University Press of Florida

Biology

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Title:
Biology
Physical Description:
Textbook
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OpenStax College
Publisher:
OpenStax College
Rice University
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Houston, TX
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University Press of Florida
Holding Location:
University Press of Florida
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This item is licensed with the Creative Commons Attribution License. This license lets others distribute, remix, tweak, and build upon this work, even commercially, as long as they credit the author for the original creation.
Resource Identifier:
isbn-10 - 1938168097
isbn-13 - 978-1-938168-09-3
System ID:
AA00026950:00001


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Biology

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OpenStax College Rice University 6100 Main Street MS380 Houston, Texas 77005 To learn more about OpenStax College, visit http://openstaxcollege.org Individual print copies and bulk orders can be purchased through our website. 2013 by Rice University. T he t extbook content was produced by OpenStax College and is licensed under a Creative Commons Attribution 3.0 Unported License. Under this license, any user of this textbook or the textbook contents herein must provide proper attribution as follows: If you redistribute this textbook in a digital format (including but not limited to EPUB, PDF, and HTML), then you must dis play on every page view the following attribution: Download for free at http://cnx.org/content/col11448/latest/ If you redistribute this textbook in a print format, then you must display on every phy sical p age the following attribution: Download for free at http://cnx.org/content/col11448/latest/ If you redistribute part of this textbook, then you must display o n every digital format page view ( including but not limited to EPUB, PDF, and HTML) and on every physical printed page the following attribution: Download for free at http://cnx.org/content/col11448/latest/ If you use this textbook as a bibliographic reference, then you should cite this textbook as follows: OpenStax College, Biology OpenStax College. 30 May 2013 < http://cnx.org/content/col11448/latest/ > Trademarks The OpenSta x College name, OpenStax College logo, OpenStax College book covers, Connexions name, and Connexions logo are not subject to the license and may not be reproduced without the prior and express written consent of Rice University. For questions regarding thi s licensing, please contact partners@openstaxcollege.org. ISBN 10 1938168097 ISBN 13 9781 938168093 Revision BM 1 001DW

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OpenStax College OpenStax College is a non profit organization committed to improving student access to quality learning materials. Our free textbooks are developed and peer reviewed by educators to ensure they are readable, accurate, and meet the scope and sequence requirements of modern college courses Through our partnerships with companies and foundations committed to reducing costs for students, OpenStax College is working to improve access to higher education for all. Connexions The technology platform supporting OpenStax College is Connexions ( http://cnx.org ), one of the worlds first and largest open education projects. Connexions provides students with free online and low cost print editions of the OpenStax College library and provides instructors with tools to customize the content so that they can have th e perfect book for their course. Rice University OpenStax College and Connexions are initiative s of Rice University. As a leading research university with a distinctive commitment to undergraduate education, Rice University aspires to path breaking research, unsurpassed teaching, and contributions to the betterment of our world. It seeks to fulfill this mission by cultivating a diverse community of learning and discovery that produces leaders across the spectrum of human endeavor. Foundation Support OpenStax College is grateful for the tremendous support of our sponsors. Without their strong engagement, the goal of free access to highquality textbooks would remain just a dream The William and Flora Hewlett Foundation has been making grants since 1967 to help solve social and environmental problems at home and around the world. The Foundation concentrates its resources on activities in education, the environment, global development and population, performing arts, and philanthropy, and makes grants to support disadvantaged communities in the San Francisco Bay Area. Guided by the belief that every life has equal value, the Bill & Melinda Gates Foundation works to help all people lead healthy, productive lives. In developing countries, it focuses on improving peoples health with vaccines and other life saving tools and giving them the chance to lift themselves out of hunger and ex treme poverty. In the United States, it seeks to significantly improve education so that all young people have the opportunity to reach their full potential. Based in Seattle, Washington, the foundation is led by CEO Jeff Raikes and Co chair William H. Gat es Sr., under the direction of Bill and Melinda Gates and Warren Buffett. Our mission at the Twenty Million Minds Foundation is to grow access and success by eliminating unnecessary hurdles to affordability. We support the creation, sharing, and proliferation of more effective, more affordable educational content by leveraging disruptive technologies, open educational resources, and new models for collaboration between for profit, nonprofit, and public entities. The Maxfield Foundation supports projects with potential for high impact in science, education, sustainability, and other areas of social importance.

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chemicalequation rUFCPCGLRFCLSK@CPMD?RMKQMDC?AFCJCKCLRGQRFCQ?KCMLC?AFQGBCMDRFC COS?RGML"AAMPBGLERMRFCJ?UMDAMLQCPT?RGMLMDK?RRCPrRFCLSK@CPMD?RMKQ@CDMPC?LB?DRCP? AFCKGA?JPC?ARGMLQFMSJB@CCOS?JrQSAFRF?RLM?RMKQ?PCrSLBCPLMPK?JAGPASKQR?LACQrAPC?RCBMP BCQRPMWCB &TCLRFMSEF?JJMDRFCPC?AR?LRQ?LBNPMBSARQMDRFGQPC?ARGML?PCKMJCASJCQtC?AF?RMKPCK?GLQ@MLBCB RM?RJC?QRMLCMRFCP?RMKnrGLRFGQPC?ARGMLMLJWFWBPMECLNCPMVGBC?LBU?RCP?PCPCNPCQCLR?RGTCMD? QS@AJ?QQMDKMJCASJCQILMUL?Q compounds RFCWAMLR?GL?RMKQMDKMPCRF?LMLCRWNCMDCJCKCLR .MJCASJ?PMVWECLrMLRFCMRFCPF?LBr?QQFMULGL Figure2.10rAMLQGQRQMDRUMBMS@JW@MLBCBMVWECL ?RMKQ?LBGQLMRAJ?QQGDGCB?Q?AMKNMSLB@SR?Q?LCJCKCLR !@>LIQLJPFK>K/ JLIB@RIB>OBGLFKBA?V>ALR?IB?LKA 4MKCAFCKGA?JPC?ARGMLQrQSAF?QRFCMLCQFMUL?@MTCrA?LNPMACCBGLMLCBGPCARGMLSLRGJRFCPC?AR?LRQ ?PC?JJSQCBSN5FCCOS?RGMLQRF?RBCQAPG@CRFCQCPC?ARGMLQAMLR?GL?SLGBGPCARGML?J?PPMU?LB?PC irreversible Reversiblereactions ?PCRFMQCRF?RA?LEMGLCGRFCPBGPCARGML*LPCTCPQG@JCPC?ARGMLQr PC?AR?LRQ?PCRSPLCBGLRMNPMBSARQr@SRUFCLRFCAMLACLRP?RGMLMDNPMBSAREMCQ@CWMLB?ACPR?GLRFPCQFMJB tAF?P?ARCPGQRGAMDRFCN?PRGASJ?PPC?ARGMLnrQMKCMDRFCQCNPMBSARQUGJJ@CAMLTCPRCB@?AIGLRMPC?AR?LRQ ?RRFGQNMGLRrRFCBCQGEL?RGMLQMDNPMBSARQ?LBPC?AR?LRQ?PCPCTCPQCB5FGQ@?AI?LBDMPRFAMLRGLSCQSLRGJ ?ACPR?GLPCJ?RGTC@?J?LAC@CRUCCLPC?AR?LRQ?LBNPMBSARQMAASPQq?QR?RCA?JJCB equilibrium 5FCQC QGRS?RGMLQMDPCTCPQG@JCPC?ARGMLQ?PCMDRCLBCLMRCB@W?AFCKGA?JCOS?RGMLUGRF?BMS@JCFC?BCB?PPMU NMGLRGLERMU?PBQ@MRFRFCPC?AR?LRQ?LBNPMBSARQ 'MPCV?KNJCrGLFSK?L@JMMBrCVACQQFWBPMECLGMLQt) f n@GLBRM@GA?P@ML?RCGMLQt)$0 nDMPKGLE?L COSGJG@PGSKQR?RCUGRFA?P@MLGA?AGBt) $0 n*DA?P@MLGA?AGBUCPC?BBCBRMRFGQQWQRCKrQMKCMDGR UMSJB@CAMLTCPRCBRM@GA?P@ML?RC?LBFWBPMECLGMLQ *L@GMJMEGA?JPC?ARGMLQrFMUCTCPrCOSGJG@PGSKGQP?PCJWM@R?GLCB@CA?SQCRFCAMLACLRP?RGMLQMDRFC PC?AR?LRQMPNPMBSARQMP@MRF?PCAMLQR?LRJWAF?LEGLErMDRCLUGRF?NPMBSARMDMLCPC?ARGML@CGLE? PC?AR?LRDMP?LMRFCP5MPCRSPLRMRFCCV?KNJCMDCVACQQFWBPMECLGMLQGLRFC@JMMBrRFCDMPK?RGMLMD A?P@MLGA?AGBUGJJ@CRFCK?HMPBGPCARGMLMDRFCPC?ARGML)MUCTCPrRFCA?P@MLGA?AGBA?L?JQMJC?TCRFC @MBW?QA?P@MLBGMVGBCE?QtTG?CVF?J?RGMLnGLQRC?BMD@CGLEAMLTCPRCB@?AIRM@GA?P@ML?RCGMLrRFSQ BPGTGLERFCPC?ARGMLRMRFCPGEFR@WRFCAFCKGA?JJ?UILMUL?Q lawofmassaction 5FCQCPC?ARGMLQ?PC GKNMPR?LRDMPK?GLR?GLGLERFCFMKCMQR?QGQMDMSP@JMMB $FEJE;$FE@:FE;J 4MKC?RMKQ?PCKMPCQR?@JCUFCLRFCWE?GLMPJMQC?LCJCARPMLtMPNMQQG@JWRUMn?LBDMPKGMLQ5FGQ DGJJQRFCGPMSRCPKMQRCJCARPMLQFCJJ?LBK?ICQRFCKCLCPECRGA?JJWKMPCQR?@JC#CA?SQCRFCLSK@CPMD CJCARPMLQBMCQLMRCOS?JRFCLSK@CPMDNPMRMLQrC?AFGMLF?Q?LCRAF?PEC Cations ?PCNMQGRGTCGMLQRF?R ?PCDMPKCB@WJMQGLECJCARPMLQ/CE?RGTCGMLQ?PCDMPKCB@WE?GLGLECJCARPMLQ?LB?PCA?JJCB?LGMLQ Anions ?PCBCQGEL?RCB@WRFCGPCJCKCLR?JL?KC@CGLE?JRCPCBRMCLBGLtGBCuRFC?LGMLMDAFJMPGLCGQ A?JJCBAFJMPGBCr?LBRFC?LGMLMDQSJDSPGQA?JJCBQSJDGBCrDMPCV?KNJC 5FGQKMTCKCLRMDCJCARPMLQDPMKMLCCJCKCLRRM?LMRFCPGQPCDCPPCBRM?Q electrontransfer "Q Figure 2.11 GJJSQRP?RCQrQMBGSKt/?nMLJWF?QMLCCJCARPMLGLGRQMSRCPCJCARPMLQFCJJ*RR?ICQJCQQCLCPEWDMP QMBGSKRMBML?RCRF?RMLCCJCARPMLRF?LGRBMCQRM?AACNRQCTCLKMPCCJCARPMLQRMDGJJRFCMSRCPQFCJJ*D QMBGSKJMQCQ?LCJCARPMLrGRLMUF?QNPMRMLQrLCSRPMLQr?LBMLJWCJCARPMLQrJC?TGLEGRUGRF?L MTCP?JJAF?PECMDf*RGQLMUPCDCPPCBRM?Q?QMBGSKGML$FJMPGLCt$JnGLGRQJMUCQRCLCPEWQR?RCtA?JJCB RFCEPMSLBQR?RCnF?QQCTCLCJCARPMLQGLGRQMSRCPQFCJJ"E?GLrGRGQKMPCCLCPEWCDDGAGCLRDMPAFJMPGLC RME?GLMLCCJCARPMLRF?LRMJMQCQCTCL5FCPCDMPCrGRRCLBQRME?GL?LCJCARPMLRMAPC?RC?LGMLUGRF NPMRMLQrLCSRPMLQr?LBCJCARPMLQrEGTGLEGR?LCRLCE?RGTCtpnAF?PEC*RGQLMUPCDCPPCBRM?Q? AFJMPGBCGML*LRFGQCV?KNJCrQMBGSKUGJJBML?RCGRQMLCCJCARPMLRMCKNRWGRQQFCJJr?LBAFJMPGLCUGJJ #(!04%2X(%#(%-)#!,&/5.$!4)/./&,)&%

