‹‘ ˆ”‘ ƒ—†ƒŽ †‹‡ ‡’ŠƒŽ‘ –‘ –‡Ž‡ ‡’ŠƒŽ‘ƒ†Žƒ–‡”ƒŽ

ϵϯϱ Š›’‘–ŠƒŽƒ—•Ǥ—”‘’‡ƒ ‘—”ƒŽ‘ˆ‡—”‘• ‹‡ ‡ʹͺǣͳͻͶͳȂͳͻͷͷǤ

ϵϯϲ 

ϵϯϳ

ϯϳ 

ϵϯϴ Figure Legends

ϵϯϵ Figure 1: Early migration routes to the olfactory cortex (E10-12). ǡ˜‡–”‹ —Žƒ”œ‘‡Ǣšǡ

ϵϰϬ ‘Žˆƒ –‘”› ‘”–‡šǢ ǡ ’‹”‹ˆ‘” ‘”–‡šǢ ǡ ‘Žˆƒ –‘”› –—„‡” Ž‡Ǣ † ǡ †‘”•‘ǦŽƒ–‡”ƒŽ ‰ƒ‰Ž‹‘‹ 

ϵϰϭ ‡‹‡ ‡Ǣ˜ ǡ˜‡–”‘ǦŽƒ–‡”ƒŽ‰ƒ‰Ž‹‘‹ ‡‹‡ ‡Ǣǡ•‡’–‘Ǧ‡‹‡–‹ƒŽ•—Ž —•Ǣǡ”‘•–”‘Ǧ

ϵϰϮ ‡†‹ƒŽ –‡Ž‡ ‡’ŠƒŽ‘Ǣ ǡ ‡†‹‘Ǧ†‘”•ƒŽ –‡Ž‡ ‡’ŠƒŽ‘Ǣ ǡ ƒ—†‘Ǧ†‘”•ƒŽ –‡Ž‡ ‡’ŠƒŽ‘Ǣ

ϵϰϯ ǡ”‘•–”‘Ǧ‡†‹ƒŽ–‡Ž‡ ‡’ŠƒŽ‹ ™ƒŽŽǤ „Ž—‡ǡ•—„’ƒŽŽ‹ƒŽ‘”‹‰‹•Ǣ‹”‡†ǡ’ƒŽŽ‹ƒŽ‘”‹‰‹•Ǥ

ϵϰϰ †ƒ’–‡†ˆ”‘ ƒ” ‹ƒǦ‘”‡‘‡–ƒŽǤǡʹͲͲͺǤ

ϵϰϱ Figure 2: Comparison of developing neocortex and piriform cortex. Aǡ ƒ‹ ‹‰”ƒ–‘”›

ϵϰϲ ”‘—–‡•–‘–Š‡‡‘ ‘”–‡šƒ†–‘–Š‡’‹”‹ˆ‘” ‘”–‡šǤ‹ǡƒ–‡”ƒŽ ‘”–‹ ƒŽ •–”‡ƒ ƒ† ‘†‡• ‘ˆ

ϵϰϳ ‹‰”ƒ–‹‘–‘–Š‡’‹”‹ˆ‘” ‘”–‡šǤBǡ”‡ƒ–ƒŽ†‡˜‡Ž‘’‡–‘ˆŽƒ›‡”•‹–Š‡‡‘ ‘”–‡šƒ†‹–Š‡

ϵϰϴ ’‹”‹ˆ‘” ‘”–‡š ‹ŽŽ—•–”ƒ–‡† „› ”‡•›Ž ˜‹‘Ž‡– •–ƒ‹‹‰•Ǥ šǡ ‡‘ ‘”–‡šǢšǡ‘Žˆƒ –‘”› ‘”–‡šǢǡ

