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8/19/2019 Biology Textbook RICE University _Part4
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%SPGLE RFGQ CQRCP @MLB DMPK?RGML, RFPCC U?RCP KMJCASJCQ ?PC PCJC?QCB. FC RFPCC D?RRW ?AGBQ GL
RFC RPG?AWJEJWACPMJ K?W @C QGKGJ?P MP BGQQGKGJ?P. ?RQ ?PC ?JQM A?JJCB triacylglycerols MP triglycerides
@CA?SQC MD RFCGP AFCKGA?J QRPSARSPC. MKC D?RRW ?AGBQ F?TC AMKKML L?KCQ RF?R QNCAGDW RFCGP MPGEGL. MP
CV?KNJC, N?JKGRGA ?AGB, ? saturated fatty acid, GQ BCPGTCB DPMK RFC N?JK RPCC. P?AFGBGA ?AGB GQ BCPGTCB
DPMK A9(*/0: /@76.(, RFC QAGCLRGDGA L?KC DMP EPMSLBLSRQ MP NC?LSRQ.
?RRW ?AGBQ K?W @C Q?RSP?RCB MP SLQ?RSP?RCB. *L ? D?RRW ?AGB AF?GL, GD RFCPC ?PC MLJW QGLEJC @MLBQ
@CRUCCL LCGEF@MPGLE A?P@MLQ GL RFC FWBPMA?P@ML AF?GL, RFC D?RRW ?AGB GQ Q?GB RM @C Q?RSP?RCB. ?RSP?RCB
D?RRW ?AGBQ ?PC Q?RSP?RCB UGRF FWBPMECL; GL MRFCP UMPBQ, RFC LSK@CP MD FWBPMECL ?RMKQ ?RR?AFCB RM RFC
A?P@ML QICJCRML GQ K?VGKGXCB. RC?PGA ?AGB GQ ?L CV?KNJC MD ? Q?RSP?RCB D?RRW ?AGB (Figure 3.14)
!@>LI< 3.14 QB>OF@ >@FA FP > @LJJLK P>QRO>QBA C>QQV >@FA.
FCL RFC FWBPMA?P@ML AF?GL AMLR?GLQ ? BMS@JC @MLB, RFC D?RRW ?AGB GQ Q?GB RM @C unsaturated. JCGA
?AGB GQ ?L CV?KNJC MD ?L SLQ?RSP?RCB D?RRW ?AGB (Figure 3.15).
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!@>LI< 3.15 IBF@ >@FA FP > @LJJLK RKP>QRO>QBA C>QQV >@FA.
MQR SLQ?RSP?RCB D?RQ ?PC JGOSGB ?R PMMK RCKNCP?RSPC ?LB ?PC A?JJCB MGJQ. *D RFCPC GQ MLC BMS@JC @MLB
GL RFC KMJCASJC, RFCL GR GQ ILMUL ?Q ? KMLMSLQ?RSP?RCB D?R (C.E., MJGTC MGJ), ?LB GD RFCPC GQ KMPC RF?L MLC
BMS@JC @MLB, RFCL GR GQ ILMUL ?Q ? NMJWSLQ?RSP?RCB D?R (C.E., A?LMJ? MGJ).FCL ? D?RRW ?AGB F?Q LM BMS@JC @MLBQ, GR GQ ILMUL ?Q ? Q?RSP?RCB D?RRW ?AGB @CA?SQC LM KMPC
FWBPMECL K?W @C ?BBCB RM RFC A?P@ML ?RMKQ MD RFC AF?GL. D?R K?W AMLR?GL QGKGJ?P MP BGDDCPCLR D?RRW
?AGBQ ?RR?AFCB RM EJWACPMJ. MLE QRP?GEFR D?RRW ?AGBQ UGRF QGLEJC @MLBQ RCLB RM ECR N?AICB RGEFRJW ?LB ?PC
QMJGB ?R PMMK RCKNCP?RSPC. LGK?J D?RQ UGRF QRC?PGA ?AGB ?LB N?JKGRGA ?AGB (AMKKML GL KC?R) ?LB RFC D?R
UGRF @SRWPGA ?AGB (AMKKML GL @SRRCP) ?PC CV?KNJCQ MD Q?RSP?RCB D?RQ. ?KK?JQ QRMPC D?RQ GL QNCAG?JGXCB
ACJJQ A?JJCB ?BGNMAWRCQ, UFCPC EJM@SJCQ MD D?R MAASNW KMQR MD RFC ACJJsQ TMJSKC. *L NJ?LRQ, D?R MP MGJ GQ
QRMPCB GL K?LW QCCBQ ?LB GQ SQCB ?Q ? QMSPAC MD CLCPEW BSPGLE QCCBJGLE BCTCJMNKCLR. LQ?RSP?RCB D?RQ
MP MGJQ ?PC SQS?JJW MD NJ?LR MPGEGL ?LB AMLR?GL *0: SLQ?RSP?RCB D?RRW ?AGBQ. C0: ?LB ;9(5: GLBGA?RC RFC
AMLDGESP?RGML MD RFC KMJCASJC ?PMSLB RFC BMS@JC @MLB. *D FWBPMECLQ ?PC NPCQCLR GL RFC Q?KC NJ?LC, GR GQ
PCDCPPCB RM ?Q ? AGQ D?R; GD RFC FWBPMECL ?RMKQ ?PC ML RUM BGDDCPCLR NJ?LCQ, GR GQ PCDCPPCB RM ?Q ? trans
fat.FC *0: BMS@JC @MLB A?SQCQ ? @CLB MP ? tIGLIu RF?R NPCTCLRQ RFC D?RRW ?AGBQ DPMK N?AIGLE RGEFRJW,
ICCNGLE RFCK JGOSGB ?R PMMK RCKNCP?RSPC (Figure 3.16). JGTC MGJ, AMPL MGJ, A?LMJ? MGJ, ?LB AMB JGTCP
MGJ ?PC CV?KNJCQ MD SLQ?RSP?RCB D?RQ. LQ?RSP?RCB D?RQ FCJN RM JMUCP @JMMB AFMJCQRCPMJ JCTCJQ UFCPC?Q
Q?RSP?RCB D?RQ AMLRPG@SRC RM NJ?OSC DMPK?RGML GL RFC ?PRCPGCQ.
84 #!% 3 X "'#! !#%#%
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!@>LI< 3.16 >QRO>QBA C>QQV >@FAP E>SB EVAOL@>O?LK @E>FKP @LKKB@QBA ?V PFKDIB ?LKAP LKIV.
KP>QRO>QBA C>QQV >@FAP E>SB LKB LO JLOB ALR?IB ?LKAP. %>@E ALR?IB ?LKA J>V ?B FK > *0: LO ;9(5:
@LKCFDRO>QFLK. K QEB *0: @LKCFDRO>QFLK, ?LQE EVAOLDBKP >OB LK QEB P>JB PFAB LC QEB EVAOL@>O?LK
@E>FK. K QEB ;9(5: @LKCFDRO>QFLK, QEB EVAOLDBKP >OB LK LMMLPFQB PFABP. ! *0: ALR?IB ?LKA @>RPBP >HFKH FK QEB @E>FK.
"7'38 F'98
*L RFC DMMB GLBSQRPW, MGJQ ?PC ?PRGDGAG?JJW FWBPMECL?RCB RM K?IC RFCK QCKG-QMJGB ?LB MD ? AMLQGQRCLAW
BCQGP?@JC DMP K?LW NPMACQQCB DMMB NPMBSARQ. GKNJW QNC?IGLE, FWBPMECL E?Q GQ @S@@JCB RFPMSEF MGJQ
RM QMJGBGDW RFCK. %SPGLE RFGQ FWBPMECL?RGML NPMACQQ, BMS@JC @MLBQ MD RFC *0:- AMLDMPK?RGML GL RFC
FWBPMA?P@ML AF?GL K?W @C AMLTCPRCB RM BMS@JC @MLBQ GL RFC RP?LQ- AMLDMPK?RGML.
?PE?PGLC, QMKC RWNCQ MD NC?LSR @SRRCP, ?LB QFMPRCLGLE ?PC CV?KNJCQ MD ?PRGDGAG?JJW FWBPMECL?RCB
RP?LQ D?RQ. CACLR QRSBGCQ F?TC QFMUL RF?R ?L GLAPC?QC GL RP?LQ D?RQ GL RFC FSK?L BGCR K?W JC?B RM ?L
GLAPC?QC GL JCTCJQ MD JMU-BCLQGRW JGNMNPMRCGLQ (%), MP t@?Bu AFMJCQRCPMJ, UFGAF GL RSPL K?W JC?B RM
NJ?OSC BCNMQGRGML GL RFC ?PRCPGCQ, PCQSJRGLE GL FC?PR BGQC?QC. ?LW D?QR DMMB PCQR?SP?LRQ F?TC PCACLRJW
@?LLCB RFC SQC MD RP?LQ D?RQ, ?LB DMMB J?@CJQ ?PC PCOSGPCB RM BGQNJ?W RFC RP?LQ D?R AMLRCLR.