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!@>LI< 0BMQFAB?LKACLOJ>QFLKFP>ABEVAO>QFLKPVKQEBPFPOB>@QFLK4EB@>O?LUVIDOLRMLCLKB >JFKL>@FAFPIFKHBAQLQEB>JFKLDOLRMLCQEBFK@LJFKD>JFKL>@FA)KQEBMOL@BPPr>JLIB@RIBLC T>QBOFPOBIB>PBA 5FCNPMBSARQDMPKCB@WQSAFJGLI?ECQ?PCA?JJCBNCNRGBCQ"QKMPC?KGLM?AGBQHMGLRMRFGQEPMUGLE AF?GLrRFCPCQSJRGLEAF?GLGQILMUL?Q?NMJWNCNRGBC&?AFNMJWNCNRGBCF?Q?DPCC?KGLMEPMSN?RMLCCLB 5FGQCLBGQA?JJCBRFC/RCPKGL?JrMPRFC?KGLMRCPKGL?Jr?LBRFCMRFCPCLBF?Q?DPCCA?P@MVWJEPMSNr ?JQMILMUL?QRFC$MPA?P@MVWJRCPKGL?J8FGJCRFCRCPKQNMJWNCNRGBC?LBNPMRCGL?PCQMKCRGKCQSQCB GLRCPAF?LEC?@JWr?NMJWNCNRGBCGQRCAFLGA?JJW?NMJWKCPMD?KGLM?AGBQrUFCPC?QRFCRCPKNPMRCGLGQSQCB DMP?NMJWNCNRGBCMPNMJWNCNRGBCQRF?RF?TCAMK@GLCBRMECRFCPrMDRCLF?TC@MSLBLMLNCNRGBCNPMQRFCRGA EPMSNQrF?TC?BGQRGLARQF?NCr?LBF?TC?SLGOSCDSLARGML"DRCPNPMRCGLQWLRFCQGQtRP?LQJ?RGMLnrKMQR NPMRCGLQ?PCKMBGDGCB5FCQC?PCILMUL?QNMQRRP?LQJ?RGML?JKMBGDGA?RGMLQ5FCWK?WSLBCPEMAJC?T?ECr NFMQNFMPWJ?RGMLrMPK?WPCOSGPCRFC?BBGRGMLMDMRFCPAFCKGA?JEPMSNQ0LJW?DRCPRFCQCKMBGDGA?RGMLQGQ RFCNPMRCGLAMKNJCRCJWDSLARGML?J $JGAIRFPMSEFRFCQRCNQMDNPMRCGLQWLRFCQGQGLRFGQ interactivetutorial(http://openstaxcollege.org/l/ protein_synth) #(!04%2X")/,/')#!,-!#2/-/,%#5,%3 4EFP@LKQBKQFP>S>FI>?IBCLOCOBB>QEQQM@KULOD@LKQBKQ@LI

