О возможной роли хаотического поведения системы генной регуляции в старении организма by KrioRus - issuu\nYCHEXOFEPOHT0/101/11/1 2001 • Bun. 8 YcnexM repowronormm.-2001.-Bun. 8.-C. 27-33\n© M.B. CO.TIOBbeB, 2001 r. YAK 612.67:575.113.001.573+577.23\n0 BO3MONCHOPI P0.11M XAOTWIECK0r0 1110BEAEHMSI CMCTEMbl rEHHOPI PErYllSILAYM B CTAPEHMM OPFAHM3MA Ht/11/1 OHK0/1011414 14M. npocp. H.H. fleTposa MinHaspaBa\nCaHKT-ilerepoypr, 197758, flecovnuri, yn. flennnrpascxas, 68; an. nc-ra: msoloviev@mail.ru\nHa ocHoae: 1) (1)3KTOB H3 6vionortui crapemis, caltaeTenbcTerowinx o TOM, 4TO momilmiaawis aKTHBHOCTH HeCK011b KHX KI1104eBbIX reHOB moweT apHBOCTH K 3Ha41/1TerlbHOMy HsmeHemao npoppnaorreabHocTvi )1013HH; 2) sacnepHmeHTOR C KOMablOTepHb1MH MO/leflAMH reHHOA perynawni, nokasbusatowamm, 4T0 TaKWM cHcremam CBOACTBeHHO meTa-\n(ppisnoAorwlecimx HameHeHmil, ynpasAeHrie\nKOTOpbIX B KOHer-IHOM cgeTe peaAnsyeTcs 9epe3 mexamis-\nMbl reHHoil peryAgumi (HX CB)IBb CO cTapetmem 06cy2Kita.ens! ynce AOCTaTOMHO ZIABHO\n[3, 10, 11]). C\nsperms!, ripeAcTaBAseTcs miTepecribim Hp0aHaAH3Hp0BaTb,\ncTa6HnbHoe noseAeHme, xapakTepHoe Ana xaoTtivecioix clacTem; 3) Bima ypasHeima (3KCHOHeHliHafibHblii pocT co\nHBAFIeTCH AH cncTema rermori peryAsann cncTemoii, KOTO-\napemeHem), ornicbtaalowyto Bo3pacTHylo miHaminKy CHabl\npoii npncytge xaoTnnecsoe nose4eHne, H los Taxoe nose-\ncmepTHocTii (no romnepT4Y), BbImmiraercs rwnOTe3a o TOM, LITO, no KpaAtiek mepe, Ha reHoperynaTopHom ypoaHe cTapeHme MCHKeT 6121Tb cywecTeeHHo xaorwlecainm npoaeccom (3,aecb xaoc noHmmaerca He Kai( oTcyrcTsae ynoPciRogeHHocTm, a KaK THn AHHaMI4K14 CaC)ICHbIX cam-am, yn-\niteHme COOTHOCHTCH C cpeHOMeHOM cTapeHrts OpraHH3Ma.\nCyLURIOCTb A wHammmecKoro XCIOCCI HH/Ke\npasnsemboc crporo AeTepmaHliponaHHbimm saKoHamii).\nTa KON asrnsw nosaonseT onticarb cTapeHme C nosawirt Ta-\nnpnee4eHo xpancoe HsAomeme\nCBOHCTB XaOTIVieCKPIX cncTem, B3SITOe H3 pa6OT seargnx\noism/ HHcbopmamvai — K3K 3a6b1BaHHe npawinbHoro 43yHawioHapoaaHms opraHmsma (Hnvi 3a6bwaHme COCTOAHOSI 3140 POBbA) noj BIIHSIHHeM ctonytcryawik (cna6bm Hecne14114)mvectsix BosAericTmal Ha ciicTemy)• npH 3TOM CpeAHAA CKOpOCTb TaK01-0 3a6bwaHms, xapaaTeptisyemas BeIM4H-\ncnegliaAncTos B 3T0r1 06AaCTH. Xa0C\nHemHoro pa3AP-MbIX nosn! nii, FIO3BOMIeT 6oAee\nHOAHO HOHATb Cy11I. HOCTb aToro cpeHomeHa.\n1. Xaoc no H. 17puzawcuriy .\nHOP", Hasmaaemoil noKasaTenem IlanyHosa, moweT 6b1Tb\nTeMbl MOHSHO pa3.ae.AnTb Ha ycTorsipmble H HeycTorsmBble.\noTomcsecrsneHa C noaasaTenem CHabl cmepTHoctii B ypaaHamm romneprio, a BenwanHa, o6parHas noKaaaTenlo 11snyHoaa (spems IlanyHosa) H xapaKTeprisytowas neptio,q, 4epe3 KOTOpbIN cacTema 3a6bisaeT ceoe HavanbHoe (3Roposoe)COCTOSIHHe, HaXOAHTCA B xopowem COOTBeTCTBMW C sospacTom f10/101301-0 cospeearms (T.e. H01-0 sospacTa, AO KOT0p01-0 AOJ1)KHO coxpaHaTbcs 3,qopoBoa COCTOREMe) ABA •-ienoiseaa, MbIUM H AP030431111b1 (opramismoe, 4b1/1 napaMeTpbl cTapeHma 143BeCTHbl C 6036-\nnplirieM CBOHCTBO yCTCIT-IYIBOCTH CHFI3aHO C OTBeTOM CHCTeMbI Ha sosmytEeHnn pasArsmoro THIIa.\nCHCTeMax cAa6bie sosmytgemis (cpAyKTyagmt) OKa3bIBatOT maAoe sos,aeficTime\n06bI9H0 OnHCbIBaeTCH KaK TpaexTopms B npocTpaHcTse, KOOp.OHHaTbI xoToporo COOTBeTCTBrOT nepememisim, Xa-\npawreplisyl)wm COCTOHHHe CHCTeMbI).\nB cAriae me Heyc-\nTaInnBbIX CHCTeM cAa6ble Bosmytgemis Kn 10 4 e a bi e cnoaa: crapeHae, reHHas perynawas, xaoc.\nKpaiilmm cAynaem HeyCTOAHHBbIX CHCTeM ABASTIOTCSI Xa0TH'IeCKHe CHCTeMbI, AASI\nKOTOpbIX OrIFICaHFIFI B Tepmimax\nCorAacHo BarAsmam, passmsaemstm 6eAbrniicsmm ma-\nOTAeAbHbIX TpaeKTopHM CTaHOBPITC51He/FOCTaT0t1HbIMH, n0-\nTeMaTHKOM, AaypeaTom Ho6eAescicori npemnn, H. npnro-\nCK0AbKy TpaeKTopHH, nepsoHa9aAsHo KaK yrozmo 6An3-\n2KP1Hb1M, HanpasAermocTs spemem4 o6ycAoBAeHa TeM, ,ITO\nme, co BpemeHem McnollerignaAbHo pacxopsacs (T.e. Ta-\n60AbLIIHM IIHHaMPFIeCKHM cHcTemam (KaK KAaCCIP-IeCKHM,\nme CHCTeMbI HrKHO OFIHCbIBaTb CTaTHCTIV-IeCKH, B Tepmn-\nHax aucam6Aeil TpaeKTOpH1). Pa36erarme TpaexTopA\nECAFI HCX04HTb H3 3TOH TO'IKH sperms, TO BOOAHe A0F119-\nornicbmaeTcs cpym(unefi exp(Xt), r.ae X — no onpe4eAe-\nHO IIpeAFIOA02KHTb, H HarIpaBAeHHOCTb 6HoAorw-lecxo-\nHMO, OOAO?KHTeAbHaH BeAHT-1141-la 4A31 XaOTIVIeCKFIX cucTem.\nTaK H KBaHTOBbIM) npncytEe xaoTPmecKoe noseAeHne\nro spemerm, Aematgas B ocHose cpeHomella cTaperms, Talc-\nmomeT 6bITb 06ycAosAeHa TeM, 'ITO 3aKOHbI Xa0Ca ynpaBASIOT noBeAennem 6H0A0r14,1eCKIIX CHCTeM.\nHa3bIBaeTCFI noKasaTeAem hirlyHOBa, a 06-\npaniasi BeArsimia t — spemeHem AnnyHoBa. Bpems Asmy-\nHOSa nossoAneT ssecm sHyTpeumiii «MaCILITa6 BpemeHH»\nTe cTaperms opraindsma noBeiteHne 6nortormecknx cncTem\nAAA XaOTH'IeCKHX cncTem, T.e. PiliTepBaA spemem, B Tette-\nHHe KOT0p0F0 stapameHne «ABe OAHHaKOBble» (00AHH H Te\nB FIOCAeA0BaTeAbHOCTH 6HOXHMINeCKYIX H\ncoomeTcTsylowHe ozunim H TM ace HacoxpaHRer CMbICA (AonycxaeT B onpeAeAeHHoii mepe npeacxasaHHe). ,11eAcTsnTeAbHo, nocAe ,11,0CTaTO9H0 nportoAxinTeAbHoro no cpasHeHHto co apemeHem ARnyHosa nepHop,a 9B0A1-0QH14, HaMATb 0 HaHaAbHOM COCTOAH1411 CHCTeMbI 110AHOCTb10 yTpaRHHaeTcR: 3HaHHe HagaAbH0r0 COCTOAHHA He nossoAReT 6oAee onpezteARTb TpaeKTOpH10. 