This Accepted Manuscript has not been copyedited and formatted. The final version may differ from this version.
Research Article: New Research | Integrative Systems
Long-lasting visuo-vestibular mismatch in freely-behaving mouse reduces the
vestibulo-ocular reflex and leads to neural changes in the direct vestibular pathway
Neural changes following long-term VOR reduction
Julie Carcaud1, Filipa França de Barros1, Erwin Idoux1, Daniel Eugène1, Lionel Reveret2, Lee E. Moore3, PierrePaul Vidal3 and Mathieu Beraneck1,*
1
Center for Neurophysics, Physiology, Pathology, CNRS UMR 8119, Université Paris Descartes, Sorbonne
Paris Cité, Paris, France
2
INRIA Grenoble, Rhône-Alpes, LJK, UMR 5224, France
3
Cognition and Action Group, CNRS UMR 8257, Université Paris Descartes, Sorbonne Paris Cité, Paris, France
DOI: 10.1523/ENEURO.0290-16.2017
Received: 26 September 2016
Revised: 5 January 2017
Accepted: 10 January 2017
Published: 16 January 2017
Author contributions: J.C. and M.B. designed research; J.C., F.F.D.B., E.I., D.E., L.R., and M.B. performed
research; J.C., F.F.D.B., E.I., L.R., L.E.M., and M.B. analyzed data; J.C., F.F.D.B., E.I., D.E., L.E.M., P.-P.V.,
and M.B. wrote the paper.
Funding: Centre National d'Etudes Spatiales (CNES)
501100002830
DAR n{degree sign} 4800000860
Funding: Centre National de la Recherche Scientifique (CNRS)
501100004794
Funding: Agence Nationale pour la Recherche (ANR)
ANR-13-CESA-0005-02
Funding: Université Paris Descartes
Funding: Agence National pour le Recherche (ANR)
ANR-15-CE32-0007
The authors declare no competing financial interests.
This research was supported by the Centre National de la Recherche Scientifique and the University Paris
Descartes. EI and MB receive support from the Centre National des Etudes Spatiales. JC and MB receive
