YAK 578.1 BBK 28.072873 H49 Cepus ocuonana « 2006 2 Heapcon JI. H49 ——Ocnosst Guoxusmn Jlenmuvxepa : 8 37. T.3: Myr nepeaaun umopmaunn / Jl. Heascon, M. Koxe ; nep. ¢ aura. —M. : BAHOM. JlaGoparopua anannii, 2015.—448 c. > wn. —(Iyaunit sapyGexnenit yueOunk). ISBN 978-5-94774-367-8 (T. 3) ISBN 978-5-94774-364-7 B yweGwom anaiii, HaNHcaHHOM aMepHKAHCKHMit Y¥eHBIMH, KOTOPLIE 0- AYMWJIM MpHOHAMNE KaK TATAHTAMBMIE MpenonaBATENH YHHBEPCHTETCKOTO YpOBHA, PaCCMOTpeHD COBPeMeHHBIC KOMMLCTIUNH GHOXHMNH 8 COOTBETCTBHH ¢ HMeHIE luefich uaeonorneli 9Toi Hay. B tom 3 noua dacre III «flyra nepenasn niopwauii, orBerst Ha Bonpo- hl, PeleHHs 3azla H NIpeaMeTHO-HMENHOH yRAXATeN’ MO MaTepHasIy TOMOB 4 TaKKE PHARTRIE COKPAULeHMR M cTOBapb TepMHHoB, OOcyxIaIOTER OCHOF Hux JOTMa MoAeKyAEpHOH OHosIOTHH H ee COBPeMenHoe HoHNMaHHe, NpoUeccEt ‘epenadit W XpaHeHH reHeTHIeCKO!L HHIopMalLnH KAK y GaxTepHii, TaK HY 3y- kapMoT (pemLaKattna, TpakeKpHIIMHR, TPAHCARUNR, PenapauutA H PEKOMGHHALLNA), CTpOeHHe xpoMocos, MexaHitsMsl GepMeHTATHBHAIX Mpotieccos, YHKLMK pa: suwnnix PHK s xierke, pHGosuMel, cmuialicMHl, anbTepHarHBHslli crutaiicHer, npotteccunr. MoapoGuo oniican GiocnHTes Genka, ero TpaxcnopruposKa K Me CTY McnOmIORAHH H AANBHCTiMIce paxpyuIeHHe, PeryastiNA >KCMPECCHH reHOR. B xaxkaofi mane (kak 8 Tomax | H 2) MpMBcieHA NpHMepbL 13 MesULAH Montexy:IspHofi GHONOTHH H CMEKHBIX OOaCTeH, a TAKKE HNTepECHBLE sAAMHA i Bonpocta ZTlas ctyaeHTon GHOROTHNCCKHX, XHMIMICCKHX H MCAHUMHCKHX ayIOR, a TAKE haysMbix pa6oTHHKOS, YAK 578.1 BBK 28.072873 VueGnoe usdanue Cepua: lysumnii sapyGexnsiit yseGunK» Heascow Jasna, Koxe Maiixs OCHOBbI BHOXHMHH JIEHHHJDKEPA, B tpex towax To 3 TYTH NEPEAAMH HH@OPMALHE Beayuit peaaxtop kana. xam. Hay 7-H. [Touxaesa. Peaaxrop Kawa. 6oa. wayk TE, Toxcmuxuna Xynoxunk H. A. Hosax. Texunuecknit peaaxtop E, B. lemoxosa. Kosnbiorepiaa seperka: 7. 