2. Xaoc no F. Lllycmepy . Axxammecxxii xaOC — 9T0 HenepHosakriecxoe ,L113412KeHHe B iteTepmHHHpoBaHliblX cHcTemax (T.e. B TaKHX, rite 6y,argee 0/1H03HaT4H0 Hmelowee KOHeT4Hblr4 rOpH3014T onpe,aemeTcR nporHosa (onpeikeAsiemmii BpeMeHeM ARnyHosa). ripmRem, KaK npaBHAo, IIpOCTpaHCTBO COCTOAH14174 XaOTHTieCKI4X cHcTem Heo,n,Hopop,Ho. B Hem moryT cyLgecTsosaTb o6Aac TH, r.ae cpAylcryammg mwkeT cxaqkom nepesecni CHCTM B Apyrro 06AaCTb npocTpaHcTsa COCTORHHil. BmecTe C TeM, eCTb 06AaCTH HOBbILLICHHOli yCTOligHBOCTH K cpAyxTyagaRm. 3. Xaoc no 43fc. Hukaaucy . 41,A51 xapaxTepHcTHK14 nose,aeHHA XaOTH‘leCK0ii CHCTeMbI peataiolgee sHaReHme HmeeT ,lyBCTBHTeAbHOCTb K MaAbIM CPAyKTyaUHAM, T.e. K HexoToport Hcxo4Hoii Heonpe,aeAeHHocTH B Ha4aAbHbIX ycA0BHFIX (B 3a4aHHH HAH HsmepeHHH COCTOAHHA). 4A51 TaKHX CHCTeM HagaAbHaff Heonpe4eAeHHocTb 9KCII0HeHUHaAbH0 BospacTaeT co spemeHem AO Tex nop, noxa cAeztylowee COCTOAHHe CHCTeMbl He CTaHOBHTC51 Henpe,acxasyeMbIM. FIOTepA HHCPOpMaUHH 0 HagaAbHOM COCTOAHHH CHCTeMbI nponcxo,n,HT sa xoHenHoe spew' (HpemR ARnyHosa), T.e. no HcTeReHHH aToro HpemeHH CHCTeMa CTaHOEHTC51 npaKTHIleCKH Henpe,ackasyemoii (mo?xHo cxasaTb, Wc0 owH6xa npe,acKasaHas HexoToporo napameTpa cTaHospiTcR comsmepHma co sHameHHem camoro napameTpa). napameTpom, 011HCb1Bal0LEHM cpe,aincao AOKaAbHp0 yCT0174'4HBOCTb, HAH Cpe4111010 CKOpOCTb, C KOTOpOH yrpaRHHaeTcR HHcpopMaWili 0 HagaAbHOM COCTOAHHH, ABAAeTCA 1-10Ka3aTeAb AAnynosa. .me»)\nHeKoTopme COOKTLII 113\nAHaAH3 ,aaHHbIX, OTHOCHIMHXCA K 6HoAorHH cTapeHH.R, crnmeTeAbcTsyeT B 410Ab3y Toro, ‘iTo He6oAbinoe tHICAO reHOB moaceT ABAATbCA '-laCaMH cTapeHHR — T.e. MCVI,HCPYIKaU1451 oilHoro HAM HeCK0AbKHX KAIOTieBbIX reHos momeT npH-\nCpaBH1413aTb C geA0BeKOM, TO 9CIXPeKT TaKI4X e11,14HWIHbIX\nnpesocxo,aHT acpcpeKT OT 110AHOH 9AHMHHaU1414 cepaeRHo-cocyn,HcTbix sa6oAeHaHHil, paxa H caxapHoro Ha6eTa nolKHAblX . KpoMe npRmbix 9KCIlepHMeHTOB no HsmeneHHio aKTHBHOCTH OT,a,eAbHbIX reHoH, liMelOTCA H secombie KOCBeHHbIe AaHHbie B 110Ab3y Toro, 'ITO 11p04,0ATKHTeAbHOCTb MyTaQHil\n214143H14 MC/?MT 3HatIliTeAbH0 143MeHATbCA BCAe,EICTBHe 143-\nmeHeHm aKTI4BHOCTH He60Ab1110r0 KOAH'ieCTBa\nHam6oAee sipicHm npHmepom mo2xeT CAy2.14HTb orpaHHKaA0p1411HOCTI4 rIliTaHl4A (CM., 143BeCTHO, 'ITO\nHanpHMep, [6, 29]). 30-40% BeAeT K\nnpo4OA?KHTeAbHOCTH 214113H11 MbniteM H KpbIC\nHe TOAbKO yMeHbIllaeTCA\nH3MeHAeTCA rIp0,(10A2KHTeAbHOCTb 214143HH H\nplICK B03HHKHOBeHHH 3,360.AeBaHHII, HO H BOSHHKaeT LSeAbIH\npR.a. scpcpexToH, o6ecneRHHarokgrix same.a.AeHHe CTa moAexywHom ypoaHe: Hsmemercs, npeccHR reHOB B sLape, 06opoT 6eAx0s B gHTOrlAa3Me, AHrieCTBO nonepenHba CBBBCH 6eAxos BHe KAeTOK H xapaxTep Me1KKAeT0911b1X B3aHMOOTHOUleHHil. KpaiiHe maAoHepOATHO, 9T06b1 ace 9TH 3C14eKTb1 BO3HHKaAH napaAAeAbHo H He3aBHCHMO, a 3HaYHT, OHM yrIpaBASHOTCH 04HHM reHom Ha KAeT0t1HOM H\nrexamx. Apyrxm npxmepom MO2KeT CAy2KHTb TOT cpaKT,\npomomtimagme BH,LI,b1 moryT 3130AIOUHOH14p0BaTb B /1,0.1r02m4ByLEHe npH cmeHe 3K0A0149eCKHX ycAosHil 3a OTHOCHTeAbHO KOpOTKHA rIepH041, BpeMeHH. HanpHMep, B H3OAHposaHHoli nonyAR4HH OHOCCyMOB rIp0,40A2KHTeAbHOCTb ?KH3HH 3HaT-114TeAbH0 yHeAHRHAacb nprimepHo 3a 4000 noKoAeHHil . npeAcTaHAReTcsi xpailHe MaA0Bep0ATHb1M, YTO npH yHeAkrieHHH 11p04,0A2KHTeAbHOCTH 214113HH B xoile 9130A101111H npoHcxo,asiT CHI4XpOHHbIe MyTaIAHH BO MHOrHX reHax. Be.a,b opraHHsm ymHpaeT OT pa3HbIX IlpHMHH — ecAI! He cpa6aTbisaeT o,aHa, HanpHMep, pax, TO B03HHKaeT ApyraR, HanpHMep, ceppoRHo-cocy,amTbie sa6oAeRaHHR. TaKHM o6pasoM, ycTpaHeHme 0AHOH 143 9THX IlpH9HH He npHse,aeT K 3HatIHTeAbH0My yseAprieHino 11p0.a0A2f414TeAbHOCTI4 214143HH. 3Ha.'4HT, AAA TaK0r0 yBeA144eH1451 ,a0A11111b1 6b1Tb\nA14 MOTKeT 11p0HCX011,HTb H\n. HanpHMep, e,a11H11914bIe myTaqHH BeAyT K 3HaYHTeAbHomy (Ha 30- 200%) yHeAHRemno ElpOZIOATEETeAbHOCTH 211.113HH y xpyrAblx RepHeil (reHbi daf 2 14 age-1, xo,amproIgHe pegenTop HHcyAHHa H mo.aymproigHe cTpeccopHbie peaxgHH [1, 20]), AposociniA (rex mth, mem6paHHbni pegenTopHblii 6eAoK, yRacTsylotgkul B peryASIUHH cTpeccopxba peaKgml ) H mbnneil (reHbi dw H\nnpoHsoiiiteT op,Ha TaKa5I myragHR, TO, cxopee scero, oHa He AacT 3HaT414TeAbH0r0 scpcpexTa H 6yaeT yCTpaHeHa H3 11011y.A51U11H. HaH6oAee Bep051THO, YTO MyragHH, B pesyAbTaTe KOTOpbIX 11p0HCX0AHT ybeAHReHne r1p021,0A2KHTeAbHOCTI4 TKH3HH, nporicxoART B HeCK0AbKHX KAIOYeBbIX reHax, KOTOpbie yripasAinoT ApyrHmH reHaMH. HHTepecHo, tITO B HeKO" TOpb1X cAyRasix yHeAHReHrie 11p0,40A2KHTeAbHOCTH 2KH3HH AocmraeTcH sa cReT myTaullii B reHax, xoTophie cynpeccHpyioT 6HoxiimHHecime npogeccu, cnoco6cTspoigHe\ndf, KowrpoAxpytotgxe paamTme 91114143a [18, 24], rex p66shc ,yYaCTBylou H13 B pery.n.sigm anouroBa ). ECAH\nBeCTH K 3Ha'114TeAbHOMy H3MeHeHH10 r1p0,a0A24414TeAbHOCTH 214113HH\nBA14.5110IgHX Ha 11p14414Hb1 CMepTHOCTH. ECAH\n11p0140A21414TeAbHOCT14 1KH3HH, T.e.