Early life experience : neuroendocrine adaptations to maternal absence
Hele tekst
(2) 5IFQJUVJUBSZBESFOBMBYJTPGUIF $%NPVTFJOGBOUEFTFOTJUJTFT UPSFQFBUFENBUFSOBM TFQBSBUJPOT CVUSFNBJOT IJHIMZSFTQPOTJWFUPTUSFTT. $IBQUFS. -&OUIPWFO .40JU[M /,POJOH .WBOEFS.BSL&3EF,MPFU.
(3)
(4) Chapter 2. "CTUSBDU 1SFWJPVTTUVEJFTIBWFTIPXOUIBUBTJOHMFFQJTPEFPGIPVSTTFQBSBUJPOGSPNUIFNPUIFSQSPEVDFT JO UIF NPVTF QVQ B QSPGPVOE IZQPUIBMBNJDQJUVJUBSZBESFOBM )1" SFTQPOTF *O UIJT TUVEZ XFFYBNJOFEJO$%NJDFXIFUIFSSFQFBUFEEBJMZCPVUTPGIPVSTNBUFSOBMTFQBSBUJPOXPVME SFTVMUJOBQFSTJTUFOUFMFWBUJPOPGDPSUJDPTUFSPOF'PSUIJTQVSQPTFUIFFĒFDUPGSFQFBUFEIPVST TFQBSBUJPOTGSPNQPTUOBUBMEBZTUPXBTNFBTVSFEPOCBTBMBOETUSFTTJOEVDFEMFWFMTPG"$5) BOEDPSUJDPTUFSPOFBTXFMMBTPOUIFFYQSFTTJPOPG)1"NBSLFSTBOEDGPTN3/"JOUIFCSBJO $JSDVMBUJOHMFWFMTPGHMVDPTFBOEHISFMJOXFSFBMTPNFBTVSFE . ѮFEBUBTIPXUIBUUIFJOGBOUTJOJUJBMJNNFEJBUF)1"SFTQPOTFUPIPVSTTFQBSBUJPOXBT. FMJNJOBUFEXIFONBUFSOBMTFQBSBUJPOTBSFSFQFBUFEUIFOFYUUXPEBZT%FTQJUFUIFBCTFODFPGBO )1"SFTQPOTFUPSFQFBUFETFQBSBUJPOT UIFNBUFSOBMMZEFQSJWFENPVTFDPOUJOVFEUPSFTQPOEUP OPWFMUZFYQPTVSF*GUIFSFQFBUFENBUFSOBMTFQBSBUJPOTXFSFDPNCJOFEFBDIUJNFXJUIOPWFMUZ FYQPTVSFUIFSFTQPOTFPGDPSUJDPTUFSPOFTFDSFUJPOSFMBUJWFUP"$5)XBTFOIBODFEѮFTFFĒFDUTPG TFQBSBUJPOPOUIF)1"BYJTXFSFSFëFDUFECZDGPTN3/"FYQSFTTJPOJOUIFQBSBWFOUSJDVMBSOVDMFVT PGUIFIZQPUIBMBNVT 17/ CVUOPUJODPSUFYPSUIBMBNVTDGPTN3/"JOUIF17/TIPXFEB QSPGPVOESFTQPOTFUPBTJOHMFTFQBSBUJPOBOETVCTFRVFOUEFTFOTJUJTBUJPOUPSFQFBUFETFQBSBUJPOT CVUSFNBJOFESFTQPOTJWFUPOPWFMUZFYQPTVSF#BTBMDJSDVMBUJOHMFWFMTPGDPSUJDPTUFSPOFBOE"$5) XFSFQFSTJTUFOUMZTVQQSFTTFEIPVSTBѫFSTFQBSBUJPO BOFĒFDUUIBUBMTPUSBOTJFOUMZPDDVSSFEGPS $3)N3/"JOUIF17/1JUVJUBSZ10.$N3/"BOEUIFHMVDPDPSUJDPJESFDFQUPS (3 N3/" FYQSFTTJPOJOIJQQPDBNQVTPSIZQPUIBMBNVTEJEOPUDIBOHF CVUUIFNJOFSBMPDPSUJDPJESFDFQUPS .3 N3/"FYQSFTTJPOJOUIFIJQQPDBNQBMEFOUBUFHZSVTTIPXFEBQSPHSFTTJWFJODSFBTFXJUI SFQFBUFETFQBSBUJPOT$JSDVMBUJOHHISFMJOJODSFBTFEBOEHMVDPTFMFWFMTEFDSFBTFEBѫFSUIFTJOHMF BTXFMMBTUIFUIJSETFQBSBUJPOBOEUIVTEJEOPUSFëFDUUIFEFTFOTJUJTBUJPOPGUIFDPSUJDPTUFSPOF SFTQPOTF . *O DPODMVTJPO XIJMF UIF JOGBOUT JOJUJBM )1" BYJT SFTQPOTF UP SFQFBUFE NBUFSOBM BCTFODF. SFBEJMZ EFTFOTJUJTFT BOE B TUBUF PG IZQPDPSUJDJTN JT QSPEVDFE UIF )1" BYJT SFNBJOT IJHIMZ SFTQPOTJWFUPNJMETUSFTTPST. *OUSPEVDUJPO %VSJOHFBSMZQPTUOBUBMMJGFSBUTBOENJDFIBWFBTUSFTTIZQPSFTQPOTJWFQFSJPE 4)31 QPTUOBUBM EBZ QOE JOSBUT< >BOEQOEJONJDF<> ѮJTQFSJPEJTDIBSBDUFSJTFECZ TUBCMFBOEMPXDJSDVMBUJOHCBTBMMFWFMTPGDPSUJDPTUFSPOFBOENPTU NJME TUSFTTPSTUIBUUSJHHFSB QSPGPVOE"$5)BOEDPSUJDPTUFSPOFSFTQPOTFBUBEVMUIPPEEPTPPOMZXFBLMZEVSJOHUIF4)31 ѮFIZQPSFTQPOTJWFOFTTPGUIF)1"BYJTJTEVFUPUIFQSFTFODFPGUIFNPUIFS< > XIJDIDBO CFEFNPOTUSBUFECZTFQBSBUJPOPGQVQTGSPNNBUFSOBMDBSFGPSBQSPMPOHFEQFSJPEPGUJNF"ѫFS GPSFYBNQMF BTJOHMFIPVSTNBUFSOBMEFQSJWBUJPO CBTBMMFWFMTPG"$5)BOEDPSUJDPTUFSPOFBSF JODSFBTFE.PSFPWFS UIFTFDSFUJPOPGQJUVJUBSZBESFOBMIPSNPOFTIBTOPXCFDPNFSFTQPOTJWFUP .
(5) $IBQUFS NJMETUSFTTPST SFTVMUJOHJOBGVSUIFSSJTFJOQMBTNB"$5)BOEDPSUJDPTUFSPOF< >ѮFTF OFVSPFOEPDSJOFFĒFDUTJOSFTQPOTFUPNBUFSOBMTFQBSBUJPOBSFBMSFBEZEFUFDUBCMFBTFBSMZBTBѫFS IPVSTPGNBUFSOBMTFQBSBUJPO CVUTJHOJêDBOUMZHSFBUFSBUBOEIPVST< > . &WFS TJODF BEWFSTF FBSMZ MJGF FWFOUT IBWF CFFO JEFOUJêFE BT B NBKPS SJTL GBDUPS GPS UIF. EFWFMPQNFOU PG EFQSFTTJPO BOE BOYJFUZ EJTPSEFST JO IVNBOT < > SPEFOUT TFQBSBUFE BT QVQT GSPN NBUFSOBM DBSF IBWF CFFO XJEFMZ VTFE BT MBCPSBUPSZ NPEFM UP TUVEZ UIF VOEFSMZJOH NFDIBOJTN< >*OQBSUJDVMBS UIFCSJFGEBJMZTFQBSBUJPOPGNPUIFSBOEQVQT BQSPDFEVSF DBMMFEIBOEMJOH XBTTIPXOUPSFTVMUJOBQFSTJTUFOUBUUFOVBUJPOPG)1"SFTQPOTJWFOFTTJOMBUFS MJGF QSFTVNBCMZ UISPVHI B NFDIBOJTN JOWPMWJOH NBUFSOBM DBSF FĒFDUT PO NFUIZMBUJPO PG UIF HMVDPDPSUJDPJESFDFQUPS (3 JOUIFEFWFMPQJOHCSBJO< >*ODPOUSBTU SFQFBUFETFQBSBUJPOT GPS BQQSPYJNBUFMZ IPVST BOE MPOHFS VTVBMMZ XFSF TIPXO UP QSPEVDF JODSFBTFE )1" BYJT SFTQPOTJWFOFTTBOECFIBWJPVSBMGFBSGVMOFTTJOBEVMUSBUT< >BOENJDF< > . ѮF NFDIBOJTN VOEFSMZJOH UIFTF MBTUJOH FĒFDUT PG QSPMPOHFE EBJMZ TFQBSBUJPOT JT TUJMM. QPPSMZ VOEFSTUPPE .BUFSOBM DBSF JT JNQPSUBOU CVU DBOOPU CF UIF POMZ GBDUPS < > %BJMZ BDUJWBUJPOPGUIF)1"BYJT BTJOEVDFECZSFQFBUFEBOEQSPMPOHFETFQBSBUJPOPGNPUIFSBOEQVQT IBTJOEFFECFFODPOTJEFSFEBOBEEJUJPOBMGBDUPS CFDBVTFJUXJMMSFTVMUJOFYQPTVSFUPJODSFBTFE MFWFMTPGDPSUJDPTUFSPOF QBSUJDVMBSMZBѫ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ѫFSIPVSTPGNBUFSOBMTFQBSBUJPO<>ѮFSFGPSF XFVTFEIPVSTPGNBUFSOBM BCTFODFUPEFUFSNJOFUIFJNNFEJBUFFĒ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
(6) Chapter 2. .BUFSJBMTBOE.FUIPET "OJNBMT 0ĒTQSJOHPG$%NJDF PCUBJOFEGSPN$IBSMFT3JWFS ѮF/FUIFSMBOET XBTVTFE"ѫFSBIBCJUVBUJPO QFSJPEPGUXPXFFLT UISFFGFNBMFTXFSFNBUFEXJUIPOFNBMFJOUZQFQPMZDBSCPOBUFDBHFT DN DPOUBJOJOHTBXEVTUCFEEJOHBOEUJTTVFGPPE 43." )PQF'BSNT ѮF/FUIFSMBOET BOE XBUFS DPOUBJOJOH)$M BEMJCJUVNMJHIUTPOGSPNUPIPVSTJOBUFNQFSBUVSF ± °$ BOEIVNJEJUZ DPOUSPMMFESPPN1SFHOBOUGFNBMFTXFSFJOEJWJEVBMMZUSBOTGFSSFEUP DMFBOUZQFQPMZDBSCPOBUFDBHFTDPOUBJOJOHTBXEVTUCFEEJOHBOEUJTTVFUPQSPWJEFOFTUCVJMEJOH NBUFSJBMEVSJOHUIFMBTUXFFLPGHFTUBUJPO'FNBMFTXFSFDIFDLFEGPSMJUUFSTEBJMZCFUXFFO BOEIPVST*GMJUUFSTXFSFQSFTFOU UIFEBZPGCJSUIGPSUIBUMJUUFSXBTUIFOEFêOFEBTQPTUOBUBM EBZ QOE 0OUIFEBZBѫFSQBSUVSJUJPO QOE MJUUFSTXFSFDVMMFEUPFJHIUIFBMUIZQVQT GPVS NBMFTBOEGPVSGFNBMFT . "OJNBM FYQFSJNFOUT XFSF BQQSPWFE CZ UIF -PDBM $PNNJUUFF GPS "OJNBM )FBMUI &UIJDT. BOE3FTFBSDIPG-FJEFO6OJWFSTJUZBOEDBSSJFEPVUJOBDDPSEBODFXJUI&VSPQFBO$PNNVOJUJFT $PVODJM%JSFDUJWF&&$ѮFQSPUPDPMTXFSFBQQSPWFECZUIF"OJNBM$BSF$PNNJUUFFPG UIF'BDVMUZPG.FEJDJOF -FJEFO6OJWFSTJUZ -FJEFO ѮF/FUIFSMBOET 4FQBSBUJPOQSPDFEVSFT .PUIFSTOVSTJOHMJUUFSTTFMFDUFEGPSNBUFSOBMTFQBSBUJPOXFSFSFNPWFEGSPNUIFJSDBHFBOEQMBDFE JODMFBOUZQFQPMZDBSCPOBUFDBHFTBUIPVSTѮFIPNFDBHFDPOUBJOJOHUIFQVQTSFNBJOFE JOUIFBEKBDFOUSPPNPOBIFBUJOHQBE °$ UPDPOUSPMGPSQVQCPEZUFNQFSBUVSF"ѫFS IPVST BUIPVST UIFNPUIFSTXFSFSFVOJUFEXJUIUIFJSQVQTBOEMFѫVOEJTUVSCFEVOUJMUIF OFYUTFQBSBUJPOQFSJPEPSVOUJMUFTUJOH$POUSPMMJUUFSTXFSFMFѫVOEJTUVSCFE /PWFMUZFYQPTVSF 'PSOPWFMUZFYQPTVSFêSTUUIFNPUIFSXBTSFNPWFEGSPNUIFIPNFDBHFѮFO JOBOBEKBDFOU SPPN UIFQVQTXFSFJOEJWJEVBMMZQMBDFEJODMFBOOPWFMDBHFTUPJOEVDFOPWFMUZTUSFTTѮFTFOPWFM DBHFT XFSF QMBDFE PO IFBUJOH QBET $ UP DPOUSPM GPS QVQ CPEZ UFNQFSBUVSF "ѫFS NJOVUFT UIFQVQTXFSFFJUIFSQMBDFECBDLXJUIUIFJSNPUIFSTJOUIFIPNFDBHFPSTBDSJêDFE . *GOPWFMUZFYQPTVSFUPPLQMBDFPOBUFTUJOHEBZ GPVSQVQT UXPNBMFTBOEUXPGFNBMFT XFSF. TBDSJêDFEJNNFEJBUFMZBѫFSSFNPWBMPGUIFEBNGSPNUIFIPNFDBHFѮFSFNBJOJOHQVQTXFSF UIFOUSBOTGFSSFEUPOPWFMDBHFTBOETBDSJêDFENJOVUFTMBUFSGPSUFTUJOH"MMPUIFSQSPDFEVSFT SFNBJOFEDPOTUBOU &YQFSJNFOUBMEFTJHOT &YQFSJNFOU*5PJOWFTUJHBUFUIFJNNFEJBUF DVNVMBUJWFFĒFDUTPGSFQFBUFETFQBSBUJPOTNJDFXFSF TFQBSBUFEPODF QOE UXJDF QOETBOE PSUISFFUJNFT QOET BOE GSPNNBUFSOBMDBSF GPSBQFSJPEPGIPVST#FGPSF BUIPVST BTXFMMBTBUUIFFOEPGFBDITFQBSBUJPOTFTTJPO BU IPVST NJDF XFSF TBDSJêDFE GPS EFUFSNJOBUJPO PG QFSJQIFSBM BOE DFOUSBM NBSLFST PG .