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/?< MFCLK@FE8IP.@>E@=@:8E:KFJMLOQ>KQ@LJMLKBKQLCQEBBIB@QOLKQO>KPMLOQ@E>FKr>M>OQLC@BIIRI>O OBPMFO>QFLKr>KAFQFPKLOJ>IIVCLRKAFKQEB@BIIRI>OLOD>KBIIBrQEBJFQL@ELKAOFLK4EFP MOLQBFKE>P>EBJBMOLPQEBQF@DOLRMr>KAQEB@BKQO>IFLKLCQEBEBJBDBQP>IQBOK>QBIV OBAR@BA>KALUFAFWBAAROFKDBIB@QOLKQO>KPCBO"B@>RPBQEFPBPPBKQF>IMOLQBFKwPOLIBFK MOLAR@FKD@BIIRI>OBKBODVFP@OR@F>IrFQE>P@E>KDBASBOVIFQQIBLSBOJFIIFLKPLCVB>OP0OLQBFK PBNRBK@FKDE>PPELTKQE>QQEBOBFP>@LKPFABO>?IB>JLRKQLC@VQL@EOLJB@>JFKL>@FA PBNRBK@BELJLILDV>JLKDAFCCBOBKQPMB@FBPFKLQEBOTLOAPrBSLIRQFLK>OVHFKPEFM@>K?B >PPBPPBA?VJB>PROFKDQEBPFJFI>OFQFBPLOAFCCBOBK@BP>JLKDS>OFLRPPMB@FBPw$.!LO MOLQBFKPBNRBK@BP 3@FBKQFPQPE>SBABQBOJFKBAQE>QERJ>K@VQL@EOLJB@@LKQ>FKP>JFKL>@FAP&LOB>@E @VQL@EOLJB@JLIB@RIBCOLJAFCCBOBKQLOD>KFPJPQE>QE>P?BBKPBNRBK@BAQLA>QBr LCQEBPB>JFKL>@FAP>MMB>OFKQEBP>JBMLPFQFLKFK>IIP>JMIBPLC@VQL@EOLJB@4EFP FKAF@>QBPQE>QQEBOBJ>VE>SB?BBK>@LJJLK>K@BPQLO/K@LJM>OFKDQEBERJ>K>KA @EFJM>KWBBMOLQBFKPBNRBK@BPrKLPBNRBK@BAFCCBOBK@BT>PCLRKA7EBKERJ>K>KA OEBPRPJLKHBVPBNRBK@BPTBOB@LJM>OBArQEBPFKDIBAFCCBOBK@BCLRKAT>PFKLKB>JFKL >@FA)K>KLQEBO@LJM>OFPLKrERJ>KQLVB>PQPBNRBK@FKDPELTP>AFCCBOBK@BFKQEBQE MLPFQFLK +IFK<@E.KIL:KLI< "QBGQASQQCBC?PJGCPrRFCQF?NCMD?NPMRCGLGQAPGRGA?JRMGRQDSLARGML'MPCV?KNJCr?LCLXWKCA?L@GLBRM ?QNCAGDGAQS@QRP?RC?R?QGRCILMUL?QRFC?ARGTCQGRC*DRFGQ?ARGTCQGRCGQ?JRCPCB@CA?SQCMDJMA?JAF?LECQ MPAF?LECQGLMTCP?JJNPMRCGLQRPSARSPCrRFCCLXWKCK?W@CSL?@JCRM@GLBRMRFCQS@QRP?RC5MSLBCPQR?LB FMURFCNPMRCGLECRQGRQDGL?JQF?NCMPAMLDMPK?RGMLrUCLCCBRMSLBCPQR?LBRFCDMSPJCTCJQMDNPMRCGL QRPSARSPCNPGK?PWrQCAMLB?PWrRCPRG?PWr?LBOS?RCPL?PW /2'7>!97:)9:7+ 5FCSLGOSCQCOSCLACMD?KGLM?AGBQGL?NMJWNCNRGBCAF?GLGQGRQ primarystructure 'MPCV?KNJCr RFCN?LAPC?RGAFMPKMLCGLQSJGLF?QRUMNMJWNCNRGBCAF?GLQr"?LB#r?LBRFCW?PCJGLICBRMECRFCP@W BGQSJDGBC@MLBQ5FC/RCPKGL?J?KGLM?AGBMDRFC"AF?GLGQEJWAGLCrUFCPC?QRFC$RCPKGL?J?KGLM?AGB GQ?QN?P?EGLCt Figure 3.25 nFCQCOSCLACQMD?KGLM?AGBQGLRFC"?LB#AF?GLQ?PCSLGOSCRMGLQSJGL !@>LI< "LSFKBPBORJFKPRIFKFP>MOLQBFKELOJLKBJ>ABLCQTLMBMQFAB@E>FKPr!t>JFKL >@FAPILKDn>KA"t>JFKL>@FAPILKDn)KB>@E@E>FKrMOFJ>OVPQOR@QROBFPFKAF@>QBA?VQEOBBIBQQBO >??OBSF>QFLKPQE>QOBMOBPBKQQEBK>JBPLCQEB>JFKL>@FAPFKQEBLOABOQEBV>OBMOBPBKQ4EB>JFKL >@FA@VPQBFKBt@VPnE>P>PRICEVAOVIt3(nDOLRM>P>PFAB@E>FK4TLPRICEVAOVIDOLRMP@>KOB>@QFK QEBMOBPBK@BLCLUVDBKQLCLOJ>AFPRICFABt3n?LKA4TLAFPRICFAB?LKAP@LKKB@QQEB!>KA" @E>FKPQLDBQEBOr>KA>QEFOAEBIMPQEB!@E>FKCLIAFKQLQEB@LOOB@QPE>MB.LQBQE>Q>IIAFPRICFAB?LKAP >OBQEBP>JBIBKDQEr?RQ>OBAO>TKAFCCBOBKQPFWBPCLO@I>OFQV #(!04%2X")/,/')#!,-!#2/-/,%#5,%3

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5FCSLGOSCQCOSCLACDMPCTCPWNPMRCGLGQSJRGK?RCJWBCRCPKGLCB@WRFCECLCCLAMBGLERFCNPMRCGL" AF?LECGLLSAJCMRGBCQCOSCLACMDRFCECLCsQAMBGLEPCEGMLK?WJC?BRM?BGDDCPCLR?KGLM?AGB@CGLE ?BBCBRMRFCEPMUGLENMJWNCNRGBCAF?GLrA?SQGLE?AF?LECGLNPMRCGLQRPSARSPC?LBDSLARGML*LQGAIJC ACJJ?LCKG?rRFCFCKMEJM@GL D AF?GLt?QK?JJNMPRGMLMDUFGAFGQQFMULGL Figure3.26nF?Q?QGLEJC ?KGLM?AGBQS@QRGRSRGMLrA?SQGLE?AF?LECGLNPMRCGLQRPSARSPC?LBDSLARGML4NCAGDGA?JJWrRFC?KGLM ?AGBEJSR?KGA?AGBGQQS@QRGRSRCB@WT?JGLCGLRFC D AF?GL8F?RGQKMQRPCK?PI?@JCRMAMLQGBCPGQ RF?R?FCKMEJM@GLKMJCASJCGQK?BCSNMDRUM?JNF?AF?GLQ?LBRUM@CR?AF?GLQRF?RC?AFAMLQGQRMD ?@MSR?KGLM?AGBQ5FCKMJCASJCrRFCPCDMPCrF?Q?@MSR?KGLM?AGBQ5FCQRPSARSP?JBGDDCPCLAC @CRUCCL?LMPK?JFCKMEJM@GLKMJCASJC?LB?QGAIJCACJJKMJCASJCqUFGAFBP?K?RGA?JJWBCAPC?QCQJGDC CVNCAR?LAWqGQ?QGLEJC?KGLM?AGBMDRFC8F?RGQCTCLKMPCPCK?PI?@JCGQRF?RRFMQC?KGLM ?AGBQ?PCCLAMBCB@WRFPCCLSAJCMRGBCQC?AFr?LBRFCKSR?RGMLGQA?SQCB@W?QGLEJC@?QCAF?LECtNMGLR KSR?RGMLnrGL@?QCQ !@>LI< 4EB?BQ>@E>FKLCEBJLDIL?FKFPOBPFARBPFKIBKDQErVBQ>PFKDIB>JFKL>@FA PR?PQFQRQFLKIB>APQLPF@HIB@BII>KBJF>)KKLOJ>IEBJLDIL?FKrQEB>JFKL>@FA>QMLPFQFLKPBSBKFP DIRQ>J>QB)KPF@HIB@BIIEBJLDIL?FKrQEFPDIRQ>J>QBFPOBMI>@BA?V>S>IFKB #CA?SQCMDRFGQAF?LECMDMLC?KGLM?AGBGLRFCAF?GLrFCKMEJM@GLKMJCASJCQDMPKJMLEDG@CPQRF?R BGQRMPRRFC@GAMLA?TCrMPBGQAQF?NCBrPCB@JMMBACJJQ?LB?QQSKC?APCQACLRMPtQGAIJCuQF?NCrUFGAF AJMEQ?PRCPGCQtFigure3.27n5FGQA?LJC?BRMKWPG?BQCPGMSQFC?JRFNPM@JCKQQSAF?Q@PC?RFJCQQLCQQr BGXXGLCQQrFC?B?AFCQr?LB?@BMKGL?JN?GLDMPRFMQC?DDCARCB@WRFGQBGQC?QC !@>LI< )KQEFP?ILLAPJB>OrSFPR>IFWBA>QUJ>DKFCF@>QFLKRPFKD?OFDEQCFBIAJF@OLP@LMVrPF@HIB @BIIP>OB@OBP@BKQPE>MBArTEFIBKLOJ>I@BIIP>OBAFP@PE>MBAt@OBAFQJLAFCF@>QFLKLCTLOH?V%A 5QEJ>KP@>IB?>OA>Q>COLJ->QQRPPBIIn !+)43*'7>!97:)9:7+ 5FCJMA?JDMJBGLEMDRFCNMJWNCNRGBCGLQMKCPCEGMLQEGTCQPGQCRMRFC secondarystructure MDRFCNPMRCGL 5FCKMQRAMKKML?PCRFC -helix ?LB -pleatedsheet QRPSARSPCQtFigure3.28n#MRFQRPSARSPCQ?PCRFC C FCJGVQRPSARSPCqRFCFCJGVFCJBGLQF?NC@WFWBPMECL@MLBQ5FCFWBPMECL@MLBQDMPK@CRUCCLRFC MVWECL?RMKGLRFCA?P@MLWJEPMSNGLMLC?KGLM?AGB?LB?LMRFCP?KGLM?AGBRF?RGQDMSP?KGLM?AGBQ D?PRFCP?JMLERFCAF?GL #(!04%2X")/,/')#!,-!#2/-/,%#5,%3 4EFP@LKQBKQFP>S>FI>?IBCLOCOBB>QEQQM@KULOD@LKQBKQ@LI

