Early life experience : neuroendocrine adaptations to maternal absence

Hele tekst

(1)Early life experience : neuroendocrine adaptations to maternal absence Enthoven, L.. Citation Enthoven, L. (2007, October 4). Early life experience : neuroendocrine adaptations to maternal absence. Retrieved from https://hdl.handle.net/1887/12379 Version:. Corrected Publisher’s Version. License:. Licence agreement concerning inclusion of doctoral thesis in the Institutional Repository of the University of Leiden. Downloaded from:. https://hdl.handle.net/1887/12379. Note: To cite this publication please use the final published version (if applicable)..

(2) 1MBTNBDPSUJDPTUFSPOFSFTQPOTFT SFnFDUUIFEFHSFFPGOPWFMUZJO NBMFBOEGFNBMF$%NJDF. "QQFOEJY*. -&OUIPWFO 4%BMN &3EF,MPFU.40JU[M.

(3)

(4) Appendix I ѮFTUVEJFTEFTDSJCFEJOUIJTBQQFOEJYQSPWJEFNFUIPEPMPHJDBMDPOTJEFSBUJPOTGPSCMPPETBNQMJOH WJBUBJMJODJTJPOJONJDF BUFDIOJRVFUIBUXBTBMTPVTFEGPSUIFEFUFSNJOBUJPOPGUIFDPSUJDPTUFSPOF SFTQPOTFTUPTXJNTUSFTTJONBUFSOBMMZEFQSJWFEBOEDPOUSPMNJDFEFTDSJCFEJO$IBQUFS. *"CTUSBDU ѮF IZQPUIBMBNJDQJUVJUBSZBESFOBM BYJT SFTQPOTF UP OPWFMUZ TUSFTT JT EFQFOEFOU PO UIF UZQF BOEEVSBUJPOPGBDFSUBJODIBMMFOHF)FSF XFTIPXJO$%NJDFUIBUDPSUJDPTUFSPOFTFDSFUJPO JODSFBTFESFëFDUJOHUIFiEFHSFFPGOPWFMUZu IPNFDBHFNJOVUFTJODZMJOEFSNJOVUFT JO DZMJOEFS NJOVUF GPSDFE TXJN ѮF NBHOJUVEF PG UIF DPSUJDPTUFSPOF SFTQPOTF BMSFBEZ EJĒFSFENJOVUFTBѫFSPOTFUPGUIFTUSFTTPS XJUIUIFNPSFTFWFSFDIBMMFOHFDBVTJOHBGBTUFS JOJUJBUJPOPGUIFHMVDPDPSUJDPJESFMBUFETUSFTTSFTQPOTF'VSUIFSNPSF GFNBMFNJDFTFDSFUFENPSF DPSUJDPTUFSPOFUIBONBMFNJDFVOEFSCBTBMDPOEJUJPOTBOETIPXFEBGBTUFSBOEIJHIFSJODSFBTFJO SFTQPOTFUPFBDIOPWFMUZ. **OUSPEVDUJPO " WBSJFUZ PG QIZTJPMPHJDBM < > BOE FNPUJPOBM < > TUJNVMJ SFTVMU JO EJĒFSFOU CFIBWJPVSBM DPQJOH SFTQPOTFT < > "OZ DIBMMFOHF FJUIFS SFBM PS JNBHJOFE FWPLFT B HMVDPDPSUJDPJESFMBUFE TUSFTT SFTQPOTF XIJDI TFSWFT UP SFTUPSF IPNFPTUBTJT BOE GBDJMJUBUF CFIBWJPVSBMBEBQUBUJPO<>4PNFDIBMMFOHFT IPXFWFS FMJDJUTUSPOHFSDPSUJDPTUFSPOFSFTQPOTFT UIBOPUIFST< >*OPVSMBC XFHFOFSBMMZQMBDFSBUTPSNJDFJOBOPWFMFOWJSPONFOUBOE FTUJNBUFQMBTNBDPSUJDPTUFSPOFDPODFOUSBUJPOTBUTFWFSBMUJNFQPJOUTEVSJOHUIJTOPWFMUZFYQPTVSF < >ѮFBJNPGUIFQSFTFOUTUVEZXBTUPEFêOFOPWFMUZDPOEJUJPOTDIBSBDUFSJTFECZ EJĒFSFOUJBM DPSUJDPTUFSPOF TFDSFUJPO *O GVUVSF FYQFSJNFOUT UIFTF TIPVME BMMPX UP DIPPTF UIF DIBMMFOHFBDDPSEJOHUPUIFIZQPUIFTJTFEDIBOHFJODPSUJDPTUFSPOFTFDSFUJPO UIVTBWPJEJOHDFJMJOH FĒFDUT . )FSF XFWBSJFECPUIUIFLJOEBOEEVSBUJPOPGOPWFMUZ.BMFBOEGFNBMF$%NJDFXFSF. QMBDFEJOUPBDZMJOEFSGPSPSNJOVUFT TXBNJOBMBSHFQPPMXJUIXBSNXBUFSGPSNJOVUF PSSFNBJOFEJOUIFJSIPNFDBHF#MPPETBNQMFTXFSFUBLFOTFRVFOUJBMMZ BOENJOVUFT BѫFSPOTFUPGOPWFMUZ JFJOUIFQSPBDUJWFQIBTFPGUIFHMVDPDPSUJDPJESFMBUFETUSFTTSFTQPOTF< >%FQFOEJOHPOUIFDIBMMFOHFXFFYQFDUFEBEJĒFSFOUBNQMJUVEFPSTMPQFPGUIFDPSUJDPTUFSPOF SFTQPOTFBOEPWFSBMMIJHIFSDPSUJDPTUFSPOFDPODFOUSBUJPOTJOGFNBMFUIBOJONBMFNJDF. *.BUFSJBMTBOE.FUIPET *"OJNBMT .BMFBOEGFNBMF$%NJDF NPOUITPGBHF XFSFIPVTFEGPVSQFSDBHFJOTBNFTFYHSPVQTXJUI GPPEBOEXBUFSBEMJCJUVNVOEFSBêYFEIPVSTMJHIUEBSLDZDMF MJHIUTPOBUIPVST BU .