(7) $IBQUFS UIF)1"BYJT"UQOEPOFHSPVQPGBOJNBMTTFSWFEBTCBTBMNFBTVSFNFOUGPSCPUIUIFOPO TFQBSBUFEBOEêSTUUJNFTFQBSBUFEBOJNBMT/POTFQBSBUFEBOJNBMTXFSFPOMZNFBTVSFEBU IPVST TJODFOPDJSDBEJBOFĒFDUTXFSFEFUFDUFEGPSDPSUJDPTUFSPOFJOi&YQFSJNFOU*uѮFêYFE GBDUPSTXFSF53&"5.&/5 OPOTFQBSBUFETFQBSBUFE CBTBM TFQBSBUFE
(8) IPVST. BOE"(& QOET BOE . &YQFSJNFOU**5PUFTUXIFUIFSQVQTBUQOEFYIJCJUFEBOBEVMUMJLFSFTQPOTFUPUIFUIJSE. TFQBSBUJPO TFTTJPO BU IPVST NPUIFST XFSF SFNPWFE GSPN UIFJS OFTUT UP JOJUJBUF NBUFSOBM TFQBSBUJPOBOEUIFIPNFDBHFTDPOUBJOJOHUIFMJUUFSTXFSFQMBDFEPOBIFBUJOHQBEѮFSFBѫFS FWFSZIPVSCMPPETBNQMFTXFSFUBLFOVQUPIPVSTPGNBUFSOBMBCTFODF'PSUIJTQVSQPTF UP NJOJNJTFOFTUFĒFDUT GSPNFBDIMJUUFSGPVSQVQT UXPNBMFTBOEUXPGFNBMFT XFSFTBDSJêDFE BUUXPEJĒFSFOUUJNFQPJOUT"UIPVST CBTBMUJNFQPJOU POFHSPVQPGBOJNBMTTFSWFEBTB CBTBMNFBTVSFNFOUGPSCPUIOPOTFQBSBUFEBOEêSTUUJNFTFQBSBUFEBOJNBMT'JYFEGBDUPSTXFSF 53&"5.&/5 OPOTFQBSBUFEBUQOETJOHMFTFQBSBUJPOBUQOEUSJQMFTFQBSBUJPOBUQOET BOE UFTUFEBUQOE BOE5*.& CBTBM IPVST BOE BOEIPVSTPGNBUFSOBM TFQBSBUJPO . &YQFSJNFOU***5PEFUFSNJOFXIFUIFSUIFEFTFOTJUJTBUJPOUPNBUFSOBMTFQBSBUJPOPCTFSWFE. JOi&YQFSJNFOU*BOE**uXBTBOBHFTQFDJêDFĒFDUXFSFQFBUFEUIJTFYQFSJNFOU CVUUIJTUJNF JODMVEFE BO IPVST NBUFSOBM TFQBSBUJPO BQQMJFE UP TFQBSBUF HSPVQT PG OBÐWF BOJNBMT BU QOET BOE0OUFTUEBZT GPVSQVQTGSPNFBDIMJUUFS UXPNBMFTBOEUXPGFNBMFT XFSFTBDSJêDFE JNNFEJBUFMZQSPWJEJOHBCBTBMTBNQMF BUIPVST ѮFPUIFSGPVSQVQTXFSFTBDSJêDFEBѫFS IPVSTPGNBUFSOBMTFQBSBUJPO BU ѮFêYFEGBDUPSTXFSF53&"5.&/5 TJOHMFTFQBSBUJPO BUQOET PSEPVCMFTFQBSBUJPOBUQOETBOE UFTUFEBUQOEUSJQMFTFQBSBUJPOTBUQOE BOE UFTUFEBUQOE BOE5*.& CBTBMBOEIPVSTNBUFSOBMTFQBSBUJPO . &YQFSJNFOU*7XBTEFTJHOFEUPUFTUXIFUIFSQVQTXFSFBCMFUPSFTQPOEUPOPWFMUZTUSFTTXJUI. JODSFBTFEDPSUJDPTUFSPOFBOE"$5)EJSFDUMZBѫFSBêSTU TFDPOEPSUIJSEQFSJPEPGTFQBSBUJPO 0OUFTUEBZTGPVSQVQTGSPNFBDIOFTU UXPNBMFTBOEUXPGFNBMFT XFSFTBDSJêDFEIPVSTBѫFS NBUFSOBMTFQBSBUJPO QSPWJEJOHBiTFQBSBUFEuTBNQMF BUIPVST ѮFSFNBJOJOHQVQTXFSF UIFOFYQPTFEUPOPWFMUZBOETBDSJêDFENJOVUFTMBUFS'JYFEGBDUPSTXFSF53&"5.&/5 OPO TFQBSBUFEBUQOETJOHMFTFQBSBUJPOBUQOEUSJQMFTFQBSBUJPOBUQOET BOE UFTUFEBUQOE BOE5*.& CBTBM FJUIFSBUIPVSTPSBѫFSIPVSTPGNBUFSOBMTFQBSBUJPOBUIPVST NJOVUFTJOEJWJEVBMOPWFMUZFYQPTVSF . &YQFSJNFOU7ѮFPCKFDUJWFTXFSF UPEFUFSNJOFUIFSFTQPOTFPGDPSUJDPTUFSPOF HISFMJO. BOEHMVDPTFVOEFSUIFDPOEJUJPOTPGEFTFOTJUJTBUJPOUPSFQFBUFENBUFSOBMTFQBSBUJPOTBOE UP EFUFSNJOFBDUJWBUJPOPGDFOUSBMIZQPUIBMBNJDCSBJOBSFBTJOSFTQPOTFUPNBUFSOBMTFQBSBUJPOBOE OPWFMUZCZNFBTVSJOHDGPTN3/"FYQSFTTJPO1VQTXFSFTBDSJêDFEBѫFSUIFêSTUPSUIFUIJSE QFSJPEPGIPVSTPGTFQBSBUJPOGSPNUIFEBNJOUIFBCTFODFPSQSFTFODFPGBEEJUJPOBMOPWFMUZ FYQPTVSF/POTFQBSBUFEBOJNBMTTBDSJêDFEBUUIFTFUJNFQPJOUTTFSWFEBTCBTBMDPOUSPMT1MBTNB DPSUJDPTUFSPOFMFWFMTXFSFEFUFSNJOFEJOBMMUIFTFHSPVQT#MPPEHMVDPTFBOEHISFMJOMFWFMTXFSF NFBTVSFEPOMZVOEFSCBTBMDPOEJUJPOTBOEBѫFSIPVSTPGNBUFSOBMBCTFODF'JYFEGBDUPSTXFSF .
(9) Chapter 2 UIFO53&"5.&/5 TJOHMFTFQBSBUJPOBUQOEBOEUSJQMFTFQBSBUJPOBUQOET BOE UFTUFE BU QOE BOE 5*.& CBTBM BOE TFQBSBUFE &YQSFTTJPO MFWFMT PG DGPT BOE $3) N3/" XFSF NFBTVSFEJOUIF17/BOEPG/1:N3/"JOUIFBSDVBUFOVDMFVTJOBMMUSFBUNFOUHSPVQT'JYFE GBDUPST TJNJMBSBTGPSDPSUJDPTUFSPOF XFSFUIFO53&"5.&/5 TU4&1BOESE4&1 BOE5*.& CBTBM TFQBSBUFE OPWFMUZ $PMMFDUJPOPGCMPPEQMBTNBBOECSBJOT "UTQFDJêFEUJNFQPJOUTEFTDSJCFEJO&YQFSJNFOUT*UP*7 TFFi&YQFSJNFOUBMEFTJHOTu BOJNBMT XFSF TBDSJêDFE CZ EFDBQJUBUJPO BOE USVOL CMPPE XBT DPMMFDUFE JOEJWJEVBMMZ JO NM &%5" DPBUFENJDSPDFOUSJGVHFUVCFT#MPPETBNQMFTXFSFLFQUPOJDFBOEDFOUSJGVHFEGPSNJOVUFTBU SQNBU$1MBTNBXBTUIFOUSBOTGFSSFEUPNM&QQFOEPSGUVCFTBOETUPSFEGSP[FOBU $VOUJMEFUFSNJOBUJPOPGDPSUJDPTUFSPOFBOE"$5)DPODFOUSBUJPOT . #MPPE HMVDPTF MFWFMT XFSF NFBTVSFE "DDV$IFDL $PNQBDU 3PDIF (FSNBOZ VTJOH B. ESPQMFUPGUSVOLCMPPEMFѫPOUIFIFBEPSCPEZ"ѫ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êYFE JO QBSBGPSNBMEFIZEF HMVUBSBMEFIZEF BOE UIFSFBѫFS BDFUZMBUFE JO .