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!@>LI< 4EB CEBIFU>KA DMIB>QBAPEBBQ>OBPB@LKA>OVPQOR@QROBPLCMOLQBFKPQE>QCLOJ?B@>RPB LCEVAOLDBK?LKAFKD?BQTBBK@>O?LKVI>KA>JFKLDOLRMPFKQEBMBMQFAB?>@H?LKB#BOQ>FK>JFKL >@FAPE>SB>MOLMBKPFQVQLCLOJ>K CEBIFUrTEFIBLQEBOPE>SB>MOLMBKPFQVQLCLOJ> DMIB>QBAPEBBQ &TCPWFCJGA?JRSPLGL?L?JNF?FCJGVF?Q?KGLM?AGBPCQGBSCQ5FC3EPMSNQtRFCT?PG?LREPMSNQnMD RFCNMJWNCNRGBCNPMRPSBCMSRDPMKRFC C FCJGVAF?GL*LRFC DNJC?RCBQFCCRrRFCtNJC?RQu?PCDMPKCB@W FWBPMECL@MLBGLE@CRUCCL?RMKQMLRFC@?AI@MLCMDRFCNMJWNCNRGBCAF?GL5FC3EPMSNQ?PC?RR?AFCB RMRFCA?P@MLQ?LBCVRCLB?@MTC?LB@CJMURFCDMJBQMDRFCNJC?R5FCNJC?RCBQCEKCLRQ?JGELN?P?JJCJMP ?LRGN?P?JJCJRMC?AFMRFCPr?LBFWBPMECL@MLBQDMPK@CRUCCLRFCN?PRG?JJWNMQGRGTCLGRPMECL?RMKGLRFC ?KGLMEPMSN?LBRFCN?PRG?JJWLCE?RGTCMVWECL?RMKGLRFCA?P@MLWJEPMSNMDRFCNCNRGBC@?AI@MLC5FC C FCJGV?LB DNJC?RCBQFCCRQRPSARSPCQ?PCDMSLBGLKMQREJM@SJ?P?LBDG@PMSQNPMRCGLQ?LBRFCWNJ?W?L GKNMPR?LRQRPSARSP?JPMJC "+79/'7>!97:)9:7+ 5FCSLGOSCRFPCCBGKCLQGML?JQRPSARSPCMD?NMJWNCNRGBCGQGRQ tertiarystructure tFigure3.29n5FGQ QRPSARSPCGQGLN?PRBSCRMAFCKGA?JGLRCP?ARGMLQ?RUMPIMLRFCNMJWNCNRGBCAF?GL1PGK?PGJWrRFC GLRCP?ARGMLQ?KMLE3EPMSNQAPC?RCQRFCAMKNJCVRFPCCBGKCLQGML?JRCPRG?PWQRPSARSPCMD?NPMRCGL5FC L?RSPCMDRFC3EPMSNQDMSLBGLRFC?KGLM?AGBQGLTMJTCBA?LAMSLRCP?ARRFCDMPK?RGMLMDRFCFWBPMECL @MLBQBCQAPG@CBDMPQR?LB?PBQCAMLB?PWQRPSARSPCQ'MPCV?KNJCr3EPMSNQUGRFJGICAF?PECQ?PCPCNCJJCB @WC?AFMRFCP?LBRFMQCUGRFSLJGICAF?PECQ?PC?RRP?ARCBRMC?AFMRFCPtGMLGA@MLBQn8FCLNPMRCGL DMJBGLER?ICQNJ?ACrRFCFWBPMNFM@GA3EPMSNQMDLMLNMJ?P?KGLM?AGBQJ?WGLRFCGLRCPGMPMDRFCNPMRCGLr UFCPC?QRFCFWBPMNFGJGA3EPMSNQJ?WMLRFCMSRQGBC5FCDMPKCPRWNCQMDGLRCP?ARGMLQ?PC?JQMILMUL ?QFWBPMNFM@GAGLRCP?ARGMLQ*LRCP?ARGML@CRUCCLAWQRCGLCQGBCAF?GLQDMPKQBGQSJDGBCJGLI?ECQGLRFC NPCQCLACMDMVWECLrRFCMLJWAMT?JCLR@MLBDMPKGLEBSPGLENPMRCGLDMJBGLE #(!04%2X")/,/')#!,-!#2/-/,%#5,%3

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!@>LI< 4EBQBOQF>OVPQOR@QROBLCMOLQBFKPFPABQBOJFKBA?V>S>OFBQVLC@EBJF@>IFKQBO>@QFLKP 4EBPBFK@IRABEVAOLMEL?F@FKQBO>@QFLKPrFLKF@?LKAFKDrEVAOLDBK?LKAFKD>KAAFPRICFABIFKH>DBP "JJMDRFCQCGLRCP?ARGMLQrUC?I?LBQRPMLErBCRCPKGLCRFCDGL?JRFPCCBGKCLQGML?JQF?NCMDRFCNPMRCGL 8FCL?NPMRCGLJMQCQGRQRFPCCBGKCLQGML?JQF?NCrGRK?WLMJMLECP@CDSLARGML?J :'9+73'7>!97:)9:7+ *LL?RSPCrQMKCNPMRCGLQ?PCDMPKCBDPMKQCTCP?JNMJWNCNRGBCQr?JQMILMUL?QQS@SLGRQr?LBRFC GLRCP?ARGMLMDRFCQCQS@SLGRQDMPKQRFC quaternarystructure 8C?IGLRCP?ARGMLQ@CRUCCLRFCQS@SLGRQ FCJNRMQR?@GJGXCRFCMTCP?JJQRPSARSPC'MPCV?KNJCrGLQSJGLt?EJM@SJ?PNPMRCGLnF?Q?AMK@GL?RGMLMD FWBPMECL@MLBQ?LBBGQSJDGBC@MLBQRF?RA?SQCGRRM@CKMQRJWAJSKNCBGLRM?@?JJQF?NC*LQSJGLQR?PRQ MSR?Q?QGLEJCNMJWNCNRGBC?LBJMQCQQMKCGLRCPL?JQCOSCLACQGLRFCNPCQCLACMDNMQRRP?LQJ?RGML?J KMBGDGA?RGML?DRCPRFCDMPK?RGMLMDRFCBGQSJDGBCJGLI?ECQRF?RFMJBRFCPCK?GLGLEAF?GLQRMECRFCP4GJIt? DG@PMSQNPMRCGLnrFMUCTCPrF?Q? DNJC?RCBQFCCRQRPSARSPCRF?RGQRFCPCQSJRMDFWBPMECL@MLBGLE@CRUCCL BGDDCPCLRAF?GLQ 5FCDMSPJCTCJQMDNPMRCGLQRPSARSPCtNPGK?PWrQCAMLB?PWrRCPRG?PWr?LBOS?RCPL?PWn?PCGJJSQRP?RCBGL Figure 3.30 #(!04%2X")/,/')#!,-!#2/-/,%#5,%3 4EFP@LKQBKQFP>S>FI>?IBCLOCOBB>QEQQM@KULOD@LKQBKQ@LI