2. Buykoaa Tloanneano 8 nesart 23.10.14. Dopmar 84x 108/16. Yea. nex. a. 47,04. Waaareascrso «BHHOM. JlaGoparopna sHaniiin 125167, Mocxsa, npoesa A3ponopra, a. 3 Tenedon: (499) 157-5272, e-mail: binom@Lbz.ru, http:/www.Lbz.ru Ornevaraio ® tanorpapun «BAJITO npunr>, Turaa First published in the United States by W.H. FREEMAN AND COMPANY, New York Copyright © 2008 by W.H. Freeman and Cormpany Al tights reserved. ISBN 978-5-94774-367-8 (T. 3) © Mepesoa wa pyccnuii sex, ISBN 978-5-94774-364-7 BHHOM. Jla6oparopua ananuii, 2015YACTD III fytn nepegaun NvHqopmaunn PCTbA, 3akMONTETbHAA, YaCTh KNUTH M10- cnautena GHOXHMINNeCKHM — MexaititaMaM, obecresnpaiouliM mpowteccn! mepenayit 1a- CaeACTBeHHO! HMOpMalH H 9BOOMHH KMBbIX cymecrs Kakosa Moekyapnat ocHopa reneriecko- ro Matepnama? Kak reerueckaa uncpopsatia c BHICOKO!i TOMNOCTHIO MepemaeTCA 13. TOKOACHHA B noxoaenne? Kak nosnukaior penkie HaMeHenia B TeHeTHUCCKOM MaTepHame, KOTOpHIe CAY:KAT HCXOA- HbIM MaTepHasiom Juin sBomouMH? Kak renerHue- ckas Mn opmaltiA MpespaulaeTcA B AMUHOKHCIOT- ble nociledoRaTembHOCTH GerKOBBIX MOmeKY.1? Coppemeniioe nonumanne meTaGomnueckux nyteli meperaun mnopmaniit ccpopmuponastoce Ha CTbIKe reHeTHKH, QDUSHKIL H XHMUIH — OCHOB CO- Bpemennoii 6uoxuatm, Jbxeiime Yorcon « Dpercuc Kpuk p 1953 r. chopmyzmponain rinotesy aByx- uertovesnoii crpyktypet J[HK (em. puc, 8-15 #7. 1), Tenierieckas Teopita nomorna ccpopmuponat Kon- Lemiuno KoANpoRANHA WpopMaluIM B renax. Bol- AaioecA OTKPEITHA B UBHKe MOsBOANIM yeTaHO- BHTb MozeKy.tapHyo cTpystypy rena, Biataa xuMit B reneTHYecKyIO TeopHio cocToAN B oMpeneseHtH coctasa HK. Oco6aa uennocts rinotesst Yor- cona—Kpnka sakiiovanach B TOM, 4TO oa cMorsIa o6o6uitTs pasHoobpasiite naS.sogemmta, Momysen- HbIe B pasAHYHBIX O6s1ACTAX Hay 24 Texbi 4 xpomocomb! 7 25 Meta6onu3m JIHK 43 26 Meta6onu3m PHK 105 27 Meta6onvam6enka 165 28 Perynayna aKcnpeccum reHoB §=229 Penomouna B Naulnx mpeacraprennax o crpykrype JHK nensGexxo nopostiiia Bonpocst o cbynxuim HK. J[pyxuenowenaa erpyerypa cama HojcKasbiBaeT MexaHu3M KonMpoBanna AHK, no- SBOMOULUIE MepeaBaTh 13 HOKOIEHHA B TOKOTE- HHe BakOZMpOBaHNyIO HHpopMawitio. OrKpEITHe MatpHunoii 4 TpancnoptHoit PHK, a TakxKe pac- umdpopka TeHeTHYecKoro Kosa NOsBONHAN TO- natb, kak HK npeobpasyerca 8 dbynkunonasy- ble Geaku. OTH HM Apyrue OTKpBITHA NosBomHAK ccbopMy- AIMOBAT OCHOBHYIO JOrMy MomeKysIApHOli GHO- HOTHM, OTPaxkaloulyIo TPH TAABHBIX Mportecca o6- paGorkH reveTH4eckoii Mipopmaunn B KAeTKe. Tlepspiii mponecc — penankauna, cyTh