(10) $IBQUFS BDFUJDBOIZESJEFJO.USJFUIBOPMBNJOF)$M4VCTFRVFOUMZ CSBJOTFDUJPOTXFSFEFIZESBUFEJO JODSFBTJOHDPODFOUSBUJPOTPGFUIBOPM5JTTVFTFDUJPOT CSBJOTFDUJPOTQFSTMJEF XFSFTBUVSBUFE XJUIʍMPGIZCSJEJTBUJPOCVĒFS N.5SJT)$M Q) GPSNBNJEF N./B$M N.&%5" Q) ¤%FJOIBSEUT ʍHNMZFBTUUSBOTGFS3/" ʍMNMUPUBM3/" NHNM TBMNPO TQFSN %/" EFYUSBO TVMGBUF N. EJUIJPUISFJUPM 4%4 BOE TPEJVN UIJPTVMGBUF DPOUBJOJOH BQQSPYJNBUFMZ Y DQN 4VMQIVS MBCFMMFE SJCPOVDMFPUJEF QSPCF#SBJOTFDUJPOTXFSFDPWFSTMJQQFEBOEJODVCBUFEPWFSOJHIUBU$ѮFGPMMPXJOHEBZUIF TFDUJPOTXFSFSJOTFEJO¤44$ USFBUFEXJUI3/"TF" NHNM BOEXBTIFEJOJODSFBTJOHMZ TUSJOHFOU44$TPMVUJPOTBUSPPNUFNQFSBUVSF'JOBMMZ TFDUJPOTXFSFXBTIFEJO¤44$GPS NJOVUFTBU$BOEEFIZESBUFEUISPVHIJODSFBTJOHDPODFOUSBUJPOTPGFUIBOPM 'PSEFUFSNJOJOHDGPTN3/"FYQSFTTJPO 17/ JOTJUVIZCSJEJTBUJPOTVTJOH1IPTQIPSMBCFMMFE PMJHPOVDMFPUJEF QSPCFT XFSF QFSGPSNFE BT EFTDSJCFE QSFWJPVTMZ <> 4MJEFT XFSF PQQPTFE UP ,PEBL#JPNBY.3êMN &BTUNBO,PEBL$P 3PDIFTUFS /: BOEEFWFMPQFE . "VUPSBEJPHSBQITXFSFEJHJUJTFEBOESFMBUJWFMFWFMTPGN3/"FYQSFTTJPOXFSFEFUFSNJOFECZ. DPNQVUFSBTTJTUFEPQUJDBMEFOTJUPNFUSZ "OBMZ4*4 4Pѫ*NBHJOH4ZTUFN(NC) ѮFBWFSBHF EFOTJUZPGNFBTVSFNFOUTXBTUBLFOGPSFBDIBOJNBM 4UBUJTUJDBMBOBMZTJT *OFBDIFYQFSJNFOUFWFSZUSFBUNFOUDPOTJTUFEPGUISFFMJUUFSTQFSBHFHSPVQ O %BUBXFSF BOBMZTFE CZ BOBMZTJT PG WBSJBODF "/07" ѮF MFWFM PG TJHOJêDBODF XBT TFU BU 1 8IFO BQQSPQSJBUF UIJT XBT GPMMPXFE CZ 5VLFZT QPTU IPD DPNQBSJTPOT ѮF JOJUJBM BOBMZTFT JO FBDI FYQFSJNFOUJODMVEFETFYBTBGBDUPS CVUCFDBVTFJUXBTOPUBTJHOJêDBOUGBDUPSJOBOZFYQFSJNFOU EBUBXFSFDPMMBQTFEBDSPTTUIJTWBSJBCMF%BUBBSFQSFTFOUFEBTNFBO±4&.. 3FTVMUT *O&YQFSJNFOU*UIFJNNFEJBUF DVNVMBUJWF FĒFDUTXFSFJOWFTUJHBUFEPGPOF QOE UXP QOETBOE PSUISFF QOET BOE UJNFTIPVSTEBJMZNBUFSOBMTFQBSBUJPO T POUIFCBTBM EFWFMPQNFOUPGCPUIDFOUSBMBOEQFSJQIFSBM)1"BYJTNBSLFST $PSUJDPTUFSPOF 'JHVSF". 8FPCTFSWFEBNBJOBĒFDUPGUSFBUNFOU ' 1 BOEBOJOUFSBDUJPOCFUXFFOBHFBOE USFBUNFOU ' 1 JOEJDBUJOHUIBUUIFFĒFDUPGBEEJUJPOBMTFQBSBUJPOT USFBUNFOU EFQFOEFEPOUIFQSFWJPVTEBJMZFYQFSJFODFTPGUIFBOJNBMTѮFêSTUQFSJPEPGTFQBSBUJPOBUQOE SFTVMUFEJOBSPCVTUJODSFBTFJODPSUJDPTUFSPOF 1 "ѫFSUIFTFDPOE QOE BOEUIJSE QOE QFSJPEPGTFQBSBUJPOUIJTDPSUJDPTUFSPOFJODSFBTFXBTDPNQMFUFMZBCPMJTIFE#BTBMDPSUJDPTUFSPOF UIFOFYUNPSOJOHIPVSTBѫFSFBDIDPOTFDVUJWFTFQBSBUJPO BUQOETBOE XBTTJHOJêDBOUMZ MPXFSDPNQBSFEUPOPOTFQBSBUFEBOJNBMT 1 .
(11) Chapter 2 A. B. 'JHVSF /POTFQBSBUFE /4&1 CBTBMBOENBUFSOBMTFQBSBUJPOJOEVDFEQMBTNBDPSUJDPTUFSPOF " BOE"$5) # MFWFMTJO NPVTFQVQTUFTUFEBUEJĒFSFOUBHFT OPOTFQBSBUFEBUIPVSTCBTBMCBTBMMFWFMTBUIPVSTPGQVQTNBUFSOBMMZ TFQBSBUFE GPS IPVST PO UIF QSFDFEJOH EBZ T TFQBSBUFE TFQBSBUJPO JOEVDFE MFWFMT BU IPVST PG BOJNBMT TFQBSBUFEGPSIPVST .BUFSOBMTFQBSBUJPOUPPLQMBDFBUQOET BOESFTVMUJOHJONJDFUIBUXFSFTFQBSBUFEGPS UIFêSTUUJNFBUQOE GPSBTFDPOEUJNFBUQOE BOEGPSBUIJSEUJNFBUQOEѮFDPOUSPMHSPVQBUQOEBMTP TFSWFTBTBACBTBMHSPVQGPSNBUFSOBMTFQBSBUJPOBUUIJTBHF%BUBSFQSFTFOUNFBO4&. *1 TJHOJêDBOU GSPNiOPOTFQBSBUFEuBUUIFTBNFEBZ #1 TJHOJêDBOUGSPNiCBTBMu . "$5) 'JHVSF#. 8FPCTFSWFEBNBJOFĒFDUPGUSFBUNFOU ' 1 ѮFêSTUQFSJPEPGTFQBSBUJPOBU QOESFTVMUFEJOBSPCVTUJODSFBTFJO"$5) 1 "ѫFSUIFTFDPOEQFSJPEPGTFQBSBUJPOBU QOE"$5)XBTJODSFBTFEDPNQBSFEUPCPUICBTBMMFWFMT 1 BOEOPOTFQBSBUFEBOJNBMT 1 BMUIPVHIUIFSFTQPOTFXBTMFTTUIBOBUQOEѮFUIJSEQFSJPEPGTFQBSBUJPO QOE EJE OPUEJĒFSGSPNOPOTFQBSBUFEBOJNBMTѮFNBHOJUVEFPGUIFSFTQPOTFXBTTNBMMFSUIBOBѫFS BêSTU QOE PSTFDPOE QOE QFSJPEPGTFQBSBUJPODPNQBSFEUPCBTBM"$5)#BTBM"$5) XBTTJHOJêDBOUMZMPXFSBѫFSFBDIDPOTFDVUJWFQFSJPEPGTFQBSBUJPODPNQBSFEUPOPOTFQBSBUFE BOJNBMT BUQOETBOE1 $3)N3/"JO17/ 5BCMF. $3)N3/"FYQSFTTJPOMFWFMTJODSFBTFEXJUIBHF ' 1 "ѫFSUIFêSTUTFQBSBUJPO .
(12) $IBQUFS QFSJPEBUQOECBTBMMFWFMTPG$3)N3/"FYQSFTTJPOXFSFEFDSFBTFEDPNQBSFEUPOPOTFQBSBUFE BOJNBMT 1 CVUBѫFSBTFDPOEQFSJPEPGEFQSJWBUJPO QOE CBTBMQOE UIJTEJĒFSFODF XBTBCPMJTIFE 10.$N3/"JOQJUVJUBSZ 5BCMF. /FJUIFSCBTBM OPSTFQBSBUJPOJOEVDFEMFWFMTPG10.$N3/"FYQSFTTJPOJOUIFQJUVJUBSZXFSF BĒFDUFECZNBUFSOBMTFQBSBUJPO JSSFTQFDUJWFPGUIFOVNCFSPGSFQFUJUJWFTFQBSBUJPOT .3N3/"JOIJQQPDBNQVT 5BCMF. .3 N3/" FYQSFTTJPO XBT NFBTVSFE JO UIF $" $" $" BOE EFOUBUF HZSVT %( IJQQPDBNQBMTVCêFMET8FEFUFDUFETVCêFMETQFDJêDFĒFDUTJOUIF$"BSFBNBUFSOBMTFQBSBUJPO EJEOPUBĒFDUFYQSFTTJPO OFJUIFSBѫFSIPVSTOPSBUCBTBMMFWFMT8FPCTFSWFEBNBJOFĒFDUPG BHFJOUIF$" ' 1 BOE%( ' 1 BOEBOJOUFSBDUJPOCFUXFFO 5BCMF&YQSFTTJPOPGN3/"GPSDFOUSBM)1"BYJTNBSLFSTJOTFMFDUFECSBJOSFHJPOT. CRH. PVN. NSEP basal separated POMC pituitary NSEP basal separated MR hippocampus CA1 NSEP basal separated hippocampus CA2 NSEP basal separated hippocampus CA3-4 NSEP basal separated hippocampus DG NSEP basal separated GR PVN NSEP basal separated pituitary NSEP basal separated hippocampus CA1 NSEP basal separated. pnd 3 35.34 ±2.72 34.00 ±3.29 46.16 ±1.75 44.81 ±1.51 39.99 ±2.09 41.46 ±3.49 95.01 ±4.10 103.02 ±2.15 58.21 ±3.24 58.59 ±1.43 57.35 ±2.95 65.42 ±2.00 37.73 ±2.13 37.10 ±2.13 59.65 ±1.83 57.83 ±1.06 38.42 ±2.49 40.05 ±1.85. pnd 4 40.57 ±2.59 31.92 ±1.28* 35.32 ±1.55 47.53 ±3.24 49.14 ±2.36 43.17 ±2.12 47.81 ±4.18 44.29 ±2.09 41.52 ±2.46 107.56 ±3.80 103.96 ±2.88 111.49 ±4.95 62.74 ±2.71 59.57 ±1.70 64.06 ±4.52 65.53 ±3.92 64.93 ±1.91 70.28 ±4.93 39.68 ±2.17 44.77 ±2.39 44.30 ±1.87 56.67 ±1.08 57.18 ±0.83 60.12 ±2.05 36.46 ±2.47 37.58 ±2.64 40.44 ±5.11. pnd 5 43.88 ±2.19 43.46 ±1.65 41.43 ±3.07 46.08 ±1.64 47.96 ±2.82 47.18 ±2.20 42.39 ±2.66 50.62 ±3.63* 45.10 ±2.37 98.64 ±5.54 111.26 ±3.15 112.44 ±5.72 58.07 ±3.71 63.36 ±3.23 63.24 ±3.84 68.85 ±3.19 69.82 ±3.37 80.88 ±3.39*# 39.11 ±1.97 45.14 ±2.25 39.98 ±1.97 56.53 ±1.63 56.26 ±1.60 58.63 ±1.23 38.76 ±2.54 40.77 ±2.23 38.70 ±1.82. /POTFQBSBUFE /4&1 CBTBM BOE NBUFSOBM TFQBSBUJPO JOEVDFE N3/" FYQSFTTJPO MFWFMT PG DFOUSBM )1" BYJT NBSLFSTJONPVTFQVQTUFTUFEBUEJĒFSFOUBHFT 4FF'JHVSFGPSBOFYQMBOBUJPOPGUIFEJĒFSFOUUSFBUNFOUHSPVQT 3FMBUJWFPQUJDBMEFOTJUZMFWFMTPGN3/"FYQSFTTJPOBSFFYQSFTTFEJOBSCJUSBSZVOJUT"MMEBUBQSFTFOUFEBTNFBO 4&. *1 TJHOJêDBOUGSPNi/4&1uBUUIFTBNFEBZ #1 WFSTVTiCBTBMuBUUIFTBNFEBZ . .