O2+-' F'99> A)/*8
&QQCLRG?J D?RRW ?AGBQ ?PC D?RRW ?AGBQ PCOSGPCB @SR LMR QWLRFCQGXCB @W RFC FSK?L @MBW. $MLQCOSCLRJW, RFCW
F?TC RM @C QSNNJCKCLRCB RFPMSEF GLECQRGML TG? RFC BGCR. Omega-3 D?RRW ?AGBQ (JGIC RF?R QFMUL GL Figure
3.17) D?JJ GLRM RFGQ A?RCEMPW ?LB ?PC MLC MD MLJW RUM ILMUL DMP FSK?LQ (RFC MRFCP @CGLE MKCE?-6 D?RRW
?AGB). FCQC ?PC NMJWSLQ?RSP?RCB D?RRW ?AGBQ ?LB ?PC A?JJCB MKCE?-3 @CA?SQC RFC RFGPB A?P@ML DPMK RFC
CLB MD RFC FWBPMA?P@ML AF?GL GQ AMLLCARCB RM GRQ LCGEF@MPGLE A?P@ML @W ? BMS@JC @MLB.
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!@>LI< 3.17 !IME>-IFKLIBKF@ >@FA FP >K BU>JMIB LC >K LJBD>-3 C>QQV >@FA. Q E>P QEOBB *0: ALR?IB
?LKAP >KA, >P > OBPRIQ, > @ROSBA PE>MB. &LO @I>OFQV, QEB @>O?LKP >OB KLQ PELTK. %>@E PFKDIV ?LKABA
@>O?LK E>P QTL EVAOLDBKP >PPL@F>QBA TFQE FQ, >IPL KLQ PELTK.
FC D?PRFCQR A?P@ML ?U?W DPMK RFC A?P@MVWJ EPMSN GQ LSK@CPCB ?Q RFC MKCE? (I) A?P@ML, ?LB GD
RFC BMS@JC @MLB GQ @CRUCCL RFC RFGPB ?LB DMSPRF A?P@ML DPMK RF?R CLB, GR GQ ILMUL ?Q ?L MKCE?-3 D?RRW
?AGB. SRPGRGML?JJW GKNMPR?LR @CA?SQC RFC @MBW BMCQ LMR K?IC RFCK, MKCE?-3 D?RRW ?AGBQ GLAJSBC ?JNF?-
JGLMJCGA ?AGB (), CGAMQ?NCLR?CLMGA ?AGB (&), ?LB BMAMQ?FCV?CLMGA ?AGB (%), ?JJ MD UFGAF ?PC
NMJWSLQ?RSP?RCB. ?JKML, RPMSR, ?LB RSL? ?PC EMMB QMSPACQ MD MKCE?-3 D?RRW ?AGBQ. CQC?PAF GLBGA?RCQ
RF?R MKCE?-3 D?RRW ?AGBQ PCBSAC RFC PGQI MD QSBBCL BC?RF DPMK FC?PR ?RR?AIQ, PCBSAC RPGEJWACPGBCQ GL RFC
@JMMB, JMUCP @JMMB NPCQQSPC, ?LB NPCTCLR RFPMK@MQGQ @W GLFG@GRGLE @JMMB AJMRRGLE. FCW ?JQM PCBSAC
GLDJ?KK?RGML, ?LB K?W FCJN PCBSAC RFC PGQI MD QMKC A?LACPQ GL ?LGK?JQ.
GIC A?P@MFWBP?RCQ, D?RQ F?TC PCACGTCB ? JMR MD @?B NS@JGAGRW. *R GQ RPSC RF?R C?RGLE ?L CVACQQ MD DPGCB
DMMBQ ?LB MRFCP tD?RRWu DMMBQ JC?BQ RM UCGEFR E?GL. MUCTCP, D?RQ BM F?TC GKNMPR?LR DSLARGMLQ. ?LW
TGR?KGLQ ?PC D?R QMJS@JC, ?LB D?RQ QCPTC ?Q ? JMLE-RCPK QRMP?EC DMPK MD D?RRW ?AGBQ: ? QMSPAC MD CLCPEW.
FCW ?JQM NPMTGBC GLQSJ?RGML DMP RFC @MBW. FCPCDMPC, tFC?JRFWu D?RQ GL KMBCP?RC ?KMSLRQ QFMSJB @C
AMLQSKCB ML ? PCESJ?P @?QGQ.
OLI< 3.18 >UV @LSBOFKDP LK PLJB IB>SBP >OB J>AB LC IFMFAP. (@OBAFQ: LDBO 'OFCCFQE)
+?FJG?F@G@;J
Phospholipids ?PC K?HMP AMLQRGRSCLRQ MD RFC NJ?QK? KCK@P?LC, RFC MSRCPKMQR J?WCP MD ?LGK?J ACJJQ. GIC
D?RQ, RFCW ?PC AMKNMQCB MD D?RRW ?AGB AF?GLQ ?RR?AFCB RM ? EJWACPMJ MP QNFGLEMQGLC @?AI@MLC. *LQRC?B MD
RFPCC D?RRW ?AGBQ ?RR?AFCB ?Q GL RPGEJWACPGBCQ, FMUCTCP, RFCPC ?PC RUM D?RRW ?AGBQ DMPKGLE BG?AWJEJWACPMJ,
?LB RFC RFGPB A?P@ML MD RFC EJWACPMJ @?AI@MLC GQ MAASNGCB @W ? KMBGDGCB NFMQNF?RC EPMSN (Figure 3.19).
NFMQNF?RC EPMSN ?JMLC ?RR?AFCB RM ? BG?EJWACPMJ BMCQ LMR OS?JGDW ?Q ? NFMQNFMJGNGB; GR GQ NFMQNF?RGB?RC
86 #!% 3 X "'#! !#%#%
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(BG?AWJEJWACPMJ 3-NFMQNF?RC), RFC NPCASPQMP MD NFMQNFMJGNGBQ. FC NFMQNF?RC EPMSN GQ KMBGDGCB @W ?L
?JAMFMJ. FMQNF?RGBWJAFMJGLC ?LB NFMQNF?RGBWJQCPGLC ?PC RUM GKNMPR?LR NFMQNFMJGNGBQ RF?R ?PC DMSLB GL
NJ?QK? KCK@P?LCQ.
!@>LI< 3.19 ! MELPMELIFMFA FP > JLIB@RIB TFQE QTL C>QQV >@FAP >KA > JLAFCFBA MELPME>QB DOLRM
>QQ>@EBA QL > DIV@BOLI ?>@H?LKB. EB MELPME>QB J>V ?B JLAFCFBA ?V QEB >AAFQFLK LC @E>ODBA LO
MLI>O @EBJF@>I DOLRMP. TL @EBJF@>I DOLRMP QE>Q J>V JLAFCV QEB MELPME>QB, @ELIFKB >KA PBOFKB, >OB
PELTK EBOB. "LQE @ELIFKB >KA PBOFKB >QQ>@E QL QEB MELPME>QB DOLRM >Q QEB MLPFQFLK I>?BIBA SF> QEB
EVAOLUVI DOLRM FKAF@>QBA FK DOBBK.
NFMQNFMJGNGB GQ ?L ?KNFGN?RFGA KMJCASJC, KC?LGLE GR F?Q ? FWBPMNFM@GA ?LB ? FWBPMNFGJGA N?PR.
FC D?RRW ?AGB AF?GLQ ?PC FWBPMNFM@GA ?LB A?LLMR GLRCP?AR UGRF U?RCP, UFCPC?Q RFC NFMQNF?RC-AMLR?GLGLE
EPMSN GQ FWBPMNFGJGA ?LB GLRCP?ARQ UGRF U?RCP (Figure 3.20).
!@>LI< 3.20 EB MELPMELIFMFA ?FI>VBO FP QEB J>GLO @LJMLKBKQ LC >II @BIIRI>O JBJ?O>KBP. EB
EVAOLMEFIF@ EB>A DOLRMP LC QEB MELPMELIFMFAP C>@B QEB >NRBLRP PLIRQFLK. EB EVAOLMEL?F@ Q>FIP >OB
PBNRBPQBOBA FK QEB JFAAIB LC QEB ?FI>VBO.
FC FC?B GQ RFC FWBPMNFGJGA N?PR, ?LB RFC R?GJ AMLR?GLQ RFC FWBPMNFM@GA D?RRW ?AGBQ. *L ? KCK@P?LC,
? @GJ?WCP MD NFMQNFMJGNGBQ DMPKQ RFC K?RPGV MD RFC QRPSARSPC, RFC D?RRW ?AGB R?GJQ MD NFMQNFMJGNGBQ D?AC
GLQGBC, ?U?W DPMK U?RCP, UFCPC?Q RFC NFMQNF?RC EPMSN D?ACQ RFC MSRQGBC, ?OSCMSQ QGBC (Figure 3.20).