(5) "QQFOEJY* 5BCMF*0WFSWJFXPGUIFDIBMMFOHFTXJUIUIFJSSFTQFDUJWFTBNQMJOHUJNFQPJOUT. Basal: Novel environment:. Control:. 5’CYL 60’CYL SWIM RSC RHC. 0 x -. sample time point (minutes) 15 30 x x x x -. x x x x -. 60 x x x x x. male 20 16 16 20 16. n=. female 20 16 16 20 16 13. *OUIJTFYQFSJNFOUBOJNBMTXFSFEJWJEFEPWFSEJĒFSFOUUSFBUNFOUHSPVQT#BTBM/PWFMFOWJSPONFOU$:- NJOVUFTFYQPTVSFUPBDZMJOEFS UIFSFBѫFSSFQMBDFNFOUJOIPNFDBHF$:-NJOVUFTFYQPTVSFUPBDZMJOEFS 48*.NJOVUFGPSDFETXJNNJOH UIFSFBѫFSQMBDFNFOUVOEFSBIFBUJOHMJHIUGPSNJOVUFTBOESFQMBDFNFOUJO IPNFDBHF$POUSPM34$SFQFBUFETBNQMJOHDPOUSPM3)$SFQFBUFEIBOEMJOHDPOUSPM"MMUSFBUNFOUTTUBSUFEBU UNJOVUFTBOEBMMIBOEMJOHQSPDFEVSFT XJUIUIFFYDFQUJPOPGBDUVBMCMPPETBNQMJOH XFSFFRVBMJOHSPVQT$:- $:- 48*. 34$BOE3)$ iYuJOEJDBUFTBCMPPETBNQMFUBLFO iuJOEJDBUFTOPCMPPETBNQMFUBLFO. UIFBOJNBMGBDJMJUJFTPGUIF4ZMWJVT-BCPSBUPSZBU-FJEFO6OJWFSTJUZ ѮF/FUIFSMBOET5FTUJOHXBT CFUXFFOIPVST.JDFXFSFTJOHMZIPVTFEPOFXFFLCFGPSFUIFFYQFSJNFOUTUBSUFE &YQFSJNFOUT XFSF BQQSPWFE CZ UIF -PDBM $PNNJUUFF PG "OJNBM )FBMUI &UIJDT BOE 3FTFBSDI PG UIF 6OJWFSTJUZ PG -FJEFO "OJNBM DBSF XBT DPOEVDUFE JO BDDPSEBODF XJUI UIF &$ $PVODJM %JSFDUJWFPG/PWFNCFS &&$ *&WBMVBUJPOPGOPWFMUZJOUFOTJUZ ѮF USFBUNFOU HSPVQT BSF TVNNBSJTFE JO 5BCMF * #BTBM 5P EFUFSNJOF CBTBM NPSOJOH DPSUJDPTUFSPOFDPODFOUSBUJPOT CMPPETBNQMFTXFSFDPMMFDUFEGSPNNBMFBOEGFNBMFNJDFXJUIJO UXPIPVSTBѫFSMJHIUTPO OQFSTFY /PWFMFOWJSPONFOU.JDF OQFSTFYBOEDPOEJUJPO XFSFQMBDFEJOBOVQSJHIUDZMJOEFS HSBZ17$DNIJHI DNJOEJBNFUFS GPSNJOVUFT CFGPSF UIFZ XFSF SFUVSOFE UP UIFJS IPNF DBHF PS GPS NJOVUFT HSPVQT $:- BOE $:- SFTQFDUJWFMZ "OPUIFSHSPVQPGNJDF OQFSTFY XBTQMBDFEJOUIFNJEEMFPGBQPPMXJUIXBSN XBUFS DNJOEJBNFUFS$ BOEIBEUPTXJNGPSNJOVUF HSPVQ48*. ѮFSFBѫFS UIFNPVTFXBTQMBDFEJOBDMFBODBHFXJUIUJTTVFCFEEJOHVOEFSBIFBUJOHMBNQGPSNJOVUFT BOESFUVSOFEUPJUTIPNFDBHF*OBMMDPOEJUJPOT PGFBDINPVTFUISFFTFRVFOUJBMCMPPETBNQMFT XFSFUBLFO BOENJOVUFTBѫFSUIFTUBSUPGUIFDIBMMFOHF"ѫFSFBDICMPPETBNQMF NJDF SFUVSOFEUPUIFJSIPNFDBHF $:- 48*. PSDZMJOEFS $:- $POUSPM5PDPOUSPMGPSUIF FĒFDUTPGSFQFBUFECMPPETBNQMJOHXFVTFEUXPIPNFDBHFDPOUSPMHSPVQT3FQFBUFETBNQMJOH DPOUSPMNJDF HSPVQ34$ OQFSTFY XFSFSFNPWFETIPSUMZGSPNUIFJSIPNFDBHF TFRVFOUJBM CMPPETBNQMFTXFSFUBLFOBUU BOENJOVUFT BOEBѫFSTBNQMJOHNJDFSFUVSOFEUPUIFJS IPNFDBHFUPNJNJDUIFIBOEMJOHFĒFDUTJOWPMWFEGPSUIFJOUSPEVDUJPOUPBOPWFMFOWJSPONFOU *OUIFSFQFBUFEIPNFDBHFDPOUSPMHSPVQ HSPVQ3)$ OGFNBMFT XFQFSGPSNFEUIFTBNF IBOEMJOH QSPDFEVSFT CVU XJUIPVU BDUVBM UBJM JODJTJPOT BU U BOE NJOVUFT 0OMZ BU U NJOVUFTBCMPPETBNQMFXBTDPMMFDUFE .