support from the French ANR-13-CESA-0005-02. FFB and MB receive support from the French ANR-15CE32-0007.
*Corresponding author: M. Beraneck. Center for Neurophysics, Physiology, Pathology, CNRS UMR 8119, 45 rue
des St-Pères 75270, Université Paris Descartes, Paris, France. E-mail: [email protected]
Cite as: eNeuro 2017; 10.1523/ENEURO.0290-16.2017
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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
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ŶŐĞůĂŬŝ ;ϮϬϬϰͿ LJĞƐ ŽŶ ƚĂƌŐĞƚ͗ ǁŚĂƚ ŶĞƵƌŽŶƐ ŵƵƐƚ ĚŽ ĨŽƌ ƚŚĞ ǀĞƐƚŝďƵůŽŽĐƵůĂƌ ƌĞĨůĞdž ĚƵƌŝŶŐ ůŝŶĞĂƌ
ŵŽƚŝŽŶ͘:EĞƵƌŽƉŚLJƐŝŽůϵϮ͗ϮϬͲϯϱ͘
ŶnjĂŝ D͕ <ŝƚĂnjĂǁĂ ,͕ EĂŐĂŽ ^ ;ϮϬϭϬͿ ĨĨĞĐƚƐ ŽĨ ƌĞǀĞƌƐŝďůĞ ƉŚĂƌŵĂĐŽůŽŐŝĐĂů ƐŚƵƚĚŽǁŶ ŽĨ ĐĞƌĞďĞůůĂƌ
ĨůŽĐĐƵůƵƐŽŶƚŚĞŵĞŵŽƌLJŽĨůŽŶŐͲƚĞƌŵŚŽƌŝnjŽŶƚĂůǀĞƐƚŝďƵůŽͲŽĐƵůĂƌƌĞĨůĞdžĂĚĂƉƚĂƚŝŽŶŝŶŵŽŶŬĞLJƐ͘
EĞƵƌŽƐĐŝZĞƐϲϴ͗ϭϵϭͲϭϵϴ͘
ĂďĂůŝĂŶ͕sŝďĞƌƚE͕ƐƐŝĞ'͕^ĞƌĂĨŝŶD͕DƵŚůĞƚŚĂůĞƌD͕sŝĚĂůWW;ϭϵϵϳͿĞŶƚƌĂůǀĞƐƚŝďƵůĂƌŶĞƚǁŽƌŬƐŝŶ
ƚŚĞ ŐƵŝŶĞĂͲƉŝŐ͗ ĨƵŶĐƚŝŽŶĂů ĐŚĂƌĂĐƚĞƌŝnjĂƚŝŽŶ ŝŶ ƚŚĞ ŝƐŽůĂƚĞĚ ǁŚŽůĞ ďƌĂŝŶ ŝŶ ǀŝƚƌŽ͘ EĞƵƌŽƐĐŝĞŶĐĞ
ϴϭ͗ϰϬϱͲϰϮϲ͘
ĂďĂůŝĂŶ >͕ sŝĚĂů WW ;ϮϬϬϬͿ &ůŽĐĐƵůĂƌ ŵŽĚƵůĂƚŝŽŶ ŽĨ ǀĞƐƚŝďƵůŽŽĐƵůĂƌ ƉĂƚŚǁĂLJƐ ĂŶĚ ĐĞƌĞďĞůůƵŵͲƌĞůĂƚĞĚ
ƉůĂƐƚŝĐŝƚLJ͗ŶŝŶǀŝƚƌŽǁŚŽůĞďƌĂŝŶƐƚƵĚLJ͘:EĞƵƌŽƉŚLJƐŝŽůϴϰ͗ϮϱϭϰͲϮϱϮϴ͘
ĂŐŶĂůů Dt͕ ^ƚĞǀĞŶƐ Z:͕ ĚƵ >ĂĐ ^ ;ϮϬϬϳͿ dƌĂŶƐŐĞŶŝĐ ŵŽƵƐĞ ůŝŶĞƐ ƐƵďĚŝǀŝĚĞ ŵĞĚŝĂů ǀĞƐƚŝďƵůĂƌ ŶƵĐůĞƵƐ
ŶĞƵƌŽŶƐŝŶƚŽĚŝƐĐƌĞƚĞ͕ŶĞƵƌŽĐŚĞŵŝĐĂůůLJĚŝƐƚŝŶĐƚƉŽƉƵůĂƚŝŽŶƐ͘:EĞƵƌŽƐĐŝϮϳ͗ϮϯϭϴͲϮϯϯϬ͘
ĂŬĞƌZ͕ǀŝŶŐĞƌ͕DĐƌĞĂZ;ϭϵϴϭͿ^ŽŵĞƚŚŽƵŐŚƚƐĂďŽƵƚƚŚĞƚŚƌĞĞŶĞƵƌŽŶƐŝŶƚŚĞǀĞƐƚŝďƵůĂƌŽĐƵůĂƌ
ƌĞĨůĞdž͘ŶŶEzĐĂĚ^Đŝϯϳϰ͗ϭϳϭͲϭϴϴ͘
ĞƌĂŶĞĐŬD͕/ĚŽƵdž;ϮϬϭϮͿZĞĐŽŶƐŝĚĞƌŝŶŐƚŚĞƌŽůĞŽĨŶĞƵƌŽŶĂůŝŶƚƌŝŶƐŝĐƉƌŽƉĞƌƚŝĞƐĂŶĚŶĞƵƌŽŵŽĚƵůĂƚŝŽŶ
ŝŶǀĞƐƚŝďƵůĂƌŚŽŵĞŽƐƚĂƐŝƐ͘&ƌŽŶƚEĞƵƌŽůϯ͗Ϯϱ͘
ĞƌĂŶĞĐŬ D͕ DĐ<ĞĞ :>͕ ůĞŝƐĂ D͕ ƵůůĞŶ < ;ϮϬϬϴͿ ƐLJŵŵĞƚƌŝĐ ƌĞĐŽǀĞƌLJ ŝŶ ĐĞƌĞďĞůůĂƌͲĚĞĨŝĐŝĞŶƚ ŵŝĐĞ
ĨŽůůŽǁŝŶŐƵŶŝůĂƚĞƌĂůůĂďLJƌŝŶƚŚĞĐƚŽŵLJ͘:EĞƵƌŽƉŚLJƐŝŽůϭϬϬ͗ϵϰϱͲϵϱϴ͘
ĞƌĂŶĞĐŬD͕/ĚŽƵdž͕hŶŽ͕sŝĚĂůWW͕DŽŽƌĞ>͕sŝďĞƌƚE;ϮϬϬϰͿhŶŝůĂƚĞƌĂůůĂďLJƌŝŶƚŚĞĐƚŽŵLJŵŽĚŝĨŝĞƐƚŚĞ