FMQNFMJGNGBQ ?PC PCQNMLQG@JC DMP RFC BWL?KGA L?RSPC MD RFC NJ?QK? KCK@P?LC. *D ? BPMN MD
NFMQNFMJGNGBQ GQ NJ?ACB GL U?RCP, GR QNMLR?LCMSQJW DMPKQ ? QRPSARSPC ILMUL ?Q ? KGACJJC, UFCPC RFC
FWBPMNFGJGA NFMQNF?RC FC?BQ D?AC RFC MSRQGBC ?LB RFC D?RRW ?AGBQ D?AC RFC GLRCPGMP MD RFGQ QRPSARSPC.
K
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? QFMPR R?GJ (Figure 3.21). ?LW QRCPMGBQ ?JQM F?TC RFC p DSLARGML?J EPMSN, UFGAF NSRQ RFCK GL RFC
?JAMFMJ AJ?QQGDGA?RGML (QRCPMJQ).
!@>LI< 3.21 QBOLFAP PR@E >P @ELIBPQBOLI >KA @LOQFPLI >OB @LJMLPBA LC CLRO CRPBA EVAOL@>O?LK OFKDP.
$FMJCQRCPMJ GQ RFC KMQR AMKKML QRCPMGB. $FMJCQRCPMJ GQ K?GLJW QWLRFCQGXCB GL RFC JGTCP ?LB GQ RFC
NPCASPQMP RM K?LW QRCPMGB FMPKMLCQ QSAF ?Q RCQRMQRCPMLC ?LB CQRP?BGMJ, UFGAF ?PC QCAPCRCB @W RFC EML?BQ
?LB CLBMAPGLC EJ?LBQ. *R GQ ?JQM RFC NPCASPQMP RM GR?KGL %. $FMJCQRCPMJ GQ ?JQM RFC NPCASPQMP MD @GJC Q?JRQ,
UFGAF FCJN GL RFC CKSJQGDGA?RGML MD D?RQ ?LB RFCGP QS@QCOSCLR ?@QMPNRGML @W ACJJQ. JRFMSEF AFMJCQRCPMJ GQ
MDRCL QNMICL MD GL LCE?RGTC RCPKQ @W J?W NCMNJC, GR GQ LCACQQ?PW DMP NPMNCP DSLARGMLGLE MD RFC @MBW. *R GQ ?
AMKNMLCLR MD RFC NJ?QK? KCK@P?LC MD ?LGK?J ACJJQ ?LB GQ DMSLB UGRFGL RFC NFMQNFMJGNGB @GJ?WCP. #CGLE
RFC MSRCPKMQR QRPSARSPC GL ?LGK?J ACJJQ, RFC NJ?QK? KCK@P?LC GQ PCQNMLQG@JC DMP RFC RP?LQNMPR MD K?RCPG?JQ
?LB ACJJSJ?P PCAMELGRGML ?LB GR GQ GLTMJTCB GL ACJJ-RM-ACJJ AMKKSLGA?RGML.
MP ?L ?BBGRGML?J NCPQNCARGTC ML JGNGBQ, CVNJMPC RFC GLRCP?ARGTC ?LGK?RGML “Biomolecules: The Lipids”
(http://openstaxcollege.org/l/lipids)
3.4 R +IFK
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Proteins ?PC MLC MD RFC KMQR ?@SLB?LR MPE?LGA KMJCASJCQ GL JGTGLE QWQRCKQ ?LB F?TC RFC KMQR BGTCPQC
P?LEC MD DSLARGMLQ MD ?JJ K?APMKMJCASJCQ. PMRCGLQ K?W @C QRPSARSP?J, PCESJ?RMPW, AMLRP?ARGJC, MP
NPMRCARGTC; RFCW K?W QCPTC GL RP?LQNMPR, QRMP?EC, MP KCK@P?LCQ; MP RFCW K?W @C RMVGLQ MP CLXWKCQ. &?AF
ACJJ GL ? JGTGLE QWQRCK K?W AMLR?GL RFMSQ?LBQ MD NPMRCGLQ, C?AF UGRF ? SLGOSC DSLARGML. FCGP QRPSARSPCQ,
JGIC RFCGP DSLARGMLQ, T?PW EPC?RJW. FCW ?PC ?JJ, FMUCTCP, NMJWKCPQ MD ?KGLM ?AGBQ, ?PP?LECB GL ? JGLC?P
QCOSCLAC.
/PGKPMLOQ BJLDIL?FK, >I?RJFK #>OOV PR?PQ>K@BP FK QEB ?ILLA LO IVJME
QEOLRDELRQ QEB ?LAV
QOR@QRO>I !@QFK, QR?RIFK, HBO>QFK #LKPQOR@Q AFCCBOBKQ PQOR@QROBP, IFHB QEB
@VQLPHBIBQLK
LOJLKBP KPRIFK, QEVOLUFKB #LLOAFK>QB QEB >@QFSFQV LC AFCCBOBKQ ?LAV PVPQBJP
$BCBKPB JJRKLDIL?RIFKP OLQB@Q QEB ?LAV COLJ CLOBFDK M>QELDBKP
#LKQO>@QFIB !@QFK, JVLPFK %CCB@Q JRP@IB @LKQO>@QFLK
QLO>DB BDRJB PQLO>DB MOLQBFKP,
BDD TEFQB (>I?RJFK)
OLSFAB KLROFPEJBKQ FK B>OIV ABSBILMJBKQ LC
QEB BJ?OVL >KA QEB PBBAIFKD
/< 3.1
PMRCGLQ F?TC BGDDCPCLR QF?NCQ ?LB KMJCASJ?P UCGEFRQ; QMKC NPMRCGLQ ?PC EJM@SJ?P GL QF?NC UFCPC?Q
MRFCPQ ?PC DG@PMSQ GL L?RSPC. MP CV?KNJC, FCKMEJM@GL GQ ? EJM@SJ?P NPMRCGL, @SR AMJJ?ECL, DMSLB GL MSP
QIGL, GQ ? DG@PMSQ NPMRCGL. PMRCGL QF?NC GQ APGRGA?J RM GRQ DSLARGML, ?LB RFGQ QF?NC GQ K?GLR?GLCB @W K?LW
BGDDCPCLR RWNCQ MD AFCKGA?J @MLBQ. $F?LECQ GL RCKNCP?RSPC, N, ?LB CVNMQSPC RM AFCKGA?JQ K?W JC?B RM
NCPK?LCLR AF?LECQ GL RFC QF?NC MD RFC NPMRCGL, JC?BGLE RM JMQQ MD DSLARGML, ILMUL ?Q denaturation. JJ
NPMRCGLQ ?PC K?BC SN MD BGDDCPCLR ?PP?LECKCLRQ MD RFC Q?KC 20 RWNCQ MD ?KGLM ?AGBQ.
A@EF A@;J
Amino acids ?PC RFC KMLMKCPQ RF?R K?IC SN NPMRCGLQ. &?AF ?KGLM ?AGB F?Q RFC Q?KC DSLB?KCLR?J
QRPSARSPC, UFGAF AMLQGQRQ MD ? ACLRP?J A?P@ML ?RMK, ?JQM ILMUL ?Q RFC ?JNF? ( ) A?P@ML, @MLBCB RM ?L
?KGLM EPMSN (2), ? A?P@MVWJ EPMSN ($), ?LB RM ? FWBPMECL ?RMK. &TCPW ?KGLM ?AGB ?JQM F?Q
?LMRFCP ?RMK MP EPMSN MD ?RMKQ @MLBCB RM RFC ACLRP?J ?RMK ILMUL ?Q RFC EPMSN (Figure 3.22).
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!@>LI< 3.22 !JFKL >@FAP E>SB > @BKQO>I >PVJJBQOF@ @>O?LK QL TEF@E >K >JFKL DOLRM, > @>O?LUVI
DOLRM, > EVAOLDBK >QLJ, >KA > PFAB @E>FK ( DOLRM) >OB >QQ>@EBA.
FC L?KC "?KGLM ?AGB" GQ BCPGTCB DPMK RFC D?AR RF?R RFCW AMLR?GL @MRF ?KGLM EPMSN ?LB A?P@MVWJ-
?AGB-EPMSN GL RFCGP @?QGA QRPSARSPC. Q KCLRGMLCB, RFCPC ?PC 20 ?KGLM ?AGBQ NPCQCLR GL NPMRCGLQ. CL MD
RFCQC ?PC AMLQGBCPCB CQQCLRG?J ?KGLM ?AGBQ GL FSK?LQ @CA?SQC RFC FSK?L @MBW A?LLMR NPMBSAC RFCK ?LB
RFCW ?PC M@R?GLCB DPMK RFC BGCR. MP C?AF ?KGLM ?AGB, RFC EPMSN (MP QGBC AF?GL) GQ BGDDCPCLR (Figure
3.23).