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!@>LIK?BL?PBOSBAFKQEBPBFIIRPQO>QFLKPt@OBAFQ JLAFCF@>QFLKLCTLOH?V.>QFLK>I(RJ>K'BKLJBBPB>O@E)KPQFQRQBn &?AFNPMRCGLF?QGRQMULSLGOSCQCOSCLAC?LBQF?NCRF?R?PCFCJBRMECRFCP@WAFCKGA?JGLRCP?ARGMLQ *DRFCNPMRCGLGQQS@HCARRMAF?LECQGLRCKNCP?RSPCrN)rMPCVNMQSPCRMAFCKGA?JQrRFCNPMRCGLQRPSARSPC K?WAF?LECrJMQGLEGRQQF?NCUGRFMSRJMQGLEGRQNPGK?PWQCOSCLACGLUF?RGQILMUL?QBCL?RSP?RGML %CL?RSP?RGMLGQMDRCLPCTCPQG@JC@CA?SQCRFCNPGK?PWQRPSARSPCMDRFCNMJWNCNRGBCGQAMLQCPTCBGLRFC NPMACQQGDRFCBCL?RSPGLE?ECLRGQPCKMTCBr?JJMUGLERFCNPMRCGLRMPCQSKCGRQDSLARGML4MKCRGKCQ BCL?RSP?RGMLGQGPPCTCPQG@JCrJC?BGLERMJMQQMDDSLARGML0LCCV?KNJCMDGPPCTCPQG@JCNPMRCGLBCL?RSP?RGML GQUFCL?LCEEGQDPGCB5FC?J@SKGLNPMRCGLGLRFCJGOSGBCEEUFGRCGQBCL?RSPCBUFCLNJ?ACBGL?FMRN?L /MR?JJNPMRCGLQ?PCBCL?RSPCB?RFGEFRCKNCP?RSPCQDMPGLQR?LACr@?ARCPG?RF?RQSPTGTCGLFMRQNPGLEQF?TC NPMRCGLQRF?RDSLARGML?RRCKNCP?RSPCQAJMQCRM@MGJGLE5FCQRMK?AFGQ?JQMTCPW?AGBGArF?Q?JMUN)r?LB BCL?RSPCQNPMRCGLQ?QN?PRMDRFCBGECQRGMLNPMACQQFMUCTCPrRFCBGECQRGTCCLXWKCQMDRFCQRMK?AFPCR?GL RFCGP?ARGTGRWSLBCPRFCQCAMLBGRGMLQ 1PMRCGLDMJBGLEGQAPGRGA?JRMGRQDSLARGML*RU?QMPGEGL?JJWRFMSEFRRF?RRFCNPMRCGLQRFCKQCJTCQUCPC PCQNMLQG@JCDMPRFCDMJBGLENPMACQQ0LJWPCACLRJWU?QGRDMSLBRF?RMDRCLRFCWPCACGTC?QQGQR?LACGLRFC DMJBGLENPMACQQDPMKNPMRCGLFCJNCPQILMUL?Q chaperones tMPAF?NCPMLGLQnRF?R?QQMAG?RCUGRFRFCR?PECR NPMRCGLBSPGLERFCDMJBGLENPMACQQ5FCW?AR@WNPCTCLRGLE?EEPCE?RGMLMDNMJWNCNRGBCQRF?RK?ICSNRFC AMKNJCRCNPMRCGLQRPSARSPCr?LBRFCWBGQ?QQMAG?RCDPMKRFCNPMRCGLMLACRFCR?PECRNPMRCGLGQDMJBCB #(!04%2X")/,/')#!,-!#2/-/,%#5,%3

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RMUMPIGLRF?RNPMACQQ4MKCCV?KNJCQMDNSKNQDMP?ARGTCRP?LQNMPR?PC/? f f "51?QCrUFGAFA?PPGCQ QMBGSK?LBNMR?QQGSKGMLQr?LB) f f "51?QCrUFGAFA?PPGCQFWBPMECL?LBNMR?QQGSKGMLQ#MRFMD RFCQC?PC?LRGNMPRCPA?PPGCPNPMRCGLQ5UMMRFCPA?PPGCPNPMRCGLQ?PC$? f "51?QC?LB) f "51?QCrUFGAF A?PPWMLJWA?JAGSK?LBMLJWFWBPMECLGMLQrPCQNCARGTCJW#MRF?PCNSKNQ !@>LI< !RKFMLOQBO@>OOFBPLKBJLIB@RIBLOFLK!PVJMLOQBO@>OOFBPQTLAFCCBOBKQJLIB@RIBPLO FLKPr?LQEFKQEBP>JBAFOB@QFLK!K>KQFMLOQBO>IPL@>OOFBPQTLAFCCBOBKQJLIB@RIBPLOFLKPr?RQFK AFCCBOBKQAFOB@QFLKPt@OBAFQJLAFCF@>QFLKLCTLOH?Vx,RM>PHyFHFJBAF>#LJJLKPn +I@D8IP:K@MLI< 0OFJ>OV>@QFSBQO>KPMLOQJLSBPFLKP>@OLPP>JBJ?O>KBr@OB>QFKD>KBIB@QOL@EBJF@>I DO>AFBKQtBIB@QOLDBKF@QO>KPMLOQnt@OBAFQJLAFCF@>QFLKLCTLOH?V->OF>K>RFWFII>OB>In 0LCMDRFCKMQRGKNMPR?LRNSKNQGL?LGK?JQACJJQGQRFCQMBGSKNMR?QQGSKNSKNt/? f f "51?QCnr UFGAFK?GLR?GLQRFCCJCARPMAFCKGA?JEP?BGCLRt?LBRFCAMPPCARAMLACLRP?RGMLQMD/? f ?LB, f nGLJGTGLE ACJJQ5FCQMBGSKNMR?QQGSKNSKNKMTCQ, f GLRMRFCACJJUFGJCKMTGLE/? f MSR?RRFCQ?KCRGKCr?R? P?RGMMDRFPCC/? f DMPCTCPWRUM, f GMLQKMTCBGL5FC/? f f "51?QCCVGQRQGLRUMDMPKQrBCNCLBGLE MLGRQMPGCLR?RGMLRMRFCGLRCPGMPMPCVRCPGMPMDRFCACJJ?LBGRQ?DDGLGRWDMPCGRFCPQMBGSKMPNMR?QQGSKGMLQ 5FCNPMACQQAMLQGQRQMDRFCDMJJMUGLEQGVQRCNQ 8GRFRFCCLXWKCMPGCLRCBRMU?PBQRFCGLRCPGMPMDRFCACJJrRFCA?PPGCPF?Q?FGEF?DDGLGRWDMPQMBGSK GMLQFPCCGMLQ@GLBRMRFCNPMRCGL "51GQFWBPMJWXCB@WRFCNPMRCGLA?PPGCP?LB?JMUCLCPEWNFMQNF?RCEPMSN?RR?AFCQRMGR "Q?PCQSJRrRFCA?PPGCPAF?LECQQF?NC?LBPCMPGCLRQGRQCJDRMU?PBQRFCCVRCPGMPMDRFCKCK@P?LC 5FCNPMRCGLsQ?DDGLGRWDMPQMBGSKBCAPC?QCQ?LBRFCRFPCCQMBGSKGMLQJC?TCRFCA?PPGCP 5FCQF?NCAF?LECGLAPC?QCQRFCA?PPGCPsQ?DDGLGRWDMPNMR?QQGSKGMLQr?LBRUMQSAFGMLQ?RR?AFRMRFC NPMRCGLS@QCOSCLRJWrRFCJMUCLCPEWNFMQNF?RCEPMSNBCR?AFCQDPMKRFCA?PPGCP 8GRFRFCNFMQNF?RCEPMSNPCKMTCB?LBNMR?QQGSKGMLQ?RR?AFCBrRFCA?PPGCPNPMRCGLPCNMQGRGMLQGRQCJD RMU?PBQRFCGLRCPGMPMDRFCACJJ #(!04%2X#452%!.$&5.#4)/./&,!3-!-%-"2!.%3

