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Hs ER CFR. att: MF Tb ih AL, 2010. 1 4/JKX: An Introduction to Systems Biology: Design Principles of Biological Circuits ISBN 978-7-122-07025-8 Ml. Be RSE ” ea REAR PA +B OH CIP BRHF (2009) 9B 200154 F An Introduction to Systems Biology; Design Principles of Biological Circuits, by Uri Alon ISBN 1-58488-642-0 Copyright © 2007 by CRC Press, Part of the Taylor & Francis Group LLC. All rights reserved. Authorized translation from the English language edition published by the Taylor & Francis Group LLC. This edition is authorized for sale throughout Mainland of China. AHH ICHAEL the Taylor®. Francis Group LLC #24046°% Mb th PEM RRA AT ARAL MLE HH TI A Sh BK 4 ABA, AE LAE tol TE Hil SEP EAC BYE TAS}. TN. ABB AMAT Taylor & Francis SSG RE, BHR. ACHE TH PALA A Fe HEAL A BEGG +. 01-2009-0415 BEER: OB DIA RF: KER BER. TEL ewe. Ae ERSTE: ACE Tae HSL CGC AE GF 18 FREES 100011) El Bl: Abe ak ED RA TIES AL e i: =SWHARHRAMAA 720mmX1000mm 1/16 ENGR 7 Fak 335 FF 2010 26 4 ALIGARH 1 Uk Ei WAP. 010-64518888(fFH: 010-64519685) IRS: 010-64518899 Hts http://www. cip. com. en FUNG IG ASS 5 Ny RAT ER aL a, AL AE HO Oh BETA. Bt: 79.00% OLR ae .VELA WK = RK Pnina 49 Hanan!BaF REM AREDE FE —AMA BOR RE) PERRARRE LT. RREWE (Systems Biology) E-NREKALAMRKHRLEA, EF MRT EY. ORE. BO. TAMAS MARAAS SSA KOM, B- SAS ER a ERLE A Ly TAB AT AE BL. RB 2000 4 LAC AARR BF NB—-KARE WF ARAVA 10 F, RRA WEN BADE HAL EA HE LAA SR HE EB HAT PRS BE Be BE Ps Be A UT RREDEARUM. RREMEILE RT RAMPS —. U.K BEA WE ESE BIG Hn fa FE HR, HART SE BR, LPN Sh ABH HE Sh FR Ot BR. 2001 AF, Hiroaki Kitano (JU#f#81) WARS “Foundations of Systems Biology” (AAW Sea) — i CULE PRA F 2007 FM), KERR EME ARH BRE Pe MR REA ET SY AE ST A, CRB EO HGR MDRFE LE AL ABP A PER A RE. 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CAMA ArT AT A A — BEAR BO BARS THT EE A 4 — PAB HSE 9 ARBAB ABS PF ETE IE (principle of robustness); 4: 47 MANE HT RES A REAR A JES RB A Bee a OR. REI LE ALBEE Ht LAT LADAAT A ME AUDA IER HB PAR SETAE. EAR > HTL BT HE SR A 5 TAT ES AR be ED AEE. BATHE TE EE A RE IT HE HR tt, Ellas ARB LtE (bacterial chemotaxis) WRMA AME MEAL. AAD IRIE BRAY DY AT 0 fa FL 20 ERO A a BB HS Be AE DEAR A Nb BL Be fF HF BET BME 8 Ay — 2A A ED LB PH SE DE BH 1 ty FRO REE Ce PE A TB SIR BE, A A FRE Th. RULES, EM RSS TERRA Ih eS TE, GIN (modu- larity) ASE. REWER RFE - TEHRAN, ET UR- RAN MEHR Lit RROD, RREMCMSRAR. RBM RRA TIL. OG SH SEE GAKHAEER PRATER A. BABI. w#-F HR Fall, C., Marland E., Wagner