(13) Chapter 2 BHFBOEUSFBUNFOUJOUIF$"BSFB ' 1 )PXFWFS POMZJOUIF%(BѫFSUIFUIJSE QFSJPEPGTFQBSBUJPO BUQOE .3N3/"FYQSFTTJPOTJHOJêDBOUMZJODSFBTFEDPNQBSFEUPOPO TFQBSBUFEBOJNBMT 1 ѮFSFXBTBMTPBUSFOEUPXBSETBTJHOJêDBOUJODSFBTFDPNQBSFEUP CBTBMMFWFMT 1 #BTBM.3N3/"FYQSFTTJPOBѫFSFBDITVDDFTTJWFQFSJPEPGTFQBSBUJPOXBT OPUBĒFDUFE (3N3/"JO17/ QJUVJUBSZBOEIJQQPDBNQVT 5BCMF. *OUIF17/BOEUIFQJUVJUBSZ DPOTFDVUJWFIPVSTNBUFSOBMTFQBSBUJPOTBUQOET BOEEJEOPU MFBEUPBOBMUFSFE(3N3/"FYQSFTTJPO OFJUIFSEJEJUBĒFDUCBTBMFYQSFTTJPO . *OUIFIJQQPDBNQVTXFXFSFPOMZBCMFUPNFBTVSF(3N3/"FYQSFTTJPOJOUIF$"TVCêFME. BUUIFTFBHFT QOETm TFFBMTP<> *OUIF$"BOE%(TVCêFMETPGUIFIJQQPDBNQVT(3 N3/"FYQSFTTJPOXBTWFSZMPXBOESFNBJOFECFMPXUIFEFUFDUJPOMJNJU(3N3/"FYQSFTTJPO XBTOPUBMUFSFECZDPOTFDVUJWFQFSJPETPGNBUFSOBMTFQBSBUJPOPSBHF OPSEJETFQBSBUJPOBĒFDU CBTBMFYQSFTTJPOMFWFMT. &YQFSJNFOU**XBTEFTJHOFEUPDPNQBSFUIFUJNFDPVSTFPGUIFFĒFDUPGBTJOHMFIPVST NBUFSOBM TFQBSBUJPO BU QOE XJUI UIF PVUDPNF PG UISFF DPOTFDVUJWF EBJMZ IPVST NBUFSOBM TFQBSBUJPOTBUQOET BOE NFBTVSFEBUQOE $PSUJDPTUFSPOF 'JHVSF". 8FPCTFSWFEBNBJOFĒFDUPGUSFBUNFOU ' 1 UJNF ' 1 BOE BO JOUFSBDUJPO CFUXFFO USFBUNFOU BOE UJNF ' 1 JOEJDBUJOH UIBU BT B DPOTFRVFODFPGQSFWJPVTFYQFSJFODFTPGUIFQVQT UIFUJNFDPVSTFPGDPSUJDPTUFSPOFTFDSFUJPOPWFS IPVSTPGNBUFSOBMBCTFODFXBTDIBOHFEѮFUISFFUJNFTTFQBSBUFEHSPVQOPMPOHFSSFTQPOEFE UPTFQBSBUJPOXJUIBOJODSFBTFJODPSUJDPTUFSPOFBOEIBEMPXFSQMBTNBDPODFOUSBUJPOTBU BOEIPVSTDPNQBSFEUPOPOTFQBSBUFEBOJNBMT 1 BOEBUBMMUJNFQPJOUTDPNQBSFE UPêSTUUJNFTFQBSBUFEBOJNBMT 1 0OUIFPUIFSIBOE BOJNBMTTFQBSBUFEGPSUIFêSTUUJNF TIPXFEBOJODSFBTFJODPSUJDPTUFSPOFUIJTJODSFBTFPOMZTUBSUFEBѫFSIPVSTPGNBUFSOBMBCTFODF BUIPVST . 1PTU IPD BOBMZTFT BMTP TIPXFE UIBU DPSUJDPTUFSPOF NFBTVSFE BU IPVST JO BOJNBMT. TFQBSBUFEGPSUIFêSTUPSUIFUIJSEUJNFXFSFTJHOJêDBOUMZIJHIFSUIBUUIPTFNFBTVSFEBU IPVST 1XJUIJOFBDIUSFBUNFOUHSPVQ "$5) 'JHVSF#. 8FPCTFSWFEBNBJOFĒFDUPGUSFBUNFOU ' 1 UJNF ' 1 BOE BO JOUFSBDUJPO CFUXFFO USFBUNFOU BOE UJNF ' 1 JOEJDBUJOH UIBU UIF SFTQPOTFUPIPVSTNBUFSOBMTFQBSBUJPOBUQOEEFQFOEFEPOUIFQSFWJPVTFYQFSJFODFPGUIF QVQT1VQTTFQBSBUFEGPSUIFUIJSEUJNFEJEOPUEJĒFSGSPNOPOTFQBSBUFEBOJNBMTBUBOZUJNF QPJOU "OJNBMT TFQBSBUFE GPS UIF êSTU UJNF IPXFWFS EJE TIPX BO JODSFBTF JO QMBTNB "$5) .
(14) $IBQUFS A. B. 'JHVSF $PSUJDPTUFSPOF " BOE"$5) # SFTQPOTFUPIPVSTNBUFSOBMTFQBSBUJPOBUQOEJOBOJNBMTXJUIOPQSFWJPVT IJTUPSZPGTFQBSBUJPO TUTFQBSBUJPO BOEJOBOJNBMTQSFWJPVTMZFYQPTFEUPIPVSTTFQBSBUJPOBUQOETBOE SE TFQBSBUJPO /POTFQBSBUFEBOJNBMT /4&1 XFSFMFѫVOEJTUVSCFEVOUJMUIFNPNFOUPGUFTUJOH"UIPVST TUBSU PGTFQBSBUJPOQFSJPE POFHSPVQPGBOJNBMTTFSWFEBTBCBTBMNFBTVSFNFOUGPSCPUIUIFiOPOTFQBSBUFEuBOEiTU TFQBSBUJPOuHSPVQ%BUBSFQSFTFOUNFBO4&.*1 TJHOJêDBOUGSPNiTUTFQBSBUJPOu #1 TJHOJêDBOU GSPN CPUI DMPTFTUiOPOTFQBSBUFEuHSPVQ T. †1 TJHOJêDBOUGSPNIPVST XJUIJOUSFBUNFOUHSPVQ . MFWFMTDPNQBSFEUPOPOTFQBSBUFEBOJNBMT"$5)TUBSUFEUPSJTFUPIPVSTBѫFSUIFPOTFUPG TFQBSBUJPO BUmIPVST. . 1PTUIPDBOBMZTFTBMTPTIPXFEUIBU"$5)WBMVFTNFBTVSFEBUIPVSTJOFJUIFSHSPVQ. XFSFTJHOJêDBOUMZIJHIFSUIBOBUIPVST 1XJUIJOFBDIUSFBUNFOUHSPVQ &YQFSJNFOU***XBTEFTJHOFEUPEFUFSNJOFXIFUIFSUIFBCTFODFPGUIFFOEPDSJOFSFTQPOTFUP BUIJSEQFSJPEPGNBUFSOBMTFQBSBUJPOBTPCTFSWFEJO&YQFSJNFOU*XBTBHFTQFDJêD CZSFQFBUJOH UIJTFYQFSJNFOUOPXJODMVEJOHBTJOHMFNBUFSOBMTFQBSBUJPOBUQOETBOE $PSUJDPTUFSPOF 'JHVSF". 8FPCTFSWFEBNBJOFĒFDUPGUSFBUNFOU ' 1 UJNF ' 1 BOE BOJOUFSBDUJPOCFUXFFOUSFBUNFOUBOEUJNF ' 1 JOEJDBUJOHUIBUUIFSFTQPOTFUP NBUFSOBMTFQBSBUJPO UJNF DIBOHFE EFQFOEJOHPOUIFBNPVOUPGDPOTFDVUJWFTFQBSBUJPOQFSJPET .
(15) Chapter 2 USFBUNFOU *SSFTQFDUJWFPGUIFBHFPGUIFBOJNBMT QVQTSFBDUFEXJUIBOJODSFBTFJODPSUJDPTUFSPOF UPUIFêSTUQFSJPEPGNBUFSOBMTFQBSBUJPO BMMBHFT1 'VSUIFSNPSF JOSFTQPOTFUPFBDI TVDDFTTJWF QFSJPE PG TFQBSBUJPO UIF NBHOJUVEF PG DPSUJDPTUFSPOF TFDSFUJPO EFDSFBTFE TFDPOE TFQBSBUJPOBUQOE 1 VOUJMJUEJTBQQFBSFE UIJSETFQBSBUJPOBUQOE "$5) 'JHVSF#. 8FPCTFSWFEBNBJOFĒFDUPGUSFBUNFOU ' 1 UJNF ' 1 BOE BOJOUFSBDUJPOCFUXFFOUSFBUNFOUBOEUJNF ' 1 JOEJDBUJOHUIBUEFQFOEJOHPO UIFBNPVOUPGDPOTFDVUJWFTFQBSBUJPOQFSJPETUIF"$5)SFTQPOTFUPIPVSTNBUFSOBMBCTFODF DIBOHFE5PUIFêSTUUJNFNBUFSOBMTFQBSBUJPOXFEFUFDUFEBOJODSFBTFJO"$5) JSSFTQFDUJWFPG BHF BMMBHFT1 'VSUIFSNPSF UPUIFTFDPOEQFSJPEPGTFQBSBUJPOUIFNBHOJUVEFPG"$5) SFTQPOTFEFDSFBTFE QOE 1 XIFSFBTBUIJSEFYQPTVSFHBWFBOFWFOTNBMMFS UIPVHITUJMM TJHOJêDBOUSFTQPOTF QOE 1 A. B. 'JHVSF /POTFQBSBUFE /4&1 CBTBMBOENBUFSOBMTFQBSBUJPOJOEVDFEQMBTNBDPSUJDPTUFSPOF " BOE"$5) # MFWFMT JONPVTFQVQTUFTUFEBUEJĒFSFOUBHFT iTUuêSTUIPVSTPGNBUFSOBMTFQBSBUJPOBUQOET PSiOE uTFQBSBUFE BUQOETBOEiSE uTFQBSBUFEBUQOET BOETFFBMTP'JHVSFGPSBEFUBJMFEFYQMBOBUJPOPOUIFUSFBUNFOU HSPVQT %BUBSFQSFTFOUNFBO4&. *1 TJHOJêDBOUGSPNi/4&1uPSiCBTBMuXJUIJOUIFTBNFUSFBUNFOU HSPVQBUUIFTBNFEBZ . .