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!@>LI< !KBIB@QOL@EBJF@>IDO>AFBKQr@OB>QBA?VMOFJ>OV>@QFSBQO>KPMLOQr@>KJLSBLQEBO PR?PQ>K@BP>D>FKPQQEBFO@LK@BKQO>QFLKDO>AFBKQPr>MOL@BPP@>IIBA@LQO>KPMLOQLOPB@LKA>OV >@QFSBQO>KPMLOQt@OBAFQJLAFCF@>QFLKLCTLOH?V->OF>K>RFWFII>OB>In )CQEBM(LRQPFABQEB@BIIAB@OB>PBPrTLRIAVLRBUMB@QQEB>JLRKQLC>JFKL>@FAP QO>KPMLOQBAFKQLQEB@BIIQLFK@OB>PBLOAB@OB>PB R LCB/I8EJGFIK #WRFCCLBMDRFGQQCARGMLrWMSUGJJ@C?@JCRM z %CQAPG@CCLBMAWRMQGQrGLAJSBGLENF?EMAWRMQGQrNGLMAWRMQGQr?LBPCACNRMPKCBG?RCBCLBMAWRMQGQ z 6LBCPQR?LBRFCNPMACQQMDCVMAWRMQGQ *L?BBGRGMLRMKMTGLEQK?JJGMLQ?LBKMJCASJCQRFPMSEFRFCKCK@P?LCrACJJQ?JQMLCCBRMPCKMTC?LB R?ICGLJ?PECPKMJCASJCQ?LBN?PRGAJCQtQCC Table5.2 DMPCV?KNJCQn4MKCACJJQ?PCCTCLA?N?@JCMD CLESJDGLECLRGPCSLGACJJSJ?PKGAPMMPE?LGQKQ:MSKGEFRF?TCAMPPCARJWFWNMRFCQGXCBRF?RRFCSNR?IC?LB PCJC?QCMDJ?PECN?PRGAJCQ@WRFCACJJPCOSGPCQCLCPEW"J?PECN?PRGAJCrFMUCTCPrA?LLMRN?QQRFPMSEFRFC KCK@P?LCrCTCLUGRFCLCPEWQSNNJGCB@WRFCACJJ E;F:PKFJ@J Endocytosis GQ?RWNCMD?ARGTCRP?LQNMPRRF?RKMTCQN?PRGAJCQrQSAF?QJ?PECKMJCASJCQrN?PRQMDACJJQr ?LBCTCLUFMJCACJJQrGLRM?ACJJ5FCPC?PCBGDDCPCLRT?PG?RGMLQMDCLBMAWRMQGQr@SR?JJQF?PC?AMKKML AF?P?ARCPGQRGA5FCNJ?QK?KCK@P?LCMDRFCACJJGLT?EGL?RCQrDMPKGLE?NMAICR?PMSLBRFCR?PECRN?PRGAJC 5FCNMAICRNGLAFCQMDDrPCQSJRGLEGLRFCN?PRGAJC@CGLEAMLR?GLCBGL?LCUJWAPC?RCBGLRP?ACJJSJ?PTCQGAJC DMPKCBDPMKRFCNJ?QK?KCK@P?LC .'-4)>948/8 1F?EMAWRMQGQtRFCAMLBGRGMLMDtACJJC?RGLEunGQRFCNPMACQQ@WUFGAFJ?PECN?PRGAJCQrQSAF?QACJJQMP PCJ?RGTCJWJ?PECN?PRGAJCQr?PCR?ICLGL@W?ACJJ'MPCV?KNJCrUFCLKGAPMMPE?LGQKQGLT?BCRFCFSK?L @MBWr?RWNCMDUFGRC@JMMBACJJA?JJCB?LCSRPMNFGJUGJJPCKMTCRFCGLT?BCPQRFPMSEFRFGQNPMACQQr QSPPMSLBGLE?LBCLESJDGLERFCKGAPMMPE?LGQKrUFGAFGQRFCLBCQRPMWCB@WRFCLCSRPMNFGJt Figure 5.20 n #(!04%2X#452%!.$&5.#4)/./&,!3-!-%-"2!.%3

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!@>LI< )KMFKL@VQLPFPrQEB@BIIJBJ?O>KBFKS>DFK>QBPrPROOLRKAP>PJ>IISLIRJBLCCIRFAr>KA MFK@EBPLCCt@OBAFQ->OF>K>RFWFII>OB>In "T?PG?RGMLMDNGLMAWRMQGQGQA?JJCB potocytosis 5FGQNPMACQQSQCQ?AM?RGLENPMRCGLrA?JJCB caveolin rML RFCAWRMNJ?QKGAQGBCMDRFCNJ?QK?KCK@P?LCrUFGAFNCPDMPKQ?QGKGJ?PDSLARGMLRMAJ?RFPGL5FCA?TGRGCQ GLRFCNJ?QK?KCK@P?LCRF?RDMPKRFCT?ASMJCQF?TCKCK@P?LCPCACNRMPQ?LBJGNGBP?DRQGL?BBGRGML RMA?TCMJGL5FCT?ASMJCQMPTCQGAJCQDMPKCBGLA?TCMJ?CtQGLESJ?PA?TCMJ?n?PCQK?JJCPRF?LRFMQCGL NGLMAWRMQGQ1MRMAWRMQGQGQSQCBRM@PGLEQK?JJKMJCASJCQGLRMRFCACJJ?LBRMRP?LQNMPRRFCQCKMJCASJCQ RFPMSEFRFCACJJDMPRFCGPPCJC?QCMLRFCMRFCPQGBCMDRFCACJJr?NPMACQQA?JJCBRP?LQAWRMQGQ +)+5947+*/'9+**4)>948/8 "R?PECRCBT?PG?RGMLMDCLBMAWRMQGQCKNJMWQPCACNRMPNPMRCGLQGLRFCNJ?QK?KCK@P?LCRF?RF?TC? QNCAGDGA@GLBGLE?DDGLGRWDMPACPR?GLQS@QR?LACQt Figure 5.22 n #(!04%2X#452%!.$&5.#4)/./&,!3-!-%-"2!.%3

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1$ 2,0 ./$*). 5FC"6$?LB"6"AMBMLQGLK3/"@MRF QNCAGDWGQMJCSAGLCF?RDC?RSPCMDRFCECLCRGA AMBCCVNJ?GLQRFGQ > AMKNJCKCLR?PGRW ? LMLQCLQCAMBMLQ @ SLGTCPQ?JGRW A BCECLCP?AW )MUK?LWLSAJCMRGBCQ?PCGLK3/" AMBMLQ > ? @ A 8FGAFQS@SLGRMDRFC b*630 NMJWKCP?QC AMLDCPQQNCAGDGAGRWRMRP?LQAPGNRGML > C ? D @ Db A H 5FC?LBPCEGMLQMDNPMI?PWMRGA NPMKMRCPQ?PCA?JJCBAMLQCLQSQQCOSCLACQ@CA?SQC >>>>>>>> > RFCW?PCGBCLRGA?JGL?JJ@?ARCPG?JQNCAGCQ ? RFCW?PCQGKGJ?PGL?JJ@?ARCPG?JQNCAGCQ @ RFCWCVGQRGL?JJMPE?LGQKQ A RFCWF?TCRFCQ?KCDSLARGMLGL?JJ MPE?LGQKQ 8FGAFDC?RSPCMDNPMKMRCPQA?L@CDMSLBGL @MRFNPMI?PWMRCQ?LBCSI?PWMRCQ > ($@MV ? 5"5"@MV @ MAR?KCP@MV A ?LBQCOSCLACQ 8F?RRP?LQAPGNRQUGJJ@CKMQR?DDCARCB@WJMU JCTCJQMDi?K?LGRGL > ?LBP3/"Q ? NPCK3/"Q @ P3/"Q?LBR3/"Q A MRFCPQK?JJLSAJC?P/"Q r 8FGAFNPCK3/"NPMACQQGLEQRCNGQ GKNMPR?LRDMPGLGRG?RGLERP?LQJ?RGML > NMJW"R?GJ ? 3/"CBGRGLE @ QNJGAGLE A KCRFWJES?LMQGLCA?N 8F?RNPMACQQGLEQRCNCLF?LACQRFCQR?@GJGRWMD NPCR3/"Q?LBNPCP3/"Q > KCRFWJ?RGML ? LSAJCMRGBCKMBGDGA?RGML @ AJC?T?EC A QNJGAGLE 5FC/"AMKNMLCLRQMDPG@MQMKCQ?PC QWLRFCQGXCBGLRFC>>>>>>>> > AWRMNJ?QK ? LSAJCSQ @ LSAJCMJSQ A CLBMNJ?QKGAPCRGASJSK *L?LWEGTCLQNCAGCQrRFCPC?PC?RJC?QRFMU K?LWRWNCQMD?KGLM?AWJR3/"QWLRFCR?QCQ > ? @ A -$/$'/#$)&$)",0 ./$*). *K?EGLCGDRFCPCUCPCAMKKMLJW MAASPPGLE?KGLM?AGBQGLQRC?BMD(GTCLUF?R WMSILMU?@MSRRFCECLCRGAAMBCrUF?RUMSJB@C RFCQFMPRCQRNMQQG@JCAMBMLJCLERF &VNJ?GL %GQASQQFMUBCECLCP?AWMDRFCECLCRGAAMBC K?ICQACJJQKMPCPM@SQRRMKSR?RGMLQ *DK3/"GQAMKNJCKCLR?PWRMRFC%/" RCKNJ?RCQRP?LB?LBRFC%/"RCKNJ?RCQRP?LBGQ AMKNJCKCLR?PWRMRFC%/"LMLRCKNJ?RCQRP?LBr RFCLUFW?PC@?QCQCOSCLACQMDK3/"?LBRFC %/"LMLRCKNJ?RCQRP?LBLMRGBCLRGA?J $MSJB RFCWCTCP@C *LWMSPMULUMPBQrBCQAPG@CRFCBGDDCPCLAC @CRUCCLPFMBCNCLBCLR?LBPFMGLBCNCLBCLR RCPKGL?RGMLMDRP?LQAPGNRGMLGLNPMI?PWMRCQ 5P?LQAPG@C?LBRP?LQJ?RCRFCDMJJMUGLE%/" QCOSCLACtLMLRCKNJ?RCQRP?LBnb "5(($$((55"55""($"b &VNJ?GLFMUQGLEJCLSAJCMRGBCAF?LECQA?L F?TCT?QRJWBGDDCPCLRCDDCARQMLNPMRCGLDSLARGML #(!04%2X'%.%3!.$/4%).3 4EFP@LKQBKQFP>S>FI>?IBCLOCOBB>QEQQM@KULOD@LKQBKQ@LI