ŵĞŵďƌĂŶĞƉƌŽƉĞƌƚŝĞƐŽĨĐŽŶƚƌĂůĞƐŝŽŶĂůǀĞƐƚŝďƵůĂƌŶĞƵƌŽŶƐ͘:EĞƵƌŽƉŚLJƐŝŽůϵϮ͗ϭϲϲϴͲϭϲϴϰ͘
ĞƌĂŶĞĐŬD͕,ĂĐŚĞŵĂŽƵŝD͕/ĚŽƵdž͕ZŝƐ>͕hŶŽ͕'ŽĚĂƵdž͕sŝĚĂůWW͕DŽŽƌĞ>͕sŝďĞƌƚE;ϮϬϬϯͿ>ŽŶŐͲ
ƚĞƌŵƉůĂƐƚŝĐŝƚLJŽĨŝƉƐŝůĞƐŝŽŶĂůŵĞĚŝĂůǀĞƐƚŝďƵůĂƌŶƵĐůĞƵƐŶĞƵƌŽŶƐĂĨƚĞƌƵŶŝůĂƚĞƌĂůůĂďLJƌŝŶƚŚĞĐƚŽŵLJ͘
:EĞƵƌŽƉŚLJƐŝŽůϵϬ͗ϭϴϰͲϮϬϯ͘
ĞƌƚŚŽnj ͕ :ŽŶĞƐ 'D͕ ĞŐƵĞ ;ϭϵϴϭͿ ŝĨĨĞƌĞŶƚŝĂů ǀŝƐƵĂů ĂĚĂƉƚĂƚŝŽŶ ŽĨ ǀĞƌƚŝĐĂů ĐĂŶĂůͲĚĞƉĞŶĚĞŶƚ
ǀĞƐƚŝďƵůŽͲŽĐƵůĂƌƌĞĨůĞdžĞƐ͘džƉƌĂŝŶZĞƐϰϰ͗ϭϵͲϮϲ͘
ŝĞƐĚŽƌĨ ^͕ DĂůŝŶǀĂƵĚ ͕ ZĞŝĐŚĞŶďĞƌŐĞƌ /͕ WĨĂŶnjĞůƚ ^͕ ^ƚƌĂŬĂ , ;ϮϬϬϴͿ ŝĨĨĞƌĞŶƚŝĂů ŝŶŚŝďŝƚŽƌLJ ĐŽŶƚƌŽů ŽĨ
ƐĞŵŝĐŝƌĐƵůĂƌ ĐĂŶĂů ŶĞƌǀĞ ĂĨĨĞƌĞŶƚͲĞǀŽŬĞĚ ŝŶƉƵƚƐ ŝŶ ƐĞĐŽŶĚͲŽƌĚĞƌ ǀĞƐƚŝďƵůĂƌ ŶĞƵƌŽŶƐ ďLJ
ŐůLJĐŝŶĞƌŐŝĐĂŶĚ'ĞƌŐŝĐĐŝƌĐƵŝƚƐ͘:EĞƵƌŽƉŚLJƐŝŽůϵϵ͗ϭϳϱϴͲϭϳϲϵ͘
ůĂnjƋƵĞnj WD͕ ,ŝƌĂƚĂ z͕ ,ŝŐŚƐƚĞŝŶ ^D ;ϮϬϬϰͿ dŚĞ ǀĞƐƚŝďƵůŽͲŽĐƵůĂƌ ƌĞĨůĞdž ĂƐ Ă ŵŽĚĞů ƐLJƐƚĞŵ ĨŽƌ ŵŽƚŽƌ
ůĞĂƌŶŝŶŐ͗ǁŚĂƚŝƐƚŚĞƌŽůĞŽĨƚŚĞĐĞƌĞďĞůůƵŵ͍ĞƌĞďĞůůƵŵϯ͗ϭϴϴͲϭϵϮ͘
ŽLJĚĞŶ ^͕ ZĂLJŵŽŶĚ :> ;ϮϬϬϯͿ ĐƚŝǀĞ ƌĞǀĞƌƐĂů ŽĨ ŵŽƚŽƌ ŵĞŵŽƌŝĞƐ ƌĞǀĞĂůƐ ƌƵůĞƐ ŐŽǀĞƌŶŝŶŐ ŵĞŵŽƌLJ
ĞŶĐŽĚŝŶŐ͘EĞƵƌŽŶϯϵ͗ϭϬϯϭͲϭϬϰϮ͘
ŽLJĚĞŶ ^͕ <ĂƚŽŚ ͕ ZĂLJŵŽŶĚ :> ;ϮϬϬϰͿ ĞƌĞďĞůůƵŵͲĚĞƉĞŶĚĞŶƚ ůĞĂƌŶŝŶŐ͗ ƚŚĞ ƌŽůĞ ŽĨ ŵƵůƚŝƉůĞ ƉůĂƐƚŝĐŝƚLJ
ŵĞĐŚĂŶŝƐŵƐ͘ŶŶƵZĞǀEĞƵƌŽƐĐŝϮϳ͗ϱϴϭͲϲϬϵ͘
ƌŽƵƐƐĂƌĚ D͕ >ŝƐďĞƌŐĞƌ ^' ;ϭϵϵϮͿ sĞƐƚŝďƵůĂƌ ŝŶƉƵƚƐ ƚŽ ďƌĂŝŶ ƐƚĞŵ ŶĞƵƌŽŶƐ ƚŚĂƚ ƉĂƌƚŝĐŝƉĂƚĞ ŝŶ ŵŽƚŽƌ
ůĞĂƌŶŝŶŐŝŶƚŚĞƉƌŝŵĂƚĞǀĞƐƚŝďƵůŽŽĐƵůĂƌƌĞĨůĞdž͘:EĞƵƌŽƉŚLJƐŝŽůϲϴ͗ϭϵϬϲͲϭϵϬϵ͘
ƌŽƵƐƐĂƌĚD͕<ĂƐƐĂƌĚũŝĂŶ;ϮϬϬϰͿ>ĞĂƌŶŝŶŐŝŶĂƐŝŵƉůĞŵŽƚŽƌƐLJƐƚĞŵ͘>ĞĂƌŶDĞŵϭϭ͗ϭϮϳͲϭϯϲ͘
ƌŽƵƐƐĂƌĚ D͕ dŝƚůĞLJ ,<͕ ŶƚĨůŝĐŬ :͕ ,ĂŵƉƐŽŶ Z ;ϮϬϭϭͿ DŽƚŽƌ ůĞĂƌŶŝŶŐ ŝŶ ƚŚĞ sKZ͗ ƚŚĞ ĐĞƌĞďĞůůĂƌ
ĐŽŵƉŽŶĞŶƚ͘džƉƌĂŝŶZĞƐϮϭϬ͗ϰϱϭͲϰϲϯ͘
ĂůĂďƌĞƐĞ Z͕ ,ƵůůĂƌ d ;ϮϬϬϲͿ WůĂŶĂƌ ƌĞůĂƚŝŽŶƐŚŝƉƐ ŽĨ ƚŚĞ ƐĞŵŝĐŝƌĐƵůĂƌ ĐĂŶĂůƐ ŝŶ ƚǁŽ ƐƚƌĂŝŶƐ ŽĨ ŵŝĐĞ͘ :
ƐƐŽĐZĞƐKƚŽůĂƌLJŶŐŽůϳ͗ϭϱϭͲϭϱϵ͘
ĂŵƉ:͕ĂůůŝƐƚĞƌZ:͕ƌŝĐŚƚĂD;ϮϬϬϲͿ/ŶŚŝďŝƚŽƌLJƐLJŶĂƉƚŝĐƚƌĂŶƐŵŝƐƐŝŽŶĚŝĨĨĞƌƐŝŶŵŽƵƐĞƚLJƉĞĂŶĚ
ŵĞĚŝĂůǀĞƐƚŝďƵůĂƌŶƵĐůĞƵƐŶĞƵƌŽŶƐŝŶǀŝƚƌŽ͘:EĞƵƌŽƉŚLJƐŝŽůϵϱ͗ϯϮϬϴͲϯϮϭϴ͘
ĂƌĞLJDZ;ϮϬϭϭͿ^LJŶĂƉƚŝĐŵĞĐŚĂŶŝƐŵƐŽĨƐĞŶƐŽƌŝŵŽƚŽƌůĞĂƌŶŝŶŐŝŶƚŚĞĐĞƌĞďĞůůƵŵ͘ƵƌƌKƉŝŶEĞƵƌŽďŝŽů
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ĂĚĂƉƚĂƚŝŽŶŝŶǁŝůĚͲƚLJƉĞĂŶĚŵƵƚĂŶƚŵŝĐĞ͘:EĞƵƌŽƐĐŝϯϰ͗ϳϮϬϯͲϳϮϭϱ͘
ƵůůĞŶ < ;ϮϬϭϮͿ dŚĞ ǀĞƐƚŝďƵůĂƌ ƐLJƐƚĞŵ͗ ŵƵůƚŝŵŽĚĂů ŝŶƚĞŐƌĂƚŝŽŶ ĂŶĚ ĞŶĐŽĚŝŶŐ ŽĨ ƐĞůĨͲŵŽƚŝŽŶ ĨŽƌ ŵŽƚŽƌ
ĐŽŶƚƌŽů͘dƌĞŶĚƐEĞƵƌŽƐĐŝϯϱ͗ϭϴϱͲϭϵϲ͘
ƵƌƚŚŽLJƐ/^;ϮϬϬϬͿsĞƐƚŝďƵůĂƌĐŽŵƉĞŶƐĂƚŝŽŶĂŶĚƐƵďƐƚŝƚƵƚŝŽŶ͘ƵƌƌKƉŝŶEĞƵƌŽůϭϯ͗ϮϳͲϯϬ͘
ĞĞĞƵǁ/͕dĞŶƌŝŶŬĞDD;ϮϬϭϱͿDŽƚŽƌ>ĞĂƌŶŝŶŐĂŶĚƚŚĞĞƌĞďĞůůƵŵ͘ŽůĚ^ƉƌŝŶŐ,ĂƌďWĞƌƐƉĞĐƚŝŽů
ϳ͗ĂϬϮϭϲϴϯ͘
ŝĞƚƌŝĐŚ ,͕ ^ƚƌĂŬĂ , ;ϮϬϭϲͿ WƌŽůŽŶŐĞĚ ǀĞƐƚŝďƵůĂƌ ƐƚŝŵƵůĂƚŝŽŶ ŝŶĚƵĐĞƐ ŚŽŵĞŽƐƚĂƚŝĐ ƉůĂƐƚŝĐŝƚLJ ŽĨ ƚŚĞ
ǀĞƐƚŝďƵůŽͲŽĐƵůĂƌƌĞĨůĞdžŝŶůĂƌǀĂůyĞŶŽƉƵƐůĂĞǀŝƐ͘Ƶƌ:EĞƵƌŽƐĐŝϰϰ͗ϭϳϴϳͲϭϳϵϲ͘
ĚƵ >ĂĐ ^ ;ϭϵϵϲͿ ĂŶĚŝĚĂƚĞ ĐĞůůƵůĂƌ ŵĞĐŚĂŶŝƐŵƐ ŽĨ ǀĞƐƚŝďƵůŽͲŽĐƵůĂƌ ƌĞĨůĞdž ƉůĂƐƚŝĐŝƚLJ͘ ŶŶ E z ĐĂĚ ^Đŝ
ϳϴϭ͗ϰϴϵͲϰϵϴ͘
ĚƵ >ĂĐ ^͕ ZĂLJŵŽŶĚ :>͕ ^ĞũŶŽǁƐŬŝ d:͕ >ŝƐďĞƌŐĞƌ ^' ;ϭϵϵϱͿ >ĞĂƌŶŝŶŐ ĂŶĚ ŵĞŵŽƌLJ ŝŶ ƚŚĞ ǀĞƐƚŝďƵůŽͲŽĐƵůĂƌ
ƌĞĨůĞdž͘ŶŶƵZĞǀEĞƵƌŽƐĐŝϭϴ͗ϰϬϵͲϰϰϭ͘
ƵƚŝĂD͕:ŽŚŶƐƚŽŶZ͕DĐYƵĞĞŶ^;ϭϵϵϮͿdŽŶŝĐĂĐƚŝǀŝƚLJŽĨƌĂƚŵĞĚŝĂůǀĞƐƚŝďƵůĂƌŶƵĐůĞƵƐŶĞƵƌŽŶĞƐŝŶ
ǀŝƚƌŽĂŶĚŝƚƐŝŶŚŝďŝƚŝŽŶďLJ'͘džƉƌĂŝŶZĞƐϴϴ͗ϰϲϲͲϰϳϮ͘
ƵƚŝĂ D͕ >ŽƚƚŽ Z͕ :ŽŚŶƐƚŽŶ Z ;ϭϵϵϱͿ WŽƐƚͲŶĂƚĂů ĚĞǀĞůŽƉŵĞŶƚ ŽĨ ƚŽŶŝĐ ĂĐƚŝǀŝƚLJ ĂŶĚ ŵĞŵďƌĂŶĞ
ĞdžĐŝƚĂďŝůŝƚLJŝŶŵŽƵƐĞŵĞĚŝĂůǀĞƐƚŝďƵůĂƌŶƵĐůĞƵƐŶĞƵƌŽŶĞƐ͘ĐƚĂ KƚŽůĂƌLJŶŐŽů^ƵƉƉůϱϮϬWƚϭ͗ϭϬϭͲ
ϭϬϰ͘
ƵŐĞŶĞ ͕ /ĚŽƵdž ͕ ĞƌĂŶĞĐŬ D͕ DŽŽƌĞ >͕ sŝĚĂů WW ;ϮϬϭϭͿ /ŶƚƌŝŶƐŝĐ ŵĞŵďƌĂŶĞ ƉƌŽƉĞƌƚŝĞƐ ŽĨ ĐĞŶƚƌĂů
ǀĞƐƚŝďƵůĂƌŶĞƵƌŽŶƐŝŶƌŽĚĞŶƚƐ͘džƉƌĂŝŶZĞƐϮϭϬ͗ϰϮϯͲϰϯϲ͘
&ĂƵůƐƚŝĐŚD͕ǀĂŶůƉŚĞŶD͕>ƵŽ͕ĚƵ>ĂĐ^͕ĞĞĞƵǁ/;ϮϬϬϲͿKĐƵůŽŵŽƚŽƌƉůĂƐƚŝĐŝƚLJĚƵƌŝŶŐǀĞƐƚŝďƵůĂƌ
ĐŽŵƉĞŶƐĂƚŝŽŶĚŽĞƐŶŽƚĚĞƉĞŶĚŽŶĐĞƌĞďĞůůĂƌ>d͘:EĞƵƌŽƉŚLJƐŝŽůϵϲ͗ϭϭϴϳͲϭϭϵϱ͘
'ŝƚƚŝƐ,͕ĚƵ>ĂĐ^;ϮϬϬϲͿ/ŶƚƌŝŶƐŝĐĂŶĚƐLJŶĂƉƚŝĐƉůĂƐƚŝĐŝƚLJŝŶƚŚĞǀĞƐƚŝďƵůĂƌƐLJƐƚĞŵ͘ƵƌƌKƉŝŶEĞƵƌŽďŝŽů
ϭϲ͗ϯϴϱͲϯϵϬ͘
'ƌĂƐƐŝ ^͕ WĞƚƚŽƌŽƐƐŝ s ;ϮϬϬϭͿ ^LJŶĂƉƚŝĐ ƉůĂƐƚŝĐŝƚLJ ŝŶ ƚŚĞ ŵĞĚŝĂů ǀĞƐƚŝďƵůĂƌ ŶƵĐůĞŝ͗ ƌŽůĞ ŽĨ ŐůƵƚĂŵĂƚĞ
ƌĞĐĞƉƚŽƌƐĂŶĚƌĞƚƌŽŐƌĂĚĞŵĞƐƐĞŶŐĞƌƐŝŶƌĂƚďƌĂŝŶƐƚĞŵƐůŝĐĞƐ͘WƌŽŐEĞƵƌŽďŝŽůϲϰ͗ϱϮϳͲϱϱϯ͘
,ĂŶƐĞů ͕ >ŝŶĚĞŶ :͕ ΖŶŐĞůŽ ;ϮϬϬϭͿ ĞLJŽŶĚ ƉĂƌĂůůĞů ĨŝďĞƌ >d͗ ƚŚĞ ĚŝǀĞƌƐŝƚLJ ŽĨ ƐLJŶĂƉƚŝĐ ĂŶĚ ŶŽŶͲ
ƐLJŶĂƉƚŝĐƉůĂƐƚŝĐŝƚLJŝŶƚŚĞĐĞƌĞďĞůůƵŵ͘EĂƚEĞƵƌŽƐĐŝϰ͗ϰϲϳͲϰϳϱ͘