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!@>LI< 3.23 EBOB >OB 20 @LJJLK >JFKL >@FAP @LJJLKIV CLRKA FK MOLQBFKP, B>@E TFQE >
AFCCBOBKQ DOLRM (S>OF>KQ DOLRM) QE>Q ABQBOJFKBP FQP @EBJF@>I K>QROB.
EF@E @>QBDLOFBP LC >JFKL >@FA TLRIA VLR BUMB@Q QL CFKA LK QEB PROC>@B LC > PLIR?IB
MOLQBFK, >KA TEF@E TLRIA VLR BUMB@Q QL CFKA FK QEB FKQBOFLO E>Q AFPQOF?RQFLK LC >JFKL >@FAP
TLRIA VLR BUMB@Q QL CFKA FK > MOLQBFK BJ?BAABA FK > IFMFA ?FI>VBO
FC AFCKGA?J L?RSPC MD RFC QGBC AF?GL BCRCPKGLCQ RFC L?RSPC MD RFC ?KGLM ?AGB (RF?R GQ, UFCRFCP GR
GQ ?AGBGA, @?QGA, NMJ?P, MP LMLNMJ?P). MP CV?KNJC, RFC ?KGLM ?AGB EJWAGLC F?Q ? FWBPMECL ?RMK ?Q RFC
EPMSN. KGLM ?AGBQ QSAF ?Q T?JGLC, KCRFGMLGLC, ?LB ?J?LGLC ?PC LMLNMJ?P MP FWBPMNFM@GA GL L?RSPC, UFGJC
?KGLM ?AGBQ QSAF ?Q QCPGLC, RFPCMLGLC, ?LB AWQRCGLC ?PC NMJ?P ?LB F?TC FWBPMNFGJGA QGBC AF?GLQ. FC QGBC
AF?GLQ MD JWQGLC ?LB ?PEGLGLC ?PC NMQGRGTCJW AF?PECB, ?LB RFCPCDMPC RFCQC ?KGLM ?AGBQ ?PC ?JQM ILMUL ?Q
@?QGA ?KGLM ?AGBQ. PMJGLC F?Q ?L EPMSN RF?R GQ JGLICB RM RFC ?KGLM EPMSN, DMPKGLE ? PGLE-JGIC QRPSARSPC.
PMJGLC GQ ?L CVACNRGML RM RFC QR?LB?PB QRPSARSPC MD ?L ?LGKM ?AGB QGLAC GRQ ?KGLM EPMSN GQ LMR QCN?P?RC
DPMK RFC QGBC AF?GL (Figure 3.23).
KGLM ?AGBQ ?PC PCNPCQCLRCB @W ? QGLEJC SNNCP A?QC JCRRCP MP ? RFPCC-JCRRCP ?@@PCTG?RGML. MP CV?KNJC,T?JGLC GQ ILMUL @W RFC JCRRCP MP RFC RFPCC-JCRRCP QWK@MJ T?J. SQR ?Q QMKC D?RRW ?AGBQ ?PC CQQCLRG?J
RM ? BGCR, QMKC ?KGLM ?AGBQ ?PC LCACQQ?PW ?Q UCJJ. FCW ?PC ILMUL ?Q CQQCLRG?J ?KGLM ?AGBQ, ?LB GL
FSK?LQ RFCW GLAJSBC GQMJCSAGLC, JCSAGLC, ?LB AWQRCGLC. &QQCLRG?J ?KGLM ?AGBQ PCDCP RM RFMQC LCACQQ?PW DMP
AMLQRPSARGML MD NPMRCGLQ GL RFC @MBW, ?JRFMSEF LMR NPMBSACB @W RFC @MBW; UFGAF ?KGLM ?AGBQ ?PC CQQCLRG?J
T?PGCQ DPMK MPE?LGQK RM MPE?LGQK.
FC QCOSCLAC ?LB RFC LSK@CP MD ?KGLM ?AGBQ SJRGK?RCJW BCRCPKGLC RFC NPMRCGL'Q QF?NC, QGXC, ?LB
DSLARGML. &?AF ?KGLM ?AGB GQ ?RR?AFCB RM ?LMRFCP ?KGLM ?AGB @W ? AMT?JCLR @MLB, ILMUL ?Q ? peptide
bond, UFGAF GQ DMPKCB @W ? BCFWBP?RGML PC?ARGML. FC A?P@MVWJ EPMSN MD MLC ?KGLM ?AGB ?LB RFC ?KGLM
EPMSN MD RFC GLAMKGLE ?KGLM ?AGB AMK@GLC, PCJC?QGLE ? KMJCASJC MD U?RCP. FC PCQSJRGLE @MLB GQ RFC
NCNRGBC @MLB (Figure 3.24).
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!@>LI< 3.24 BMQFAB ?LKA CLOJ>QFLK FP > ABEVAO>QFLK PVKQEBPFP OB>@QFLK. EB @>O?LUVI DOLRM LC LKB
>JFKL >@FA FP IFKHBA QL QEB >JFKL DOLRM LC QEB FK@LJFKD >JFKL >@FA. K QEB MOL@BPP, > JLIB@RIB LC
T>QBO FP OBIB>PBA.
FC NPMBSARQ DMPKCB @W QSAF JGLI?ECQ ?PC A?JJCB NCNRGBCQ. Q KMPC ?KGLM ?AGBQ HMGL RM RFGQ EPMUGLE
AF?GL, RFC PCQSJRGLE AF?GL GQ ILMUL ?Q ? NMJWNCNRGBC. &?AF NMJWNCNRGBC F?Q ? DPCC ?KGLM EPMSN ?R MLC CLB.
FGQ CLB GQ A?JJCB RFC RCPKGL?J, MP RFC ?KGLM RCPKGL?J, ?LB RFC MRFCP CLB F?Q ? DPCC A?P@MVWJ EPMSN,
?JQM ILMUL ?Q RFC $ MP A?P@MVWJ RCPKGL?J. FGJC RFC RCPKQ NMJWNCNRGBC ?LB NPMRCGL ?PC QMKCRGKCQ SQCB
GLRCPAF?LEC?@JW, ? NMJWNCNRGBC GQ RCAFLGA?JJW ? NMJWKCP MD ?KGLM ?AGBQ, UFCPC?Q RFC RCPK NPMRCGL GQ SQCBDMP ? NMJWNCNRGBC MP NMJWNCNRGBCQ RF?R F?TC AMK@GLCB RMECRFCP, MDRCL F?TC @MSLB LML-NCNRGBC NPMQRFCRGA
EPMSNQ, F?TC ? BGQRGLAR QF?NC, ?LB F?TC ? SLGOSC DSLARGML. DRCP NPMRCGL QWLRFCQGQ (RP?LQJ?RGML), KMQR
NPMRCGLQ ?PC KMBGDGCB. FCQC ?PC ILMUL ?Q NMQR-RP?LQJ?RGML?J KMBGDGA?RGMLQ. FCW K?W SLBCPEM AJC?T?EC,
NFMQNFMPWJ?RGML, MP K?W PCOSGPC RFC ?BBGRGML MD MRFCP AFCKGA?J EPMSNQ. LJW ?DRCP RFCQC KMBGDGA?RGMLQ GQ
RFC NPMRCGL AMKNJCRCJW DSLARGML?J.
$JGAI RFPMSEF RFC QRCNQ MD NPMRCGL QWLRFCQGQ GL RFGQ interactive tutorial (http://openstaxcollege.org/l/
protein_synth) .
92 #!% 3 X "'#! !#%#%
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http://openstaxcollege.org/l/protein_synthhttp://openstaxcollege.org/l/protein_synthhttp://openstaxcollege.org/l/protein_synthhttp://openstaxcollege.org/l/protein_synth
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/?< MFLK@FEIP @>E@=@E< F= CPKF?IF< #VQL@EOLJB @ FP >K FJMLOQ>KQ @LJMLKBKQ LC QEB BIB@QOLK QO>KPMLOQ @E>FK, > M>OQ LC
@BIIRI>O OBPMFO>QFLK, >KA FQ FP KLOJ>IIV CLRKA FK QEB @BIIRI>O LOD>KBIIB, QEB JFQL@ELKAOFLK.