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R ) 3+..$*) !@>LI< 4EBDBKBQF@@LKQBKQLCB>@EPLJ>QF@@BIIFK>KLOD>KFPJFPQEBP>JBr?RQKLQ>IIDBKBP >OBBUMOBPPBAFKBSBOV@BII4EB@LKQOLILCTEF@EDBKBP>OBBUMOBPPBAAF@Q>QBPTEBQEBO>@BIIFPt>n >KBVB@BIILOt?n>IFSBO@BII)QFPQEBAFCCBOBKQF>IDBKBBUMOBPPFLKM>QQBOKPQE>Q>OFPBFKAFCCBOBKQ@BIIP QE>QDFSBOFPBQLt@n>@LJMIBQBLOD>KFPJ ?8GKLC8K@FEF="LC8K@FE LB8IPFK@: G@>LC8K@FE LB8IPFK@:/I8EJ:I@GK@FE"LC8K@FE LB8IPFK@:+FJKnKI8EJ:I@GK@FE8C"LC8K@FE LB8IPFK@:/I8EJC8K@FE8CE;+FJKnKI8EJC8K@FE8C"LC8K@FE E:LC8K@FE $EKIF;L:K@FE &?AFQMK?RGAACJJGLRFC@MBWECLCP?JJWAMLR?GLQRFCQ?KC%/""DCUCVACNRGMLQGLAJSBCPCB@JMMB ACJJQrUFGAFAMLR?GLLM%/"GLRFCGPK?RSPCQR?RCr?LBQMKCGKKSLCQWQRCKACJJQRF?RPC?PP?LECRFCGP %/"UFGJCNPMBSAGLE?LRG@MBGCQ*LECLCP?JrFMUCTCPrRFCECLCQRF?RBCRCPKGLCUFCRFCPWMSF?TCEPCCL CWCQr@PMULF?GPr?LBFMUD?QRWMSKCR?@MJGXCDMMB?PCRFCQ?KCGLRFCACJJQGLWMSPCWCQ?LBWMSPJGTCPr CTCLRFMSEFRFCQCMPE?LQDSLARGMLOSGRCBGDDCPCLRJW*DC?AFACJJF?QRFCQ?KC%/"rFMUGQGRRF?RACJJQMP MPE?LQ?PCBGDDCPCLR FWBMACJJQGLRFCCWCBGDDCPQMBP?K?RGA?JJWDPMKACJJQGLRFCJGTCP 8FCPC?QC?AFACJJQF?PCQRFCQ?KCECLMKC?LB%/"QCOSCLACrC?AFACJJBMCQLMRRSPLMLrMPCVNPCQQr RFCQ?KCQCRMDECLCQ&?AFACJJRWNCLCCBQ?BGDDCPCLRQCRMDNPMRCGLQRMNCPDMPKGRQDSLARGML5FCPCDMPCr MLJW?QK?JJQS@QCRMDNPMRCGLQGQCVNPCQQCBGL?ACJJ'MPRFCNPMRCGLQRM@CCVNPCQQCBrRFC%/"KSQR @CRP?LQAPG@CBGLRM3/"?LBRFC3/"KSQR@CRP?LQJ?RCBGLRMNPMRCGL*L?EGTCLACJJRWNCrLMR?JJECLCQ CLAMBCBGLRFC%/"?PCRP?LQAPG@CBGLRM3/"MPRP?LQJ?RCBGLRMNPMRCGL@CA?SQCQNCAGDGAACJJQGLMSP @MBWF?TCQNCAGDGADSLARGMLQ4NCAG?JGXCBNPMRCGLQRF?RK?ICSNRFCCWCtGPGQrJCLQr?LBAMPLC?n?PCMLJW CVNPCQQCBGLRFCCWCrUFCPC?QRFCQNCAG?JGXCBNPMRCGLQGLRFCFC?PRtN?ACK?ICPACJJQrFC?PRKSQAJCr?LB T?JTCQn?PCMLJWCVNPCQQCBGLRFCFC?PR"R?LWEGTCLRGKCrMLJW?QS@QCRMD?JJMDRFCECLCQCLAMBCB @WMSP%/"?PCCVNPCQQCB?LBRP?LQJ?RCBGLRMNPMRCGLQ5FCCVNPCQQGMLMDQNCAGDGAECLCQGQ?FGEFJW #(!04%2X'%.%%802%33)/.

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MFCLK@FEF="LC8K@FE 0OLH>OVLQF@@BIIP@>KLKIVOBDRI>QBDBKBBUMOBPPFLK?V@LKQOLIIFKDQEB>JLRKQLC QO>KP@OFMQFLK!PBRH>OVLQF@@BIIPBSLISBArQEB@LJMIBUFQVLCQEB@LKQOLILCDBKBBUMOBPPFLK FK@OB>PBA&LOBU>JMIBrTFQEQEBBSLIRQFLKLCBRH>OVLQF@@BIIP@>JB@LJM>OQJBKQ>IFW>QFLK LCFJMLOQ>KQ@BIIRI>O@LJMLKBKQP>KA@BIIRI>OMOL@BPPBP!KR@IB>OOBDFLKQE>Q@LKQ>FKP QEB$.!T>PCLOJBA4O>KP@OFMQFLK>KAQO>KPI>QFLKTBOBMEVPF@>IIVPBM>O>QBAFKQLQTL AFCCBOBKQ@BIIRI>O@LJM>OQJBKQP)QQEBOBCLOB?B@>JBMLPPF?IBQL@LKQOLIDBKBBUMOBPPFLK?V OBDRI>QFKDQO>KP@OFMQFLKFKQEBKR@IBRPr>KA>IPL?V@LKQOLIIFKDQEB2.!IBSBIP>KAMOLQBFK QO>KPI>QFLKMOBPBKQLRQPFABQEBKR@IBRP 3LJB@BIIRI>OMOL@BPPBP>OLPBCOLJQEBKBBALCQEBLOD>KFPJQLABCBKAFQPBIC#BIIRI>O MOL@BPPBPPR@E>PDBKBPFIBK@FKDABSBILMBAQLMOLQB@QQEB@BIICOLJSFO>ILOM>O>PFQF@ FKCB@QFLKP)CQEB@BII@LRIANRF@HIVPERQLCCDBKBBUMOBPPFLKCLO>PELOQMBOFLALCQFJBr FQTLRIA?B>?IBQLPROSFSB>KFKCB@QFLKTEBKLQEBOLOD>KFPJP@LRIAKLQ4EBOBCLOBrQEB LOD>KFPJBSLISBA>KBTMOL@BPPQE>QEBIMBAFQPROSFSBr>KAFQT>P>?IBQLM>PPQEFPKBT ABSBILMJBKQQLLCCPMOFKD R +IFB8IPFK@:"LC8K@FE #WRFCCLBMDRFGQQCARGMLrWMSUGJJ@C?@JCRM z %CQAPG@CRFCQRCNQGLTMJTCBGLNPMI?PWMRGAECLCPCESJ?RGML z &VNJ?GLRFCPMJCQMD?ARGT?RMPQrGLBSACPQr?LBPCNPCQQMPQGLECLCPCESJ?RGML 5FC%/"MDNPMI?PWMRCQGQMPE?LGXCBGLRM?AGPASJ?PAFPMKMQMKCQSNCPAMGJCBGLRFCLSAJCMGBPCEGML MDRFCACJJAWRMNJ?QK1PMRCGLQRF?R?PCLCCBCBDMP?QNCAGDGADSLARGMLrMPRF?R?PCGLTMJTCBGLRFCQ?KC @GMAFCKGA?JN?RFU?Wr?PCCLAMBCBRMECRFCPGL@JMAIQA?JJCB operons'MPCV?KNJCr?JJMDRFCECLCQ LCCBCBRMSQCJ?ARMQC?Q?LCLCPEWQMSPAC?PCAMBCBLCVRRMC?AFMRFCPGLRFCJ?ARMQCtMP 3(* nMNCPML *LNPMI?PWMRGAACJJQrRFCPC?PCRFPCCRWNCQMDPCESJ?RMPWKMJCASJCQRF?RA?L?DDCARRFCCVNPCQQGMLMD MNCPMLQPCNPCQQMPQr?ARGT?RMPQr?LBGLBSACPQ Repressors ?PCNPMRCGLQRF?RQSNNPCQQRP?LQAPGNRGMLMD? ECLCGLPCQNMLQCRM?LCVRCPL?JQRGKSJSQrUFCPC?Q activators ?PCNPMRCGLQRF?RGLAPC?QCRFCRP?LQAPGNRGML MD?ECLCGLPCQNMLQCRM?LCVRCPL?JQRGKSJSQ'GL?JJWrGLBSACPQ?PCQK?JJKMJCASJCQRF?RCGRFCP?ARGT?RCMP PCNPCQQRP?LQAPGNRGMLBCNCLBGLEMLRFCLCCBQMDRFCACJJ?LBRFC?T?GJ?@GJGRWMDQS@QRP?RC /?< 975 *GS>FI>?IBCLOCOBB>QEQQM@KULOD@LKQBKQ@LI