\n6brrb OCKpbITO>) B HaLIMX\nYCilEX1/1 rEPOHT01101 - 1/11/1 • 2001 • Bbin. 8\nnpeAr1oAo2KeHtie, 9TO eAHHH9HaA peryAHropHast myragxx moxceT Bb1313aTb 09eHb 60AbLIIHe 143MeHeHHA B xapaKTepe aKcnpeccHH rexox. 143-3a Happuemix cKoopAHHHposaktHoro noseAeHHA cHcrembi peryisigHH reHoma.\nmomer CAyN4I4Tb Bum AHU.lb OAHH reH) H aKTI4BHp0BaHO B peayAbrare eAHH119HbIX myrauHii. AAA AaAbHetiwero aHaAH3a BaN4140 OTMeTHTb, 9TO !NIA 3aBHCHMOCTer4, OTHOCALWIXCA K reHeTHKe cTapemix, HOCAT\nxapaKTep, 6A1431C14i1 K 3KCTIOHeH13Na.AbHOMy . TaK, B pa6oTe  Ha KHLUe9410r1 naAo9Ke 6bIAa npoitemoHcTpHportaHa npHmepHo 3KCHOHeHl1HaAbHaA 3aBHCHMOCTb mexcAy KOA14•1eCTBOM CAriariHO HHAMI4p0BaHHbIX myTagmil H npHcnoco6AexHoc-rbio. Y HeAosexa. H MbIUIH npoptcxoAHr 9KCLIOHeHlaHaAbHbIrl pOCT C BO3paCTOM KOAH.leCTBa xpomoCOMHbIX a6eppauml . 4aCTOTa TO9140BbIX myragHrt B I109e9HbIX KAeTKaX (rAe MaAa CKOpOCTb o6HomeHHH KAeTOK) TaKNie\nTa6nmua 1 rIptimep enyvailuoil 6ynettoii 4yulututi: cocromie rexa A onpegenexo 4epe3 Kor461•Kautmo COCTOAHHil reHoe X, Y, Z HeKOTOpbIM npotiatonbumm enoco6om reH\nCIO 3aKOHy, 6M43KOMy K 3KCII0HeHUH-\nMeTacTa6mnbmoe noeememme moitenbiumx remoperynaTopmmx cmcTem\nC. Kay(pMaHa cAeAyeT\nOTHCCTH TO, 9TO 6HHapHble 3HageHliA COCTOAHHA MHOS H\nremiori peryAstum y BbICIIIHX opraHHamou HeAOCTaT09140 xopouio H3y9eH81, 9T06bI BbICKa3b1BaTb onpeAeAemble cy2K4exput 0 MaTeMaTH9eCKHX 3aBHCHMOCTAX, xapaKrepHspoLptx HX o6wHe CBOHCTBa. 0A,HaK0 cyLgecrBytOT KomnbtorepHbte mo„AeAH, B KOTOpbIX BOCI1p0H3B0A,SIT4eTaAH\nCA 14X OCHOBHbIe HaCTHbIe CBCAICTBa, 9TO II03BOASIeT AeAaTb\nnpmigHne moryr BeCTH ce6si CHreHHoH pery.A.RUHH (T.e. o61.0-te CBOACTBa BbIBOARTCA I43 9aCTHbIX nyTem HccAeAosaHHA moAeAeii). HaH6oAee NOAH° moAe.AH perymturni B CHCTeMaX C 60AbILIHM KOAH9eCTBOM reHOB 6bIAH HCCAeA0BaHbI C. Kaycpmamom . B ero moAeAstx reHom 61)1A ripeAcratmeH KaK AorHmecicast CeTb. kaNiAbIfl reH mor HaX0AHTbCA B Axyx COCTOAHHAX: «awrHseH» (cocrxercrayeT 'AorrtecKomy 3aK.M09eHHA 0 TOM, KaK B\n3HatleHI440 «liCTHHa») 14 «HeaKTHBell» (COOTBeTCTBeHHO — «.A0N4b»). FeHbI 6bIAH CB.513aHb1 MewsAy co6oil CAriaiiHbIM o6paaoM: AAR KaN4A0r0 rexa cArtacato B816paHHoil Kom6HHagHH COCTOAHHil Apyrmx reHob TaK2Ke CArlaiiHbIM o6pa30M COHOCTaBASIAOCb COCTOAHHe «14CTHHa» HAH «AO/Kb»\n(T.e. Kamttomy rerum CONOCTaBAAAaCb CAriaiiHaA 6yAeBa npHmep B Ta6A. 1). KOAH.leCTBeHHbIe Llapamerpbt TaKHX C011OCTaBAeHHil 681.AH upeztmerom HccAe,A0BaHHA. B MaCTHOCTH, C. KaycpmaH II0Ka3aA, 9TO KOTAa414CAO cmaeli me may reHamH 6oAee 2, Tame reHoperyAxroptibte CeTH moryr rtepexoAHTb B pexcHmbi, AAA KOTOpbIX xapaKrepxo noxe,AeHme, CB0i4CTBeHHOe XaOTH.leCKHM CHCCpyHKQPIA — CM.\nHanplimep, ero aKcnepHmewrbi n0 mo,aewoBaHHIO AwnepeligHposicH (B KOTOpbIX npe,anoAaraAocb, TeMaM.\npa3HbIM yCT011914BbIM COCTOBHHAM\noAHoro reHa 6131.Ao B KageCTBeHHO\n06061gaA AaHHbre CB011X\nHeHHe» (corn, rAe com(x)=1—x). FlpHmepbt COCTOAHHA\nreHa A, castaaHHoro C reHamit X, Y, Z moryT\nA=max[X, min(Y, com(Z)]\nA=min[X, max(Y, Z)].\nC mo.aeAblo 6blA npoae.aeH cAe.apoloil mcnepHmeHT: gepes onpe.aeAeHHbie npome)KyricH apemem Ha mo.aeAb BosileiicTsosaAa cArtarmast cpAywryagm — nptiHyAmeA8-\noAHoro HAH HeCK0AbKHX reHos. npH 3TOM H3y9.3.AHCb HameHemut AHHaMHKH moiteAH. 314 aKcnepHmeHrbt HoAraepAHAH pesyAbramt, HoAy9eHH8te C. KayclimaHom. npH onpe,aeAemori CBS13140CTH HO H3MeHAA0Cb COCTOAHHe\nOTBeTCTBy}OT pasAmmme THnbi KAeTOK) LIOKa3aAH, 9TO nocAe 80% BbIl4yNtAeHHbIX H3MeHeHHil aKTHBHOCTH reHOB K.AerKa BO3BpalEaAaCb B npe2KHee COCTOAHHe, HO npH 9TOM cylgecruosaAH TaKHe H3MeHeHHA, npH KOTOpbIX Hamel-tem-Ix\ncArtaritible 6yAexbi 433,14KQHH TpyAno parrepnperHposam. AAA yAyqutexHB ruirepnperripyemocTH pe3yAbTaTOB MOzteAb C. Kaycpmaxa HaMH 6bIAa momcpHuHposaHa cAeArowpm o6paBoM. ByAeBbl LpyHKUHH Bbl6HpaAHCb He 143 xcero BO3MO1SHOr0 mHomecrsa TaKHX (pp-n(4mA, a TOAbK0 143 'Tex BO3MONCliblX BapHaHTOB: «H», «HAM», «He». ilpH 3TOM BO3M01KHa raKast Hwrepnperawm 3THX (13),HKaHil: «H» — rem A aKTHBeH, eCAH aKTFIBHbl o6a rem X H Y; «HAM» — reH A aKTHBeH, ecAH aKTHBeH OAHH H3 reHOB X HAM Y; «He» — reH A aKTHBeH, eCAH reH X HeaKTI4BeH. Haptizty C 143rIeHHeM cereil C 6HHapHb1M14 COCTOAHHAMH renou, 6bIAM HCCAeA0BaHIDI cerH, rAe COCTOBHHA reHou. MOPAl4 MeHATbCA HenpeptaaHo B AHanaaoHe OT 0 AO 1 (MONieT 6bITb HHTepnpemposarto KaK pa3AH9HaA creneHb DKCnpecc“H reHa). 