(16) $IBQUFS A. B. 'JHVSF #BTBM TFQBSBUJPOBOEOPWFMUZ NJOVUFTJTPMBUFEFYQPTVSFUPBOFXFOWJSPONFOU JOEVDFEQMBTNBDPSUJDPTUFSPOF " BOE"$5) # MFWFMTBUQOE/4&1BOETU4&1IBEOPQSFWJPVTIJTUPSZPGUSFBUNFOUTSE4&1BOJNBMTXFSF FYQPTFEUPIPVSTTFQBSBUJPOBUQOETBOESE4&1
(17) /BOJNBMTXFSFFYQPTFEUPIPVSTNBUFSOBMTFQBSBUJPO EJSFDUMZGPMMPXFECZNJOVUFTOPWFMUZBUQOETBOEi4FQBSBUJPOuMFWFMTPGTU4&1 SE4&1BOESE4&1
(18) /XFSF NFBTVSFEBUUIFFOEPGIPVSTNBUFSOBMTFQBSBUJPO%BUBSFQSFTFOUNFBO4&.*1 TJHOJêDBOUGSPN CBTBMPSTFQBSBUFE #1 TJHOJêDBOUGSPN/4&1 . &YQFSJNFOU*7XBTEFTJHOFEUPEFUFSNJOFXIFUIFSDPSUJDPTUFSPOFBOE"$5)TUJMMSFTQPOEFE UPFYQPTVSFPGBOBEEJUJPOBMOPWFMUZTUSFTTPSEJSFDUMZBѫFSNBUFSOBMTFQBSBUJPO $PSUJDPTUFSPOF 'JHVSF". 8F PCTFSWFE BO FĒFDU PG USFBUNFOU ' 1 UJNF ' 1 BOE BOJOUFSBDUJPOCFUXFFOUSFBUNFOUBOEUJNF ' 1 %FQFOEJOHPOUIFUSFBUNFOU UIFQVQTSFDFJWFE UIFJSSFTQPOTFUPOPWFMUZDIBOHFE"TFYQFDUFE OPOTFQBSBUFEBOJNBMTXFSF OPUBCMFUPSFTQPOEUPOPWFMUZXJUIBSJTFJODPSUJDPTUFSPOF XIFSFBTQVQTTFQBSBUFEGSPNUIFJS NPUIFSGPSUIFêSTUUJNFXFSFBCMFUPSFTQPOEUPOPWFMUZFWFOUIPVHIDPSUJDPTUFSPOFMFWFMTXFSF BMSFBEZJODSFBTFEEVFUPUIFQSFDFEJOHTFQBSBUJPO . ѮPVHIQVQTUIBUXFSFTFQBSBUFEGPSUISFFUJNFTIBEDPSUJDPTUFSPOFMFWFMTTJNJMBSUPUIPTF. PCTFSWFE GPS VOUSFBUFE BOJNBMT UIFZ XFSF TUJMM DBQBCMF UP SFTQPOE UP OPWFMUZ "MTP QVQT UIBU XFSF SFQFBUFEMZ TFQBSBUFE JO DPNCJOBUJPO XJUI OPWFMUZ FYQPTVSF BѫFS FBDI TFQBSBUJPO QFSJPE SE4&1
(19) / TUJMMSFTQPOEFEUPOPWFMUZѮFJSDPSUJDPTUFSPOFMFWFMTBѫFSUIFUIJSETFQBSBUJPOXFSF .
(20) Chapter 2 TJHOJêDBOUMZIJHIFSUIBOUIPTFPGOPOTFQBSBUFEBOJNBMT /4&1 CVUOPUDPNQBSFEUPUIPTFPG UIFSFQFBUFETFQBSBUJPOHSPVQXJUIPVUOPWFMUZFYQPTVSF SE4&1 "$5) 'JHVSF#. 8F PCTFSWFE B NBJO FĒFDU PG USFBUNFOU ' 1 BOE UJNF ' 1 "MUIPVHI "$5) MFWFMT PG QVQT TFQBSBUFE GPS UIF êSTU UJNF XFSF BMSFBEZ IJHIFS UIBO UIPTF PG OPOTFQBSBUFEQVQT BOJNBMTJOCPUIHSPVQTXFSFVOBCMFUPSFTQPOEUPOPWFMUZXJUIBOJODSFBTF JO"$5) . "OJNBMT TFQBSBUFE GPS UISFF UJNFT SE 4&1 IBE IJHIFS "$5) MFWFMT DPNQBSFE UP OPO. TFQBSBUFEBOJNBMTBOESFTQPOEFEUPOPWFMUZXJUIBGVSUIFSJODSFBTFJO"$5)*OQVQTSFDFJWJOH NBUFSOBMTFQBSBUJPOJODPNCJOBUJPOXJUIOPWFMUZFBDIUJNF SE4&1
(21) / TIPXFE BѫFSUIFUIJSE TFQBSBUJPOQFSJPE "$5)MFWFMTTJNJMBSUPUIPTFPGOPOTFQBSBUFEBOJNBMTBOEUIFFYQPTVSFUPB OPWFMUZTUSFTTPSEJEOPUFWPLFBSFTQPOTF &YQFSJNFOU7ѮFPCKFDUJWFXBTUPEFUFSNJOFDJSDVMBUJOHHISFMJOBOEHMVDPTFDPODFOUSBUJPOT JO SFTQPOTF UP SFQFBUFE NBUFSOBM TFQBSBUJPOT $PSUJDPTUFSPOF BOE N3/" FYQSFTTJPO PG DGPT N3/"XFSFBMTPNFBTVSFEVOEFSCBTBMDPOEJUJPOTBOEJOSFTQPOTFUPNBUFSOBMTFQBSBUJPOTXJUI BOEXJUIPVUOPWFMUZFYQPTVSF $PSUJDPTUFSPOF 'JHVSF. "NBJOFĒFDUXBTPCTFSWFEPGUSFBUNFOU ' 1 UJNF ' 1 BOEBOJOUFSBDUJPOCFUXFFOUIFTFGBDUPST ' 1 *OSFTQPOTFUPBêSTUTFQBSBUJPO QFSJPE DPSUJDPTUFSPOF JODSFBTFE 1 BOE UIFTF MFWFMT JODSFBTFE FWFO GVSUIFS XIFO QVQT XFSFTVCKFDUFEUPBEEJUJPOBMOPWFMUZTUSFTT 1 *OSFTQPOTFUPBUIJSEQFSJPEPGTFQBSBUJPO IPXFWFS DPSUJDPTUFSPOFWBMVFTEJEOPUBMUFS UIPVHIFYQPTVSFUPBOPWFMFOWJSPONFOUXBTBCMF UPBDUJWBUFUIF)1"BYJT 1 #BTBMDPSUJDPTUFSPOFBUUIFTUBSUPGFJUIFSUIFêSTUPSUIJSE TFQBSBUJPOQFSJPEEJEOPUEJĒFS. 'JHVSF $PSUJDPTUFSPOF JO BOJNBMT BU QOE XJUI OP QSFWJPVT IJTUPSZ PG TFQBSBUJPO TU 4&1 BOE JO BOJNBMT QSFWJPVTMZ FYQPTFEUPIPVSTTFQBSBUJPOTBUQOETBOE SE4&1 $PSUJDPTUFSPOFSFTQPOTFXBTNFBTVSFEBѫFSIPVSTPG NBUFSOBMTFQBSBUJPOBOEBѫFSIPVSTPGNBUFSOBMTFQBSBUJPOXJUIBOBEEJUJPOBMNJOVUFTPGOPWFMUZTUSFTT%BUB SFQSFTFOUNFBO4&.*1 TJHOJêDBOUGSPNCBTBM #1 TJHOJêDBOUGSPNTFQBSBUFE . .
(22) $IBQUFS A. B. 'JHVSF (MVDPTF " BOE HISFMJO # SFTQPOTF UP NBUFSOBM TFQBSBUJPO BU QOE JO BOJNBMT XJUI OP QSFWJPVT IJTUPSZ PG TFQBSBUJPOT TU 4&1 BOE JO BOJNBMT QSFWJPVTMZ FYQPTFE UP IPVST TFQBSBUJPO BU QOET BOE SE 4&1 %BUB SFQSFTFOUNFBO4&.*1 TJHOJêDBOUGSPNCBTBM . (MVDPTF 'JHVSF". "NBJOFĒFDUPGUJNFXBTEFUFSNJOFE ' 1 CVUOPUPGUSFBUNFOU*OSFTQPOTF UPBêSTUNBUFSOBMTFQBSBUJPOQFSJPECMPPEHMVDPTFMFWFMTEFDSFBTFEUPBMNPTUIBMGUIFJSPSJHJOBM MFWFMT 1 "ѫFSSFQFBUFETFQBSBUJPOTUIJTTBNFSFTQPOTFXBTTUJMMQSFTFOU 1 #BTBM HMVDPTFWBMVFTXFSFTJNJMBSCFUXFFOUIFBOJNBMTTFQBSBUFEGPSUIFêSTUPSUIJSEUJNF. 5BCMFN3/"FYQSFTTJPOPGDGPTJOTFMFDUFECSBJOBSFBT. Cortex pPVTh. 1st SEP 3rd SEP 1st SEP 3rd SEP. basal 14.63 ±2.09 12.34 ±2.56 40.76 ±2.56 39.76 ±4.50. separated 13.92 ±2.22 11.70 ±2.15 39.35 ±4.31 31.85 ±2.55. novelty 15.10 ±3.28 12.00 ±0.92 40.08 ±3.61 32.39 ±1.7. 5PEFUFSNJOFDFOUSBMBDUJWBUJPOPGOFVSPOTJOSFTQPOTFUPNBUFSOBMTFQBSBUJPOBOEBEEJUJPOBMOPWFMUZTUSFTTDGPT N3/"FYQSFTTJPOXBTEFUFSNJOFEJOUIF17/ 'JHVSF DPSUFYBOEQ17Ѯ QBSBWFOUSJDVMBSUIBMBNJDOVDMFVT %BUBBSFFYQSFTTFEBTPQUJDBMEFOTJUZNFBTVSFEJOBSCJUSBSZVOJUTBOEQSFTFOUFEBTNFBO4&.. .
(23) Chapter 2. 'JHVSF $GPTN3/"FYQSFTTJPONFBTVSFEJOUIF17/BUQOEJOBOJNBMTXJUIOPQSFWJPVTIJTUPSZPGTFQBSBUJPO TU4&1 BOEJOBOJNBMTQSFWJPVTMZFYQPTFEUPIPVSTTFQBSBUJPOTBUQOETBOE SE4&1 &YQSFTTJPOXBTNFBTVSFEBѫFS IPVSTPGNBUFSOBMTFQBSBUJPOBOEBѫFSIPVSTPGNBUFSOBMTFQBSBUJPOXJUIBOBEEJUJPOBMNJOVUFTPGOPWFMUZ TUSFTT3FMBUJWFPQUJDBMEFOTJUZMFWFMTPGN3/"FYQSFTTJPOBSFFYQSFTTFEJOBSCJUSBSZVOJUT%BUBSFQSFTFOUNFBO 4&.*1 TJHOJêDBOUGSPNCBTBM #1 TJHOJêDBOUGSPNTFQBSBUFE . (ISFMJO 'JHVSF#. "NBJOFĒFDUPGUJNFXBTNFBTVSFE ' 1 CVUOPUPGUSFBUNFOU(ISFMJOMFWFMT XFSF UXJDF BT IJHI BѫFS B êSTU 1 CVU BMTP BѫFS B UIJSE QFSJPE PG NBUFSOBM TFQBSBUJPO 1 BTDPNQBSFEUPCBTBMMFWFMT#BTBMMFWFMTXFSFJOEJTUJOHVJTIBCMFCFUXFFOCPUIUSFBUNFOU HSPVQT DGPTN3/"FYQSFTTJPO 'JHVSF 5BCMF. ѮFFYQSFTTJPOPGDGPTN3/"JOUIF17/TIPXFEBNBJOFĒFDUPGUJNF ' 1 *OSFTQPOTFUPNBUFSOBMTFQBSBUJPODGPTN3/"FYQSFTTJPOJODSFBTFEJOUIFBOJNBMTTFQBSBUFE GPSUIFêSTUUJNF 1 XIFSFBTJUEJEOPUSFTQPOEJOUIFUIJSEUJNFTFQBSBUJPOHSPVQ#BTBM MFWFMTXFSFDPNQBSBCMFCFUXFFOCPUIUSFBUNFOUHSPVQT6QPOFYQPTVSFUPBOPWFMFOWJSPONFOU GPSNJOVUFTDGPTN3/"FYQSFTTJPOJODSFBTFEUPDPNQBSBCMFMFWFMTJOBOJNBMTTFQBSBUFECPUI GPSUIFêSTUBOEUIJSEUJNF 1 $GPTN3/"FYQSFTTJPOJOUIFDPSUFYBOEJOUIFQ17Ѯ QBSBWFOUSJDVMBSUIBMBNJDOVDMFVT XBTVOBĒFDUFECZUSFBUNFOUPSUJNF. %JTDVTTJPO ѮF QSFTFOU TUVEZ TIPXT JO $% NJDF UIBU UIF JOGBOU NPVOUT B MBSHF SFTQPOTF PG "$5) BOE DPSUJDPTUFSPOFBѫFSIPVSTPGTFQBSBUJPOGSPNUIFNPUIFS$POUSBSZUPPVSFYQFDUBUJPOT EBJMZ SFQFBUFE TFQBSBUJPOT PG IPVST SFBEJMZ BCPMJTIFE UIF QJUVJUBSZBESFOBM SFTQPOTF QSPEVDJOH DPSUJDPTUFSPOFMFWFMTUIBUXFSFFWFODPOTJTUFOUMZMPXFSIPVSTBѫFSSFVOJPO)PXFWFS EFTQJUF UIFBCTFODFPGBO)1"SFTQPOTFUPSFQFBUFETFQBSBUJPOT UIFEFQSJWFEQVQDPOUJOVFEUPSFTQPOE UPOPWFMUZѮFêOEJOHUIBUUIFSFTQPOTFUPNVMUJQMFTFQBSBUJPOTXBTSFEVDFECFMPXDPOUSPMMFWFMT XIJMF TUSFTT SFTQPOTJWFOFTT SFNBJOFE FOIBODFE JOEJDBUFT QSPOPVODFE EJĒFSFOU FĒFDUT PO UIF .