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!@>LI< 4EBCFSBDBKBPQE>Q>OBKBBABAQLPVKQEBPFWBQOVMQLME>KFK *630 >OBIL@>QBAKBUQ QLB>@ELQEBOFKQEB ;97 LMBOLK7EBKQOVMQLME>KFPMIBKQFCRIrQTLQOVMQLME>KJLIB@RIBP?FKAQEB OBMOBPPLOMOLQBFK>QQEBLMBO>QLOPBNRBK@B4EFPMEVPF@>IIV?IL@HPQEB2.!MLIVJBO>PBCOLJ QO>KP@OF?FKDQEBQOVMQLME>KDBKBP7EBKQOVMQLME>KFP>?PBKQrQEBOBMOBPPLOMOLQBFKALBPKLQ?FKA QLQEBLMBO>QLO>KAQEBDBKBP>OBQO>KP@OF?BA "%/"QCOSCLACRF?RAMBCQDMPNPMRCGLQGQPCDCPPCBRM?QRFCAMBGLEPCEGML5FCDGTCAMBGLEPCEGMLQDMP RFCRPWNRMNF?L@GMQWLRFCQGQCLXWKCQ?PC?PP?LECBQCOSCLRG?JJWMLRFCAFPMKMQMKCGLRFCMNCPML+SQR @CDMPCRFCAMBGLEPCEGMLGQRFC transcriptionalstartsite 5FGQGQRFCPCEGMLMD%/"RMUFGAF3/" NMJWKCP?QC@GLBQRMGLGRG?RCRP?LQAPGNRGML5FCNPMKMRCPQCOSCLACGQSNQRPC?KMDRFCRP?LQAPGNRGML?JQR?PR QGRCC?AFMNCPMLF?Q?QCOSCLACUGRFGLMPLC?PRFCNPMKMRCPRMUFGAFNPMRCGLQt?ARGT?RMPQMPPCNPCQQMPQn A?L@GLB?LBPCESJ?RCRP?LQAPGNRGML "%/"QCOSCLACA?JJCBRFCMNCP?RMPQCOSCLACGQCLAMBCB@CRUCCLRFCNPMKMRCPPCEGML?LBRFCDGPQR ;97 AMBGLEECLC5FGQ operator AMLR?GLQRFC%/"AMBCRMUFGAFRFCPCNPCQQMPNPMRCGLA?L@GLB8FCL RPWNRMNF?LGQNPCQCLRGLRFCACJJrRUMRPWNRMNF?LKMJCASJCQ@GLBRMRFC ;97 PCNPCQQMPrUFGAFAF?LECQ QF?NCRM@GLBRMRFC ;97 MNCP?RMP#GLBGLEMDRFCRPWNRMNF?LpPCNPCQQMPAMKNJCV?RRFCMNCP?RMPNFWQGA?JJW NPCTCLRQRFC/"NMJWKCP?QCDPMK@GLBGLEr?LBRP?LQAPG@GLERFCBMULQRPC?KECLCQ 8FCLRPWNRMNF?LGQLMRNPCQCLRGLRFCACJJrRFCPCNPCQQMP@WGRQCJDBMCQLMR@GLBRMRFCMNCP?RMPRFCPCDMPCr RFCMNCPMLGQ?ARGTC?LBRPWNRMNF?LGQQWLRFCQGXCB#CA?SQCRFCPCNPCQQMPNPMRCGL?ARGTCJW@GLBQRMRFC MNCP?RMPRMICCNRFCECLCQRSPLCBMDDrRFC ;97 MNCPMLGQLCE?RGTCJWPCESJ?RCB?LBRFCNPMRCGLQRF?R@GLBRM RFCMNCP?RMPRMQGJCLAC ;97 CVNPCQQGML?PC negative regulators 8?RAF this video (http://openstaxcollege.org/l/trp_operon) RMJC?PLKMPC?@MSRRFC ;97 MNCPML K89FC@K<:K@M8KFI+IFK<@E+E:K@M8KFI-<>LC8KFI +SQR?QRFC ;97 MNCPMLGQLCE?RGTCJWPCESJ?RCB@WRPWNRMNF?LKMJCASJCQrRFCPC?PCNPMRCGLQRF?R@GLBRM RFCMNCP?RMPQCOSCLACQRF?R?AR?Q? positiveregulator RMRSPLECLCQML?LB?ARGT?RCRFCK'MPCV?KNJCr UFCLEJSAMQCGQQA?PACr b*630 @?ARCPG?A?LRSPLRMMRFCPQSE?PQMSPACQDMPDSCJ5MBMRFGQrLCUECLCQRM NPMACQQRFCQC?JRCPL?RCECLCQKSQR@CRP?LQAPG@CB8FCLEJSAMQCJCTCJQBPMNrAWAJGA".1tA".1n@CEGLQ RM?AASKSJ?RCGLRFCACJJ5FCA".1KMJCASJCGQ?QGEL?JGLEKMJCASJCRF?RGQGLTMJTCBGLEJSAMQC?LB CLCPEWKCR?@MJGQKGL b*630 8FCLEJSAMQCJCTCJQBCAJGLCGLRFCACJJr?AASKSJ?RGLEA".1@GLBQRMRFC #(!04%2X'%.%%802%33)/.

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NMQGRGTCPCESJ?RMP cataboliteactivatorprotein(CAP) r?NPMRCGLRF?R@GLBQRMRFCNPMKMRCPQMDMNCPMLQ RF?RAMLRPMJRFCNPMACQQGLEMD?JRCPL?RGTCQSE?PQ8FCLA".1@GLBQRM$"1rRFCAMKNJCV@GLBQRMRFC NPMKMRCPPCEGMLMDRFCECLCQRF?R?PCLCCBCBRMSQCRFC?JRCPL?RCQSE?PQMSPACQtFigure16.4n*LRFCQC MNCPMLQr?$"1@GLBGLEQGRCGQJMA?RCBSNQRPC?KMDRFC3/"NMJWKCP?QC@GLBGLEQGRCGLRFCNPMKMRCP 5FGQGLAPC?QCQRFC@GLBGLE?@GJGRWMD3/"NMJWKCP?QCRMRFCNPMKMRCPPCEGML?LBRFCRP?LQAPGNRGMLMDRFC ECLCQ !@>LI< 7EBKDIR@LPBIBSBIPC>IIr *630 J>VRPBLQEBOPRD>OPCLOCRBI?RQJRPQQO>KP@OF?BKBT DBKBPQLALPL!PDIR@LPBPRMMIFBP?B@LJBIFJFQBAr@!-0IBSBIPFK@OB>PB4EFP@!-0?FKAPQLQEB #!0MOLQBFKr>MLPFQFSBOBDRI>QLOQE>Q?FKAPQL>KLMBO>QLOOBDFLKRMPQOB>JLCQEBDBKBPOBNRFOBAQL RPBLQEBOPRD>OPLRO@BP /?< 1') *GS>FI>?IBCLOCOBB>QEQQM@KULOD@LKQBKQ@LI

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!@>LI< 4O>KP@OFMQFLKLCQEB 3(* LMBOLKFP@>OBCRIIVOBDRI>QBAPLQE>QFQPBUMOBPPFLKLKIV L@@ROPTEBKDIR@LPBFPIFJFQBA>KAI>@QLPBFPMOBPBKQQLPBOSB>P>K>IQBOK>QFSBCRBIPLRO@B )K *630rQEB ;97 LMBOLKFPLK?VABC>RIQrTEFIBQEB 3(* LMBOLKFPLCC7EVALVLRQEFKHQEFP FPQEB@>PB *DEJSAMQCGQ?@QCLRrRFCL$"1A?L@GLBRMRFCMNCP?RMPQCOSCLACRM?ARGT?RCRP?LQAPGNRGML*DJ?ARMQCGQ ?@QCLRrRFCLRFCPCNPCQQMP@GLBQRMRFCMNCP?RMPRMNPCTCLRRP?LQAPGNRGML*DCGRFCPMDRFCQCPCOSGPCKCLRQGQ KCRrRFCLRP?LQAPGNRGMLPCK?GLQMDD0LJWUFCL@MRFAMLBGRGMLQ?PCQ?RGQDGCBGQRFC 3(* MNCPMLRP?LQAPG@CB tTable 16.2 n .@>E8CJK?8K$E;L:
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