ϵϰϵ ‘Žˆƒ –‘”›„—Ž„Ǣ ‹’ǡŠ‹’’‘ ƒ’—•Ǣ ǡŽƒ–‡”ƒŽ‰ƒ‰Ž‹‘‹ ‡‹‡ ‡Ǣ ǡ‡†‹ƒŽ‰ƒ‰Ž‹‘‹‰

ϵϱϬ ‡‹‡ ‡Ǣǡ’ƒŽŽ‹—Ǧ•—„’ƒŽŽ‹—„‘—†ƒ”›Ǣǡ‡š–”ƒ ‡ŽŽ—Žƒ”ƒ–”‹šǢǡƒ”‰‹ƒŽœ‘‡Ǣǡ

ϵϱϭ ‘”–‹ ƒŽ’Žƒ–‡Ǣǡ•—„’Žƒ–‡Ǣǡ•—„˜‡–”‹ —Žƒ”œ‘‡Ǣǡ˜‡–”‹ —Žƒ”œ‘‡Ǣ ǡ‹–‡”‡†‹ƒ–‡œ‘‡Ǣ

ϵϱϮ ǡ ™Š‹–‡ ƒ––‡”Ǣ ǡ Žƒ›‡”Ǣ ǡ Žƒ–‡”ƒŽ ‘Žˆƒ –‘”› –”ƒ –Ǣ •–”ǡ •–”‹ƒ–—Ǣ ȗǡ ƒ–‡”‹‘” ‘‹••—”‡Ǥ

ϵϱϯ  ƒŽ‡„ƒ”•ǡʹͲͲρǤ

ϵϱϰ Figure 3: Piriform cortex neuron identities. Aǡ š’”‡••‹‘ ‘ˆ ’”‹ ‹’ƒŽ ‡‘ ‘”–‹ ƒŽ Žƒ›‡”

ϵϱϱ ƒ”‡”•‹–Š‡’‹”‹ˆ‘” ‘”–‡šǤIn situŠ›„”‹†‹œƒ–‹‘•ˆ”‘ŽŽ‡”ƒ‹–Žƒ•†ƒ–ƒ„ƒ•‡ȋ’‘•–ƒ–ƒŽ

ϵϱϲ †ƒ›ͷ͸ȌǤǡ ‘”’—• ƒŽŽ‘•—Ǣǡƒ–‡”‹‘” ‘‹••—”‡Ǣ–”ǡ•–”‹ƒ–—ǢǡŽƒ›‡”Ǥ ƒŽ‡„ƒ”ǡʹͲͲρǤ

ϵϱϳ Bǡ ”‰ƒ‹œƒ–‹‘ǡ ‘Ž‡ —Žƒ”‹†‡–‹–‹‡•ƒ†‘™–ƒ”‰‡–•‘ˆ’”‘Œ‡ –‹‘‡—”‘• ‹–Š‡’‹”‹ˆ‘”

ϵϱϴ ‘”–‡šǤǡŽƒ–‡”ƒŽ‘Žˆƒ –‘”›–”ƒ –Ǣ‘ǡ’‘•–‡”‘‡†‹ƒŽ ‘”–‹ ƒŽƒ›‰†ƒŽƒ— Ž‡—•ǢŽǡŽƒ–‡”ƒŽ

ϵϱϵ ‡–‘”Š‹ƒŽ ‘”–‡šǢǡ‘Žˆƒ –‘”›„—Ž„Ǣ Ǧ ǡ‹ˆ”ƒŽ‹„‹ •—„†‹˜‹•‹‘‘ˆ–Š‡‡†‹ƒŽ’”‡ˆ”‘–ƒŽ

ϵϲϬ ‘”–‡šǤ†ƒ’–‡†ˆ”‘—œ—‹Ƭ‡‡”•ǡʹͲͳͳƬ‹‘†ƒ–‘‡–ƒŽǤǡʹͲͳ͸Ǥ

ϵϲϭ 

ϯϴ Cx

OCx D Origin in VZ Migration Destination in OCx Cell identity in OCx OB APA V subpallial VZ origins Tbr1 (74.1%) dLGE radial piriform cortex Calretinin (50%) Reelin (23.5%) vz dLGE PC vLGE Calbindin (11.3%) OT vLGE radial Calretinin (1.6%) Tbr1 (1.2%)