(24) $IBQUFS OFVSPFOEPDSJOFTZTUFNTPGUIFOFXCPSOUIBUSFHVMBUFCBTBMTFDSFUJPOTBTPQQPTFEUPUIPTFUIBU SFRVJSFNJMETUSFTTPST . ѮFSFTVMUTPGUIJTTUVEZSFKFDUUIFIZQPUIFTJTUIBUEBJMZTFQBSBUJPOTXPVMEMFBEUPBOFOIBODFE. BESFOBM DPSUJDPTUFSPOF PVUQVU 4VSQSJTJOHMZ UIF )1" BYJT SBQJEMZ EFTFOTJUJTFE .PSFPWFS IPVST BѫFS SFVOJPO DPSUJDPTUFSPOF MFWFMT XFSF DPOTJTUFOUMZ MPXFS BOE UIJT TUBUF PG SFEVDFE DPSUJDPTUFSPOFQFSTJTUFEPWFSUIFOFYUUXPEBZT3FDFOUMZ UIFSFIBTCFFOBOJODSFBTFEBXBSFOFTT PG UIF TJHOJêDBODF PG IZQPDPSUJTPMJTN TJODF UIFSF JT B SFEVDFE DPSUJTPM TFDSFUJPO PCTFSWFE JO DIJMESFOFYQPTFEUPBEWFSTFFBSMZMJGFFYQFSJFODF< >ѮJTTUVEZJTUIFêSTUEFNPOTUSBUJPO UIBU UIF TBNF QIFOPNFOPO BMTP PDDVST JO SPEFOUT *U SBJTFT UIF JOUSJHVJOH QPTTJCJMJUZ UIBU B AOPSNBMHMVDPDPSUJDPJEMFWFMJTSFRVJSFEGPSCSBJOEFWFMPQNFOUBOEUIBUFJUIFSUPPMPXPSUPP IJHIMFWFMTNBZIBWFEFUSJNFOUBMFĒFDUT . *O&YQFSJNFOU* BOIPVSTFQJTPEFPGNBUFSOBMTFQBSBUJPOXBTVTFEUPFOTVSFBOBDUJWBUJPO. PG UIF QVQT )1" BYJT < > 8IFO JU BQQFBSFE UIBU BѫFS UISFF EBJMZ QFSJPET PG IPVST NBUFSOBM TFQBSBUJPO UIF FOEPDSJOF SFTQPOTF BTTPDJBUFE XJUI NBUFSOBM BCTFODF XBT OP MPOHFS QSFTFOU BU êSTU B QPTTJCMF TIJѫ JO UJNF DPVSTF PG UIF SFTQPOTF XBT JOWFTUJHBUFE &YQFSJNFOU ** ѮFSFBTPOJOHXBTUIBUSFQFBUFETFQBSBUJPOTNJHIUIBWFGBDJMJUBUFEUIFPOTFUPGBUSBOTJFOU )1"SFTQPOTFUPBOFBSMJFSUJNFQPJOUXJUIDPSUJDPTUFSPOFMFWFMTSFUVSOJOHBMSFBEZUPCBTFMJOFBU UIFIPVSTJOUFSWBMSFDPSEFEJOUIFêSTUFYQFSJNFOUѮFUJNFDPVSTFTUVEZEFNPOTUSBUFEUIBU UIFSFXBTOPFBSMJFSSFTQPOTFBOEUIBUUIFIPVSJOUFSWBMXBTBQQSPQSJBUFUPSFMJBCMZNPOJUPS EFTFOTJUJTBUJPOPGUIF"$5)BOEDPSUJDPTUFSPOFSFTQPOTFUPSFQFBUFETFQBSBUJPOT . ѮFUJNFDPVSTFFYQFSJNFOUJOWPMWJOHIPVSFQJTPEFTBMTPTIPXFEBѫFSUIFêSTUTFQBSBUJPO. UIBUUIFJOJUJBM"$5)FMFWBUJPOPDDVSFEBѫFSIPVST XIFSFBTDPSUJDPTUFSPOFTUBSUFEUPSJTFPOMZ BѫFSIPVSTѮJTJOEJDBUFEUIBUUIF)1"BYJTSFTQPOEFETMPXMZUPNBUFSOBMTFQBSBUJPOBOEUIBU JUUPPLBGFXIPVSTPG"$5)TUJNVMBUJPOCFGPSFUIFTUSPOHMZSFEVDFEBESFOBMTFOTJUJWJUZUP"$5) EVSJOHUIF4)31XBTPWFSDPNFѮJTPCTFSWBUJPOJTJOTVQQPSUPGQSFWJPVTTUVEJFT< > . &YQFSJNFOU ** BMTP EFNPOTUSBUFE UIBU JG JNNFEJBUFMZ BѫFS TFQBSBUJPO UIF JOGBOUT XFSF. BMTPFYQPTFEUPBNJOVUFTOPWFMUZTUSFTTPS BQSPGPVOE"$5)BOEDPSUJDPTUFSPOFSFTQPOTF PDDVSFEVOEFSDPOEJUJPOTUIBUUIFQVQT)1"SFTQPOTFUPSFQFBUFETFQBSBUJPOTXBTBCPMJTIFE Ѯ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ѮFEBUBTIPXFEUIBUUIFFOEPDSJOFSFTQPOTFUPNBUFSOBMTFQBSBUJPO HSBEVBMMZEJNJOJTIFEXJUIFBDITVDDFTTJWFTFQBSBUJPOGPSCPUIDPSUJDPTUFSPOFBOE"$5)JFUIF .
(25) Chapter 2 SFTQPOTFTBUQOEXFSFMPXFSBTDPNQBSFEUPQOEѮFFYQFSJNFOUDPOêSNFEUIBUCBTBMMFWFMT PG DPSUJDPTUFSPOF BOE "$5) BU IPVST BѫFS SFVOJPO XFSF FWFO MPXFS JG DPNQBSFE UP OPO TFQBSBUFEBOJNBMT'VSUIFSNPSF XFPCTFSWFEUIBUUIFFOEPDSJOFSFTQPOTFUPBTJOHMFTFQBSBUJPO XBTQSFTFOUBUQOET BOE MJLFQSFWJPVTMZXBTTIPXOGPSQOE<>ѮFTFSFTVMUTJOEJDBUF UIBUUIFQVQTUSVMZEFTFOTJUJTFUPEBJMZNBUFSOBMBCTFODFBOEUIBUUIJTEFTFOTJUJTBUJPOJTOPUBHF TQFDJêD 4JODFJUXBTBMSFBEZTIPXOBUQOEUIBUN3/"FYQSFTTJPOPGDFOUSBM)1"BYJTNBSLFST XFSFSFTQPOTJWFUPIPVSTPGNBUFSOBMBCTFODF TFWFSBMPGUIFTFNBSLFSTXFSFFYBNJOFEJOPVS QBSBEJHNJOPSEFSUPêOEDMVFTUPXBSETUIFDBVTFPGUIFPCTFSWFEEFTFOTJUJTBUJPO"UQOE$3) N3/"FYQSFTTJPOJOUIF17/EFDSFBTFEBMSFBEZBѫFSIPVSTPGNBUFSOBMBCTFODF< >*O PVSQBSBEJHNVTJOHEBZTPMENJDF$3)N3/"JOUIF17/POMZSFTQPOEFEUPUIFêSTUQFSJPE PGNBUFSOBMBCTFODFBUIPVSTBѫFSSFVOJPO4JODF$3)N3/"FYQSFTTJPOJTDPSUJDPTUFSPOF SFTQPOTJWF< > UIJTEPXOSFHVMBUJPOJO$3)N3/"FYQSFTTJPONJHIUCFBDPOTFRVFODFPGUIF IJHIDPSUJDPTUFSPOFDPODFOUSBUJPOTJOEVDFECZUIFêSTUTFQBSBUJPOQFSJPE$PODPNJUBOUMZ UIF QSPHSFTTJWFBUUFOVBUJPOPGUIFDPSUJDPTUFSPOFSFTQPOTFEVSJOHUIFTFDPOEBOEUIJSETFQBSBUJPO SFTQFDUJWFMZ NJHIUIBWFCFFOQFSNJTTJWFGPS$3)N3/"FYQSFTTJPOMFWFMTUPSFTUPSF"MUFSOBUJWFMZ MPXFS$3)N3/"FYQSFTTJPOSFëFDUTUIFMPXFSCBTBM"$5)BOEDPSUJDPTUFSPOFMFWFMTBUUIFTUBSU PGUIFTFDPOETFQBSBUJPOQFSJPE)PXFWFS UIFEFDSFBTFE$3)N3/"MFWFMSFDPWFSFEBѫFSUIF TFDPOETFQBSBUJPOXIJMFUIJTJTOPUUIFDBTFXJUIUIFIZQPDPSUJDJTN TVHHFTUJOHUIBUUIFQFSTJTUFOU IZQPSFTQPOTJWFOFTTJOUIFSFQFBUFEMZTFQBSBUFENPVTFPDDVSTBUUIFMFWFMPGUIFBESFOBMJUTFMG .PTUDFOUSBM)1"NBSLFSTSFNBJOFETUBCMFEVSJOHSFQFBUFETFQBSBUJPOT XJUIUIFFYDFQUJPO PGUIF.3N3/"FYQSFTTJPOJOUIFEFOUBUFHZSVT %( PGUIFIJQQPDBNQVT XIJDITIPXFEB HSBEVBMJODSFBTF"OBMUFSFEGVODUJPOBMJUZPGCPUISFDFQUPSTIBTCFFOJNQMJDBUFEEVSJOHAOPSNBM QPTUOBUBMEFWFMPQNFOUJOSFTQPOTFUPQSPMPOHFENBUFSOBMBCTFODF< >"MSFBEZVOEFS VOEJTUVSCFEDPOEJUJPOTN3/"FYQSFTTJPOPG.3JOUIF%(HSBEVBMMZJODSFBTFEXJUIBHF<> )FSF UIFEFWFMPQNFOUBMJODSFBTFPG.3N3/"FYQSFTTJPOJOUIF%(PGUIFIJQQPDBNQVTTFFNFE GBDJMJUBUFEEVSJOHSFQFBUFETFQBSBUJPOT.3JTVOEFSQPTJUJWFSFHVMBUJPOPG$3)JOBEVMUT JF IJHIFS$3)MFWFMTUSBOTMBUFJOUPIJHIFSIJQQPDBNQBM.3MFWFMT<>*OTVQQPSUPGUIJTSFBTPOJOH JOOFPOBUBM$3)SMJNCJDCSBJOTQFDJêDLOPDLPVUNJDF.3N3/"FYQSFTTJPOJTEFDSFBTFE<> ѮPVHIXFEJEOPUNFBTVSF$3)MFWFMTJOMJNCJDSFHJPOTCFZPOEUIF17/ UIF$3)STZTUFN QSFTFOUTBHPPEDBOEJEBUFGPSUIFGBDJMJUBUFEEFWFMPQNFOUBMJODSFBTFJO.3N3/"FYQSFTTJPOJO SFQFBUFEMZTFQBSBUFENJDF Ѯ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ѫFSiSFQFBUFETFQBSBUJPOTXJUI .