(6) Appendix I *#MPPEDPMMFDUJPOBOEDPSUJDPTUFSPOFEFUFSNJOBUJPO #MPPE TBNQMFT XFSF DPMMFDUFE CZ UBJM JODJTJPO B NFUIPE UIBU BMMPXT FTUJNBUJPO PG CBTBM DPODFOUSBUJPOTPGDPSUJDPTUFSPOFBOENVMUJQMFTBNQMFTGSPNUIFTBNFBOJNBM< >#SJFëZ UIF NPVTFJTQMBDFEPOUIFHSJEUPQPGJUTIPNFDBHF"TNBMMJODJTJPOBUUIFCBTFPGUIFUBJMXJUIB SB[PSCMBEFBMMPXTDPMMFDUJPOPGMCMPPEXJUIJOTFDPOET XJUIPVUBOFTUIFTJB . #MPPEXBTDPMMFDUFEJOμMDBQJMMBSJFT DPBUFEXJUIQPUBTTJVN&%5" BOEDFOUSJGVHFE. GPSNJOVUFTBUSQNBU$1MBTNBXBTTFQBSBUFEGSPNUIFQFMMFUBOETUPSFEBU$ $PSUJDPTUFSPOF XBT NFBTVSFE XJUI B *DPSUJDPTUFSPOF SBEJP JNNVOPBTTBZ 3*" LJU *$/ 1IBSNBDFVUJDBMT*OD /FX:PSL 64" *%BUBBOBMZTJT %BUB BSF QSFTFOUFE BT NFBO 4&. 1MBTNB DPSUJDPTUFSPOF DPODFOUSBUJPOT XFSF BOBMZTFE CZ "/07" GBDUPS TFY XIFO BQQSPQSJBUF XJUI SFQFBUFE NFBTVSFT BU BOE NJOVUFT GBDUPST4&9$0/%*5*0/HSPVQT34$ $:- $:- 48*. BOECZ-4%QPTUIPDUFTU 4UBUJTUJDBMTJHOJêDBODFXBTBDDFQUFEBU1. *3FTVMUT #BTBMDPSUJDPTUFSPOFDPODFOUSBUJPOTXFSFMPXJOCPUINBMFBOEGFNBMF$%NJDF BOEBTFYQFDUFE TJHOJêDBOUMZIJHIFSJOGFNBMFUIBOJONBMFNJDF ' 1NBMFOHNM GFNBMFOHNM . .BMF NJDF TFDSFUFE MFTT DPSUJDPTUFSPOF UIBO GFNBMF NJDF JO SFTQPOTF UP FBDI DIBMMFOHF. ' 1'JHVSF* ѮFNBHOJUVEFPGUIFDPSUJDPTUFSPOFTFDSFUJPOEFQFOEFE PO UIF DIBMMFOHF ' 1 $PSUJDPTUFSPOF DPODFOUSBUJPOT JODSFBTFE XJUI UJNF ' 1 TIPXFEBEJĒFSFOUDPVSTFJONBMFBOEGFNBMFNJDF JOUFSBDUJPOUJNFY TFY' 1 EFQFOEJOHPOUIFFYQFSJNFOUBMDPOEJUJPO JOUFSBDUJPOUJNFYTFYY DPOEJUJPO' 1 ѮFIJHIFTUDPSUJDPTUFSPOFDPODFOUSBUJPOXBTPCTFSWFEGPSUIF 48*.DPOEJUJPOBOEUIFMPXFTUDPODFOUSBUJPOGPSUIFIPNFDBHFDPOUSPM HSPVQ34$ *.BMFNJDF "UUNJOVUFT DPSUJDPTUFSPOFTFDSFUJPOXBTTJHOJêDBOUMZJODSFBTFEJOSFMBUJPOUPUIFEFHSFF PG OPWFMUZ MPXFTU JO UIF IPNF DBHF DPOUSPM 34$ OHNM WT $:- $:- PS 48*. 1 GPMMPXFE CZ NJDF FYQPTFE GPS NJOVUFT UP UIF DZMJOEFS $:- WT$:-PS48*.1 BOENJDFDPOUJOVPVTMZFYQPTFEUPUIFDZMJOEFS $:- OHNM XJUIUIFMBUUFSTJNJMBSUPUIF48*.HSPVQ OHNM "UUNJOVUFT DPSUJDPTUFSPOFDPODFOUSBUJPOTXFSFGVSUIFSJODSFBTFEFRVBMMZIJHIJOHSPVQT34$BOE$:- BOECPUITJHOJêDBOUMZMPXFSUIBO$:-BOE48*. 1 "UUNJOVUFT DPSUJDPTUFSPOF DPODFOUSBUJPOTXFSFBHBJOGVSUIFSJODSFBTFEBOEEJĒFSFECFUXFFOBMMHSPVQT 1UP1 XJUIIJHIFTUDPODFOUSBUJPOTJOUIF48*.HSPVQ .