3TO COOTBeTCTByeT TaK Haabmaemoii HegeTKOii (HAH Hertpepbtaxort) AorHKe. B HeH 6HHapnbte AOni9eckHe CPyHKUHI4 3aMeHAFOTCA Ha HX Henpepbtaxbie 3KBHBaAeHTbI. HamH 6bIA HCCACA0BaH Hart6oAee npoc-ro BapHaHT, B KOTOpOM «H» aamefuleTcx Ha «MHHI4MyM» (min), «HAN» — Ha «maKcHmym» (max), H «He» — Ha oitonoA-\nCBA3aHlibIX C AaHHbIM —\nH CAWKHOCTH (HCCAeA0BaAHCb CHCTeMbl OT OAHOrt AO\nreHOB) B OTBeT Ha AericrEsHe cpAyKTyagHil citcTema reHHOH peryAsigHH AemoHcrimposaAa HeyCT0i4911meracra6HAbitoe noBeAeHme, xapaKrepHoe AAA xamrptAeCATH TbI019\nApyroe COCTORHI4e. axcriepHmeHToB, OH BbIC14.33aA 29\nCHCTeM — He6oAbwHe Boammemx reHHOH aKTHBHOCTH npwao,AHAH K nepexoay B Apyroe CTa611AbH0e COCTOAHHe. npH 9TOM He 6bIAo o6HappKeHo KatleCTBeHHbIX pa36yAeBbIMH H HetieTKHMH morteAamH. AHYHH\nBI421.140 H3 BbILLIeH3A0>KeHHOCO, C 0411140r4 CT0p0HbI,\ncyulecTaamo H3MeHeHa npocTbnw B03/1eACTBHAMH (HanpHmep, 143114eHeHmem aKTHBHOCTH HeCK0AbKHX reHoB). 3T0 KOCBCHHO CBHliereAbcrayeT o TOM, tiTO B KOHTeKCTe 11HHaMHKI4 (T.e. 143meHefflia CBOHCTB BO apemem) OpraHH3M KaK QeAoe ABAAeTCA HeycTorimaoci, MeTaCTa611AbH0r4 CHCTeMOH. C Apyroil CT0p0Hb1, HCCAeA0BaHHA morteAeil CHCTeMbI reHHoii peryARUHH (KOTOpall ABABeTCA KOHetIFIbIM 3BeHOM 60AbILIHHCT11p0/1,0A/KHTeAbHOCTb /14143H14 MO?KeT 6bITb\nBa (1)H3HOAOCHtleCKHX H 6HOXI4MH'IeCKHX 143MeHeHHA, CBA-\nco cTapeHHem) noKaabmaioT, ATO oHa momeT ynpaBASITb03 3aKOHaMH xaoca. nO3TOMy npeAcTaaAaeTcx anoAHe 060CHOBaHHbIM npe,anoAoHmTb, ATO 3aKOHbI Xa0Ca HrpalOT KA109eBy10 p0Ab B CTapeHHH Opta1843Ma. BoAee IteTanbno aTa rHnoTe3a momeT 6-bITb 01114CaHa cAe4pww4m o6paaom. 1. HopmaAkHosgHorniposame opramama H, 13 Oco6eHHOCTH, ero peaKgHH Ha cTpeccopnbie B03/1eACTBHA C011150B0Maa1-OTC51 6110X1,1M1,19eCKI4M14 cpAyKTyagaxmH. VHH moryT 6bITb KpancoapemeHHbima (o6paTHmbimH): HanpHmep, AOKaAbHOe II0BbILLIeHHe TemnepaTypial HAM KOHgeHTpagHH HcAeAcTaHe TenAosoro ,aBH111eH1451 H «n06OYHbIX» XHMHtleCKHX H itoAroapemeHHblmu (Heo6paTHMbIMH HAH tlaCTHAHO 06paTI4MbIMH — penamyembimm): HartpHmep, MyTaW4H, HecnegHcimmecKHe mo,n,mimagHH cTpywrypb1 6eAKOB H AH11144,0B acAeAcTeme BO3ael4CTBKH CB0604HbIX paAHKaA0B 14 -aA - ucrpomarmiTHoro 143ArieHHA. Tame qmywryagHH MOfyT 8p14801114Tb K HecnegwpmecKomy 1,13MeHeH1-110 reHHoll aKTHBHOCTH, HanpHmep, BCAeACTNaxe noapeacileom cpparmeErra 4HK HAH OTC0e,(84HeHHA (113-3a AOKaAbHOil TeI1A0B011 (pAyKTyagHH), moAeKy.Abi 6eAKa, 6AompylotEero TpaHcKpHrugHto 4HK. B CBOIO ogepe4b, /twice KpaTKoapememoe Hama-lame reHHoil aKTHBHOCTH MO?KeT npimecTH K CTOHKHM OTKAOHeHHAM B naTTepHe aKTHBHOCTH rams — T.e. ycToilmaomy OTKAOHeHHIO KAeTKH OT HopmaAbHoro COCTOAHHA, K yCT0114HBOMy «HeHopmaAbHomy» CPyl4KUHOHI4p0BaHHIO KAeTKH. no mepe HaK011AeH1451 TaKHX «OTKAOHHBLIIHXCA» KAeTOK nocTenamo yxy4waeTcx cpywcumnwposame TKaHeci, opraHoe. H opraHH3Ma B UeA0M. npH 9TOM OTKAOHeHHA B aKTHBHOCTH Tex reHoa, KOTO3aHlIbIX\npme 110A111HbI 6bITb yCTOrriHBbl K OTKAOHeHHIO (TaK KaK\nB TO apemst\nTex reHax, tlbA HOpMe He noiwep2Kena OTKAOHeH1410, MO?KeT «ysecTH» KAeTtcy B 06AaCTb Xa0Ca. 2. Hs npe,abuytmero nyuKTa, B tlaCTHOCTH, cAe,ayeT, YTO itaxie npH a6COAIOTHO TO4H0f4 penapagm HAH npH OTCyTCTBHH 3H,a0reHHbIX H 91(301THHbIX noapemaemii MaKpomoAeHyA cTapeme B °pram-fame 6y4eT 11p0HCX0/1,HTb. 3. B HOXHMxveCKHe (pAywryawm He mom 6bITb I4CKAIOtleHbI (HanpHmep, nort ztelicTaHem ecrecTseHHoro OT6opa), TaK KaK 40A1KHa 14MeTbCA onpelteAeHHail 110/41314?KHOCTb moAeKyA 14 14X gacTert (a TaKafl 110,1/13142EHOCTb ae4eT K Bocnomm‘wocTH K ,AdicTakno (pArcryawn1). Be/Lb ecAH 01114 6y,ayT <<CHaAHHbI HamepTaoD, 3TO npimeiteT K «3aCTbIBaH1410>> CHCTeMbl, HeB03MO2KHOCTI4 ee cpyrmun0mposaHeCII0C06HOCTH ocytgecTaAemm ee peryAmpoaamm, KoTopoe Bo MHoroM 06ecnemaaeTcs CAa6bIMH 3.AeKTpoMarHHTHb1MH B3aHM0/1,ekTBIIAMH (HanpHmep, CB513131BaHHe pery.wprowx (PaKTOpOB C 6eAKOM HAH reHom) — T.e. K Hecnoco6HocTH noLa,epxcamx romeocTasa, a Taime ocyWeCTBACHHA oHToreHesa H 9BOA1OUHH. 143 aToro cAeAyeT, tITO apemeHHbie H3MeHeHHA noA 4alcTimem 80TOKa CPAyKTyaUHil HMMaHeHTHbl cHcTeme reHHol-I per-yAxgm. 4pyroe cAeltcTme — He cytgecTayeT oCTHMyA0BD (3130A104HOHH01-0 AaBAeH1451), HanpaaAeHHbot Ha TO, ATO6b1 MHHHMH3Hp0BaTb nospeautems moAeKyA HAM yBeA14.414Tb TOYHOCTb HX penapawm, BOCKOAbKy cTapeHim ace paBHO 6y.ti,eT 8p0HCX0,414Tb H3-3a Apyrmx (HanpHMep, TenAoaboc) cpAyKTyaUHA H nocAe,a,rowero HaprueHag peryAxwin (T.e. OTKAOHelitia OT «HOpMaAbH0ii» TpaeKTopm reHHoii aKTHBHOCTH). 4. B npmgme TO, tITO cTapeme ABASIeTCA XaOTHtleCKI4M npogeccom, cAeztyeT Ha ypatmemx FomnepTga. 3anxwem era B cAe,a,pormem Bpute: µ(t)=µ 0 exp(at), rzte f.t(t) — CHAa CMepTHOCTH B MOMeHT apemeHH t, o HatlaAbHa51 CHAR CMepTHOCTH, a — 110Ka3aTeAb CHAbI cmepTHocTH. Flycrb 2 opraHxama He3HatIHTeAbH0 pa3AHtlaIOTCA B HattaAbHbni MOMeHT apeman.' no po Ha BeAHAHHy 41.0 . Torita gepes spew t pasmtga B HX cHAe CMepTHOCTH L1,1 aKTHBHOCTb B\nmexcRy xaocom N crapeemem eexoropme ee cnericreae\nHOpMaAbHOil 214143HH KAeTOK.