EFP MOLQBFK E>P > EBJB MOLPQEBQF@ DOLRM, >KA QEB @BKQO>I FLK LC QEB EBJB DBQP >IQBOK>QBIV
OBAR@BA >KA LUFAFWBA AROFKD BIB@QOLK QO>KPCBO. "B@>RPB QEFP BPPBKQF>I MOLQBFKwP OLIB FK
MOLAR@FKD @BIIRI>O BKBODV FP @OR@F>I, FQ E>P @E>KDBA SBOV IFQQIB LSBO JFIIFLKP LC VB>OP. OLQBFK
PBNRBK@FKD E>P PELTK QE>Q QEBOB FP > @LKPFABO>?IB >JLRKQ LC @VQL@EOLJB @ >JFKL >@FA
PBNRBK@B ELJLILDV >JLKD AFCCBOBKQ PMB@FBP; FK LQEBO TLOAP, BSLIRQFLK>OV HFKPEFM @>K ?B
>PPBPPBA ?V JB>PROFKD QEB PFJFI>OFQFBP LO AFCCBOBK@BP >JLKD S>OFLRP PMB@FBPw $! LO
MOLQBFK PBNRBK@BP.
@FBKQFPQP E>SB ABQBOJFKBA QE>Q ERJ>K @VQL@EOLJB @ @LKQ>FKP 104 >JFKL >@FAP. &LO
B>@E @VQL@EOLJB @ JLIB@RIB COLJ AFCCBOBKQ LOD>KFPJP QE>Q E>P ?BBK PBNRBK@BA QL A>QB,
37 LC QEBPB >JFKL >@FAP >MMB>O FK QEB P>JB MLPFQFLK FK >II P>JMIBP LC @VQL@EOLJB @. EFP
FKAF@>QBP QE>Q QEBOB J>V E>SB ?BBK > @LJJLK >K@BPQLO. K @LJM>OFKD QEB ERJ>K >KA
@EFJM>KWBB MOLQBFK PBNRBK@BP, KL PBNRBK@B AFCCBOBK@B T>P CLRKA. EBK ERJ>K >KA
OEBPRP JLKHBV PBNRBK@BP TBOB @LJM>OBA, QEB PFKDIB AFCCBOBK@B CLRKA T>P FK LKB >JFKL
>@FA. K >KLQEBO @LJM>OFPLK, ERJ>K QL VB>PQ PBNRBK@FKD PELTP > AFCCBOBK@B FK QEB 44QE
MLPFQFLK.
+IFK
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ACJJ ?LCKG?, RFC FCKMEJM@GL D AF?GL (? QK?JJ NMPRGML MD UFGAF GQ QFMUL GL Figure 3.26) F?Q ? QGLEJC
?KGLM ?AGB QS@QRGRSRGML, A?SQGLE ? AF?LEC GL NPMRCGL QRPSARSPC ?LB DSLARGML. NCAGDGA?JJW, RFC ?KGLM
?AGB EJSR?KGA ?AGB GQ QS@QRGRSRCB @W T?JGLC GL RFC D AF?GL. F?R GQ KMQR PCK?PI?@JC RM AMLQGBCP GQ
RF?R ? FCKMEJM@GL KMJCASJC GQ K?BC SN MD RUM ?JNF? AF?GLQ ?LB RUM @CR? AF?GLQ RF?R C?AF AMLQGQR MD
?@MSR 150 ?KGLM ?AGBQ. FC KMJCASJC, RFCPCDMPC, F?Q ?@MSR 600 ?KGLM ?AGBQ. FC QRPSARSP?J BGDDCPCLAC
@CRUCCL ? LMPK?J FCKMEJM@GL KMJCASJC ?LB ? QGAIJC ACJJ KMJCASJCqUFGAF BP?K?RGA?JJW BCAPC?QCQ JGDC
CVNCAR?LAWqGQ ? QGLEJC ?KGLM ?AGB MD RFC 600. F?R GQ CTCL KMPC PCK?PI?@JC GQ RF?R RFMQC 600 ?KGLM
?AGBQ ?PC CLAMBCB @W RFPCC LSAJCMRGBCQ C?AF, ?LB RFC KSR?RGML GQ A?SQCB @W ? QGLEJC @?QC AF?LEC (NMGLR
KSR?RGML), 1 GL 1800 @?QCQ.
!@>LI< 3.26 EB ?BQ> @E>FK LC EBJLDIL?FK FP 147 OBPFARBP FK IBKDQE, VBQ > PFKDIB >JFKL >@FA
PR?PQFQRQFLK IB>AP QL PF@HIB @BII >KBJF>. K KLOJ>I EBJLDIL?FK, QEB >JFKL >@FA >Q MLPFQFLK PBSBK FP
DIRQ>J>QB. K PF@HIB @BII EBJLDIL?FK, QEFP DIRQ>J>QB FP OBMI>@BA ?V > S>IFKB.
#CA?SQC MD RFGQ AF?LEC MD MLC ?KGLM ?AGB GL RFC AF?GL, FCKMEJM@GL KMJCASJCQ DMPK JMLE DG@CPQ RF?R
BGQRMPR RFC @GAMLA?TC, MP BGQA-QF?NCB, PCB @JMMB ACJJQ ?LB ?QQSKC ? APCQACLR MP tQGAIJCu QF?NC, UFGAF
AJMEQ ?PRCPGCQ (Figure 3.27). FGQ A?L JC?B RM KWPG?B QCPGMSQ FC?JRF NPM@JCKQ QSAF ?Q @PC?RFJCQQLCQQ,
BGXXGLCQQ, FC?B?AFCQ, ?LB ?@BMKGL?J N?GL DMP RFMQC ?DDCARCB @W RFGQ BGQC?QC.
!@>LI< 3.27 K QEFP ?ILLA PJB>O, SFPR>IFWBA >Q 535U J>DKFCF@>QFLK RPFKD ?OFDEQ CFBIA JF@OLP@LMV, PF@HIB
@BIIP >OB @OBP@BKQ PE>MBA, TEFIB KLOJ>I @BIIP >OB AFP@-PE>MBA. (@OBAFQ: JLAFCF@>QFLK LC TLOH ?V %A
QEJ>K; P@>IB-?>O A>Q> COLJ >QQ RPPBII)
!+)43*'7> !97:)9:7+
FC JMA?J DMJBGLE MD RFC NMJWNCNRGBC GL QMKC PCEGMLQ EGTCQ PGQC RM RFC secondary structure MD RFC NPMRCGL.
FC KMQR AMKKML ?PC RFC α -helix ?LB β-pleated sheet QRPSARSPCQ (Figure 3.28). #MRF QRPSARSPCQ ?PC RFC
-FCJGV QRPSARSPCqRFC FCJGV FCJB GL QF?NC @W FWBPMECL @MLBQ. FC FWBPMECL @MLBQ DMPK @CRUCCL RFC
MVWECL ?RMK GL RFC A?P@MLWJ EPMSN GL MLC ?KGLM ?AGB ?LB ?LMRFCP ?KGLM ?AGB RF?R GQ DMSP ?KGLM ?AGBQ
D?PRFCP ?JMLE RFC AF?GL.
94 #!% 3 X "'#! !#%#%
EFP @LKQBKQ FP >S>FI>?IB CLO C OBB >Q EQQM://@KU.LOD/@LKQBKQ/@LI11448/1.9
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!@>LI< 3.28 EB C -EBIFU >KA D-MIB>QBA PEBBQ >OB PB@LKA>OV PQOR@QROBP LC MOLQBFKP QE>Q CLOJ ?B@>RPB
LC EVAOLDBK ?LKAFKD ?BQTBBK @>O?LKVI >KA >JFKL DOLRMP FK QEB MBMQFAB ?>@H?LKB. #BOQ>FK >JFKL
>@FAP E>SB > MOLMBKPFQV QL CLOJ >K C -EBIFU, TEFIB LQEBOP E>SB > MOLMBKPFQV QL CLOJ > D-MIB>QBA PEBBQ.
&TCPW FCJGA?J RSPL GL ?L ?JNF? FCJGV F?Q 3.6 ?KGLM ?AGB PCQGBSCQ. FC EPMSNQ (RFC T?PG?LR EPMSNQ)
MD RFC NMJWNCNRGBC NPMRPSBC MSR DPMK RFC -FCJGV AF?GL. *L RFC D-NJC?RCB QFCCR, RFC tNJC?RQu ?PC DMPKCB @W
FWBPMECL @MLBGLE @CRUCCL ?RMKQ ML RFC @?AI@MLC MD RFC NMJWNCNRGBC AF?GL. FC EPMSNQ ?PC ?RR?AFCB
RM RFC A?P@MLQ ?LB CVRCLB ?@MTC ?LB @CJMU RFC DMJBQ MD RFC NJC?R. FC NJC?RCB QCEKCLRQ ?JGEL N?P?JJCJ MP
?LRGN?P?JJCJ RM C?AF MRFCP, ?LB FWBPMECL @MLBQ DMPK @CRUCCL RFC N?PRG?JJW NMQGRGTC LGRPMECL ?RMK GL RFC
?KGLM EPMSN ?LB RFC N?PRG?JJW LCE?RGTC MVWECL ?RMK GL RFC A?P@MLWJ EPMSN MD RFC NCNRGBC @?AI@MLC. FC
-FCJGV ?LB D-NJC?RCB QFCCR QRPSARSPCQ ?PC DMSLB GL KMQR EJM@SJ?P ?LB DG@PMSQ NPMRCGLQ ?LB RFCW NJ?W ?L
GKNMPR?LR QRPSARSP?J PMJC.