KoTopoii aakutouaerea B KonMporaaun HK n coanannn Ha ocHope MatepuucKoii JJHK Mostexyap1 s104ep- nei JLHK c naentHynoii nocaeqopateabHocTbIo: Bropoit npouecc — TpancKpunuAs, B peay.ibrare KOTOpoit YacTb reneTHUeCKoit HNCbopMalitH, 3aKo- supobaxxoit 8 JJHK, npespautaerca 6 mowexy.1b1 PHK. Tpernii npouece — rpaneaauna, np KoTO- poli renetHueckan Hudopmaita, 3akoaMpopaititas b PHK, nepenocures a puGocom1, re Tpanc- pyerca B noammentHa c onperexennoii mocnenopa- TEAbHOCTDIO AMMHOKHCIOT. Octiopiat ora MoseKyaspnoii GuororHn ykasbibaeT Hanpapslenne nepeaain HHqopMaitit B[6] Yacte TIT. Nyt nepenasn muopmaynn Penmixanns HK ‘Tpaxckpumyin PHK ‘Tpancasuns Bestox OcxoBvan Kojeryna (jorma) MoneKynapHom 6nonorw, KO- Topan O6tACHReT TnapHHie MeTaGonMNeckNe MyTA Tepenayn wniopmanan — permuKanpo, rpakicKpuTuO M TpaHCnANHO.. Tosopin, #70 270 «norman, He copcem wepxo. Beb ara Korert- un Kax Jorma Guia mpennonesia Opertcucom KpuKow, Korna Guino Mano OKASATERLCTS, NORTBEPATAWMMX ELYEMHYTEE we, TloaKe craBumMe xopouto OGOCKOBAHKOM TeopMelt. KuleTKe: OT pelLNkalWH K TpaNcKPHMIWHH Hi Tpattc- aan. TepMui <0rMa» He CoBCeM BepeH H Coxpa- HIWICA TOKO 10 HcTOpHYecKUM UIpHuMaM. Or 6bu1 BReAEH Dpencucom KpHKOM B TO BpeMs, KOT CYIECTBOBAIO Mas10 AOKasaresIbCTB, TOTBEPKAAB- MIX BLULBHHYTHIE IYLeH, HlowKe CTaBUIHe XopoLIO oGocHopanHoit teopneli B aacrn IT] o6cysxnatores at mt apyrite mpo- lech, cBsiganHBe ¢ nepenayei mopman. B rape 24 Mit paceMoTpuM cTpykTypy, TOHOOTIHO 1 yhaxopky xpomocom 1 revios. Hpoueccui. sexkaute B OCHOBE AOTMBI, paccmaTpuBaIOTes! B raBAx 25-27. B sakiovenne mut pacemotpiim lmpontece perysia1iit oKcHpeccun reneriueckoit Hiopmarun (rapa 28). Baxkneiiuntit Bompoc, o6cyxknaionutiies BO BEX OTUX TAABAX, KacaeTcA CTOKHBIX Mpoueccon 6110- cutrresa iipopmannonir Maxpomostexyz1. COopKa HYKNCOTHAOB MH AMIHOKUCTOT B OTIPEAEIEHHBIe 110- CieROBATEMDHOCTH HYKACHHOBDIX KCIOT H GenKOB CYT [VIA COXPaHEHMst HM TOUHOFO KONMPOBAHHst MaTPHIUbl, @ BE/b Ha 3TOM OCHOBala CaMa 2KH3HD. Moxno noaymatb, 70 o6pasonanne cocebonacbirp- ubix cpaseli 6 JUHK yi nerrriptx cBsseii B 6ex1- KaX — TPHBMAJIbHasl ayaa JA KIETOK, OOaLaIOUUX LleNLIM apCeHasio (bepMeHTATHBHDIX H_XHMHYECKUX