\nKaK He3HatIHTeAbHble OTKAOHeli1431 B\nr1100T030 0 CIMIN\nxorte T.e. OHH 40AACHb1 H3MeHHTb CBOH aKTHBHOCTb, a noTom BepHyTbca K HopMaAbH0My ypOBH10 aKTHBHOCTH), He ae4yr K OTKAOHeHHIO TaKHe OTKAOHeHHA B\ncAeApoweil: 4(0 = p.'(t) — µ„ (t) = (µ0, — µ0") exp(at) = exp(at). ria sToro ypatmemx\nSHAH°, tITO pa3HOCTb HatlaAbHbIX\npasHHge COCTOAHHil BoapacTom oTpaeKTopm CHAbI cmepTHocTH» paCX04ATCA no 3KCII0HeHUHaAbHOIVfy 3aK0Hy) — T.e. 3BOAIOUHA CHAbI CMepTHOCTH npoHcxwmT TaxK 3KCII0HeHUHaAbHal\n(MO/KHO CKa3aTb, YTO C\nKaK 3BOA1OUHA XaOTHtieCKHX CHCTeM.\n5. alccnoliempaAbtme pacxomytemte «Tpaewroppdt CHAbI CMepTHOCTI4>> 1103BOAAeT Bb1ABHHyTb ripeimoxoace }me, tITO 110Ka3aTeAb aticrrottexTbi (a) B ypasmemat Fom30\nYCF1EXV1rEPOHT0/101 - 141/1 • 2001 Bun. 8\n.A.snyHosa BeAH9HHe, KOTOpaA x.apaxTepH3yeT cpeAmwo CKOpOCTb 3a6bIBaHHH Ha9aAbH0r0 COCTOAHHA B xaoTHmecmx CHCTeMaX. B cAymae CTaPeHHA aTomy MO2KeT COOTBeTCTBOBaTb 9KCI1OHCHUHaAbH0e 3a6bIBaHHe HaYaAbH0I -0 (3Aoposoro, HOpMaAbH0 peamprowero) COCTORHHA. Pa3BHBaA aTy aHaAorHio AaAbwe, MO2KHO npeAnoAo2KHTb, 9TO opram43m He A0A2KeH 3a6bIBaTb csoe HaTlaAbH0e COCTOHHIle B Tevefuie nepHoAa, AOCTaT09/10r0 AAA peripoAyxgm (HHame roe.ops: 3a6b1BaTb COCTOAHHe 3A0p013b51 1103BOAHTeAbH0, Koma npoH3seAeHo IIOTOMCTB0). TOT-IHOe mawHHe 9Toro nepHoAa 3aBHCHT OT 6H0A0114H HowerHoro sHAa. ELAN (B IlepBOM npH6AHmeHHH) II0A02KPITb ero npHmepso paBHbIM so3pacTy noAosoro CO3peBaHHA H cpasHHTb 3TO 3i-1w9el-we CO spemeHem ABIlyHOBa (1 / ), TO MO2KHO o6HappsHTb, ‘ITO AAA meAosexa, MbILUH H Apo3oCPHAbI (OpraHH3MOB, ‘IbH napameTpbi cTapemis H3BeCTHbI C 60AbIl1011 TO9HOCTb10) 9TH BeAHLIHHbI npHmepHo cosnaAa10T (Ta6A. 2). aTO Ha6mAeHHe corAacyeTcR C AaHHbIMII 0 TOM, 'ITO 60AbLIIHHCTBO C4H31,10A0r1V9eCKHX napameTpos HaYHHalOT yxyAwaTbcs C MOMeHTa noAosoro CO3peBaHHA (CM., HanpHmep, ).\nnepTga (X) —\nspemeHHbim napameTpom, xapaxTepHapoLgfim 6H0A0IIP9eCKHA BHA) — T.e. B TemeHHe 9Toro nepHoAa HY2KHO YAeP/KHBaTb TpaexTopHH B nrixe (He AOITYCKaTb HbIM\nMO2KeT COOTBeTCTBOBaTb Il0Ka3aTeA10\n3HWIHTeAbH0r0 OTKAOHeHHA OT HOpMaAbH0r0 COCTOAHHA).\n0,087-0,118  0,023-0,038  0,093-0,131 \n8,5-11,5 26,3-43,5 7,6-10,8\nKaK, C TO9KH\npHH mwpopmagHH, npomcxoitmT 3a6bmaHHe\ngeHlie FlOKa3aTeAS B\nARnyHosa csHAeTeAbcTsyer 0\nKOAH‘leCTBO HHCPOpMaQPIH (B\n(6oAe3Hbl,naTrmecKH)plug. 6. 143 xaoTHmecxoft npmpoAbi reHHoil peryARgHH He nocpeAcTseHHo cAeAyeT, Tro cTapeHHe A0A2KHO 6bITb MHO-' FOCTaAHAFIbIM npogeccom (KaKOBbIM OHO, no 'Keil useposTHOCTH, H ABASIeTCA ). n0A, BAHHHHeM cpAperyagnii 11p0HCX0AHT 9BOAIOVA TpaexTopml reHHbix COCTOAHHH OT OAHOFO ycTortmmaoro COCTOAHHA (T.e. CTaAHH) K Apyromy. CKopee stem, 110406HbIii 2fie MHOrOCTaAH1111bIii npogecc AenniT H B ocHose pa3BHTHSI paKOBbIX 3a60AesaHHil. 7.\nKaK COBOKyIIHOCTb H3Me-\ncHcTeme reHHoli peryARgHH, ynpasAsembix 3axoHaMH xaoca, MO/KHO npeAAommTb 0673ACHeHHe H TaKOMy (\_.'+ cpeHomeHy, KaK ymeHbateHHe CHAbI cmepTHocTH B cTapumx so3pacTHbix rpynnax (6oAee Rn ACT). nO.A BAHAHmem r 4 i"p )r.,. 4 cpAyKTyagHil npoHcxoAHT DEIOAIOQHA CHCTeMbI TeHHOii peryAsigHH. npH 3TOM CHCTeMa KaK 6bI oHlMeTD 6oAee clu MHBble COCTOAHHH (meHee yCTOrP-IHBble OHa IIOKHAaeT 110A BAHAHHeM (pAyxTyagHii). C soapacTom BepOATHOCTb HaX01KAeHHSI CHCTeMbI B 6oAee yCTOiNHBOM COCTOAHHH yseAH914BaeTCH. BbIBeCTH cHcTemy H3 ycToilmHsoro COCTOAHHH Tpy4Hee. noaTomy AAA Toro, 91•06bI meAosex 3a60AeA, Hy2KHO 6oAee CHAbH0e someiicTsHe, mem paHbuie. COOTBeTCTBeHHO, KTO He HaX0AHT TaK0I-0 yCTOAIIHBOI*0 COCTOAHHA K AaHHOMy so3pacTy — yMHpaeT. C Apyroil CT0p0HbI, nepHoA pocTa momeT conposomA9TbC5i 110BbILLIeHHOff noTpe6HocTb10 B reHHoii peryugHH, *ITO BeAeT K yse.AHmeHlis) cTeneHeil cmo60,abi reHoperyAHTopHog CHCTeMbI. 3THM momeT 6bITb 06'bACHeHO nommenHas! CHAR CMepTHOCTH B M.AaAIIIHX BO3paCTHbIX rpynnaX. 8. TaxAce, C TO9K143pems xaommectwii npHpoAbi reHHOH MO2KHO nonbrraTbcs TIOHATb, nomemy CTapalm HmeeT mHoro o6wHx mepT y pa3HbIX BHAOB mHoroKAeTOHFIbIX ?KHBOTHbIX. &AMMO, BO mmorom 9T0 MO2KeT 6bITb CBH3aHO C 061MHOCTb10 B OpraHH3aUHH renoMa: CPO CAO2KHOCTH, KOMPleCTBa reHOB, crenemi C13513HOCTH, xapaxTepa C13513HOCTH. nO9TOMy npH B03,AerICTBHH cpAyicryaReHm=1 B\n12-15 30-50***  9 \n* A wamepReTcs B 1 40A, ocranbsue Ber1W41411b1 — B ropax. ** HamepRercsi 8 ocranbHble aenw-RiHbi — B cyncax. *** 30 — OT momeHra po(geHmq, 50 — OT momeHra onnoaorsopemnsi.