,ŝŵ ͕ ƵƚŝĂ D ;ϮϬϬϭͿ /ŶƚƌŝŶƐŝĐ ĞdžĐŝƚĂďŝůŝƚLJ ĐŚĂŶŐĞƐ ŝŶ ǀĞƐƚŝďƵůĂƌ ŶƵĐůĞƵƐ ŶĞƵƌŽŶƐ ĂĨƚĞƌ ƵŶŝůĂƚĞƌĂů
ĚĞĂĨĨĞƌĞŶƚĂƚŝŽŶ͘ƌĂŝŶZĞƐϵϬϴ͗ϱϴͲϲϲ͘
/ĚŽƵdž;ϮϬϭϱͿsĞƐƚŝďƵůĂƌƉůĂƐƚŝĐŝƚLJ͘/Ŷ͗dŚĞƌĂƚŶĞƌǀŽƵƐƐLJƐƚĞŵ͕ϰƚŚĚŝƚŝŽŶ;WĂdžŝŶŽƐ'͕ĞĚͿ͕ƉƉϴϯϳͲϴϰϮ͗
ůƐĞǀŝĞƌ͘
/ƚŽD;ϭϵϴϮͿĞƌĞďĞůůĂƌĐŽŶƚƌŽůŽĨƚŚĞǀĞƐƚŝďƵůŽͲŽĐƵůĂƌƌĞĨůĞdžͲͲĂƌŽƵŶĚƚŚĞĨůŽĐĐƵůƵƐŚLJƉŽƚŚĞƐŝƐ͘ŶŶƵZĞǀ
EĞƵƌŽƐĐŝϱ͗ϮϳϱͲϮϵϲ͘
<ĂƐƐĂƌĚũŝĂŶ ͕ dĂŶ z&͕ ŚƵŶŐ :z͕ ,ĞƐŬŝŶ Z͕ WĞƚĞƌƐŽŶ D:͕ ƌŽƵƐƐĂƌĚ D ;ϮϬϬϱͿ dŚĞ ƐŝƚĞ ŽĨ Ă ŵŽƚŽƌ
ŵĞŵŽƌLJƐŚŝĨƚƐǁŝƚŚĐŽŶƐŽůŝĚĂƚŝŽŶ͘:EĞƵƌŽƐĐŝϮϱ͗ϳϵϳϵͲϳϵϴϱ͘
<ĞůůĞƌ >͕ WƌĞĐŚƚ t ;ϭϵϳϵͿ DŽĚŝĨŝĐĂƚŝŽŶ ŽĨ ĐĞŶƚƌĂů ǀĞƐƚŝďƵůĂƌ ŶĞƵƌŽŶ ƌĞƐƉŽŶƐĞ ďLJ ĐŽŶĨůŝĐƚŝŶŐ ǀŝƐƵĂůͲ
ǀĞƐƚŝďƵůĂƌƐƚŝŵƵůĂƚŝŽŶ͘WƌŽŐƌĂŝŶZĞƐϱϬ͗ϳϲϯͲϳϳϬ͘
<ůĞŝŵ:͕&ƌĞĞŵĂŶ:,͕:ƌ͕͘ƌƵŶĞĂƵZ͕EŽůĂŶ͕ŽŽƉĞƌEZ͕ŽŽŬ͕tĂůƚĞƌƐ;ϮϬϬϮͿ^LJŶĂƉƐĞĨŽƌŵĂƚŝŽŶ
ŝƐĂƐƐŽĐŝĂƚĞĚǁŝƚŚŵĞŵŽƌLJƐƚŽƌĂŐĞŝŶƚŚĞĐĞƌĞďĞůůƵŵ͘WƌŽĐEĂƚůĐĂĚ^Đŝh^ϵϵ͗ϭϯϮϮϴͲϭϯϮϯϭ͘
<ŽĚĂŵĂ d͕ 'ƵĞƌƌĞƌŽ ^͕ ^ŚŝŶ D͕ DŽŐŚĂĚĂŵ ^͕ &ĂƵůƐƚŝĐŚ D͕ ĚƵ >ĂĐ ^ ;ϮϬϭϮͿ EĞƵƌŽŶĂů ĐůĂƐƐŝĨŝĐĂƚŝŽŶ ĂŶĚ
ŵĂƌŬĞƌ ŐĞŶĞ ŝĚĞŶƚŝĨŝĐĂƚŝŽŶ ǀŝĂ ƐŝŶŐůĞͲĐĞůů ĞdžƉƌĞƐƐŝŽŶ ƉƌŽĨŝůŝŶŐ ŽĨ ďƌĂŝŶƐƚĞŵ ǀĞƐƚŝďƵůĂƌ ŶĞƵƌŽŶƐ
ƐƵďƐĞƌǀŝŶŐĐĞƌĞďĞůůĂƌůĞĂƌŶŝŶŐ͘:EĞƵƌŽƐĐŝϯϮ͗ϳϴϭϵͲϳϴϯϭ͘
>ŝƐďĞƌŐĞƌ^';ϭϵϴϴͿdŚĞŶĞƵƌĂůďĂƐŝƐĨŽƌůĞĂƌŶŝŶŐŽĨƐŝŵƉůĞŵŽƚŽƌƐŬŝůůƐ͘^ĐŝĞŶĐĞϮϰϮ͗ϳϮϴͲϳϯϱ͘
>ŝƐďĞƌŐĞƌ^'͕DŝůĞƐ&͕KƉƚŝĐĂŶ>D;ϭϵϴϯͿ&ƌĞƋƵĞŶĐLJͲƐĞůĞĐƚŝǀĞĂĚĂƉƚĂƚŝŽŶ͗ĞǀŝĚĞŶĐĞĨŽƌĐŚĂŶŶĞůƐŝŶƚŚĞ
ǀĞƐƚŝďƵůŽͲŽĐƵůĂƌƌĞĨůĞdž͍:EĞƵƌŽƐĐŝϯ͗ϭϮϯϰͲϭϮϰϰ͘
>ŝƐďĞƌŐĞƌ^'͕DŝůĞƐ&͕ĞĞ^;ϭϵϴϰͿ^ŝŐŶĂůƐƵƐĞĚƚŽĐŽŵƉƵƚĞĞƌƌŽƌƐŝŶŵŽŶŬĞLJǀĞƐƚŝďƵůŽŽĐƵůĂƌƌĞĨůĞdž͗
ƉŽƐƐŝďůĞƌŽůĞŽĨĨůŽĐĐƵůƵƐ͘:EĞƵƌŽƉŚLJƐŝŽůϱϮ͗ϭϭϰϬͲϭϭϱϯ͘
ϯϬ
ϲϴϴ
ϲϴϵ
ϲϵϬ
ϲϵϭ
ϲϵϮ
ϲϵϯ
ϲϵϰ
ϲϵϱ
ϲϵϲ
ϲϵϳ
ϲϵϴ
ϲϵϵ
ϳϬϬ
ϳϬϭ
ϳϬϮ
ϳϬϯ
ϳϬϰ
ϳϬϱ
ϳϬϲ
ϳϬϳ
ϳϬϴ
ϳϬϵ
ϳϭϬ
ϳϭϭ
ϳϭϮ
ϳϭϯ
ϳϭϰ
ϳϭϱ
ϳϭϲ
ϳϭϳ
ϳϭϴ
ϳϭϵ
ϳϮϬ
ϳϮϭ
ϳϮϮ
ϳϮϯ
ϳϮϰ
ϳϮϱ
ϳϮϲ
ϳϮϳ
ϳϮϴ
ϳϮϵ
ϳϯϬ
ϳϯϭ
ϳϯϮ
ϳϯϯ
ϳϯϰ
>ŝƐďĞƌŐĞƌ ^'͕ WĂǀĞůŬŽ d͕ ƌŽƵƐƐĂƌĚ D ;ϭϵϵϰͿ EĞƵƌĂů ďĂƐŝƐ ĨŽƌ ŵŽƚŽƌ ůĞĂƌŶŝŶŐ ŝŶ ƚŚĞ ǀĞƐƚŝďƵůŽŽĐƵůĂƌ
ƌĞĨůĞdž ŽĨ ƉƌŝŵĂƚĞƐ͘ /͘ ŚĂŶŐĞƐ ŝŶ ƚŚĞ ƌĞƐƉŽŶƐĞƐ ŽĨ ďƌĂŝŶ ƐƚĞŵ ŶĞƵƌŽŶƐ͘ : EĞƵƌŽƉŚLJƐŝŽů ϳϮ͗ϵϮϴͲ
ϵϱϯ͘