(7) "QQFOEJY*. 'JHVSF* /PWFMUZJOEVDFEDPSUJDPTUFSPOFSFTQPOTFGPSNBMFBOEGFNBMF$%NJDF NFBTVSFE BOENJOVUFTBѫFS JOJUJBUJPOPGFBDITUSFTTPS'PSBCCSFWJBUJPOTTFF5BCMF*7BMVFTBSFQSFTFOUFEBTNFBO±4&.. *'FNBMFNJDF "UUNJOVUFT DPSUJDPTUFSPOFXBTTJHOJêDBOUMZMPXFSJOHSPVQ34$ OHNM1 UIBOJOHSPVQT$:- OHNM $:- OHNM BOE48*. OHNM "UUNJOVUFT DPSUJDPTUFSPOFGVSUIFSJODSFBTFE CVUXBTTUJMMMPXFTUJO34$ 1 DPNQBSFEUP$:-BOE48*."UUBOENJOVUFT DPSUJDPTUFSPOFPGHSPVQ$:-XBT TJHOJêDBOUMZIJHIFSUIBOPGHSPVQ$:- 1 CVUTJNJMBSUPHSPVQ48*."UUNJOVUFT DPSUJDPTUFSPOFXBTGVSUIFSJODSFBTFE CVUTIPXFEUIFTBNFDPODFOUSBUJPOTJOBMMGPVSHSPVQT . .JNJDLJOHUIFCMPPETBNQMJOHQSPDFEVSFXJUIPVUBDUVBMUBJMJODJTJPOBUUBOENJOVUFT . DPSUJDPTUFSPOFDPODFOUSBUJPOTNFBTVSFEJOGFNBMFNJDF3)$HSPVQBUUNJOVUFT OHNM XFSFTJNJMBSUPCBTBMDPOEJUJPOT. *%JTDVTTJPO 8FEJĒFSFOUJBUFECFUXFFOUIFiEFHSFFPGOPWFMUZuPGBDIBMMFOHFBOEHFOEFSCZBOBMZTJOHUIF QMBTNBDPSUJDPTUFSPOFDPODFOUSBUJPOTJOUIFQSPBDUJWFQIBTFPGUIFHMVDPDPSUJDPJESFMBUFETUSFTT SFTQPOTF< >ѮFEVSBUJPOPGFYQPTVSFUPUIFTBNFDIBMMFOHF JF$:-WFSTVT$:- BĒFDUFE .