J., and Tyson J. (2005). Computational Cell Biology, Springer. Fell, D., (1996). Understanding the Control of Metabolism. Portland Press. Heinrich, R. and Schuster, S. (1996). The Regulation of Cellular Systems. Kluwer Academic Publishers. Klipp, E., Herwig, R., Kowald, A., Wierling, C., and Lehrach, H. (2005). Systems Biology in Prac- tice: Concepts, Implementation and Application. Wiley. Kriete, A. and Eils, R. (2005). Computational Systems Biology. Academic Press. Palsson, B.O. (2006). Systems Biology: Properties of Reconstructed Networks. Cambridge Univer- sity Press. Savageau,, M.A. (1976). Biochemical Systems Analysis: A Study of Function and Design in Molecular ‘Biology. Addison Wesley.SHELA oh LB PAL — TE. EE SB PIB AD TY FE BE GE HTT — RES. I, OR RK HY Wi-—— AMA (Escherichia coli) RL-TRARAA SARWAN, REA ATLAS RA 4000 MAMI ESM. KAMAL REE LH 2.1), RL MBL. coli MAA, MARI WH Saccharomyces cerevisiae HEAR, yeast) FINGAL Bh MA CASE LEE A) RAE AES wae mae mf E ealé (S. cerevisine) | ARSHRAFH EINE) ra Bia #1000 #10000, SARK ae Hy Ax 108 4x 10° 4x10 RARE nm BAMK 4.6% 10% bp 1.3X10’bp 3X 10° bp. 4500 36H 6600 388 £30000 a wn T Mat #5 1obp sob» | #4 10bp Raa #5 100bp H 1000p) #4 10t~10%b— RAK 48 1000bp 4 1000bp #4 10~10%bp AREF ACES E/ Se #4 [nmol/L 44 Ipmol/L_ #4 0. 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A FEAR YE Yk — NE WALK ZEA RG A ET A ERE OK LE AL, RATT HEE He FASE AB AE (logic approximation) 3 3 (2.48 5K P28 (48) A BR % (A 2.4) (Glass fl Kauffman, 1973 4F; Thieffry #1 Thomas, 1998 4). 72% MAK, BARMAN, BX )=0, MABE, BI FX =p. WBE K. Atk, 999 GM (logic input function) giz LBC AY Hill PR SCG TE RT TT» BA CT LF YB BH (step-func- tion) 92m, 4 X* MAA K Mt, RI — TT: SOX )=—X">K) BG AA (2.4)2 es B22 ED BRAS HT 4RRARK n roof MAE A BE AM BA 7 AK . RADE EF BI TTS + Ca ak OY Bo BC HO: F(X" = po X* K,)6Y* >Ky)~X* AND Y* (2.6) MA-LEA, PEASE MAREE, ARMAS —T BM (OR)” KA: SOX" 5Y*)=B(X'>K, OR Y*>K,)~X* ORY? 2.7 HEA ipa BY aE DL AB A 3 PA A A HH A GD, A PE mB (SUM) AU A RRL, TERME A RRP A ET AAS CKalir Al Alon, 2004 AE): F(X" Y")=B.X* +BvY" (2.8) FE E. coli HY A RAF RR (lac system) 4 —* BM (B25), £Y FE A A A BR i RR (Yuh %§, 1998 4; Davidson MURSRAR NBA MA HT. WR. Hin, 2.5 PR FLARAT Cac) HARM, FER lac BF Hee a. AA ABE WRF Ht AND BE OR WER AIK (Mayo %, 2006), ix AH AY 3A AY BA BB BBO HE IE SX EE A J FP TEP HD 2.3.6 NE Se Rh EAH RTH SPEAR MER, XY RR OLMMY, SMEAR TRAE, EM Ch — TRS TEGAD ESAs mRNA E. XY RRMA X HH — SA TRAE YO — MERERRS. BARS B 1 le aah Fist ot 10 ‘cAMP/(mmoV/L) le AFB IPTG/(umol) 1 oo Ort A 10 ‘cAMP/(mmoV/L) | : a d _ ane: 1 oor Ol 1 10 ‘cAMPA(mmol/L) setae enna aaa EMS Te one MEATS RH, PARK, Ma RAMAMRAMAEERM, AFAR MT. (oO EMER” HARE. WE REMAP, ABFMR ARM Bt C5] A Setty $F, 2003) SAX @BORRATRATSRAY HearAe, All, SAX SOWRA RSA RAY AOA RAM. BAR Y WYRE RAF, EM Z RAR, WUE, RT TE FAAS. BS ABBY oA Bean Sa AG RA EA, BR AAT A HN PA. TEN ARES. CARR BS Oe ee RB Fie te. le DFE IPTG/(umoV/L)4 8 R28 BLT OERAF SRAM AIF bey DNA BSL. MITA Pe, TTI A OME AR, LA Y A KR SA RESRIN'F X* AAT ANSE. 2650S AEN I LA AN Aisi, 46 BA MAE LE RE I, EJ A AOE Ja HOE Y BaF LX MAALAM DNA EES —PSRL RAE, ORME ER WX >Y, RRA AM BS AE 2.4 FE) AG Bl DPS oy 79 °F Aly OT] FEA DBAS US A — FU SF 5 BLE BE SB HE, ALR RA CRE TA A SRT A LA BG es ERAS EL, AN EPS Ta) TG BEG) SBCA PU HP Hg eR A EFS BF SK: X+Y ete “HRA X WRAY”. BX BSR, Y MBER. WE EM ABA Wi RA Y ERATE (dynamics), RE AOR APS fa] (response time) . EMAMAMASM, XBRRA, THY MAA (A 2.2(bd]. 4S S. WOUG. XRT X*. FSRAYMROATHAA, RBA R JAY, GR mRNA, HARA Y BUR. AA ee BK AULA Y, TERIOR B CAAA REE HAY HP ORATAEN Bi YO, BIB ABER (degradation) (+H 4 HLS AFR AES LT SCE TS RSL) ALARAR (dilution) CEE KET PREHISTORY oss PRR BIW ayy, PERERA / RR GATT) = Ajeg +24 (2.9) BEY 4) Ye BEE A S/R SE, a He Fr Fe: OAMARARHS SME MMENME, ORGAN, DmRNA ORE, OBE BOK, ESSE SRM (ribosome) {4249 mRNA FEF, $9 mRNA 1 4H UH EIU BUR RE HEEY) RNA SPF HORE, EHR AUER He OG AE. EAE ch oN ETL eA MN, TREES EMR. ORPHAAK AREA FW EMME SIF COLI, Monod 4, 1952 4). Ei BIOTA T Ae SARIN AL. A BE EH SAF a 4 0 SD AR a I ME HH (Rosenfeld. 2002 Fs Rosenfeld #1 Alon, 2003 46). (27G3:RJ2. AE HMAC, RUBLE TY CHARA XD MOREE BURA RE, ie B= FOC) At. Gh RT BD 5 BOK SE Ayah Aa EO EAS LA A BEIT SE CHE 2. 2).eRe. RAMS mw 1S aY/dt=B—aY (2.10) HEROD, HAL Y UME EN Yan. MRDRM AY /de—0, MILLERS EAH. SRSA ASHI W/E Yu=Bla (2.11) SEC 5 REA A AACS A YR A / Hai, IM Yak, SUA BR ATES + GREG BY Ik AS (B=). HZ det A PE EE we 4 p=0 Bt, WAR C2. 10) HARA BE YA — 7 He a wh mB CHO 2. 6 Ca) Bir Y@=Yae# (2.12) BL, Y REMMI 7h A AEA BASEL AR 5 EL TE A AP AIR AC BIMBO]. EE AeA 2.12 AE AER LTT + i ok FE You F Re BR SE AY — 0) AG TL a YCO=Yoc/ 2 HORTA ACSOR TR UO FS EPR I a= ‘a (2.13) saa edn, BEE / ARE RI, BE He BERT VATE AE. A ARR AE B ROO AR AS KO. MELAS BY Og At Tl Be AR AAD Ca TEA. ERAS, BOR A HE BL Ee Qh T AAPA OY Yu Bla, BEAEBERE PE HK @ HEFT BE Ec SACRE AS FLO 9 A — EL 2 5 8 at, A ABAD ASB SY ERIE. SLR ILA — FI BRAT, AY 0 ASR CTE MRR Ma, BT LY FEU SR IA AB] — AL ES A MU A CO (2.10)] MAHA CH 