>ŝƐďĞƌŐĞƌ ^'͕ DŝůĞƐ &͕ KƉƚŝĐĂŶ >D͕ ŝŐŚŵLJ ;ϭϵϴϭͿ KƉƚŽŬŝŶĞƚŝĐ ƌĞƐƉŽŶƐĞ ŝŶ ŵŽŶŬĞLJ͗ ƵŶĚĞƌůLJŝŶŐ
ŵĞĐŚĂŶŝƐŵƐ ĂŶĚ ƚŚĞŝƌ ƐĞŶƐŝƚŝǀŝƚLJ ƚŽ ůŽŶŐͲƚĞƌŵ ĂĚĂƉƚŝǀĞ ĐŚĂŶŐĞƐ ŝŶ ǀĞƐƚŝďƵůŽŽĐƵůĂƌ ƌĞĨůĞdž͘ :
EĞƵƌŽƉŚLJƐŝŽůϰϱ͗ϴϲϵͲϴϵϬ͘
>ƵĞďŬĞ ͕ ZŽďŝŶƐŽŶ ;ϭϵϵϰͿ 'ĂŝŶ ĐŚĂŶŐĞƐ ŽĨ ƚŚĞ ĐĂƚΖƐ ǀĞƐƚŝďƵůŽͲŽĐƵůĂƌ ƌĞĨůĞdž ĂĨƚĞƌ ĨůŽĐĐƵůƵƐ
ĚĞĂĐƚŝǀĂƚŝŽŶ͘džƉƌĂŝŶZĞƐϵϴ͗ϯϳϵͲϯϵϬ͘
DĂůŝŶǀĂƵĚ͕sĂƐƐŝĂƐ/͕ZĞŝĐŚĞŶďĞƌŐĞƌ/͕ZŽƐƐĞƌƚ͕^ƚƌĂŬĂ,;ϮϬϭϬͿ&ƵŶĐƚŝŽŶĂůŽƌŐĂŶŝnjĂƚŝŽŶŽĨǀĞƐƚŝďƵůĂƌ
ĐŽŵŵŝƐƐƵƌĂůĐŽŶŶĞĐƚŝŽŶƐŝŶĨƌŽŐ͘:EĞƵƌŽƐĐŝϯϬ͗ϯϯϭϬͲϯϯϮϱ͘
DĂƌƌ;ϭϵϲϵͿƚŚĞŽƌLJŽĨĐĞƌĞďĞůůĂƌĐŽƌƚĞdž͘:WŚLJƐŝŽůϮϬϮ͗ϰϯϳͲϰϳϬ͘
DĂƐƵĚĂE͕ŵĂƌŝ^;ϮϬϬϴͿĐŽŵƉƵƚĂƚŝŽŶĂůƐƚƵĚLJŽĨƐLJŶĂƉƚŝĐŵĞĐŚĂŶŝƐŵƐŽĨƉĂƌƚŝĂůŵĞŵŽƌLJƚƌĂŶƐĨĞƌŝŶ
ĐĞƌĞďĞůůĂƌǀĞƐƚŝďƵůŽͲŽĐƵůĂƌͲƌĞĨůĞdžůĞĂƌŶŝŶŐ͘:ŽŵƉƵƚEĞƵƌŽƐĐŝϮϰ͗ϭϯϳͲϭϱϲ͘
DĂƚƐƵŶŽ,͕<ƵĚŽŚD͕tĂƚĂŬĂďĞ͕zĂŵĂŵŽƌŝd͕^ŚŝŐĞŵŽƚŽZ͕EĂŐĂŽ^;ϮϬϭϲͿŝƐƚƌŝďƵƚŝŽŶĂŶĚ^ƚƌƵĐƚƵƌĞ
ŽĨ ^LJŶĂƉƐĞƐ ŽŶ DĞĚŝĂů sĞƐƚŝďƵůĂƌ EƵĐůĞĂƌ EĞƵƌŽŶƐ dĂƌŐĞƚĞĚ ďLJ ĞƌĞďĞůůĂƌ &ůŽĐĐƵůƵƐ WƵƌŬŝŶũĞ
ĞůůƐ ĂŶĚ sĞƐƚŝďƵůĂƌ EĞƌǀĞ ŝŶ DŝĐĞ͗ >ŝŐŚƚ ĂŶĚ ůĞĐƚƌŽŶ DŝĐƌŽƐĐŽƉLJ ^ƚƵĚŝĞƐ͘ W>Ž^ KŶĞ
ϭϭ͗ĞϬϭϲϰϬϯϳ͘
DĐůǀĂŝŶ>͕ĂŐŶĂůůDt͕^ĂŬĂƚŽƐ͕ĚƵ>ĂĐ^;ϮϬϭϬͿŝĚŝƌĞĐƚŝŽŶĂůƉůĂƐƚŝĐŝƚLJŐĂƚĞĚďLJŚLJƉĞƌƉŽůĂƌŝnjĂƚŝŽŶ
ĐŽŶƚƌŽůƐ ƚŚĞ ŐĂŝŶ ŽĨ ƉŽƐƚƐLJŶĂƉƚŝĐ ĨŝƌŝŶŐ ƌĞƐƉŽŶƐĞƐ Ăƚ ĐĞŶƚƌĂů ǀĞƐƚŝďƵůĂƌ ŶĞƌǀĞ ƐLJŶĂƉƐĞƐ͘ EĞƵƌŽŶ
ϲϴ͗ϳϲϯͲϳϳϱ͘
DĞĚŝŶĂ:&;ϮϬϭϭͿdŚĞŵƵůƚŝƉůĞƌŽůĞƐŽĨWƵƌŬŝŶũĞĐĞůůƐŝŶƐĞŶƐŽƌŝͲŵŽƚŽƌĐĂůŝďƌĂƚŝŽŶ͗ƚŽƉƌĞĚŝĐƚ͕ƚĞĂĐŚĂŶĚ
ĐŽŵŵĂŶĚ͘ƵƌƌKƉŝŶEĞƵƌŽďŝŽůϮϭ͗ϲϭϲͲϲϮϮ͘
DĞůǀŝůů:ŽŶĞƐ'͕'ƵŝƚƚŽŶ͕ĞƌƚŚŽnj;ϭϵϴϴͿŚĂŶŐŝŶŐƉĂƚƚĞƌŶƐŽĨĞLJĞͲŚĞĂĚĐŽŽƌĚŝŶĂƚŝŽŶĚƵƌŝŶŐϲŚŽĨ
ŽƉƚŝĐĂůůLJƌĞǀĞƌƐĞĚǀŝƐŝŽŶ͘džƉƌĂŝŶZĞƐϲϵ͗ϱϯϭͲϱϰϰ͘
DĞŶnjŝĞƐ :Z͕ WŽƌƌŝůů :͕ ƵƚŝĂ D͕ ĞĂŶ W ;ϮϬϭϬͿ ^LJŶĂƉƚŝĐ ƉůĂƐƚŝĐŝƚLJ ŝŶ ŵĞĚŝĂů ǀĞƐƚŝďƵůĂƌ ŶƵĐůĞƵƐ ŶĞƵƌŽŶƐ͗
ĐŽŵƉĂƌŝƐŽŶǁŝƚŚĐŽŵƉƵƚĂƚŝŽŶĂůƌĞƋƵŝƌĞŵĞŶƚƐŽĨsKZĂĚĂƉƚĂƚŝŽŶ͘W>Ž^KŶĞϱ͘
DŝůĞƐ &͕ >ŝƐďĞƌŐĞƌ ^' ;ϭϵϴϭͿ WůĂƐƚŝĐŝƚLJ ŝŶ ƚŚĞ ǀĞƐƚŝďƵůŽͲŽĐƵůĂƌ ƌĞĨůĞdž͗ Ă ŶĞǁ ŚLJƉŽƚŚĞƐŝƐ͘ ŶŶƵ ZĞǀ
EĞƵƌŽƐĐŝϰ͗ϮϳϯͲϮϵϵ͘
DŝƚĐŚĞůů͕ĞůůĂ^ĂŶƚŝŶĂ͕ƵůůĞŶ<;ϮϬϭϲͿWůĂƐƚŝĐŝƚLJǁŝƚŚŝŶŶŽŶͲĐĞƌĞďĞůůĂƌƉĂƚŚǁĂLJƐƌĂƉŝĚůLJƐŚĂƉĞƐ
ŵŽƚŽƌƉĞƌĨŽƌŵĂŶĐĞŝŶǀŝǀŽ͘EĂƚŽŵŵƵŶϳ͗ϭϭϮϯϴ͘
EĂŐĂŽ^͕<ŝƚĂnjĂǁĂ,;ϮϬϬϯͿĨĨĞĐƚƐŽĨƌĞǀĞƌƐŝďůĞƐŚƵƚĚŽǁŶŽĨƚŚĞŵŽŶŬĞLJĨůŽĐĐƵůƵƐŽŶƚŚĞƌĞƚĞŶƚŝŽŶŽĨ