(8) Appendix I UIFTMPQFPGUIFTUSFTTSFTQPOTFBTDPSUJDPTUFSPOFTFDSFUJPOXBTJOJUJBUFEGBTUFSѮJTJOJUJBMGBTUFS TFDSFUJPO PG DPSUJDPTUFSPOF SFTVMUFE JO NPSF TFDSFUFE DPSUJDPTUFSPOF BѫFS NJOVUFT UIPVHI DFJMJOHFĒFDUTXFSFBMSFBEZPCTFSWFEBUNJOVUFT0OFNJOVUFGPSDFETXJNNJOH HSPVQ48*. GPMMPXFECZUISFFNJOVUFTVOEFSBIFBUJOHMBNQCFGPSFSFUVSOJOHUPUIFIPNFDBHFSFTVMUFEJOB TJNJMBSSFTQPOTFBTHSPVQ$:- CVUXJUIBIJHIFSBNQMJUVEFJONBMFNJDF . #FTJEFTDIBMMFOHFFĒFDUT GFNBMFNJDFBMTPSFTQPOEFEEJĒFSFOUMZUIBONBMFTUPUIFDIBMMFOHFT. 'FNBMFNJDFTIPXFEIJHIFSDPSUJDPTUFSPOFMFWFMT CBTBMBOEJOSFTQPOTFUPFBDIPGUIFDIBMMFOHFT 'VSUIFSNPSF GFNBMFNJDFTFFNUPSFBDIBQMBUFBVPGDPSUJDPTUFSPOFTFDSFUJPOXIFSFNBMFTEPOPU ѮFTFPCTFSWBUJPOTBSFDPOTJTUFOUXJUIMJUFSBUVSF TIPXJOHUIBUFTUSPHFOTJODSFBTFDPSUJDPTUFSPJE CJOEJOHHMPCVMJOWBMVFTSFTVMUJOHJOIJHIFSUPUBMQMBTNBDPSUJDPTUFSPOFDPODFOUSBUJPOT< > *SSFTQFDUJWFPGHFOEFS UIFNBJOGBDUPSPGHSPVQEJĒFSFODFTJOEJDBUJOHUIFEFHSFFPGOPWFMUZ 34$ $:-$:-48*. XBTEFUFSNJOFECZUIFTIBQFPGUIFTUSFTTSFTQPOTFJOUIFQSPBDUJWF QIBTF . .BSRVF[FUBM<>TIPXFEJOSBUTUIBUUIFSFDPWFSZTQFFEPGUIFTUSFTTTZTUFN JFSFBDUJWF. QIBTF EJĒFST BѫFS BENJOJTUSBUJPO PG FJUIFS GPPU TIPDL PS JNNPCJMJTBUJPO ѮFZ UPP GPVOE B NBYJNBMDPSUJDPTUFSPOFSFTQPOTFNJOVUFTBѫFSJOJUJBUJPOPGCPUIDIBMMFOHFT CVUXFSFVOBCMF UPEJTUJOHVJTICFUXFFOUIFUXPTUSFTTPSTJOUIFQSPBDUJWFQIBTF$PNCJOFEXJUIPVSPCTFSWBUJPOT UIFTFFYQFSJNFOUTJOEJDBUFUIBU EFTQJUFDFJMJOHFĒFDUT EJĒFSFOUTUJNVMJFMJDJUEJĒFSFOUQSPBDUJWF BOESFBDUJWFTUSFTTSFTQPOTFT'VSUIFSNPSF XFTVHHFTUUIBUCPUIQSPBDUJWFBOESFBDUJWFBTQFDUT