2.6¢b)]. Y@)=Y¥sx1—-e"") (2.14) Y AY HES 0 FFAS, SP REWUCATRASARHE Yo —p/a. RRB, CI AE, BOM oc<1 BY, TTL AMR RURETEORRA Y OY — PSR te RIN Y~pe CE RIFANT TA at<1 (2.15) AN, PERMIT, ERY Oy UREA RE IEE RR i, BE Y BEIM, BRAT — oY FAR ERE, RE Y OCP. HFG, UTS Yu /2 BAO RUA LGM ALOR A YC) = V4 /2 BAB GR, AUR RRC. 14) APLAR He REN AE — Pe om Tie = log(2)/a (2.16) © Afi ~ 1a MY4= pla.0 - ° 00s 115225 3354 455 00s 113225 3354 455 Wa Wa © O) 2.6 Ca) RRA EAE RR, A I eR A a, Be Be TAIL ATATELT), JE Tye log(2)/a, AERIEE, JA ARF aR den ah IE AR AUAR SS AR ZN EE 5 FAB KF a BR HAY ek A Be RD ym OT (Co) ML HE A A RH CAE A TSA OFAN a) 5 Be ES BIE BALE MMT, He Tie log(2)/e, BH, KARRRLBEMS—-ASAMAKH HA, Y= pe CAARERD AG IC BE 2B A FD ATT) 5d EL SD / AS FEE / PRE o MAK, HARE RR 2.41 BEZARNHMRHAS—TABHER TERA AHD Codee =O) Ph, AS AA RIE ARE FE oh EY. SE Ha BERAR (stable proteins), 5 BAMA MAL eh EC aL A A WHR ERE OH, Kiba=ai. WPRAWREEA, LMS STM ARLE AR C9 TS], TAR — A ASAP AR Ae TJ Nk (B=0). SAE KE A ARAM, PART A. Ak, BT PHU AE EBT iA] (cell generation time) c/a, RA HREM T 50%, uk: Ty =log(2)/aai= 2 MPU ERE ale —- S EAR (2.17) TARR. A OTHE ARE TE] MA 30min BI JL st HRA on tL A FI BE BEE ME, BRAT a 2 I i) SE BREREN. EPH, RTE RRRA, CH RT IMATE. ue eA 3 FALRRA. BARS w 17 Ptashne, M. (1986). A Genetic Switch, Cell Press and Scientific Publications. Ptashne, M. and Gann, A. (2002). Genes and Signals. Cold Spring Harbor Laboratory Press. ER PH SRR Alon, U. (2003). Biological networks: the tinkerer as an engineer. Science, 301: 1866-1867. Levine, M. and Davidson, E.H. (2005). Gene regulatory networks for development. Proc. Natl. Acad. Sci. U.S.A. 102: 4936-4942 “Thieffry, D., Huerta, A.M., Perez-Rueda, E., and Collado-Vides, J. (1998). From specific gene reg- ulation to genomic networks: a global analysis of transcriptional regulation in Escherichia coli, Bioessays, 20: 433-440. Ecocye S482 www.ecocyc.org FE BH Monod, J., Pappenheimer, A.M., Jr., and Cohen-Bazire, G. (1952). The kinetics of the biosynthesis of beta-galactosidase in Escherichia coli as a function of growth. Biochem. Biophys. Acta, 9: 648-660. Rosenfeld, N. and Alon, U. (2003). Response delays and the structure of transcription networks. J. Mol. Biol., 329: 645-654. 2.1 FRAME. SUPARARAYARRRE RE REGR, RAH © ae RR BALA Be 6 a HERE HAAR YO HR. b. HBB HD CBA A— FAN Bt TLD? RE Cah: a DREAM AM RA HO, AREKRLW. LRA YOO) -Y.=B/a, YRATRA. KRE, dY/dt=8, —aY (P2.1) BARHBRAY—C +e", APREARHR CHC, KA YU-0)=B/ a AHMADKE YRARHRS B/a. 