ĂĚĂƉƚĂƚŝŽŶŽĨƚŚĞŚŽƌŝnjŽŶƚĂůǀĞƐƚŝďƵůŽͲŽĐƵůĂƌƌĞĨůĞdž͘EĞƵƌŽƐĐŝĞŶĐĞϭϭϴ͗ϱϲϯͲϱϳϬ͘
EĂŐĂŽ^͕,ŽŶĚĂd͕zĂŵĂnjĂŬŝd;ϮϬϭϯͿdƌĂŶƐĨĞƌŽĨŵĞŵŽƌLJƚƌĂĐĞŽĨĐĞƌĞďĞůůƵŵͲĚĞƉĞŶĚĞŶƚŵŽƚŽƌůĞĂƌŶŝŶŐ
ŝŶŚƵŵĂŶƉƌŝƐŵĂĚĂƉƚĂƚŝŽŶ͗ĂŵŽĚĞůƐƚƵĚLJ͘EĞƵƌĂůEĞƚǁϰϳ͗ϳϮͲϴϬ͘
KŚLJĂŵĂd͕DĂƵŬD;ϮϬϬϭͿ>ĂƚĞŶƚĂĐƋƵŝƐŝƚŝŽŶŽĨƚŝŵĞĚƌĞƐƉŽŶƐĞƐŝŶĐĞƌĞďĞůůĂƌĐŽƌƚĞdž͘:EĞƵƌŽƐĐŝϮϭ͗ϲϴϮͲ
ϲϵϬ͘
KŚLJĂŵĂ d͕ EŽƌĞƐ t>͕ DĞĚŝŶĂ :&͕ ZŝƵƐĞĐŚ &͕ DĂƵŬ D ;ϮϬϬϲͿ >ĞĂƌŶŝŶŐͲŝŶĚƵĐĞĚ ƉůĂƐƚŝĐŝƚLJ ŝŶ ĚĞĞƉ
ĐĞƌĞďĞůůĂƌŶƵĐůĞƵƐ͘:EĞƵƌŽƐĐŝϮϲ͗ϭϮϲϱϲͲϭϮϲϲϯ͘
KŬĂŵŽƚŽd͕^ŚŝƌĂŽd͕^ŚƵƚŽŚ&͕^ƵnjƵŬŝd͕EĂŐĂŽ^;ϮϬϭϭͿWŽƐƚͲƚƌĂŝŶŝŶŐĐĞƌĞďĞůůĂƌĐŽƌƚŝĐĂůĂĐƚŝǀŝƚLJƉůĂLJƐĂŶ
ŝŵƉŽƌƚĂŶƚƌŽůĞĨŽƌĐŽŶƐŽůŝĚĂƚŝŽŶŽĨŵĞŵŽƌLJŽĨĐĞƌĞďĞůůƵŵͲĚĞƉĞŶĚĞŶƚŵŽƚŽƌůĞĂƌŶŝŶŐ͘EĞƵƌŽƐĐŝ
>ĞƚƚϱϬϰ͗ϱϯͲϱϲ͘
KŽŵŵĞŶ^͕^ƚĂŚů:^;ϮϬϬϴͿLJĞŽƌŝĞŶƚĂƚŝŽŶĚƵƌŝŶŐƐƚĂƚŝĐƚŝůƚƐĂŶĚŝƚƐƌĞůĂƚŝŽŶƐŚŝƉƚŽƐƉŽŶƚĂŶĞŽƵƐŚĞĂĚ
ƉŝƚĐŚŝŶƚŚĞůĂďŽƌĂƚŽƌLJŵŽƵƐĞ͘ƌĂŝŶZĞƐϭϭϵϯ͗ϱϳͲϲϲ͘
WĂƐƚŽƌ D͕ ĚĞ ůĂ ƌƵnj ZZ͕ ĂŬĞƌ Z ;ϭϵϵϰͿ ĞƌĞďĞůůĂƌ ƌŽůĞ ŝŶ ĂĚĂƉƚĂƚŝŽŶ ŽĨ ƚŚĞ ŐŽůĚĨŝƐŚ ǀĞƐƚŝďƵůŽŽĐƵůĂƌ
ƌĞĨůĞdž͘:EĞƵƌŽƉŚLJƐŝŽůϳϮ͗ϭϯϴϯͲϭϯϵϰ͘
WĞƚƚŽƌŽƐƐŝ s͕ ŝĞŶŝ s͕ ^ĐĂƌĚƵnjŝŽ D͕ 'ƌĂƐƐŝ ^ ;ϮϬϭϭͿ >ŽŶŐͲƚĞƌŵ ƉŽƚĞŶƚŝĂƚŝŽŶ ŽĨ ƐLJŶĂƉƚŝĐ ƌĞƐƉŽŶƐĞ ĂŶĚ
ŝŶƚƌŝŶƐŝĐĞdžĐŝƚĂďŝůŝƚLJŝŶŶĞƵƌŽŶƐŽĨƚŚĞƌĂƚŵĞĚŝĂůǀĞƐƚŝďƵůĂƌŶƵĐůĞŝ͘EĞƵƌŽƐĐŝĞŶĐĞϭϴϳ͗ϭͲϭϰ͘
ϯϭ
ϳϯϱ
ϳϯϲ
ϳϯϳ
ϳϯϴ
ϳϯϵ
ϳϰϬ
ϳϰϭ
ϳϰϮ
ϳϰϯ
ϳϰϰ
ϳϰϱ
ϳϰϲ
ϳϰϳ
ϳϰϴ
ϳϰϵ
ϳϱϬ
ϳϱϭ
ϳϱϮ
ϳϱϯ
ϳϱϰ
ϳϱϱ
ϳϱϲ
ϳϱϳ
ϳϱϴ
ϳϱϵ
ϳϲϬ
ϳϲϭ
ϳϲϮ
ϳϲϯ
ϳϲϰ
ϳϲϱ
ϳϲϲ
ϳϲϳ
ϳϲϴ
ϳϲϵ
ϳϳϬ
ϳϳϭ
ϳϳϮ
ϳϳϯ
ϳϳϰ
ϳϳϱ
ϳϳϲ
ϳϳϳ
ϳϳϴ
ϳϳϵ
ϳϴϬ
ϳϴϭ
ϳϴϮ
WŽƌƌŝůů:͕ĞĂŶW;ϮϬϬϳͿĞƌĞďĞůůĂƌŵŽƚŽƌůĞĂƌŶŝŶŐ͗ǁŚĞŶŝƐĐŽƌƚŝĐĂůƉůĂƐƚŝĐŝƚLJŶŽƚĞŶŽƵŐŚ͍W>Ž^ŽŵƉƵƚ
ŝŽůϯ͗ϭϵϯϱͲϭϵϱϬ͘
WƵŐŚ:Z͕ZĂŵĂŶ/D;ϮϬϬϲͿWŽƚĞŶƚŝĂƚŝŽŶŽĨŵŽƐƐLJĨŝďĞƌW^ƐŝŶƚŚĞĐĞƌĞďĞůůĂƌŶƵĐůĞŝďLJEDƌĞĐĞƉƚŽƌ
ĂĐƚŝǀĂƚŝŽŶĨŽůůŽǁĞĚďLJƉŽƐƚŝŶŚŝďŝƚŽƌLJƌĞďŽƵŶĚĐƵƌƌĞŶƚ͘EĞƵƌŽŶϱϭ͗ϭϭϯͲϭϮϯ͘
WƵLJĂů:͕'ƌĂƐƐŝ^͕ŝĞŶŝ͕&ƌŽŶĚĂƌŽůŝ͕ĞŵĞŵĞƐ͕ZĂLJŵŽŶĚ:͕WĞƚƚŽƌŽƐƐŝs;ϮϬϬϯͿĞǀĞůŽƉŵĞŶƚĂůƐŚŝĨƚ
ĨƌŽŵůŽŶŐͲƚĞƌŵĚĞƉƌĞƐƐŝŽŶƚŽůŽŶŐͲƚĞƌŵƉŽƚĞŶƚŝĂƚŝŽŶŝŶƚŚĞƌĂƚŵĞĚŝĂůǀĞƐƚŝďƵůĂƌŶƵĐůĞŝ͗ƌŽůĞŽĨ
ŐƌŽƵƉ/ŵĞƚĂďŽƚƌŽƉŝĐŐůƵƚĂŵĂƚĞƌĞĐĞƉƚŽƌƐ͘:WŚLJƐŝŽůϱϱϯ͗ϰϮϳͲϰϰϯ͘