\n06o6Lgam npeAnoAomeHme o\npef3y.AbTaTe cpAyxTyagHil) — T.e. BospacTeT mHCAO cTeneHeil CB060A,b1 CHCTeMbI . B KOHTeKCTe reHHoii peryAsgHH 3T0 0311agaeT, 9TO yBeAHYHBatTCA 9HC.A0 BO3MO2KFIbIX peaxgHii Ha OAHO H TO Am B03AefICTBHe — T.e. K ■KIIpal3flAbHb1M)) (HOpMaAbHbIM, 3Aoposbim, H3Ha‘laAbH0 3anporpamMHp0BaHHb1M) peaKUHRM 71,06aBASHOTCH «HenpasHAbHbie» e\nBo3pacr nonoaoro co3peBaHt4R\nM bi wb**\n(3A0p0B0r0) COCTOAHHHopraHH3ma. 1 I0A0HiliTeAbH0e 3Ha-\nA0110AHeHHA K AaHHOMy\nnpeA,AoncHTb o&bscHeHHe Tomy,\nCoomercreme mexpy BpemeHem 11nnyHoBa (T) H H03pacrom nonortoro co3pesaHHB BHA\nnpHATH K cAeAroigHm sbmoAam: a) C TOMKH spemis TeopHH impopmagHH, cTapefwe MONKHO omicaTb Kai( HameHeHHe peaxgHH cHcTembi Ha cnegHciammecnie CHI'HaAbI 110A, BAHAHHem Hecneguctimmecxxx cHrHaAos (cpApayagHii) — T.e. 3a6bwamie npasHAbHoro (PyHKUHOHHp0BaHHA (HAN 3a6bIBaHHe COCTOAHHA 3A0p0B1351); 6) C TemeHHem spemeHH B03M0HiHbIe TpaesTopHH reHHort peryARgsm pa36eraioTc.s (OTKAOHAIOTCH Apyr OT Apyra y pa3HbIX OpraHH3MOB HAH OT HOpMaAbH0r0 COCTOAHHA reHoperyAmTopHoil CHCTeMbI y oAHoro H Toro me OpraHH3Ma) noA AeiicTsHem cpAywryagml. CTeneHb HX pa36eraHHA peryAmpyeTcs eCTeCTBCHHbIM oT6opom; B) npH 3TOM spems AHITyHOBa ABAHeTCH nepHoAom, KorAa MO2KHO CHHTaTb, tITO TpaexTopm4 reHHoii peryAmumm HaX0ASITCH B nymice — T.e. HX pa36eraHHe He3HWIHTeAbH0. saHanasoH 3Ha9eHHA T momer onpeAemincm nepHo4om noAosoro coapeaainim (HAH KaKHM-HH6yA,b Apyrnm no.Ao63,A0p0Bbfl, MOASHO\nTaK2Ke CX0A,Hbl — BBHAy CX0ACTBa 6HOXH-\nMHH) ,zatHammxa reHHar peryABUHH B o6wem 9B0.110UHOHH-\nKOMIIIDIOTepH0r0 M0411,eAHIJOBaHHSI (B TOM HHCAe HCII0Ab3y-\npyeT 4110CTaT09H0 e4HH006pa3Ho. XOTH rippi 3TOM B HaCT-\nlowero ocHombre 110A0211eHI-151 TeopHH xaoca) H moAexy-\nHbIX cArraax KOHKpeTHbIe Aerami 1-13MeHeHHAreHHori pery-\nAapHo-reHeTpNeckoro aHaA113a, nomomeT Bb1511:314Tb (npezt-\nASIgHH (T.e. KaKHe HMeHHO reubi MeHAFOT caoe COCTOAHHe y\nCKaBaTb) HOBbIe TOHK11 (reHbI), B034,efiCTBH51 Ha aoTopbre\n4a.HHOr0 BH,aa) ripm cTaperwrx moryT pa3A1riaTbC51 BecbMa\ncmwEeT 60Aee 3C10CpeKTI3BHO H 110AHO BO3BpaIgaTb TpaekTo-\npwo reHHog peryAaww B sztopoaoe COCTOH1114e.\nCTOHT 3aMeTHTb, 4TO cyrgecTayroT Teoppm (cm. ),\no6bacHaragme aaamogwo H orrorelles mepea Teopmo xao-\nca - T.e. H 3130AIOUHOHHOe npomcxors,a,efare craperima MO-\n1. Bakaea B.B., TmxoHoaa 0.B., KOXVIH E.M. Hccnenoaaame B03,11e6CTBMS1 pa3/11,14HbIX rtarropoe Ha 11p0gOIDKVITellbHOCT6 *143Hti HemaTo,abi Caenorhabditis elegans // Ycnexv, repoHTon.-2000.-Bun. 4. - C. 153 - 159.\naceT 6bITb o6si3aHo xaocy. KpoMe Toro, xaoc HposisA.HeTcH He TOAbKO B remoii perynagmi, HO H B ApyriAx 6HOXI4MIrieCKHX peaKuHax, a TaK1Ke H Ha KAeTOHHOM ypoBHe, ripmgem\n2. raspmnoa Faapvinoaa H.C. Smonorma nponorpkviTenaHocni xmaym.-M.: HayKa, 1986.-168 c.\n3TH npogeccbi Taicaie moryT HrpaTb Ba2KHyto poAb B npo-\n3. rponakimckm BOOTDHKO B.fl., KmaxameA0B KoribToaep B.K. Ha,qexHocm M cTapemme 61,10/101- W-OCKI4X CHOTem.-Kmes: Haykoaa nymka, 1987.-176 c.\ngecce cTapeHHa H pa3WITHH aCCOUHI1p0BaHHbIX C HHM TOAOrHH [27, 31]. 143 Bcero 3Tor0 MO2KHO 3aKAIOHHTb, HTO\ncTapeHHe moaleT 11BAHTbCH cAe.acT8Hem XaOTHHeCKOil rippr-\n4. Avinbmam B.M. LleTbipe moilenvi menvimmu.-11.: MeLlyakwHa, 1987.-288 C.\npo,abr 1K1480171 maTeow B UeAoro.\n5. M0p03013 BT., XaBOHCOH B.X. flenTyguible 6,4operynsroobi (25-neTHHH 0116IT aKcnepvimeHranbHoro M KTEIHAVOCKOM MCcne,a0BaHfr1A).-C116.: HayKa, 1996.-74 c.\n6. FIVIKI4THH B.H. akcnepmmemTanembie nonxonbi K npo,arieHMO XGA3H14 //. &Armory's, c -rapeHms, / OTB. pen. B. B. Opona-\nAAA noitTHepmaeHasi rHnoTe3bi 0 XaOTH4eCK011 npHpo,ae craperwa Heo6xwwmo Bb1511311Tb OCHOBHbIe xapakTe-\nkmc.-11.: HayKa, 1982.-C. 548-568.\npliCTHKH renoperyAaTopHbix CHCTeM YeA0BeKa H 211HBOT-\n7. H144(01114C 4,4Hamywa HepapxmecKmx cmcrem: 38onioumomioe npeAcTaanerime.-M.: Mmp, 1989.-488 c.\nHbIX. aTO 1/101KHO 6y,/1eT C,I1eAaTb nyTeM co ,reTaHma KOM-\n8. 06beKT61 emonoryw paaeynAs, / OTB. pen. T. A. „fleTnacp.-M.: HayKa, 1975.-579 c.\nnbxyrepHoro moz(eAmposarwa C coapememmimH meTwiamm Bbl.HBAeHHA ypOSHA aKcnpeccHH reHoa\n9. flpinro*mm M., Crearepc H. Bpema, xaoc, KBaHT.-M.: flporpecc, 1994.-272 c.\nBoapacmpo Z1HHaMHKy rem-loci aKTIIBHOCTH) H aHaAH3a HyKAeOTH.4HbIX\n10. Oporibkpic B. B. PerynopoaaHme, npmcnoco6nemme crapeame.-11.: HayKa, 1970.-432 c.\nrlaCTKOB, OTBeTCTBeHHbIX 3a peryAsigHlo aKTHBHOCTH AaH-\n11. cIpponbkinc B. B., MypaHHH X. K. CTapaHlle, 3130/110L1H51 npo,rmeyme AcHaym.-KineH: HayKoea AyMKa, 1992.-336 c.\nHOrO reHa (3Toro morceT OKa3aTbC51 ./10CTaTO4HO, HT06b1\n12. Xaammcom B.X., Mopoaoe B.F. FlenTmamap perynpuval romeocTaaa nppi cTapeFaiiii // Ycnexm repowron.-2000.Bun. 4.-C. 75-79.\nBOCCTaHOBHTb 110AHyFO xapTHHy B3aHMOCBABH reHOB). H\neCAH moiteAb peaAbHOH CHCTeMbl reHHOLI peryArrupw 6yzteT ,aeMOHCTpHp0BaTb CBOVICTBa, xaparcreprible 4A21 xaoTwiec-\n13. WycTep F. AeTepmviHvipoeammuci xaoc.-M.: 1988.-240 c.\nKHX cHcTem, MONSHO 6yAeT flpaKTHHeCKH HasepHsno yT-\n14. Anisimov V.N. Age as a risk factor in multistage carcinogenesis // Comprehensive Geriatric Oncology / Eds.: Balducci L., Lyman G.H., Ershler W.B.-Amsterdam: Harwood Academic Publishers, 1998.