"+79/'7> !97:)9:7+
FC SLGOSC RFPCC-BGKCLQGML?J QRPSARSPC MD ? NMJWNCNRGBC GQ GRQ tertiary structure (Figure 3.29). FGQ
QRPSARSPC GQ GL N?PR BSC RM AFCKGA?J GLRCP?ARGMLQ ?R UMPI ML RFC NMJWNCNRGBC AF?GL. PGK?PGJW, RFC
GLRCP?ARGMLQ ?KMLE EPMSNQ APC?RCQ RFC AMKNJCV RFPCC-BGKCLQGML?J RCPRG?PW QRPSARSPC MD ? NPMRCGL. FC
L?RSPC MD RFC EPMSNQ DMSLB GL RFC ?KGLM ?AGBQ GLTMJTCB A?L AMSLRCP?AR RFC DMPK?RGML MD RFC FWBPMECL@MLBQ BCQAPG@CB DMP QR?LB?PB QCAMLB?PW QRPSARSPCQ. MP CV?KNJC, EPMSNQ UGRF JGIC AF?PECQ ?PC PCNCJJCB
@W C?AF MRFCP ?LB RFMQC UGRF SLJGIC AF?PECQ ?PC ?RRP?ARCB RM C?AF MRFCP (GMLGA @MLBQ). FCL NPMRCGL
DMJBGLE R?ICQ NJ?AC, RFC FWBPMNFM@GA EPMSNQ MD LMLNMJ?P ?KGLM ?AGBQ J?W GL RFC GLRCPGMP MD RFC NPMRCGL,
UFCPC?Q RFC FWBPMNFGJGA EPMSNQ J?W ML RFC MSRQGBC. FC DMPKCP RWNCQ MD GLRCP?ARGMLQ ?PC ?JQM ILMUL
?Q FWBPMNFM@GA GLRCP?ARGMLQ. *LRCP?ARGML @CRUCCL AWQRCGLC QGBC AF?GLQ DMPKQ BGQSJDGBC JGLI?ECQ GL RFC
NPCQCLAC MD MVWECL, RFC MLJW AMT?JCLR @MLB DMPKGLE BSPGLE NPMRCGL DMJBGLE.
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!@>LI< 3.29 EB QBOQF>OV PQOR@QROB LC MOLQBFKP FP ABQBOJFKBA ?V > S>OFBQV LC @EBJF@>I FKQBO>@QFLKP.
EBPB FK@IRAB EVAOLMEL?F@ FKQBO>@QFLKP, FLKF@ ?LKAFKD, EVAOLDBK ?LKAFKD >KA AFPRICFAB IFKH>DBP.
JJ MD RFCQC GLRCP?ARGMLQ, UC?I ?LB QRPMLE, BCRCPKGLC RFC DGL?J RFPCC-BGKCLQGML?J QF?NC MD RFC
NPMRCGL. FCL ? NPMRCGL JMQCQ GRQ RFPCC-BGKCLQGML?J QF?NC, GR K?W LM JMLECP @C DSLARGML?J.
Q:'9+73'7> !97:)9:7+
*L L?RSPC, QMKC NPMRCGLQ ?PC DMPKCB DPMK QCTCP?J NMJWNCNRGBCQ, ?JQM ILMUL ?Q QS@SLGRQ, ?LB RFC
GLRCP?ARGML MD RFCQC QS@SLGRQ DMPKQ RFC quaternary structure. C?I GLRCP?ARGMLQ @CRUCCL RFC QS@SLGRQ
FCJN RM QR?@GJGXC RFC MTCP?JJ QRPSARSPC. MP CV?KNJC, GLQSJGL (? EJM@SJ?P NPMRCGL) F?Q ? AMK@GL?RGML MD
FWBPMECL @MLBQ ?LB BGQSJDGBC @MLBQ RF?R A?SQC GR RM @C KMQRJW AJSKNCB GLRM ? @?JJ QF?NC. *LQSJGL QR?PRQ
MSR ?Q ? QGLEJC NMJWNCNRGBC ?LB JMQCQ QMKC GLRCPL?J QCOSCLACQ GL RFC NPCQCLAC MD NMQR-RP?LQJ?RGML?J
KMBGDGA?RGML ?DRCP RFC DMPK?RGML MD RFC BGQSJDGBC JGLI?ECQ RF?R FMJB RFC PCK?GLGLE AF?GLQ RMECRFCP. GJI (?
DG@PMSQ NPMRCGL), FMUCTCP, F?Q ? D-NJC?RCB QFCCR QRPSARSPC RF?R GQ RFC PCQSJR MD FWBPMECL @MLBGLE @CRUCCL
BGDDCPCLR AF?GLQ.
FC DMSP JCTCJQ MD NPMRCGL QRPSARSPC (NPGK?PW, QCAMLB?PW, RCPRG?PW, ?LB OS?RCPL?PW) ?PC GJJSQRP?RCB GL
Figure 3.30.
96 #!% 3 X "'#! !#%#%
EFP @LKQBKQ FP >S>FI>?IB CLO C OBB >Q EQQM://@KU.LOD/@LKQBKQ/@LI11448/1.9
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!@>LI< 3.30 EB CLRO IBSBIP LC MOLQBFK PQOR@QROB @>K ?B L?PBOSBA FK QEBPB FIIRPQO>QFLKP. (@OBAFQ:
JLAFCF@>QFLK LC TLOH ?V >QFLK>I RJ>K 'BKLJB BPB>O@E KPQFQRQB)
D
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MP ?L ?BBGRGML?J NCPQNCARGTC ML NPMRCGLQ, TGCU this animation (http://openstaxcollege.org/l/
proteins) A?JJCB t#GMKMJCASJCQ: FC PMRCGLQ.u
3.5 R L
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!@>LI< 3.31 ! KR@IBLQFAB FP J>AB RM LC QEOBB @LJMLKBKQP: > KFQOLDBKLRP ?>PB, > MBKQLPB PRD>O,
>KA LKB LO JLOB MELPME>QB DOLRMP. #>O?LK OBPFARBP FK QEB MBKQLPB >OB KRJ?BOBA 1{ QEOLRDE 5{
(QEB MOFJB AFPQFKDRFPEBP QEBPB OBPFARBP COLJ QELPB FK QEB ?>PB, TEF@E >OB KRJ?BOBA TFQELRQ RPFKD >MOFJB KLQ>QFLK). EB ?>PB FP >QQ>@EBA QL QEB 1{ MLPFQFLK LC QEB OF?LPB, >KA QEB MELPME>QB FP >QQ>@EBA
QL QEB 5{ MLPFQFLK. EBK > MLIVKR@IBLQFAB FP CLOJBA, QEB 5{ MELPME>QB LC QEB FK@LJFKD KR@IBLQFAB
>QQ>@EBP QL QEB 3{ EVAOLUVI DOLRM >Q QEB BKA LC QEB DOLTFKD @E>FK. TL QVMBP LC MBKQLPB >OB CLRKA
FK KR@IBLQFABP, ABLUVOF?LPB (CLRKA FK $!) >KA OF?LPB (CLRKA FK !). $BLUVOF?LPB FP PFJFI>O FK
PQOR@QROB QL OF?LPB, ?RQ FQ E>P >K FKPQB>A LC >K >Q QEB 2{ MLPFQFLK. ">PBP @>K ?B AFSFABA FKQL QTL
@>QBDLOFBP: MROFKBP >KA MVOFJFAFKBP. ROFKBP E>SB > ALR?IB OFKD PQOR@QROB, >KA MVOFJFAFKBP E>SB >
PFKDIB OFKD.
FC LGRPMECLMSQ @?QCQ, GKNMPR?LR AMKNMLCLRQ MD LSAJCMRGBCQ, ?PC MPE?LGA KMJCASJCQ ?LB ?PC QM
L?KCB @CA?SQC RFCW AMLR?GL A?P@ML ?LB LGRPMECL. FCW ?PC @?QCQ @CA?SQC RFCW AMLR?GL ?L ?KGLM EPMSN
RF?R F?Q RFC NMRCLRG?J MD @GLBGLE ?L CVRP? FWBPMECL, ?LB RFSQ, BCAPC?QCQ RFC FWBPMECL GML AMLACLRP?RGML GL
GRQ CLTGPMLKCLR, K?IGLE GR KMPC @?QGA. &?AF LSAJCMRGBC GL % AMLR?GLQ MLC MD DMSP NMQQG@JC LGRPMECLMSQ
@?QCQ: ?BCLGLC (), ES?LGLC () AWRMQGLC ($), ?LB RFWKGLC ().