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(Pe. 5) HERAT EP24 WY, PAW TAT HGRA RRA Es dY/de= pB../a,,—aY (P2. 6) REL, FARCRERRKE TBS mRNA HRERU + mRNA BF KE GRAS, HHRMA S—M: B= p8./a, (2.7) 2.3 ARMA E MSM. HEH RASHRA YA AH RM RR BO) Fo Pe DN PEAR Holt), RREANKRR, CARA MB, aE. QHARMRS, THETAAE AM, YO = exp[— facirae’ ][¥cop + [ace rexp( facta”) ee”) (P2.8) 2.4 QR. SREPREANRR, XYZ, CGM XAMUEREDAH ATMA, XHMAGSS, AHA HON HR. SRREXRREREERA, H SEAYWRHFHBS, MUEGRY THURS BAH, YY RR Maw EK, MH, SA ZTMRRR, PHAR RAR/RR EH Ho, HARM MEK, RE AZAR ARTES S HeRA A, HARTA? PDREPRRW EAAAAW, WHRSEH? HRMEREE LT HEHE ER, 2.5 MWe (Far-out), HRAFXABALHAY HY. HRA, RE AAA A A te A a A eK, OK, AXAM AKON NRE PK. CHUA RM/ AB, CHES RHRE EM. RUPEE E PA (RRESY, PY.) TARAMRRARAEA-F? 2.6 TRAE RAO RKR, STM 2.4 RRM, THAT Do, HA ES EHO HM, RRR. @) REYO? (b) BREA ZAR RE AR Roe eet RSD? Co) fe op EB DR, SHEAR ZANRARE A AHAB,RRA. BARS Bw 19 EBE, Os 15 sie ©) 2.7 Ca) —Ar eH E.coli A SELIM Hy RSE RR. ERE OX V4 Z) PEN GEE RE BE OY I AR, SPH Lac] WM TetR, 4 E. coli PPAF BO Me A ME AB ARAB EAA ee ea. Mabie 4A BF DNA HR 538% MEAASMED. TeR ATLA MRO REA RA, 4YRRIA HT BBA (reporter), FE, MMEMT SG TetR WH FEM MRE Ik SE HR E. coli BRA, Lac] MHRORHRARA, PARR SHR BB. Cb) OFT AML Tal Be AP LE BE Ay — AER. ED RH. RR IPTG ‘SRL Lac] RAN, MH Lac] RR, AAAI PRL TetR AY me. 2 Tet is BURSA S/R SUR 1 8 oh Fe HE AOR, ALE A A DPS A), Se AE AST ABE 9 SH A A AR] (27°C F AG 36°C Fm A He tHE AR FAVRE AG 2 48). x MH 18 Bt Ti] JA A AAR SH AC TD Lt, GHEY Rosenfeld i Alon, 2003 4#)3.1 5le ee “ie lexauABe va Fuip0e 7 ara Be” FA @ HPLC (Erdos-Renyi)ASWh: —Heeee w 21 TEMP. BOE 7 EF RP — EL EE EE £, RNKA-—T+ HAS THE AM ER 8.1). He AF, RTE AA AGHA CE. coli) BAA HH 2042 AMRMR REE (transcription interactions) [4% (Shen-Orre 4, 2002 4F). RIMABLSENRA, RN ARRAS L-HT MRE RAR, EM TY fe Ra AG. IL, A RA TEI eh A EE, PET MAM DAS. LAM, RTA MTILARAR. O PX-AERAM SE POM RR TK, BAH TCHR % AMAA (network motif) , © WHE eh fT OY LR, BL SEW (negative autoregula- © BABA A PA SLA BA FD Bs REP AP I), EM. 3.2 BER, ROBLES BABE EE. coli Re RUE HH COA AA AA AES, BRAT A Ir Ie HAY IER SRA LR. TE SE, HER 5h AE 4% (randomized network) fA UREAT AS He. Hb A lL A 2B AL 5 BK AL PIA MEE Clan, 5 SESE SR RESCH YA 7 La ZAI ERE OL OY. BB LAL oa asi BEAR — EE EME TEE RTE BOR LTE RE BI EER FER. TE aoa AS DNA FF HY 98 2 EA LE ERAS OSA. ETT i. WT ARR ROTI. aE Lm PRM AM NER. RICE RE ATMA. OK ERLE REOLE KN, BET 1OmL We PSSA POOP, NLA AUR BL AM AHL 10 ‘AUC SR, SP PRE BT 10! UR DNA SEA. FAB SLBA