-P. 157-178.\nHepmaaTb, 'TO crapeEme - BTO cyr.gecTseimo xaommecKIfil npouecc. EcAH 3Ta rHnoTesa OKa1KeTCH sepia, TO cAe,apoto-rm\n15. Anisimov V.N., Khavinson V.Kh., Mikhalski A.I., Yashin A.I. Effect of synthetic thymic and pineal peptides on biomarkers of ageing, survival and spontaneous tumour incidence in female CBA mice // Mech. Ageing Dev.-2001.-Vol. 122.-P. 41-61.\nIllarOM ,710J1KeH CTaTb HOHCK MeTOHOB B0311e1ICTEH51 Ha re-\nHoperyAsiTopfibie CHCTeMbI C UeAblo 3aMe4AeHRH, a B nepcneKTHBe H npeztoTapargeHaa crapelma. Ope./1,CTaBASIEOTC51\n16. Anisimov V.N., Mylnikov S.V., Khavinson V.Kh. Pineal peptide preparation Epithalamin increase the life span of fruit flies, mice and rats // Ibid.-1998.-Vol. 103.-P. 123-132.\n6HoperyAmppoweil TepanHH [5, 12, 16]. [Jo Hceil HepoHT-\n17. Austad S.N. Retarded senescense in aninsular population of Virginia opossums (Didelphis virginiana) // J. Zoology.-1993.-Vol. 229.-P. 695-708.\nHOCTH, aCIXIXKTHBHOCTb nenTap,Hoii 6HoperyAsigHH 110 3aMe4ACHHIO erapanw MO/KHO 06bACHHTb cAeztyboigmm o6-\n18. Brown-Borg H.M., Borg K.E., Meliska C.J., Bartke A. Dwarf mice and the ageing process // Nature.-1996.-Vol. 384.-p. 33.\naosapargarb naTrepH remwil aKTHBHOCTI4 B 06AaCTb, COOT-\n19. Elena S. F., Lenski R. E. Test of synergistic interactions among deleterious mutations in bacteria // Nature.-1997.-Vol. 390.-P. 395-398.\nBeTcTBylotgyio HopmaAbitomy ipp-murformposaiiwo opraHHBMa (HHarie rosopa, He6oAbwoe peryArropHoe B034erlCTBHe Ha reHbl abrHymileHHo <OlpHB0,11,14T>> OTKAOHeHHy10\n20. Johnson T.E. Increased life-span of age-1 mutants in Caenorhabditis elegans and lower Gompertz rate of aging // Science.-1990.-Vol. 249.-P. 908-912.\nTpaexTopmo B «szoposoe» COCTOAHHe). Ha ceroxisuumil ileHb B 6HoperyAHpyrorgeii repanHH AomHHkrpyer ammipm-\n21. Kauffman S.A. The Origins of Order: Self-Organization and Selection in Evolution.-Oxford etc.: Oxford University Press, 1993.-709 p.\nnecxHii no4xo4. itonoAHeHHe ero meTozwmH aHaA133a mercreHHbix aaammortericTaHil, 6asupyrorgHxca\nYCIIEXI'I rEPOHT01101- 101 • 2001 • Bun. 8\n27. Rossor M.N. Catastrophe, chaos and Alzheimer's disease. The F. E. Williams lecture // Journal of the Royal College of Physicians (London).-1995.-Vol. 29.-P. 412-418. 28. Rudolph K.L., Chang S., Lee H.W. et al. Longevity, stress response, and cancer in aging telomerase-deficient mice // Cell.-1999.-Vol. 96.-P. 701-712. 29. Sohal R. S., Weindruch R. Oxidative stress, caloric restriction, and aging // Science.-1996.-Vol. 273.-P. 59-63. 30. Vijg J. Somatic mutations and aging: a re-evaluation // Mutat. Res.-2000.-Vol. 447.-P. 117-135. 31. Waliszewski P., Molski N1., Konarski J. On the holistic approach in cellular and cancer biology: nonlinearity, complexity, and quasi-determinism of the dynamic cellular network // J. Surg. Oncol.-1998.-Vol. 68.-P. 70-78.\n22. Lin Y.J., Seroude L., Benzer S. Extended life-span and stress resistance in the Drosophila mutants methuselah // Science.-1998.-Vol. 282.-P. 943-946. 23. Martin G.M. Genetic and environmental modulations of chromosomal stability: their roles in aging and oncogenesis // Ann. N. Y. Acad. Sci.-1991.-Vol. 621.-P. 401-417. 24. Mattison J.M. Ames dwarf mice: A model for delayed aging // Ycnexw repoi-ron.-2000.-Bbin. 4.-C. 141-146. 25. Migliacchio E., Giorgio M., Mele S. et al. The p66shc adaptor protein controls oxidative stress response and life span in mammals // Nature.-1999.-Vol. 402.-P. 309-313. 26. Miller R.A. Kleemeier Award Lecture: Are There Genes for Aging? // J. Gerontol.: Biological Sciences.-1999.-Vol. 54A, Ns 7.-P. B297-B307.\nAdv. Gerontol.-2001.-Vol. 8.-P. 27-33\nM. V. Soloviev ON POSSIBLE ROLE OF CHAOTIC BEHAVIOR OF THE GENE REGULATION SYSTEM IN AGING N.N. Petrov Research Institute of Oncology, 197758, St. Petersburg, Pesochny-2, Leningradskaya, 68; e-mail: msoloviev@mail.ru 1. The facts of biology of aging say that the modification of activity of few key genes may lead to the significant increase of life span. 2. The experiments with computational models of gene regulation show that these models are characterized by metastable behavior typical for the chaotic systems. 3. The force of mortality in Gompertz equation grows exponentially in time. It allows to hypothesize that at least at the level of gene regulation aging can be a chaotic process (here implied that chaos is a kind of the complex systems dynamics controlled by strictly determined rules). This approach allows to describe aging from the view point of the information theory — as the forgetting of the right functioning of the Organism (i.e. forgetting the state of health) caused by fluctuations (weak nonspecific influencies on the system). Average rate of such forgetting (described by parameter, named Lyapunov exponent) could be identified with the exponent of mortality rate in Gompertz equation. And the parameter inverse to Lyapunov's exponent (Lyapunov time), that describes the time of the initial (healthy) state forgetting, well corresponds to the time of the puberty beginning (i.e. the minimal time for the healthy state to be kept) for human beings, mice, and fruit flies (organisms whose parameters of aging are most perfectly known). Key words: aging, gene regulation, chaos.\nО возможной роли хаотического поведения системы генной регуляции в старении организма\nСтатья по геронтологии М. Соловьева