radial Calbindin (54.5%) Tbr1 (15.5%) vz SES + olfactory tubercle Calretinin (12.6%) tangential Reelin (7.1%) SES

pallial VZ origins RDT RDT-MDT tangential olfactory tubercle Tbr1 (92.7%) vz Calretinin (36.7%) Reelin (18.8%) CDT tangential piriform cortex

tangential dorso-lateral route RMTW in neocortex piriform cortex vz RMTW Calretinin (50%) ventro-medial route Tbr1 (10%) in medial wall olfactory tubercle

Figure 1: Early migration routes to the olfactory cortex (E10-12). VZ, ventricular zone; OCx, olfactory cortex; PC, piriform cortex; OT, olfactory tubercle; dLGE, dorso-lateral ganglionic eminence; vLGE, ventro-lateral ganglionic eminence; SES, septo-eminential sulcus; RDT, rostro-medial telencephalon; MDT, medio-dorsal telen- cephalon; CDT, caudo-dorsal telencephalon; RMTW, rostro-medial telencephalic wall. In blue, subpallial VZ origins; in red, pallial VZ origins. Adapted from Garcia-Moreno et al., 2008. A B Neocortex Cx P0 Hip * MZ Cx LGE CP MGE OCx E16.5 L5 OB i MZ D L6 AP E14.5 CP V MZ PSB SP SP * lateral cortical stream CP IZ interneuron migration SP IZ excitatory neuron migration SVZ/ SVZ/ VZ VZ i WM

Piriform cortex LGE P0

MGE LOT radial glial cell E16.5 L1 radial migration LOT L2 in chain migration E14.5 L1 pial surface L2 LOT OC ECM L3 ? L3

WM * WM WM MGE str * str *

Figure 2: Comparison of developing neocortex and piriform cortex. A, Main migratory routes to the neocortex and to the piriform cortex. i, Lateral cortical stream and modes of migration to the piriform cortex. B, Prenatal development of layers in the neocortex and in the piriform cortex illus- trated by cresyl violet stainings. Cx, neocortex; OCx, olfactory cortex; OB, ; Hip, ; LGE, lateral ganglioninc eminence; MGE, medial ganglioning eminence; PSB, palli- um-subpallium boundary; ECM, extracellular matrix; MZ, marginal zone; CP, cortical plate; SP, subplate; SVZ, subventricular zone; VZ, ventricular zone; IZ, intermediate zone; WM, ; L, layer; LOT, lateral ; str, ; *, . Scale bars, 200 mm. A Neocortex

CC Str

AC Piriform cortex

Cux1 Cux2 Satb2 Rorb Kcnip2 Er81Ctip2 Fezf2 Sema3e Foxp2 Tbr1 Reelin L1

L2/3

L4

L5 Neocortex

L6

L1a L1b L2a L2b

L3 Piriform cortex

B Superficial Deep Semilunar cells pyramidal cells pyramidal cells L1a LOT

L1b

L2a Reelin+ Fezf2+ - &ඝච Cux1+ Cux1+ Ctip2+ L2b Ctip2- Ctip2+ Cux1-

Cux1+

L3

CoA OBOB IL-mPFC OB lENT & IL-mPFC Figure 3: Piriform cortex neuron identities. A, Expression of principal neocortical layer markers in the piriform cortex. In situ hybridizations from Allen Atlas database (postnatal day 56). CC, ; AC, anterior commissure; Str, striatum; L, layer. Scale bar, 200 mm. B, Organiza- tion, molecular identities and known targets of projection neurons in the piriform cortex. LOT, lateral olfactory tract; CoA, posteromedial cortical nucleus; lENT, lateral ; OB, olfactory bulb; IL-mP- FC, infralimbic subdivision of the medial prefrontal cortex. Adapted from Suzuki & Bekkers, 2011 & Diodato et al., 2016.