BCLGLC ?LB ES?LGLC ?PC AJ?QQGDGCB ?Q purines. FC NPGK?PW QRPSARSPC MD ? NSPGLC GQ RUM A?P@ML-LGRPMECL PGLEQ. $WRMQGLC, RFWKGLC, ?LB SP?AGJ ?PC AJ?QQGDGCB ?Q pyrimidines UFGAF F?TC ? QGLEJC A?P@ML-
LGRPMECL PGLE ?Q RFCGP NPGK?PW QRPSARSPC (Figure 3.31). &?AF MD RFCQC @?QGA A?P@ML-LGRPMECL PGLEQ F?Q
BGDDCPCLR DSLARGML?J EPMSNQ ?RR?AFCB RM GR. *L KMJCASJ?P @GMJMEW QFMPRF?LB, RFC LGRPMECLMSQ @?QCQ ?PC
QGKNJW ILMUL @W RFCGP QWK@MJQ , , , $, ?LB . % AMLR?GLQ , , , ?LB $ UFCPC?Q AMLR?GLQ
, , , ?LB $.
FC NCLRMQC QSE?P GL % GQ BCMVWPG@MQC, ?LB GL , RFC QSE?P GQ PG@MQC ( Figure 3.31). FC
BGDDCPCLAC @CRUCCL RFC QSE?PQ GQ RFC NPCQCLAC MD RFC FWBPMVWJ EPMSN ML RFC QCAMLB A?P@ML MD RFC PG@MQC
?LB FWBPMECL ML RFC QCAMLB A?P@ML MD RFC BCMVWPG@MQC. FC A?P@ML ?RMKQ MD RFC QSE?P KMJCASJC ?PC
LSK@CPCB ?Q 1v, 2v, 3v, 4v, ?LB 5v (1v GQ PC?B ?Q tMLC NPGKCu). FC NFMQNF?RC PCQGBSC GQ ?RR?AFCB RM RFC
FWBPMVWJ EPMSN MD RFC 5v A?P@ML MD MLC QSE?P ?LB RFC FWBPMVWJ EPMSN MD RFC 3v A?P@ML MD RFC QSE?P MD
RFC LCVR LSAJCMRGBC, UFGAF DMPKQ ? 5vp3v phosphodiester JGLI?EC. FC NFMQNFMBGCQRCP JGLI?EC GQ LMR
DMPKCB @W QGKNJC BCFWBP?RGML PC?ARGML JGIC RFC MRFCP JGLI?ECQ AMLLCARGLE KMLMKCPQ GL K?APMKMJCASJCQ:
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GRQ DMPK?RGML GLTMJTCQ RFC PCKMT?J MD RUM NFMQNF?RC EPMSNQ. NMJWLSAJCMRGBC K?W F?TC RFMSQ?LBQ MD
QSAF NFMQNFMBGCQRCP JGLI?ECQ.
DA DFLQFSB $! FP >K >KQFM>O>IIBI ALR?IB EBIFU. EB MELPME>QB ?>@H?LKB (FKAF@>QBA ?V QEB
@ROSV IFKBP) FP LK QEB LRQPFAB, >KA QEB ?>PBP >OB LK QEB FKPFAB. %>@E ?>PB COLJ LKB PQO>KA FKQBO>@QP
SF> EVAOLDBK ?LKAFKD TFQE > ?>PB COLJ QEB LMMLPFKD PQO>KA. (@OBAFQ: *BOLJB >IHBO/$BKKFP VQP)
LJW ACPR?GL RWNCQ MD @?QC N?GPGLE ?PC ?JJMUCB. MP CV?KNJC, ? ACPR?GL NSPGLC A?L MLJW N?GP UGRF ?
ACPR?GL NWPGKGBGLC. FGQ KC?LQ A?L N?GP UGRF , ?LB A?L N?GP UGRF $, ?Q QFMUL GL Figure 3.33. FGQ
GQ ILMUL ?Q RFC @?QC AMKNJCKCLR?PW PSJC. *L MRFCP UMPBQ, RFC % QRP?LBQ ?PC AMKNJCKCLR?PW RM C?AF
MRFCP. *D RFC QCOSCLAC MD MLC QRP?LB GQ $$, RFC AMKNJCKCLR?PW QRP?LB UMSJB F?TC RFC QCOSCLAC$$. %SPGLE % PCNJGA?RGML, C?AF QRP?LB GQ AMNGCB, PCQSJRGLE GL ? B?SEFRCP % BMS@JC FCJGV
AMLR?GLGLE MLC N?PCLR?J % QRP?LB ?LB ? LCUJW QWLRFCQGXCB QRP?LB.
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QEVJFKB, >KA DR>KFKB ?>PB M>FOP TFQE @VQLPFKB.
! JRQ>QFLK L@@ROP, >KA @VQLPFKB FP OBMI>@BA TFQE >ABKFKB. E>Q FJM>@Q AL VLR QEFKHQEFP TFII E>SB LK QEB $! PQOR@QROB
A
G@MLSAJCGA ?AGB, MP , GQ K?GLJW GLTMJTCB GL RFC NPMACQQ MD NPMRCGL QWLRFCQGQ SLBCP RFC BGPCARGML MD
%. GQ SQS?JJW QGLEJC-QRP?LBCB ?LB GQ K?BC MD PG@MLSAJCMRGBCQ RF?R ?PC JGLICB @W NFMQNFMBGCQRCP
@MLBQ. PG@MLSAJCMRGBC GL RFC AF?GL AMLR?GLQ PG@MQC (RFC NCLRMQC QSE?P), MLC MD RFC DMSP
LGRPMECLMSQ @?QCQ (, , , ?LB $), ?LB RFC NFMQNF?RC EPMSN.
FCPC ?PC DMSP K?HMP RWNCQ MD : KCQQCLECP (K), PG@MQMK?J (P), RP?LQDCP
(R), ?LB KGAPM (KG). FC DGPQR, K, A?PPGCQ RFC KCQQ?EC DPMK %, UFGAF
AMLRPMJQ ?JJ MD RFC ACJJSJ?P ?ARGTGRGCQ GL ? ACJJ. *D ? ACJJ PCOSGPCQ ? ACPR?GL NPMRCGL RM @C QWLRFCQGXCB, RFC
ECLC DMP RFGQ NPMBSAR GQ RSPLCB tMLu ?LB RFC KCQQCLECP GQ QWLRFCQGXCB GL RFC LSAJCSQ. FC @?QC
QCOSCLAC GQ AMKNJCKCLR?PW RM RFC AMBGLE QCOSCLAC MD RFC % DPMK UFGAF GR F?Q @CCL AMNGCB. MUCTCP,GL , RFC @?QC GQ ?@QCLR ?LB GQ NPCQCLR GLQRC?B. *D RFC % QRP?LB F?Q ? QCOSCLAC $$,
RFC QCOSCLAC MD RFC AMKNJCKCLR?PW GQ $$. *L RFC AWRMNJ?QK, RFC K GLRCP?ARQ UGRF
PG@MQMKCQ ?LB MRFCP ACJJSJ?P K?AFGLCPW (Figure 3.34).
!@>LI< 3.34 ! OF?LPLJB E>P QTL M>OQP: > I>ODB PR?RKFQ >KA > PJ>II PR?RKFQ. EB J! PFQP FK
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FC K GQ PC?B GL QCRQ MD RFPCC @?QCQ ILMUL ?Q AMBMLQ. &?AF AMBML AMBCQ DMP ? QGLEJC ?KGLM
?AGB. *L RFGQ U?W, RFC K GQ PC?B ?LB RFC NPMRCGL NPMBSAR GQ K?BC. Ribosomal RNA (rRNA) GQ ?
K?HMP AMLQRGRSCLR MD PG@MQMKCQ ML UFGAF RFC K @GLBQ. FC P CLQSPCQ RFC NPMNCP ?JGELKCLR
MD RFC K ?LB RFC PG@MQMKCQ; RFC P MD RFC PG@MQMKC ?JQM F?Q ?L CLXWK?RGA ?ARGTGRW (NCNRGBWJ
RP?LQDCP?QC) ?LB A?R?JWXCQ RFC DMPK?RGML MD RFC NCNRGBC @MLBQ @CRUCCL RUM ?JGELCB ?KGLM ?AGBQ. Transfer
RNA (tRNA) GQ MLC MD RFC QK?JJCQR MD RFC DMSP RWNCQ MD , SQS?JJW 70p90 LSAJCMRGBCQ JMLE. *R A?PPGCQ
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RFC AMPPCAR ?KGLM ?AGB RM RFC QGRC MD NPMRCGL QWLRFCQGQ. *R GQ RFC @?QC N?GPGLE @CRUCCL RFC R ?LB
K RF?R ?JJMUQ DMP RFC AMPPCAR ?KGLM ?AGB RM @C GLQCPRCB GL RFC NMJWNCNRGBC AF?GL. KGAPMQ ?PC RFC
QK?JJCQR KMJCASJCQ ?LB RFCGP PMJC GLTMJTCQ RFC PCESJ?RGML MD ECLC CVNPCQQGML @W GLRCPDCPGLE UGRF RFC
CVNPCQQGML MD ACPR?GL K KCQQ?ECQ. Table 3.2 QSKK?PGXCQ DC?RSPCQ MD % ?LB .