(26) $IBQUFS OPWFMUZ 4&1
(27) / u)PXFWFS GPSQVQTUSFBUFEXJUIUIJTQBSBEJHNBOPWFMFOWJSPONFOUXBTTUJMM TUSFTTGVMBUUIFUIJSEFYQPTVSF . *OUFSFTUJOHMZ UIF"$5)SFTQPOTFJOUIFUISFFUJNFTTFQBSBUJPOXJUIUISFFUJNFTOPWFMUZ. SE4&1
(28) / HSPVQXBTNVDIMFTTUIBOJOBOJNBMTUIBUXFSFPOMZTFQBSBUFEGSPNUIFJSNPSIUFST GPS UISFF UJNFT SE 4&1 "U UIF TBNF UJNF UIF DPSUJDPTUFSPOF SFTQPOTF SFNBJOFE PG TJNJMBS NBHOJUVEF "QQBSFOUMZ QSPMPOHFE NBUFSOBM TFQBSBUJPO êSTU PWFSDPNFT UIF SFEVDFE BESFOBM TFOTJUJWJUZBTTPDJBUFEXJUIUIF4)31< > XIFSFBTGVSUIFSTUJNVMBUJPOCZEBJMZFYQPTVSFUP BOPWFMFOWJSPONFOUUIFSFBѫFSFOIBODFTTFOTJUJWJUZPGUIFBESFOBMGPS"$5)Ѯ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ѮF DVSSFOU êOEJOHT SBJTF UIF RVFTUJPO XIZ UIF QJUVJUBSZBESFOBM SFTQPOTF DFBTFT BOE. CFDPNFTIZQPBDUJWFBѫFSSFQFBUFEEBJMZNBUFSOBMTFQBSBUJPOT0OFNFDIBOJTNDPVMECFSFMBUFE UPUIFQBUUFSOPGGPPEJOUBLF CFDBVTFGFFEJOHJTSFRVJSFEUPNBJOUBJOIZQPSFTQPOTJWFOFTTPG UIFBESFOBMUPTUSFTT< >&JHIUIPVSTPGGPPEEFQSJWBUJPOEFDSFBTFTDJSDVMBUJOHHMVDPTFBOE JODSFBTFTHISFMJOMFWFMT XIJMFBENJOJTUSBUJPOPGBHISFMJOBOUBHPOJTUDBOQSFWFOUUIFTFQBSBUJPO JOEVDFE)1"BDUJWBUJPO<>)FODF UIFOFYUIPVSTDPVMEIBWFMFEUPBOBMUFSFEQBUUFSOPGGPPE JOUBLFUIBUJTQFSIBQTSFëFDUFEJOUIFDJSDVMBUJOHMFWFMPGUIFTFNFUBCPMJDTJHOBMT&YQFSJNFOU 7 TIPXT UIBU BѫFS SFQFBUFE NBUFSOBM TFQBSBUJPOT HMVDPTF BOE HISFMJO MFWFMT TUJMM SFTQPOEFE TJNJMBSMZBTBѫFSUIFêSTUTFQBSBUJPOFWFOU Bê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ĒFDUTXJUIMPOHUFSNDPOTFRVFODFTGPSCSBJOEFWFMPQNFOUBOEGVODUJPOJOH <> 'VSUIFS FYQFSJNFOUT BSF OFFEFE XIJDI NBZ JODMVEF UIF BTTFTTNFOU PG HISFMJO SFDFQUPS FYQSFTTJPO JO UIF BSDVBUF OVDMFVT BT XFMM BT UIF SPMF PG MFQUJO JO TUSVDUVSBM SFNPEFMMJOH PG IZQPUIBMBNJDQBUIXBZT<> . "OPUIFSNFDIBOJTNNBZCFSFMBUFEUPUIFMBDLPGTFOTPSZTUJNVMBUJPO TJODFJUJTLOPXO. GPSTPNFUJNFUIBUUIFBDUJWBUJPOPGDFOUSBMDPNQPOFOUTPGUIF)1"BYJTPGUIFEFQSJWFEQVQ .
(29) Chapter 2 DBOCFQSFWFOUFECZTUSPLJOH<>.PSFPWFS VQPOSFVOJPOUIFQVQSFDFJWFTFYDFTTJWFNBUFSOBM DBSF XIJDI BMTP DPVME BĒFDU )1" BDUJWBUJPO UIF OFYU EBZ &YQFSJNFOU 7 EFNPOTUSBUFE UIBU BѫFSSFQFBUFETFQBSBUJPOTUIFCBTBMDGPTN3/"SFTQPOTFJOUIF17/XBTBUUFOVBUFE CFDPNJOH JOEJTUJOHVJTIBCMFGSPNDPOUSPMT:FUUIF17/DGPTSFTQPOTFUPOPWFMUZSFNBJOFEGBDJMJUBUFEJOUIF TFQBSBUFEBOJNBMT XIJDIJTSFNJOJTDFOUUPUIFSFTQPOTFQBUUFSOPGUIF)1"BYJT*OUFSFTUJOHMZ JOOFPOBUFSBUTIBOEMFEEBJMZGPSNJOVUFTGSPNQOETUP BOFOIBODFESFTQPOTFPGDGPT JNNVOPSFBDUJWJUZXBTPCTFSWFEJOUIF QBSBWFOUSJDVMBSUIBMBNJDOVDMFVT Q17ѮNFBTVSFE NJOVUFT BѫFS SFVOJPO BU QOE BT DPNQBSFE UP VOEJTUVSCFE DPOUSPMT BOE QVQT IBOEMFE PODF POMZBUQOE<>*OUIFDVSSFOUTUVEZDGPTN3/"EJEOPUSFTQPOEJOQ17Ѯ BCSBJOSFHJPO JNQPSUBOU GPS SFMBZ PG TFOTPSZ JOGPSNBUJPO "MTP FMTFXIFSF JO UIF CSBJO XF EJE OPU êOE DGPT N3/"DIBOHFT . ѮBUDGPTN3/"JOUIFIZQPUIBMBNVTEJTQMBZTBDPSUJDPTUFSPOFMJLFSFTQPOTFQBUUFSOUP. SFQFBUFE TFQBSBUJPOT TVHHFTUT UIBU B DFOUSBM NFDIBOJTN JT JOWPMWFE *U NBZ XFMM CF UIBU FWFO UIFEBZTPMEJOGBOUDBOBMSFBEZQSFEJDUBѫFSPOFFYQFSJFODFUIBUUIFNPUIFSXJMMSFUVSOJO IPVSTѮFJOGBOUUIVTSBQJEMZBEBQUTPSIBCJUVBUFTUPNBUFSOBMBCTFODF CVUUIF)1"BYJTTUBZTPO BMFSUBOEDBOCFBDUJWBUFECZTUSFTTPST3FDFOUFWJEFODFTVHHFTUTJOEFFEUIBUMFBSOJOHDBOPDDVS JOOFPOBUBMSBUT< >*OUIFTFTUVEJFTJUXBTGPVOEUIBUUIFêSTUXFFLPGMJGFBUUBDINFOU PGUIFJOGBOUUISPVHIPMGBDUPSZDVFTGSPNUIFNPUIFSEFQFOETPOBMPDVTDPFSVMFVTmPMGBDUPSZ QBUIXBZ*OUIFêSTUQPTUOBUBMXFFLJOGBOUSBUTSBQJEMZMFBSONBUFSOBMPEPSUPTVQQPSUBUUBDINFOU CFIBWJPVSBOEUPTVQQSFTTPEPSBWFSTJPOTEVSJOHUIBUUJNF*UJTPOMZJOUIFBCTFODFPGUIFNPUIFS UIBUBWFSTJWFPEPSTTUBSUUPBDUJWBUFUIFBNZHEBMBDJSDVJUUIBUHPWFSOTGFBSNPUJWBUFEBWPJEBODF ѮJT TXJUDI GSPN NBUFSOBM BUUSBDUJPO UP BWPJEBODF JT GBDJMJUBUFE CZ DPSUJDPTUFSPOF BOE DBO CF CMPDLFECZBHMVDPDPSUJDPJEBOUBHPOJTU*UJTDPODFJWBCMFUIBU TVDIBDPSUJDPTUFSPOFEFQFOEFOU NFDIBOJTNNBZGPSNUIFCBTJTGPSBEBQUJOHUPUIFUSBOTJFOUOBUVSFPGNBUFSOBMBCTFODF TJODFUIF JOGBOUJTTUJMMDBQBCMFUPNPVOUBDPSUJDPTUFSPOFSFTQPOTFVQPOFYQPTVSFUPOPWFMUZ . ѮFJNQMJDBUJPOTPGPVSêOEJOHTBSFWFSZJOUFSFTUJOH*UXPVMEJNQMZUIBUUIFPVUDPNFPG. UIFSFQFBUFENBUFSOBMTFQBSBUJPOTXPVMECF CFTJEFTHFOEFSBOETUSBJO OPUPOMZEFQFOEFOUVQPO UIFEVSBUJPOPGUIFTFQBSBUJPOBOEUIFNBUFSOBMDBSFSFDFJWFECZUIFQVQVQPOSFVOJPO CVUBMTP POUIFQVQTTUSFTTGVMFYQFSJFODFTEVSJOHUIFTFQBSBUJPOQSPDFEVSFѮBUNBUFSOBMDBSFJTOPUUIF TPMFGBDUPSEFUFSNJOJOHFQJHFOFUJDQSPHSBNNJOHBOEPVUDPNFXBTSFDFOUMZEFNPOTUSBUFEJOUIF FMFHBOUTUVEJFTPG.BDSJBOE8ÛSCFM< >ѮFSFGPSF TUBOEBSEJTBUJPOJOQSPUPDPMTVTFEGPS TUVEZPGUIFPVUDPNFPGNBUFSOBMTFQBSBUJPOTJTSFRVJSFE TJODFUIFDVSSFOUEBUBTIPXBEJĒFSFOU PVUDPNFJGUIFQVQJTSFNPWFE UIFEBN PSCPUI . *ODPODMVTJPO UIFDVSSFOUEBUBEFNPOTUSBUFUIFBNB[JOHDBQBDJUZPGUIFOFXCPSOSPEFOU. UPSFBEJMZBEBQUUPUIFBCTFODFPGUIFNPUIFSBOEQSFTFOUBTUSJLJOHFYBNQMFPGQMBTUJDJUZ TJODF UIFJOGBOUTSFNBJOSFTQPOTJWFUPOPWFMUZBOETFFNUPCFNBSLFECZBUPPMPXDJSDVMBUJOHBNPVOU PGDPSUJDPTUFSPOFѮFêOEJOHNBZCFIFMQGVMUPEFTJHOSBUJPOBMFYQFSJNFOUTUPVOEFSTUBOEUIF NFDIBOJTNVOEFSMZJOHUIFMBTUJOHFĒFDUTPGFBSMZBEWFSTJUZPOCSBJOBOECFIBWJPVS .