PGUIFTUSFTTSFTQPOTFNBZSFëFDUUIFJOUFOTJUZPGBHJWFODIBMMFOHF< > . *UXBTTUSJLJOHBOEVOFYQFDUFEUPOPUJDFUIBUFWFOUIFSFQFBUFEMZTBNQMFEDPOUSPMHSPVQ. HSPVQ 34$ FMJDJUFE B TUSFTT SFTQPOTF UIPVHI BOJNBMT SFUVSOFE UP UIFJS IPNF DBHFT 3BUT SFTQPOEFEEJĒFSFOUMZUPSFQFBUFETBNQMJOHWJBUBJMJODJTJPO BTXBTTIPXOCZ'MVUUFSUFUBM<>*O UIFJSTUVEZGSFRVFOUTBNQMJOHPGMPGCMPPEGSPNIBOEMFE8JTUBSSBUTXJUIBOJOUFSWBMPG NJOVUFTPOMZTMJHIUMZFMFWBUFEDPSUJDPTUFSPOFMFWFMT#MPPETBNQMJHWJBUBJMJODJTJPOXBTUIFSFGPSF DPOTJEFSFEOPUUPCFIBSNGVMUPUIFBOJNBM< >*UJTQPTTJCMFUIBUUIFSFTQPOTFXFPCTFSWFEXBT OPUEVFUPUIFQSPDFEVSFPGUBJMJODJTJPOJUTFMG CVUUPUIFGSFRVFODZPGCMPPETBNQMJOH.JDFIBWFB NVDITNBMMFSCMPPEWPMVNFUIBOSBUTBOEXFFTUJNBUFEUIBUPVSNJDFMPTUVQUPPGUIFJSUPUBM CMPPEWPMVNFBѫFSUISFFDPOTFDVUJWFTBNQMFTXJUIJOBUJNFTQBOPGNJOVUFT"DUJWBUJPOPG UIFIZQPUIBMBNJDQJUVJUBSZBESFOBM )1" BYJTIBTCFFOTIPXOUPPDDVSBUBQQSPYJNBUFMZ QMBTNBWPMVNFEFêDJU< >5PBTTFTTUIJTQPTTJCJMJUZBOBJWFHSPVQPGGFNBMFNJDFSFDFJWFE UIFTBNFIBOEMJOHQSPDFEVSFTBTHSPVQ34$ JODMVEJOHBWJSUVBMTBNQMJOHVTJOHUIFCMVOUTJEFPG UIFSB[PSCMBEF CVUXJUIPVUBDUVBMJODJTJPOBOECMPPEMPTTBUBOENJOVUFT HSPVQ3)$XF FYQFDUUIFTBNFSFTVMUTJONBMFT 0OMZBUNJOVUFTBCMPPETBNQMFXBTUBLFOѮFTFIBOEMJOH QSPDFEVSFTCZJUTFMGEJEOPUBĒFDUDJSDVMBUJOHDPSUJDPTUFSPOFDPODFOUSBUJPOT$POTFRVFOUMZ JUJT NPTUMJLFMZUIBUEFIZESBUJPONJHIUFYQMBJOUIFSJTFJODPSUJDPTUFSPOFPCTFSWFEJOHSPVQ34$ &BDIPGUIFDIBMMFOHFTTUJMMFMJDJUFEBGBTUFSBOEIJHIFSDPSUJDPTUFSPOFSFTQPOTFUIBOHSPVQ34$ UIFSFGPSFFĒFDUTPGEJĒFSFOUDIBMMFOHFTPOUIF)1"BYJTSFTQPOTJWFOFTTXFSFTUJMMEFUFDUBCMFVTJOH SFQFBUFETBNQMJOH)PXFWFS DPOTJEFSBUJPOTIBWFUPCFNBEFXIFOEFTJHOJOHBOFYQFSJNFOUPO .