ZĂLJŵŽŶĚ :>͕ >ŝƐďĞƌŐĞƌ ^' ;ϭϵϵϲͿ ĞŚĂǀŝŽƌĂů ĂŶĂůLJƐŝƐ ŽĨ ƐŝŐŶĂůƐ ƚŚĂƚ ŐƵŝĚĞ ůĞĂƌŶĞĚ ĐŚĂŶŐĞƐ ŝŶ ƚŚĞ
ĂŵƉůŝƚƵĚĞĂŶĚĚLJŶĂŵŝĐƐŽĨƚŚĞǀĞƐƚŝďƵůŽͲŽĐƵůĂƌƌĞĨůĞdž͘:EĞƵƌŽƐĐŝϭϲ͗ϳϳϵϭͲϳϴϬϮ͘
ZŝŶĂůĚŝ͕ĞĨƚĞƌĂůŝ͕DŝĂůŽƚ͕'ĂƌĚĞŶ>͕ĞƌĂŶĞĐŬD͕EŽůĂŶD&;ϮϬϭϯͿ,EϭĐŚĂŶŶĞůƐŝŶĐĞƌĞďĞůůĂƌ
WƵƌŬŝŶũĞ ĐĞůůƐ ƉƌŽŵŽƚĞ ůĂƚĞ ƐƚĂŐĞƐ ŽĨ ůĞĂƌŶŝŶŐ ĂŶĚ ĐŽŶƐƚƌĂŝŶ ƐLJŶĂƉƚŝĐ ŝŶŚŝďŝƚŝŽŶ͘ : WŚLJƐŝŽů
ϱϵϭ͗ϱϲϵϭͲϱϳϬϵ͘
ZŽLJ :͕ ƵůůĞŶ < ;ϮϬϬϰͿ ŝƐƐŽĐŝĂƚŝŶŐ ƐĞůĨͲŐĞŶĞƌĂƚĞĚ ĨƌŽŵ ƉĂƐƐŝǀĞůLJ ĂƉƉůŝĞĚ ŚĞĂĚ ŵŽƚŝŽŶ͗ ŶĞƵƌĂů
ŵĞĐŚĂŶŝƐŵƐŝŶƚŚĞǀĞƐƚŝďƵůĂƌŶƵĐůĞŝ͘:EĞƵƌŽƐĐŝϮϰ͗ϮϭϬϮͲϮϭϭϭ͘
^ĐĂƌĚƵnjŝŽ D͕ WĂŶŝĐŚŝ Z͕ WĞƚƚŽƌŽƐƐŝ s͕ 'ƌĂƐƐŝ ^ ;ϮϬϭϮͿ dŚĞ ƌĞƉĞƚŝƚŝŽŶ ƚŝŵŝŶŐ ŽĨ ŚŝŐŚ ĨƌĞƋƵĞŶĐLJ ĂĨĨĞƌĞŶƚ
ƐƚŝŵƵůĂƚŝŽŶ ĚƌŝǀĞƐ ƚŚĞ ďŝĚŝƌĞĐƚŝŽŶĂů ƉůĂƐƚŝĐŝƚLJ Ăƚ ĐĞŶƚƌĂů ƐLJŶĂƉƐĞƐ ŝŶ ƚŚĞ ƌĂƚ ŵĞĚŝĂů ǀĞƐƚŝďƵůĂƌ
ŶƵĐůĞŝ͘EĞƵƌŽƐĐŝĞŶĐĞϮϮϯ͗ϭͲϭϭ͘
^ĞŬŝƌŶũĂŬ͕ĚƵ>ĂĐ^;ϮϬϬϲͿWŚLJƐŝŽůŽŐŝĐĂůĂŶĚĂŶĂƚŽŵŝĐĂůƉƌŽƉĞƌƚŝĞƐŽĨŵŽƵƐĞŵĞĚŝĂůǀĞƐƚŝďƵůĂƌŶƵĐůĞƵƐ
ŶĞƵƌŽŶƐƉƌŽũĞĐƚŝŶŐƚŽƚŚĞŽĐƵůŽŵŽƚŽƌŶƵĐůĞƵƐ͘:EĞƵƌŽƉŚLJƐŝŽůϵϱ͗ϯϬϭϮͲϯϬϮϯ͘
^ĞŬŝƌŶũĂŬ͕sŝƐƐĞů͕ŽůůŝŶŐĞƌ:͕&ĂƵůƐƚŝĐŚD͕ĚƵ>ĂĐ^;ϮϬϬϯͿWƵƌŬŝŶũĞĐĞůůƐLJŶĂƉƐĞƐƚĂƌŐĞƚƉŚLJƐŝŽůŽŐŝĐĂůůLJ
ƵŶŝƋƵĞďƌĂŝŶƐƚĞŵŶĞƵƌŽŶƐ͘:EĞƵƌŽƐĐŝϮϯ͗ϲϯϵϮͲϲϯϵϴ͘
^ĞƌĂĨŝŶ D͕ ĚĞ tĂĞůĞ ͕ <ŚĂƚĞď ͕ sŝĚĂů WW͕ DƵŚůĞƚŚĂůĞƌ D ;ϭϵϵϭͿ DĞĚŝĂů ǀĞƐƚŝďƵůĂƌ ŶƵĐůĞƵƐ ŝŶ ƚŚĞ
ŐƵŝŶĞĂͲƉŝŐ͘/͘/ŶƚƌŝŶƐŝĐŵĞŵďƌĂŶĞƉƌŽƉĞƌƚŝĞƐŝŶďƌĂŝŶƐƚĞŵƐůŝĐĞƐ͘džƉƌĂŝŶZĞƐϴϰ͗ϰϭϳͲϰϮϱ͘
^ŚŝŶ D͕ DŽŐŚĂĚĂŵ ^,͕ ^ĞŬŝƌŶũĂŬ ͕ ĂŐŶĂůů Dt͕ <ŽůŬŵĂŶ <͕ :ĂĐŽďƐ Z͕ &ĂƵůƐƚŝĐŚ D͕ ĚƵ >ĂĐ ^ ;ϮϬϭϭͿ
DƵůƚŝƉůĞ ƚLJƉĞƐ ŽĨ ĐĞƌĞďĞůůĂƌ ƚĂƌŐĞƚ ŶĞƵƌŽŶƐ ĂŶĚ ƚŚĞŝƌ ĐŝƌĐƵŝƚƌLJ ŝŶ ƚŚĞ ǀĞƐƚŝďƵůŽͲŽĐƵůĂƌ ƌĞĨůĞdž͘ :
EĞƵƌŽƐĐŝϯϭ͗ϭϬϳϳϲͲϭϬϳϴϲ͘
^ŚŝŶ^>͕ŚĂŽ'Y͕ZĂLJŵŽŶĚ:>;ϮϬϭϰͿ^ŝŐŶĂůƐĂŶĚůĞĂƌŶŝŶŐƌƵůĞƐŐƵŝĚŝŶŐŽĐƵůŽŵŽƚŽƌƉůĂƐƚŝĐŝƚLJ͘:EĞƵƌŽƐĐŝ
ϯϰ͗ϭϬϲϯϱͲϭϬϲϰϰ͘
^ŚƵƚŽŚ &͕ KŚŬŝ D͕ <ŝƚĂnjĂǁĂ ,͕ /ƚŽŚĂƌĂ ^͕ EĂŐĂŽ ^ ;ϮϬϬϲͿ DĞŵŽƌLJ ƚƌĂĐĞ ŽĨ ŵŽƚŽƌ ůĞĂƌŶŝŶŐ ƐŚŝĨƚƐ
ƚƌĂŶƐƐLJŶĂƉƚŝĐĂůůLJĨƌŽŵĐĞƌĞďĞůůĂƌĐŽƌƚĞdžƚŽŶƵĐůĞŝĨŽƌĐŽŶƐŽůŝĚĂƚŝŽŶ͘EĞƵƌŽƐĐŝĞŶĐĞϭϯϵ͗ϳϲϳͲϳϳϳ͘
^ƚĂŚů:^;ϮϬϬϰͿhƐŝŶŐĞLJĞŵŽǀĞŵĞŶƚƐƚŽĂƐƐĞƐƐďƌĂŝŶĨƵŶĐƚŝŽŶŝŶŵŝĐĞ͘sŝƐŝŽŶZĞƐϰϰ͗ϯϰϬϭͲϯϰϭϬ͘
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