!QFLK KSLISBA FK MOLQBFK PVKQEBPFP
L@>QFLK BJ>FKP FK QEB KR@IBRP B>SBP QEB KR@IBRP
QOR@QROB $LR?IB EBIFU PR>IIV PFKDIB-PQO>KABA
RD>O $BLUVOF?LPB F?LPB
VOFJFAFKBP #VQLPFKB, QEVJFKB #VQLPFKB, RO>@FI
ROFKBP !ABKFKB, DR>KFKB !ABKFKB, DR>KFKB
/< 3.2
&TCL RFMSEF RFC GQ QGLEJC QRP?LBCB, KMQR RWNCQ QFMU CVRCLQGTC GLRP?KMJCASJ?P @?QC
N?GPGLE @CRUCCL AMKNJCKCLR?PW QCOSCLACQ, APC?RGLE ? NPCBGAR?@JC RFPCC-BGKCLQGML?J QRPSARSPC CQQCLRG?J
DMP RFCGP DSLARGML.Q WMS F?TC JC?PLCB, GLDMPK?RGML DJMU GL ?L MPE?LGQK R?ICQ NJ?AC DPMK % RM RM NPMRCGL.
% BGAR?RCQ RFC QRPSARSPC MD K GL ? NPMACQQ ILMUL ?Q transcription, ?LB BGAR?RCQ RFC
QRPSARSPC MD NPMRCGL GL ? NPMACQQ ILMUL ?Q translation. FGQ GQ ILMUL ?Q RFC $CLRP?J %MEK? MD GDC,
UFGAF FMJBQ RPSC DMP ?JJ MPE?LGQKQ; FMUCTCP, CVACNRGMLQ RM RFC PSJC MAASP GL AMLLCARGML UGRF TGP?J
GLDCARGMLQ.
M JC?PL KMPC ?@MSR %, CVNJMPC RFC Howard Hughes Medical Institute BioInteractive
animations (http://openstaxcollege.org/l/DNA) ML RFC RMNGA MD %.
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NO JGNGB K?BC MD ? JMLE-AF?GL D?RRW ?AGB RF?R GQ CQRCPGDGCB RM ? JMLE-AF?GL ?JAMFMJ; QCPTCQ ?Q ?
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C#A+/ A
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BGKCLQGML?J QRPSARSPC GQ RFC RCPRG?PW QRPSARSPC. FCL RUM MP KMPC NMJWNCNRGBCQ AMK@GLC RM DMPK RFC
AMKNJCRC NPMRCGL QRPSARSPC, RFC AMLDGESP?RGML GQ ILMUL ?Q RFC OS?RCPL?PW QRPSARSPC MD ? NPMRCGL. PMRCGL
QF?NC ?LB DSLARGML ?PC GLRPGA?RCJW JGLICB; ?LW AF?LEC GL QF?NC A?SQCB @W AF?LECQ GL RCKNCP?RSPC MP N
K?W JC?B RM NPMRCGL BCL?RSP?RGML ?LB ? JMQQ GL DSLARGML.
3.5 L. KMLMKCPQ
?. NMJWKCPQ
@. U?RCP ?LB NMJWKCPQ
A. LMLC MD RFC ?@MTC
5. %SPGLE RFC @PC?IBMUL MD NMJWKCPQ, UFGAF MD
RFC DMJJMUGLE PC?ARGMLQ R?ICQ NJ?AC
>. FWBPMJWQGQ
?. BCFWBP?RGML
@. AMLBCLQ?RGML
A. AMT?JCLR @MLB
6. L CV?KNJC MD ? KMLMQ?AAF?PGBC GQ >>>>>>>>.
>. DPSARMQC
?. EJSAMQC
@. E?J?ARMQC
A. ?JJ MD RFC ?@MTC
7. $CJJSJMQC ?LB QR?PAF ?PC CV?KNJCQ MD:
>. KMLMQ?AAF?PGBCQ
?. BGQ?AAF?PGBCQ
@. JGNGBQ
A. NMJWQ?AAF?PGBCQ
8. J?LR ACJJ U?JJQ AMLR?GL UFGAF MD RFC DMJJMUGLE
GL ?@SLB?LAC
>. QR?PAF
?. ACJJSJMQC
@. EJWAMECL
A. J?ARMQC
9. ?ARMQC GQ ? BGQ?AAF?PGBC DMPKCB @W RFC
DMPK?RGML MD ? >>>>>>>> @MLB @CRUCCL EJSAMQC
?LB >>>>>>>>.
>. EJWAMQGBGA; J?ARMQC
?. EJWAMQGBGA; E?J?ARMQC
@. FWBPMECL; QSAPMQC
A. FWBPMECL; DPSARMQC
10. ?RSP?RCB D?RQ F?TC ?JJ MD RFC DMJJMUGLE
AF?P?ARCPGQRGAQ CVACNR:
>. RFCW ?PC QMJGB ?R PMMK RCKNCP?RSPC
?. RFCW F?TC QGLEJC @MLBQ UGRFGL RFC
A?P@ML AF?GL
@. RFCW ?PC SQS?JJW M@R?GLCB DPMK ?LGK?J
QMSPACQ
A. RFCW RCLB RM BGQQMJTC GL U?RCP C?QGJW
11. FMQNFMJGNGBQ ?PC GKNMPR?LR AMKNMLCLRQ MD
>>>>>>>>.
>. RFC NJ?QK? KCK@P?LC MD ?LGK?J ACJJQ
?. RFC PGLE QRPSARSPC MD QRCPMGBQ@. RFC U?VW AMTCPGLE ML JC?TCQ
A. RFC BMS@JC @MLB GL FWBPMA?P@ML AF?GLQ
12. FC KMLMKCPQ RF?R K?IC SN NPMRCGLQ ?PC
A?JJCB >>>>>>>>.
>. LSAJCMRGBCQ
?. BGQ?AAF?PGBCQ
@. ?KGLM ?AGBQ
A. AF?NCPMLCQ
13. FC FCJGV ?LB RFC D-NJC?RCB QFCCR ?PC N?PR MD
UFGAF NPMRCGL QRPSARSPC
>. NPGK?PW
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?. QCAMLB?PW
@. RCPRG?PW
A. OS?RCPL?PW
14. LSAJCMRGBC MD % K?W AMLR?GL >>>>>>>>.
>. PG@MQC, SP?AGJ, ?LB ? NFMQNF?RC EPMSN
?. BCMVWPG@MQC, SP?AGJ, ?LB ? NFMQNF?RC
EPMSN
@. BCMVWPG@MQC, RFWKGLC, ?LB ? NFMQNF?RC
EPMSN
A. PG@MQC, RFWKGLC, ?LB ? NFMQNF?RC EPMSN
15. FC @SGJBGLE @JMAIQ MD LSAJCGA ?AGBQ ?PC
>>>>>>>>.
>. QSE?PQ
?. LGRPMECLMSQ @?QCQ
@. NCNRGBCQ
A. LSAJCMRGBCQ
C$/$CA' /#$&$" /$*
16. FW ?PC @GMJMEGA?J K?APMKMJCASJCQ
AMLQGBCPCB MPE?LGA
17. F?R PMJC BM CJCARPMLQ NJ?W GL BCFWBP?RGML
QWLRFCQGQ ?LB FWBPMJWQGQ
18. %CQAPG@C RFC QGKGJ?PGRGCQ ?LB BGDDCPCLACQ
@CRUCCL EJWAMECL ?LB QR?PAF.
19. FW GQ GR GKNMQQG@JC DMP FSK?LQ RM BGECQR
DMMB RF?R AMLR?GLQ ACJJSJMQC
20. &VNJ?GL ?R JC?QR RFPCC DSLARGMLQ RF?R JGNGBQ
QCPTC GL NJ?LRQ ?LB/MP ?LGK?JQ.
21. FW F?TC RP?LQ D?RQ @CCL @?LLCB DPMK QMKC
PCQR?SP?LRQ MU ?PC RFCW APC?RCB
22. &VNJ?GL UF?R F?NNCLQ GD CTCL MLC ?KGLM ?AGB
GQ QS@QRGRSRCB DMP ?LMRFCP GL ? NMJWNCNRGBC AF?GL.
PMTGBC ? QNCAGDGA CV?KNJC.
23. %CQAPG@C RFC BGDDCPCLACQ GL RFC DMSP NPMRCGL
QRPSARSPCQ.
24. F?R ?PC RFC QRPSARSP?J BGDDCPCLACQ @CRUCCL
?LB %
25. F?R ?PC RFC DMSP RWNCQ MD ?LB FMU BM
RFCW DSLARGML
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