(9) "QQFOEJY* UIFOVNCFSPGDPOTFDVUJWFUBJMTBNQMFTBOEUIFUJNFGSBNFJOXIJDIUIFTFTBNQMFTBSFUBLFO *ODPODMVTJPO UIJTTVSWFZEFNPOTUSBUFTUIBUEJĒFSFOUDIBMMFOHFTPSWBSZJOHFYQPTVSFUJNFT UPUIFTBNFDIBMMFOHFDBVTFTEJĒFSFOUSFBDUJPOQBUUFSOTJOCPUINBMFBOEGFNBMF$%NJDFѮFTF FĒFDUTXFSFPCTFSWFEJOUIFQSPBDUJWFQIBTFPGUIFDPSUJDPTUFSPOFTUSFTTSFTQPOTFBOEJOEJDBUF QMBTUJDJUZGPSDPQJOHXJUIBOPWFMFOWJSPONFOU. *"DLOPXMFEHFNFOUT ѮJTTUVEZXBTTVQQPSUFECZUIF/FUIFSMBOET0SHBOJTBUJPOGPS4DJFOUJêD3FTFBSDI /80 BOE BOE/80m45*(0//%3'5FDIOJDBMBTTJTUBODFPG .'MVUUFSU .WBOEFS.BSLBOE.7JTTFSJTHSBUFGVMMZBDLOPXMFEHFE. *3FGFSFODFT . . #BUT4 ѮPVNBT+- -PSEJ# 5POPO.$ -BMPOEF3BOE$BTUPO+ ѮFFĒFDUTPGB NJMETUSFTTPSPOTQPOUBOFPVTBMUFSOBUJPOJONJDF#FIBW#SBJO3FT EF,MPFU&3BOE3FVM+. 'FFECBDLBDUJPOBOEUPOJDJOëVFODFPGDPSUJDPTUFSPJET POCSBJOGVODUJPOBDPODFQUBSJTJOHGSPNUIFIFUFSPHFOFJUZPGCSBJOSFDFQUPSTZTUFNT 1TZDIPOFVSPFOEPDSJOPMPHZ EF,MPFU&3 7SFVHEFOIJM& 0JU[M.4BOE+PFMT. #SBJODPSUJDPTUFSPJESFDFQUPS CBMBODFJOIFBMUIBOEEJTFBTF&OEPDS3FW %VSTDIMBH. 8VSCFM) 4UBVĒBDIFS.BOE7PO)PMTU% 3FQFBUFECMPPEDPMMFDUJPO JOUIFMBCPSBUPSZNPVTFCZUBJMJODJTJPONPEJêDBUJPOPGBOPMEUFDIOJRVF1IZTJPM#FIBW 'MVUUFSU. %BMN4BOE0JU[M.4 "SFêOFENFUIPEGPSTFRVFOUJBMCMPPETBNQMJOH CZUBJMJODJTJPOJOSBUT-BC"OJN (BSDJB " .BSUJ 0 7BMMFT " %BM ;PUUP 4 BOE "SNBSJP " 3FDPWFSZ PG UIF IZQPUIBMBNJDQJUVJUBSZBESFOBM SFTQPOTF UP TUSFTT &ĒFDU PG TUSFTT JOUFOTJUZ TUSFTT EVSBUJPOBOEQSFWJPVTTUSFTTFYQPTVSF/FVSPFOEPDSJOPMPHZ (SPPUFOEPSTU+ EF,MPFU&3 %BMN4BOE0JU[M.4 3FWFSTBMPGDPHOJUJWFEFêDJU PGBQPMJQPQSPUFJO&LOPDLPVUNJDFBѫFSSFQFBUFEFYQPTVSFUPBDPNNPOFOWJSPONFOUBM FYQFSJFODF/FVSPTDJFODF (SPPUFOEPSTU + &OUIPWFO - %BMN 4 EF ,MPFU &3 BOE 0JU[M .4 *ODSFBTFE DPSUJDPTUFSPOFTFDSFUJPOBOEFBSMZPOTFUPGDPHOJUJWFEFDMJOFJOGFNBMFBQPMJQPQSPUFJO& LOPDLPVUNJDF#FIBW#SBJO3FT (SPPUFOEPSTU+ 0JU[M.4 %BMN4 &OUIPWFO- 4DIBDIOFS. EF,MPFU&3BOE4BOEJ$ 4USFTTBMMFWJBUFTSFEVDFEFYQSFTTJPOPGDFMMBEIFTJPONPMFDVMFT /$". - BOE EFêDJUTJOMFBSOJOHBOEDPSUJDPTUFSPOFSFHVMBUJPOPGBQPMJQPQSPUFJO&LOPDLPVUNJDF&VS +/FVSPTDJ )FOOFTTZ.#BOE-FWJOF4 4FOTJUJWFQJUVJUBSZBESFOBMSFTQPOTJWFOFTTUPWBSZJOH JOUFOTJUJFTPGQTZDIPMPHJDBMTUJNVMBUJPO1IZTJPM#FIBW ,BOU (+ &HHMFTUPO 5 -BOENBO3PCFSUT - ,FOJPO $$ %SJWFS ($ BOE .FZFSIPĒ +- )BCJUVBUJPO UP SFQFBUFE TUSFTT JT TUSFTTPS TQFDJêD 1IBSNBDPM #JPDIFN #FIBW ,PPMIBBT+. EF#PFS4' EF3VUUFS"+ .FFSMP1BOE4HPJGP" 4PDJBMTUSFTTJOSBUT BOENJDF"DUB1IZTJPM4DBOE4VQQM. .

