HAL Id: tel-01359194
https://tel.archives-ouvertes.fr/tel-01359194
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ldentification and characterization of epigenetically
dynamic regions in response to different environmental
stimuli in liver cells
Pierre-Benoit Ancey
To cite this version:
Pierre-Benoit Ancey. ldentification and characterization of epigenetically dynamic regions in response to different environmental stimuli in liver cells. Cancer. Université Claude Bernard - Lyon I, 2015. English. �NNT : 2015LYO10258�. �tel-01359194�
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ŶĚŽĚĞƌŵĨŽƌŵĂƚŝŽŶͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ Ϯϰ 3. &ƌŽŵĞŶĚŽĚĞƌŵƚŽĨŽƌĞŐƵƚͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ Ϯϱ 4. &ƌŽŵĨŽƌĞŐƵƚƚŽŚĞƉĂƚŽďůĂƐƚͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ Ϯϲ 5. &ƌŽŵŚĞƉĂƚŽďůĂƐƚƚŽďŝůŝĂƌLJĐĞůůƐͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ Ϯϳ 6. &ƌŽŵŚĞƉĂƚŽďůĂƐƚƚŽŚĞƉĂƚŽĐLJƚĞƐͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ Ϯϴ ,ĞƉĂƚŽĐĞůůƵůĂƌĐĂƌĐŝŶŽŵĂ ͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ Ϯϵ ĞůůŽĨŽƌŝŐŝŶŽĨ,ͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ ϯϬ 1. ,ĞƉĂƚŽĐLJƚĞ ͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ ϯϬ 2. WƌŽŐĞŶŝƚŽƌͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ ϯϭ KƌŝŐŝŶŽĨ,ĞƉĂƚŝĐƉƌŽŐĞŶŝƚŽƌĐĞůůͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ ϯϮ ,WƐƌĞĂĐƚŝǀĂƚŝŽŶŝŶŝŶũƵƌLJĐŽŶƚĞdžƚͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ ϯϰ 3. ĂŶĐĞƌƐƚĞŵĐĞůůƐͬƚƵŵŽƵƌŝŶŝƚŝĂƚŝŶŐĐĞůůƐͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ ϯϳ ZŝƐŬƐĨĂĐƚŽƌƐŽĨ,ĞƉĂƚŽĐĞůůƵůĂƌĐĂƌĐŝŶŽŵĂ ͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ ϰϬ /ŶĨůĂŵŵĂƚŝŽŶ ͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ ϰϭ ,sŝŶĨĞĐƚŝŽŶͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ ϰϮ 1. ,sǀŝƌƵƐĚĞƐĐƌŝƉƚŝŽŶ ͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ ϰϯ 2. ,sŶĂƚƵƌĂůŚŝƐƚŽƌLJͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ ϰϰ 3. ŝŽůŽŐLJŽĨƚƌĂŶƐŝĞŶƚŝŶĨĞĐƚŝŽŶƐ ͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ ϰϳ 4. ŝŽůŽŐLJŽĨĐŚƌŽŶŝĐŝŶĨĞĐƚŝŽŶƐͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ ϰϳ 5. ŝĨĨĞƌĞŶƚ,sŐĞŶŽƚLJƉĞƐͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ ϰϴ 6. DŽůĞĐƵůĂƌŵĞĐŚĂŶŝƐŵƐŽĨĐĂƌĐŝŶŽŐĞŶĞƐŝƐďLJ,s ͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ ϰϵ 7. ǀŝĚĞŶĐĞƚŚĂƚŝŶƚĞŐƌĂƚŝŽŶŽĨǀŝƌĂůEĐĂŶďĞƉƌŽͲŽŶĐŽŐĞŶŝĐ ͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ ϱϬ 8. ZŽůĞŽĨ,sƉƌŽƚĞŝŶƐŝŶŚĞƉĂƚŽĐĂƌĐŝŶŽŐĞŶĞƐŝƐͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ ϱϭ ,dž ͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ ϱϮ >ƉƌŽƚĞŝŶ ͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ ϱϯ &ƌŽŵĞŶǀŝƌŽŶŵĞŶƚƚŽŐĞŶŽŵĞ͗ƚŚĞƌŽůĞŽĨĞƉŝŐĞŶĞƚŝĐƐͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ ϱϯ /ŶƚƌŽĚƵĐƚŝŽŶŽŶĞƉŝŐĞŶĞƚŝĐŵĞĐŚĂŶŝƐŵƐͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ ϱϱ 1. EŵĞƚŚLJůĂƚŝŽŶƉŚLJƐŝŽůŽŐLJͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ ϱϲ 2. DĞƚŚLJůĂƚŝŽŶƌĞŐƵůĂƚĞƐƚƌĂŶƐĐƌŝƉƚŝŽŶĂŶĚŐĞŶŽŵĞŽƌŐĂŶŝƐĂƚŝŽŶ͘ ͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ ϱϳ ďĞƌƌĂŶƚEŵĞƚŚLJůĂƚŝŽŶƉƌŽĨŝůĞŝŶ,ͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ ϲϬ 1. ůƚĞƌĂƚŝŽŶŝŶEDdϭ͕EDdϯ͕ĂŶĚEDdϯĞdžƉƌĞƐƐŝŽŶͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ ϲϭ 2. EŵĞƚŚLJůĂƚŝŽŶĂůƚĞƌĂƚŝŽŶƐŝŶƉƌĞĐĂŶĐĞƌŽƵƐƐƚĂŐĞƐͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ ϲϮϭϴ DĞƚŚLJůĂƚŝŽŶĂŶĚĚŝĨĨĞƌĞŶƚŝĂƚŝŽŶͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ ϲϯ ,sĂŶĚEŵĞƚŚLJůĂƚŝŽŶͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ ϲϱ 1. dŚĞƉŽƚĞŶƚŝĂůƌŽůĞŽĨ,džŝŶ,sĞƉŝŐĞŶĞƚŝĐƌĞŐƵůĂƚŝŽŶͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ ϲϲ EŵĞƚŚLJůĂƚŝŽŶĚŝƐƌƵƉƚŝŽŶďLJĂƐƉĞĐŝĨŝĐĂŶƚŝͲŝŶĨůĂŵĂƚŽƌLJĐLJƚŽŬŝŶĞ͗d'&Ƙͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ ϲϳ 1. d'&ɴƉĂƚŚǁĂLJͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ ϲϳ 2. d'&ɴŝŶĚƵĐĞƐd/ƐƉŽƉƵůĂƚŝŽŶŽǀĞƌƌĞƉƌĞƐĞŶƚĂƚŝŽŶ ͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ ϲϵ 3. d'&ɴƉƌŽĚƵĐĞEŵĞƚŚLJůĂƚŝŽŶĂůƚĞƌĂƚŝŽŶƐŝŶůŝǀĞƌĐĂŶĐĞƌĐĞůůƐͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ ϳϬ IV. /D^ͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺϳϮ V. Dd Z/>^ED d,K^ ͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺϳϴ ĞůůĐƵůƚƵƌĞĂŶĚƚƌĞĂƚŵĞŶƚƐͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ ϴϬ 1. W,,ĐƵůƚƵƌĞĂŶĚ,sŝŶĨĞĐƚŝŽŶͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ ϴϬ 2. ,ĞƉĂZ'ĐĞůůĐƵůƚƵƌĞĂŶĚĚŝĨĨĞƌĞŶƚŝĂƚŝŽŶͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ ϴϬ 3. ,ƵŚϳĂŶĚ,ĞƉ'ϮĐĞůůĐƵůƚƵƌĞͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ ϴϭ 4. d'&ɴϭƚƌĞĂƚŵĞŶƚͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ ϴϭ 5. ƐŝZEƚƌĞĂƚŵĞŶƚͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ ϴϭ DŽůĞĐƵůĂƌďŝŽůŽŐLJͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ ϴϮ 1. EĞdžƚƌĂĐƚŝŽŶ ͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ ϴϮ 2. ŝƐƵůĨŝƚĞŵŽĚŝĨŝĐĂƚŝŽŶ ͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ ϴϮ 3. WLJƌŽƐĞƋƵĞŶĐŝŶŐͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ ϴϮ 4. ZEĞdžƚƌĂĐƚŝŽŶ ͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ ϴϯ 5. ZĞǀĞƌƐĞƚƌĂŶƐĐƌŝƉƚŝŽŶĂŶĚƋƵĂŶƚŝƚĂƚŝǀĞWZͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ ϴϯ 6. /ŵŵƵŶŽĨůƵŽƌĞƐĐĞŶĐĞ ͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ ϴϯ 7. /ŶƐŝƚƵWƌŽdžŝŵŝƚLJ>ŝŐĂƚŝŽŶƐƐĂLJͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ ϴϰ 8. ŚƌŽŵĂƚŝŶ/ŵŵƵŶŽƉƌĞĐŝƉŝƚĂƚŝŽŶ ͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ ϴϰ 9. &ůƵŽƌĞƐĐĞŶĐĞĐƚŝǀĂƚĞĚĞůů^ŽƌƚŝŶŐ;&^Ϳͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ ϴϰ ĞĂĚĂƌƌĂLJĂƐƐĂLJƐͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ ϴϱ 1. ĞĂĚĂƌƌĂLJŵĞƚŚLJůĂƚŝŽŶĂƐƐĂLJƐͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ ϴϱ 2. tŚŽůĞŐĞŶŽŵĞĞdžƉƌĞƐƐŝŽŶĂƌƌĂLJ ͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ ϴϲ ŝŽŝŶĨŽƌŵĂƚŝĐƐŶĂůLJƐŝƐ ͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ ϴϳ Z/^WZͬĂƐϵͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ ϴϴ 1. ĞƐŝŐŶĂŶĚƚƌĂŶƐĨĞĐƚŝŽŶ ͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ ϴϴ 2. ůŽŶĂůƐĞůĞĐƚŝŽŶͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ ϵϬ ^ƚĂƚŝƐƚŝĐĂůŶĂůLJƐŝƐͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ ϵϬ VI. Z ^h>d^ͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺϵϰ 'ĞŶŽŵŝĐƌĞƐƉŽŶƐĞƐƚŽ,ĞƉĂƚŝƚŝƐǀŝƌƵƐ;,sͿŝŶĨĞĐƚŝŽŶŝŶƉƌŝŵĂƌLJŚƵŵĂŶŚĞƉĂƚŽĐLJƚĞƐ ͺͺͺͺͺ ϵϲ 1. LJŶĂŵŝĐĐŚĂŶŐĞƐŝŶŐĞŶĞĞdžƉƌĞƐƐŝŽŶŝŶĐƵůƚƵƌĞĚƉƌŝŵĂƌLJŚĞƉĂƚŽĐLJƚĞƐͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ ϵϲ 2. ,sŝŶĨĞĐƚŝŽŶŝŶĚƵĐĞƐƚŝŵĞͲĚĞƉĞŶĚĞŶƚĐŚĂŶŐĞƐŝŶŐĞŶĞĞdžƉƌĞƐƐŝŽŶͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ ϵϴ 3. ZĞŐŝŽŶͲůĞǀĞůĚŝĨĨĞƌĞŶƚŝĂůEŵĞƚŚLJůĂƚŝŽŶŝŶƌĞƐƉŽŶƐĞƚŽ,sŝŶĨĞĐƚŝŽŶ ͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ ϭϬϯ 4. ^ŝƚĞͲƐƉĞĐŝĨŝĐĚŝĨĨĞƌĞŶƚŝĂůEŵĞƚŚLJůĂƚŝŽŶŝŶƌĞƐƉŽŶƐĞƚŽ,sŝŶĨĞĐƚŝŽŶͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ ϭϬϲ 5. &ƵŶĐƚŝŽŶĂůƐŝŐŶŝĨŝĐĂŶĐĞŽĨEŵĞƚŚLJůĂƚŝŽŶĐŚĂŶŐĞƐƵƉŽŶ,sŝŶĨĞĐƚŝŽŶͺͺͺͺͺͺͺͺͺͺͺͺͺ ϭϭϬ
ϭϵ ,E&ϰĂůƚĞƌŶĂƚŝǀĞƉƌŽŵŽƚĞƌĐŚŽŝĐĞŝƐůŝŶŬĞĚƚŽƉƌŽŐƌĞƐƐŝǀĞĚĞŵĞƚŚLJůĂƚŝŽŶĚƵƌŝŶŐŚĞƉĂƚŽĐLJƚĞ ĚŝĨĨĞƌĞŶƚŝĂƚŝŽŶ ͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ ϭϭϰ 1. ŝĨĨĞƌĞŶƚŝĂƚŝŽŶŽĨŚĞƉĂƚŝĐďŝƉŽƚĞŶƚƉƌŽŐĞŶŝƚŽƌŝŶƚŽŚĞƉĂƚŽĐLJƚĞƐͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ ϭϭϰ 2. ^ĞƋƵĞŶƚŝĂůŚĞƉĂƚŽĐLJƚĞĚŝĨĨĞƌĞŶƚŝĂƚŝŽŶŝƐĐŽƌƌĞůĂƚĞĚǁŝƚŚĂƉƌŽŐƌĞƐƐŝǀĞ,E&ϰŝŶƚƌĂŐĞŶŝĐ ĚĞŵĞƚŚLJůĂƚŝŽŶ͘ͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ ϭϭϲ 3. ,LJƉŽŵĞƚŚLJůĂƚŝŽŶŽĨ,E&ϰůŽĐƵƐŝƐĐŽƌƌĞůĂƚĞĚƚŽŝƐŽĨŽƌŵƐǁŝƚĐŚͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ ϭϮϮ 4. d dƐĞdžƉƌĞƐƐŝŽŶĚƵƌŝŶŐĚŝĨĨĞƌĞŶƚŝĂƚŝŽŶͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ ϭϮϰ ^ƚƵĚLJŽĨŐĞŶĞďŽĚLJŵĞƚŚLJůĂƚŝŽŶŝŶƌĞƐƉŽŶƐĞƚŽd'&Ƙͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ ϭϮϴ 1. EŵĞƚŚLJůĂƚŝŽŶŵĂĐŚŝŶĞƌLJŝƐƌĞƋƵŝƌĞĚĨŽƌd'&ƘƐŝŐŶĂůƚƌĂŶƐĚƵĐƚŝŽŶ ͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ ϭϮϴ 2. ŚĂƌĂĐƚĞƌŝnjĂƚŝŽŶŽĨEŵĞƚŚLJůĂƚŝŽŶĐŚĂŶŐĞƐŝŶƚŚĞŐĞŶĞďŽĚLJĂŶĚƚŚĞŝƌƉŽƚĞŶƚŝĂůŝŵƉĂĐƚŽŶŐĞŶĞ ĞdžƉƌĞƐƐŝŽŶ͘ͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ ϭϯϬ 3. ŚĂƌĂĐƚĞƌŝƐĂƚŝŽŶƐŽĨdZZWŐĞŶĞďŽĚLJŵĞƚŚLJůĂƚŝŽŶƐƚĂďŝůŝƚLJĂŶĚĚLJŶĂŵŝĐŝƚLJ͘ ͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ ϭϯϮ 4. EDdϯĂŶĚEDdϯĂƌĞƌĞƋƵŝƌĞĚĨŽƌdZZWŚLJƉĞƌŵĞƚŚLJůĂƚŝŽŶŝŶƌĞƐƉŽŶƐĞƚŽd'&ƘĂƐǁĞůůĂƐŝŶ ƚŚĞŵĞƚŚLJůĂƚŝŽŶŵĂŝŶƚĞŶĂŶĐĞ͘ ͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ ϭϯϳ 5. ZŽůĞŽĨŐĞŶĞďŽĚLJĞŶŚĂŶĐĞƌŝŶdZZWĞdžƉƌĞƐƐŝŽŶĂŶĚd'&ƘƌĞƐƉŽŶƐĞ ͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ ϭϯϵ VII. /^h^^/KEͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺϭϰϮ 'ĞŶŽŵŝĐƌĞƐƉŽŶƐĞƐƚŽ,ĞƉĂƚŝƚŝƐǀŝƌƵƐ;,sͿŝŶĨĞĐƚŝŽŶŝŶƉƌŝŵĂƌLJŚƵŵĂŶŚĞƉĂƚŽĐLJƚĞƐ ͺͺͺͺ ϭϰϰ 1. 'ĞŶĞĞdžƉƌĞƐƐŝŽŶĚLJŶĂŵŝĐƐĚƵĞƚŽW,,ŝŶǀŝƚƌŽĐƵůƚƵƌĞ͘ͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ ϭϰϰ 2. 'ĞŶĞĞdžƉƌĞƐƐŝŽŶĚLJŶĂŵŝĐƐŝŶƌĞƐƉŽŶƐĞƚŽ,sŶĂƚƵƌĂůŝŶĨĞĐƚŝŽŶ͘ͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ ϭϰϱ 3. ,sƐƚĂďůLJŝŶĚƵĐĞƐEŵĞƚŚLJůĂƚŝŽŶĐŚĂŶŐĞƐĚƵƌŝŶŐƚŚĞĨŝƌƐƚĚĂLJŽĨW,,ŶĂƚƵƌĂůŝŶĨĞĐƚŝŽŶ ͺͺͺͺ ϭϰϱ >ŽĐĂůEĚĞŵĞƚŚLJůĂƚŝŽŶĚĞĨŝŶĞƐ,E&ϰƉƌŽŵŽƚĞƌĐŚŽŝĐĞĚƵƌŝŶŐŚĞƉĂƚŽĐLJƚĞĚŝĨĨĞƌĞŶƚŝĂƚŝŽŶϭϰϴ 'ĞŶĞďŽĚLJŵĞƚŚLJůĂƚŝŽŶĨƵŶĐƚŝŽŶƐĂŶĚƉĂƌƚŶĞƌƐͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ ϭϱϯ 1. d'&ƘŝƐĞŶƌŝĐŚĞĚŝŶŝŶƚƌĂŐĞŶŝĐƌĞŐƵůĂƚŽƌLJƌĞŐŝŽŶ ͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ ϭϱϯ 2. 'ĞŶĞďŽĚLJŵĞƚŚLJůĂƚŝŽŶĞƐƚĂďůŝƐŚŵĞŶƚĂŶĚŵĂŝŶƚĞŶĂŶĐĞ ͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ ϭϱϰ 3. ZŽůĞŽĨŝŶƚƌĂŐĞŶŝĐŵĞƚŚLJůĂƚŝŽŶ͗ƚŚĞdZZWĞdžĂŵƉůĞ ͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ ϭϱϱ 4. ^ƚĂďŝůŝƚLJŽĨEŵĞƚŚLJůĂƚŝŽŶͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ ϭϱϴ 5. /ŵƉŽƌƚĂŶĐĞŽĨŐĞŶĞďŽĚLJŵĞƚŚLJůĂƚŝŽŶŝŶĐĂŶĐĞƌ ͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺ ϭϱϵ VIII. KE>h^/KE^ ͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺϭϲϬ IX. Z & Z E ^ͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺϭϲϰ X. EE y ^ ͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺͺϮϬϴ
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