SARA 10-? AP F ARTA BYE DNA SAFE. AER RUHR «AT Te eH. EE — DNA $F 7 SR AB A eG tt BUA AE a” Ate A B21 AMT ARAL KA : FRAME. RTE ARDS Y WR BF RI Le RF X AR PA AL HE 25, BAH MINA BURL. ROM ELLA REL. BLM, TT DLE ik SA HL La A PR SA A A22 @ Re DNA }r& (Shapiro, 1999 42) 4: RL. FA, AT AB ERO HC HE AB AE #8 By PAS BAP Ih BA SREB, ASR — 7 Pa A EP a BU SE EB I Be 1 S, PAC-EBRECATA WAN RA RAS Tie. PAEDIATR AEOLIELAE 3.2.1 55 MALAY Pad 28 EL 88k Se BB a WT RRR, HAS — ROLL Se ETT R.A 4/8 hh Erdos #1 Renyi 9] i A) Je fii 2 09 BA BL 4G Fl 4K (Erdos #l Renyi, 1959 46; Bollobas, 1985 4), CREAT it Se. FL AE GG 2 Ht 5 BE Oy AE A BOLL A Bet REE HEAR GOO TES DE PHT). WaT HORE HE. RAL AL eS Se A ARPA. SE AY PRAGA N PHM. ER. WICH ErdosRenyi (ER) MOUTHR, K {TV — a De BA SH LP PEG Pe ER BEAD AE XH HL Sy RUSE aR AT AAA (A HLT AY. BAAN MAR, SRA Na ea EM A WER. CATH, SRWARD APA, Abst NCN—-1) PATE ED NETAMTARAMR-RA AM. HI, —KUTETALTA-THaR. B M—H ABW (self-edge) CHIE N AUTRE A EW), AL BIRT AEN. N(N-1)-+N=N? B.D ERBH YY, ERWRMMMEEN? HRM, AST REM A Ee FAN P= E/N?, A 3.1(b) MAT —P DMB SHR. Ay FAV 3a BAA HCY EER RL 9 HB 3.3 AST: — pee Bee PEM FRA KY E. coli PRM SMALE. WA EWI K, RAHM SRERM—TAR. RIF HY E. coli M6 FFE 40 RAW CA 3.1 CD], KH AEM MAL PH SEARO RAF. BAZHKASSAPY ns IAREHEND, RARAAR ler, jive autoregtaton) (3.2). [ROL Ai, BSCR PY EL? Oy TR TE, (1G BUH ER MLAS PRA ROEM MRE. HT M-R ABW, RTE EME EN ERE, KEAN PAE. Ak, PBA: =i (3.2) NE REL Seat, EU RL RI, BTU FATE kA EIN BEEase): —Hmaea m 23 3.2 PARMA MRASIT, (a) RAX HA MMB, (b) RX WHAD WM. Ml, CRG SA ey, BOY X PA. A A fa OY. BADX SLASH POL RAE, ATA A SER. S41 RRM Ye FAL. ELSA Pe FB UAL Kae 5 BEL Ia ah F 50.76 TET REY XH WALMRKAR GPRM ERP BEA WIE): PUD=(F Phy A~ bug dP* (3.3) Fa 25 AF 9 ER PE CE BU BF BO TH 9 OB A BE TT TE BEAR AED) 3.4) HPAL ST HAMAD (LK EE 2. BUN MRK RA, KERMA rang ~ VEIN (3.5) 7ER 3.1 PRAM E.coli RMB. HR A A Hy N= 424 E=519, FR, MBARG. 4 BARGS > BAA ALE N ALE WPEBLISE Penne id (Neeit rand ~E/N~1. 24 Grana~ Ve 2~ 1. 1 3.6) SARA, CERIN RM PH 40 RAE. MMOL MERA. RES FL 9 SD PEE A PL DE EI a Neat? reat = (Neat) cand (3.7) rand AMA Z~32 CRE: RBS “~” Bis “HH”, RAGLAN KE SEM, KRAEMER SES. WER, 32 (her HERARG HRT EX. Auk, ASW MHA, KHAAR ASHTRAY. MARA RE SP Bee HH BB SAC He BRL He BES EPROM BE: AHtORADAT RAMA! CRORE sate? Fa T ABE aR, ERATE OS Se A OE Wi) MEAT HGRE CPR 3.2), WORE bRMEJE 7 HE 19 0G AZ BY IG] (response time) . TE ANBR ATE A — Bh EB wD SE YAS ED, SF TP / REE o HEN:(3.8) ST MAOH A Te]. BE RAT Sie ASE FH 3.4 SAE DS OSs Dm A UL Be HP Ta] ® 4RRAF XAMHASRRM, RSW MA ASM (A 3.2¢b)]. 4X GABA SAF LAM mRNA HAE Rt, MS SL I, HARE, X Me BERG, HADLAY mRNA AY AE RRR RRR. EAN AT HEA EAR BX BH HT AA A FO) AE / FERS. AX /di= f(X) —aX (3.9) SUP, fOO 2X BM M. 2 HA, MPS a POR, AREF HEED ER A AG AR RI CP 2. 4): B 1+(X/K)" ERMAARECR, 4X HR K MEST, FRM, A SRARAM @ AWK, MMM XMM, HRM, BD f(X)~O. KAMA, BNR K BORNE A AL, Ase LRM BERR X BUA oh F 501675 HEMT AN Hee. aT RRR ADT ROR AT ET AAS AE OR TT HB, YX>K Wt, ERB Os MA X1, Me: ERGROMHRAAAH, ¥CX/K DLA, RTL RERAR RAE 1, FRA © RMN ARR RMN HCPS. 2), ERR EE! Xe K (Blak. Blt, AE WORESDATRMRKA, MY Ale>K MH XODK. MMAR XML K AI. TH aU ae ASN A EAT. RAT PTE AA CT 130 B oR dX/de= pK" /X"—aX (P3. 2) ATRPRAS, FEAAHRUX, HRRM HHH ux, E Ral du/de=(n+ 1) X"dX/de, WARE: du/de= (n+ DBK"— Cnt Dau (P3.3) KREARARAMEOHRKATRANUR, REREAD MR: u=uy[ Ie or) (P34) RRARREXHBA, TH: X= XL eee vor (P3. 5) MAT X(T =X,/2 RH. FRB: Ty =COrt Da} ‘log 2/2? =1)] (P3. 6) Pm OLR AEA TR 14253 Bb, Tye GRR A aE BO oy eB fl CTYP*=log(2)/a] He AH HH TEA? /TYP*=0. 2, 0.06 KR 0.02, H 3.5 fre ARH RAAT RDM ATH. KERMA R REM RAD, WRAB RSHARATARHAORKMARERAR, LR RRERK, 3.2 CWRBE. AFIAR LARA, AHF XMRAKPAT SOMMER BRR CRLAGHE. HHS PHAHAARRASAAR ERK (Savageau, 1976 4; Goldbeter # Koshland, 1981 4; Heinrich 42 Schuster, 1996 ©). 5#& BAX EE A SM HERA (parameter sensitivity coefficient) XH x BY He Hh ADHBAKE DWN RA AP ABKKK, RAH SCA, BD. HSH, SCA, B)=(AA/A)/(AB/B)=(B/A)dA/4B (P3.7) BE: AEH AE (P3.2), 4 dX/de=0, AAR RE X= K(plak yer (P3. 8) ATHR ETS ARE LW RWHASHHARKMSRARER: SCX 1B) = (B/X AX y/dB=1/ (n+) (P3. 9) Bk, AEM BAR RK TBM. 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AXGHRN EEX HRN, RARARDAT AAR?4 Fal £83 EN Do] £3 TM 410518 TEAS BE, RATT HE ak Ae BR PS OR 2 A A ET HED EEN ARARR SAS BT ABER hE WAAR, RAL FH BE ER ae eee Me hf AF a BE H+ ZT FE REAP RSE — KL — Th a BK HE Ha TAA RE ABER EEN SAT 9 So seme 13 AAT A BAS TL RG ES IL, PE ROR E, RAP — eT Ha TT AREA RA es BE TE aE SA A Ha Pid A A — A ABT ABI HR ELIA PE, At Fe Ze 8 BAT AE) FPL (common type) #91447 — GT RMSE AEM AY FFL Zia, HPAP AYRE: ATS 6 SRA) FFL Bi (19 9 BO BAG 7 eT RR PT BE A SF SA > AM FFL ZAM. Ra. RAMI ee PPL TEE. 4.2 PEL CBE Bo Sp 1 HN BS 2 EMR PRN T BRAT RRB MAR, CH RAT RRR. RNRES BAA ES MR, RSC RFR (sub- graph), Hi 4.1(a) AMTE*HAFROATAT, MEA RRM AH {HER (feed-forward loop), HILA 13 HTROAAAA AMER TAR, TE A4.1(b) PEAAH. BH 199 PEMA RAF 5.5) UR 9364 TARTAR. PT RMF AP Ee EA), RATT AG BA PO EE BALAI A PO 28 Ps ASAT HALT RY BB SBE BLL PS BAC ST A SS HE