(30) $IBQUFS. "DLOPXMFEHFNFOUT ѮJT TUVEZ XBT TVQQPSUFE CZ UIF 3PZBM /FUIFSMBOET "DBEFNZ PG "SUT BOE 4DJFODFT /80 45*(0//%3'BOE/808FXPVMEMJLFUPUIBOL*&.EF+POH 1(.WBO0WFSWFMEBOE14UFFOCFSHFOGPSUFDIOJDBMBTTJTUBODFXJUIUIFBOJNBMFYQFSJNFOUT. 3FGFSFODFT . . . #PVSFU4( %SBQFS4+BOE4JNFSMZ3# 5SPQIJDBDUJPOPGMFQUJOPOIZQPUIBMBNJD OFVSPOTUIBUSFHVMBUFGFFEJOH4DJFODF $IFDLMFZ 4 ѮF OFVSPFOEPDSJOPMPHZ PG EFQSFTTJPO BOE DISPOJD TUSFTT #S .FE #VMM EF,MPFU&3BOE0JU[M.4 8IPDBSFTGPSBTUSFTTFECSBJO ѮFNPUIFS UIFLJEPS CPUI /FVSPCJPM"HJOH4VQQM44 EF,MPFU&3 7SFVHEFOIJM& 0JU[M.4BOE+PFMT. #SBJODPSUJDPTUFSPJESFDFQUPS CBMBODFJOIFBMUIBOEEJTFBTF&OEPDS3FW &EXBSET)&BOE#VSOIBN8. Ѯ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Ē$# "MUFSFEQJUVJUBSZ BESFOBM BYJT SFTQPOTFT UP QSPWPDBUJWF DIBMMFOHF UFTUT JO BEVMU TVSWJWPST PG DIJMEIPPE BCVTF"N+1TZDIJBUSZ )PGFS." &BSMZSFMBUJPOTIJQTBTSFHVMBUPSTPGJOGBOUQIZTJPMPHZBOECFIBWJPS"DUB 1BFEJBUS4VQQM ,JTIJ5BOE&MNRVJTU+, #PEZXFJHIUJTSFHVMBUFECZUIFCSBJOBMJOLCFUXFFO GFFEJOHBOEFNPUJPO.PM1TZDIJBUSZ -BEE$0 )VPU3- ѮSJWJLSBNBO,7 /FNFSPĒ$# .FBOFZ.+BOE1MPUTLZ1. -POHUFSNCFIBWJPSBMBOEOFVSPFOEPDSJOFBEBQUBUJPOTUPBEWFSTFFBSMZFYQFSJFODF1SPH #SBJO3FT -FWJOF4 ѮFPOUPHFOZPGUIFIZQPUIBMBNJDQJUVJUBSZBESFOBMBYJTѮFJOëVFODF PGNBUFSOBMGBDUPST"OO/:"DBE4DJ -FWJOF 4 1SJNBSZ TPDJBM SFMBUJPOTIJQT JOëVFODF UIF EFWFMPQNFOU PG UIF IZQPUIBMBNJDQJUVJUBSZBESFOBMBYJTJOUIFSBU1IZTJPM#FIBW -FWJOF4 )VDIUPO%. 8JFOFS4(BOE3PTFOGFME1 5JNFDPVSTFPGUIFFĒFDUPG NBUFSOBMEFQSJWBUJPOPOUIFIZQPUIBMBNJDQJUVJUBSZBESFOBMBYJTJOUIFJOGBOUSBU%FW 1TZDIPCJPM .
(31) Chapter 2 -JV% %JPSJP+ %BZ+$ 'SBODJT%%BOE.FBOFZ.+ .BUFSOBMDBSF IJQQPDBNQBM TZOBQUPHFOFTJTBOEDPHOJUJWFEFWFMPQNFOUJOSBUT/BU/FVSPTDJ .BD2VFFO (. 3BNBLSJTIOBO , 3BUOBTJOHBO 3 $IFO # BOE :PVOH -5 %FTJQSBNJOFUSFBUNFOUSFEVDFTUIFMPOHUFSNCFIBWJPVSBMBOEOFVSPDIFNJDBMTFRVFMBF PGFBSMZMJGFNBUFSOBMTFQBSBUJPO*OU+/FVSPQTZDIPQIBSNBDPM .BDSJ4 .BTPO(+BOE8VSCFM) %JTTPDJBUJPOJOUIFFĒFDUTPGOFPOBUBMNBUFSOBM TFQBSBUJPOTPONBUFSOBMDBSFBOEUIFPĒ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ĒFDUVQPOBEVMUBESFOBMDPSUJDBM .
(32) $IBQUFS. . . . . SFTQPOTFUPTUSFTTBOEBESFOBMBOEPWBSJBODPNQFOTBUPSZIZQFSUSPQIZ&OEPDSJOPMPHZ 4DIBQJSP4 (FMMFS&BOE&JEVTPO4 /FPOBUBMBESFOBMDPSUJDBMSFTQPOTFUPTUSFTT BOEWBTPQSFTTJO1SPD4PD&YQ#JPM.FE 4DINJEU .7 %FVTTJOH +. 0JU[M .4 0IM ' -FWJOF 4 8VSTU 8 )PMTCPFS ' .VMMFS .# BOE EF ,MPFU &3 %JĒFSFOUJBM EJTJOIJCJUJPO PG UIF OFPOBUBM IZQPUIBMBNJD QJUVJUBSZBESFOBMBYJTJOCSBJOTQFDJêD$3)SFDFQUPSLOPDLPVUNJDF&VS+/FVSPTDJ 4DINJEU.7 &OUIPWFO- WBOEFS.BSL. -FWJOF4 EF,MPFU&3BOE0JU[M.4 ѮFQPTUOBUBMEFWFMPQNFOUPGUIFIZQPUIBMBNJDQJUVJUBSZBESFOBMBYJTJOUIFNPVTF*OU +%FW/FVSPTDJ 4DINJEU.7 &OUIPWFO- WBO8PF[JL+) -FWJOF4 EF,MPFU&3BOE0JU[M.4 Ѯ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ѮF)1"TZTUFNEVSJOHUIF QPTUOBUBMEFWFMPQNFOUPG$%NJDFBOEUIFFĒFDUTPGNBUFSOBMEFQSJWBUJPO#SBJO3FT %FW#SBJO3FT 4NPUIFSNBO81 .FOEP[B41BOE-FWJOF4 0OUPHFOFUJDDIBOHFTJOQVQFMJDJUFE NBUFSOBMQJUVJUBSZBESFOBMBDUJWJUZQVQBHFBOETUBHFPGMBDUBUJPOFĒFDUT%FW1TZDIPCJPM 4UBOUPO.& (VUJFSSF[:3BOE-FWJOF4 .BUFSOBMEFQSJWBUJPOQPUFOUJBUFTQJUVJUBSZ BESFOBMTUSFTTSFTQPOTFTJOJOGBOUSBUT#FIBW/FVSPTDJ WBO0FST)+ EF,MPFU&3 8IFMBO5BOE-FWJOF4 .BUFSOBMEFQSJWBUJPOFĒFDUPO UIFJOGBOUTOFVSBMTUSFTTNBSLFSTJTSFWFSTFECZUBDUJMFTUJNVMBUJPOBOEGFFEJOHCVUOPUCZ TVQQSFTTJOHDPSUJDPTUFSPOF+/FVSPTDJ 7B[RVF[ %. 4USFTT BOE UIF EFWFMPQJOH MJNCJDIZQPUIBMBNJDQJUVJUBSZBESFOBM BYJT1TZDIPOFVSPFOEPDSJOPMPHZ 7FOFSPTJ " $JSVMMJ ' $BQPOF ' BOE "MMFWB & 1SPMPOHFE QFSJOBUBM ";5 BENJOJTUSBUJPOBOEFBSMZNBUFSOBMTFQBSBUJPOFĒFDUTPOTPDJBMBOEFNPUJPOBMCFIBWJPVS PGQFSJBEPMFTDFOUNJDF1IBSNBDPM#JPDIFN#FIBW 8BMLFS$% 1FSSJO. 7BMF8BOE3JWJFS$ 0OUPHFOZPGUIFTUSFTTSFTQPOTFJOUIF SBUSPMFPGUIFQJUVJUBSZBOEUIFIZQPUIBMBNVT&OEPDSJOPMPHZ 8FBWFS*$ $FSWPOJ/ $IBNQBHOF'" %"MFTTJP"$ 4IBSNB4 4FDLM+3 %ZNPW4 4[ZG .BOE.FBOFZ.+ &QJHFOFUJDQSPHSBNNJOHCZNBUFSOBMCFIBWJPS/BU/FVSPTDJ 8PSLFM+0 0JU[M.4 -FEFCPFS"BOEEF,MPFU&3 ѮF#SPXO/PSXBZSBUEJTQMBZT FOIBODFETUSFTTJOEVDFE"$5)SFBDUJWJUZBUEBZBѫFSINBUFSOBMEFQSJWBUJPOBU EBZ#SBJO3FT%FW#SBJO3FT ;IBOH-9 -FWJOF4 %FOU(8 ;IBO: 9JOH( 0LJNPUP%, ,BUIMFFO(. 1PTU3. BOE4NJUI." .BUFSOBMEFQSJWBUJPOJODSFBTFTDFMMEFBUIJOUIFJOGBOUSBUCSBJO #SBJO3FT%FW#SBJO3FT .
(33)
(34)
(35)
GERELATEERDE DOCUMENTEN
License: Licence agreement concerning inclusion of doctoral thesis in the Institutional Repository of the University of Leiden. Downloaded
Chapter 1 ZQPUIFTFTBOEBQQSPBDIFT 1BSU*i3FQFBUFENBUFSOBMTFQBSBUJPOTuѮFEBUBEFTDSJCFEJOTFDUJPOTBOEIBWFMFE
" NBJO FĒFDU XBT PCTFSWFE PG USFBUNFOU ' 1 BOE JOKFDUJPO ' 1 BT XFMM BT BO JOUFSBDUJPO CFUXFFO USFBUNFOU BOE JOKFDUJPO ' 1 JOEJDBUJOH UIBU UIF FĒFDU PG JOKFDUJPO
%JTDVTTJPO $% NJDF SFBEJMZ MFBSOFE UIJT TQBUJBM OBWJHBUJPO UBTL JOEJDBUFE CZ UIF EFDSFBTFE MBUFODZ BOE EJTUBODF TXBN UP MPDBUF UIF TVCNFSHFE QMBUGPSN .BUFSOBM EFQSJWBUJPO EJE
"CTUSBDU .BUFSOBMEFQSJWBUJPO BTFQBSBUJPOPGNPUIFSBOEQVQTGPSIPVSTJOUIFêSTUXFFLTPGMJGF IBTMPOHMBTUJOHDPOTFRVFODFTGPSUIFHMVDPDPSUJDPJETUSFTTTZTUFNѮFTFFĒFDUTBSFLOPXOUP EFQFOE PO UIF BHF
$IBQUFS 3N3/"XBTBSSFTUFEPOEBZ"MUIPVHI 3N3/"FYQSFTTJPOHSBEVBMMZJODSFBTFEBHBJO JU SFNBJOFECFIJOEUIFMFWFMTNFBTVSFEJODPOUSPMBOJNBMTVOUJMBUMFBTUQOE*OFBSMZBEPMFTDFODF QOE 3 N3/"
*OTQJUFPGUIJTBEBQUBUJPO UIF1"BYJTSFNBJOFESFTQPOTJWFBOEUIFDPSUJDPTUFSPOFTFDSFUJPO CFDBNFFWFOSFMBUJWFMZNPSFTFOTJUJTFEUPUIFFĒFDUTPGOPWFMUZFYQPTVSFѮJTêOEJOHJOEJDBUFT UIBU UIF
NBUFSOBM BCTFODF XBT HSBEVBMMZ FMJNJOBUFE XIFO UIF NBUFSOBM TFQBSBUJPOT XFSF SFQFBUFE UIF OFYUUXPEBZT%FTQJUFUIFBCTFODFPGBO1"BYJTSFTQPOTFUPUIFUIJSENBUFSOBMTFQBSBUJPO