(10) Appendix I .BSRVF[ $ #FMEB 9 BOE "SNBSJP " 1PTUTUSFTT SFDPWFSZ PG QJUVJUBSZBESFOBM IPSNPOFTBOEHMVDPTF CVUOPUUIFSFTQPOTFEVSJOHFYQPTVSFUPUIFTUSFTTPS JTBNBSLFS PGTUSFTTJOUFOTJUZJOIJHIMZTUSFTTGVMTJUVBUJPOT#SBJO3FT .D$PSNJDL$. -JOLSPVN8 4BMMJOFO#+BOE.JMMFS/8 1FSJQIFSBMBOEDFOUSBM TFYTUFSPJETIBWFEJĒFSFOUJBMFĒFDUTPOUIF)1"BYJTPGNBMFBOEGFNBMFSBUT4USFTT .D&XFO#4BOE4BQPMTLZ3. 4USFTTBOEDPHOJUJWFGVODUJPO$VSS0QJO/FVSPCJPM 1JKMNBO'5BOEWBO3FF+. 1IZTJDBMCVUOPUFNPUJPOBMTUSFTTJOEVDFTBEFMBZJO CFIBWJPVSBMDPQJOHSFTQPOTFTJOSBUT#FIBW#SBJO3FT 4BQPMTLZ3. 3PNFSP-.BOE.VODL"6 )PXEPHMVDPDPSUJDPJETJOëVFODFTUSFTT SFTQPOTFT *OUFHSBUJOH QFSNJTTJWF TVQQSFTTJWF TUJNVMBUPSZ BOE QSFQBSBUJWF BDUJPOT &OEPDS3FW 4DINJEU.7 0JU[M.4 -FWJOF4BOEEF,MPFU&3 ѮF)1"TZTUFNEVSJOHUIF QPTUOBUBMEFWFMPQNFOUPG$%NJDFBOEUIFFĒFDUTPGNBUFSOBMEFQSJWBUJPO#SBJO3FT %FW#SBJO3FT 4JJUFSJ1, .VSBJ+5 )BNNPOE(- /JTLFS+" 3BZNPVSF8+BOE,VIO38 ѮF TFSVNUSBOTQPSUPGTUFSPJEIPSNPOFT3FDFOU1SPH)PSN3FT 5BOJNVSB4.BOE8BUUT"( $PSUJDPTUFSPOFNPEVMBUJPOPG"$5)TFDSFUPHPHVF HFOFFYQSFTTJPOJOUIFQBSBWFOUSJDVMBSOVDMFVT1FQUJEFT WBO EFO #FSH $- -BNCFSUT 33 8PMUFSJOL ( 8JFHBOU 7. BOE WBO 3FF +. &NPUJPOBMBOEGPPUTIPDLTUJNVMJJOEVDFEJĒFSFOUJBMMPOHMBTUJOHCFIBWJPVSBMFĒFDUTJO SBUTJOWPMWFNFOUPGPQJPJET#SBJO3FT 8BUUT"( 4BODIF[8BUUT(BOE,FMMZ"# %JTUJODUQBUUFSOTPGOFVSPQFQUJEFHFOF FYQSFTTJPO JO UIF MBUFSBM IZQPUIBMBNJD BSFB BOE BSDVBUF OVDMFVT BSF BTTPDJBUFE XJUI EFIZESBUJPOJOEVDFEBOPSFYJB+/FVSPTDJ 8JOEMF3+ 'PSTMJOH.- 4NJUI$1BOE#BMNFOU3+ 1BUUFSOTPGOFVSPIZQPQIZTJBM IPSNPOFSFMFBTFEVSJOHEFIZESBUJPOJOUIFSBU+&OEPDSJOPM 8PSLFM +0 0JU[M .4 'MVUUFSU . -FTTDIFS ) ,BSTTFO ". BOE EF ,MPFU &3 %JĒFSFOUJBM BOE BHFEFQFOEFOU FĒFDUT PG NBUFSOBM EFQSJWBUJPO PO UIF IZQPUIBMBNJD QJUVJUBSZBESFOBMBYJTPGCSPXOOPSXBZSBUTGSPNZPVUIUPTFOFTDFODF+/FVSPFOEPDSJOPM 8PSLFM+0 0JU[M.4 -FEFCPFS"BOEEF,MPFU&3 ѮF#SPXO/PSXBZSBUEJTQMBZT FOIBODFETUSFTTJOEVDFE"$5)SFBDUJWJUZBUEBZBѫFSINBUFSOBMEFQSJWBUJPOBU EBZ#SBJO3FT%FW#SBJO3FT . .

(11)

(12)

(13)

(14)

No results found