adidas continental 80 vs nike cortez Nike Cortez G Swarovski (W) - CI2283-111 Nike Cortez Sneakers Skate shoe Adidas Nike Cortez Sneakers Shoe Adidas, nike Nike Cortez - Sneakers Nike Cortez Suede | Nike cortez nike cortez vs adidas superstar - OFF79 Nike's Cortez Sneakers and its Failed nike cortez ultra moire womens white adidas classic cortez Shop Clothing Nike Classic Cortez Sneaker Nike Cortez Shoes. Nike.com Gifts For Sneakerheads Featuring Nike Nike Cortez Shoes. Nike.com Sneakers Shoe Nike Cortez Adidas, PNG Cortez - Sneaker Freaker Nike Cortez Shoe Sneakers Adidas Samba adidas nike cortez\nThe price - by The p, 2021-10-20 02:43:13\nche costa - by che c, 2021-10-13 18:43:13\nThe shoes - by The s, 2021-10-21 23:43:13\nShopping - by Shopp, 2021-10-21 16:43:13


Pioneer MVH-AV190 : цены в Санкт-Петербурге, характеристики, фото и отзывы. | Скидки и акции на Sravni.com\nPioneer MVH-AV190 цены в Санкт-Петербурге\nКупить Pioneer MVH-AV190 в Санкт-Петербурге\nМодели Pioneer Pioneer SPH-DA120 | Pioneer FH-X730BT | Pioneer AVH-X8800BT | Pioneer AVH-X7800BT | Pioneer MVH-29BT | Pioneer AVH-290BT | Pioneer AVH-X1800DVD | Pioneer DEH-80PRS | Pioneer MVH-X580BT | Pioneer DEH-4800FD | Pioneer DEH-X2900UI | Pioneer AVH-190 | Pioneer DEH-X8700BT | Pioneer MVH-580AV | Pioneer MVH-X460UI | Pioneer DEH-9450UB | Pioneer MVH-280FD | Pioneer DEH-S100UBB | Pioneer MVH-AV290BT | Pioneer MVH-S100UI | Pioneer MVH-190UI | Pioneer MVH-S100UBA | Pioneer DEX-P99RS | Pioneer DVH-880AVBT | Pioneer DEH-S5000BT | Pioneer DEH-X9600BT | Pioneer FH-X380UB | Pioneer AVH-X4800DVD | Pioneer DEH-S100UBG | Pioneer DEH-S100UBА | Pioneer MVH-S300BT | Pioneer MVH-S100UBG | Pioneer AVH-190G | Pioneer DEH-S3000BT | Pioneer DVH-780AV | Pioneer DEH-S100UB | Pioneer AVH-Z1000DVD\nНайти характеристики и самую низкую цену на Pioneer MVH-AV190 – сравнить цены на Автомагнитолы Pioneer MVH-AV190 в интернет-магазин Санкт-Петербурга. Заказать и купить Автомагнитолы Пионер MVH-AV190 с доставкой в г. Санкт-Петербург.


Blog Music de VoIcE-BLaCk-oFFiCiaL-00 - BLaCk VoIcE - Skyrock.com\nMixTape 7aliL Wa Na9icH Vol_1\nPeace & BlaCk VoiCe .\nLA SUITE DE LA MIXTAPE 7ALIL WA NA9ICH VOL1 EST DISPONIBLE SUR : www.7alil-wa-na9ich.skyblog.com\nCréation : 18/08/2009 à 12:23\nMise à jour : 23/01/2012 à 02:37\nREMARQUE / LA SUITE DE LA...\n7ALIL WA NA9ICH / ISTI3MAR BLACK VOICY (2011)\n7ALIL WA NA9ICH ( INTRO )\n1 7ALIL WA NA9ICH ( INTRO )VoIcE-BLaCk-oFFiCiaL-00\n2 OPSY VOICEVoIcE-BLaCk-oFFiCiaL-00\n3 B.V.P MUSICVoIcE-BLaCk-oFFiCiaL-00\n4 BLaCk VoiCe - HouWa Malo_iDLane Prood VoIcE-BLaCk-oFFiCiaL-00\n6 LMKHIPSY O LMASTERVoIcE-BLaCk-oFFiCiaL-00\n7 BLaCk VoiCe - L7a9i9a_iDLane ProodVoIcE-BLaCk-oFFiCiaL-00\n8 BA2INCHITONI FEAT SLIM BLACK VoIcE-BLaCk-oFFiCiaL-00\n9 BLaCk VoiCe - Rap FuCkiNah FeaT Mr.A VoIcE-BLaCk-oFFiCiaL-00\n10 ISTI3MAR BLACK VOICYVoIcE-BLaCk-oFFiCiaL-00\nKhwi Rasek Mixtape : WWW.KHWI-RASEK-MIXTAPE-PART1.SKYBLOG.COM\nKhwi Rasek Mixtape ( Part.1 ) Bu Master Lblack Voicy\nEnjooy La Famille ,.\nle blog de khwi-rasek-mixtape-part1\nvia : khwi-rasek-mixtape-part1.skyrock.com\nModifié le lundi 23 janvier 2012 02:37\nModifié le mardi 03 mai 2011 19:44\n# Posté le lundi 25 avril 2011 18:19\nModifié le lundi 25 avril 2011 19:56\nTitre : ISTI3MAR BLACK VOICY\nAlbum : 7ALIL WA NA9ICH\n7ALIL WA NA9ICH / BA2INCHITONI FEAT SLIM BLACK (2011)\nTitre : BA2INCHITONI FEAT SLIM BLACK\n7ALIL WA NA9ICH / LMKHIPSY O LMASTER (2011)\nTitre : LMKHIPSY O LMASTER\n# Posté le lundi 25 avril 2011 15:47\n7ALIL WA NA9ICH / L2ICHKALIYA (2011)\nTitre : L2ICHKALIYA\n7ALIL WA NA9ICH / B.V.P MUSIC (2011)\nTitre : B.V.P MUSIC\n# Posté le lundi 25 avril 2011 14:58\n7ALIL WA NA9ICH / OPSY VOICE (2011)\nTitre : OPSY VOICE\n7ALIL WA NA9ICH / 7ALIL WA NA9ICH ( INTRO ) (2011)\nTitre : 7ALIL WA NA9ICH ( INTRO )\n7aliL Wa Na9iCh / BLaCk VoiCe - Rap FuCkiNah FeaT Mr.A (2011)\nTitre : BLaCk VoiCe - Rap FuCkiNah FeaT Mr.A


Union Latinos by Norlan feat Henry Mendez/Opb on MP3, WAV, FLAC, AIFF & ALAC at Juno Download\nGet new release alerts for Norlan feat Henry Mendez/Opb\n1. Union Latinos\t 03:52 120\n2. Union Latinos (instrumental)\t 03:52 120\n3. Union Latinos (Gee remix)\t 03:58 120