MNICTpyMeHtTos, onicanbix B YacT IL, OgitaKo, ¥T0- 601 YHECTS MexalMIgMBl Coxparlenia MU nepenayit HH- copmaium, nam mpigerca aHauirTeabHO paciunpitrh CHCTeMy: HALINX R3rAIOB, ChopMysmpoRAHHy!O Ha ocHoBe anaiiga MeTaGosMueckHx tyTeii. Xumvve- CKHe CBSAM JOKED ROSHUKATH MEXKy KONKpemnbLve cySvenumayat nncbopmanutonnx 6HonomMepoB ¢ MHLHMabHOMi BepostrHOCTbIO LOABZEHUS H 3aKpe- IMIeHHs OUIMGOK. Dro TpeGovarite OKasbIBAeT OVEHD CeppesHoe BIMAHNE Ha TepMOAMHaMHKy, XUMHIO 1 SHSHMOJIOFi0 Tpoleccos GuociHTesa. OGpaszoBanne nenrijwoll caasn Tpe6yeT 3arpaTHt aHepri, mpH- MepHo pannoii 21 KJbk/Monb, H MoxkeT MpoucxosHT ¢ yuactHeM OTHOCHTeNDHO MpocTbIX (bepMeHToR, BUICTYHAIONLX B KavecTBe KaTaHgaTopoB. Ho jt CHHTESA CBAIH MEKILY ABYNST OLIpeseCHHBIMI aMHt- HOKHCHOTAMH. B KOHKpeTHoit TOKE nonuMeNTHAA apeGyeres npumepiio 125 «]bxk/ Mom, IpraeM B aTOM mpontecce aaeticrnonario Gonee 200 dbepmertron, Mo- aiexyat PHI ni cnewmpuseckix 6eaiKon. XuMitsecKsii Mporece OGpasowanina NenTHAHO!i CBASH TOT 2%ke Ca~ Mblii, HO 3/€Cb NOAKMOYAIOTCA /ONOAHNTEbHBIE mpouecces, rapanrupyrontute o6pasowaite sro cBsisit CTPOTO MeXkAY ONPEAENeHHDIMH aMMHOKHCHOTAMH Viacpopmaiuts eronr joporo. Ene ofa paxktiaa Tema, satparipaewas B ¥a- ctu IIL, kacaetes AMMaMM4ecKoro BaanMogelicTBnA M@KAY HYKICHHOBLIMH KHCIOTaMM HM GeKaMH. 3a HCK/IOVEHHeEM TeX PeEAKHX CLYYAER, KOUTA B POI Ka: ‘raigaTopos BHICTYNaIoT MouleKy.1Ht PHK (ara tema o6cyaxaetes B ri. 26 1 27), MeraboruecKHe upo- hleccht, cRagamHtite ¢ nepenaveli mipopsauut, KaTa- Amaupylores uw peryazmpyiores Geant. Hayyenne THX (pepMentow 1 lpyrHx GerKoR HMeeT He TOIbKO. Hay¥HOe, HOH HpHKAaAHOe 3naNeHHe, TOCKOADKY MO3BOAACT UPHMEHATH HX B-TEXHOJOTHAX, OCHOBAH- Hix ta pexomOunantHpix JLHK (em. en. 9 BT. 1). UH prope pospparunca k Teme aposowutH. Mno- rue mponteccei, pacemotpenmise pvacrit HIT, posimctt MHLUIMapiUbl ACT Haga, a HEKOTOPbIe MPOCACKUBAIOT- CA BILIOT LO HocAeAHErO yHHBEpCasIHHOTO O6ULeTO npeamecrsennska — LUCA (or aura. last universal common ancestor). PuGocoml, TipaktiieckH Bech TpancaAiHONHDIii armapaT HHeKOTOpHIe gvleMerrTDt ‘TpancKpUMIMONHOrO anmapata ecth y BCEX 2HBLIX opranuanon na nauteii naanere. Teneriueckyio un chopmaniio Moxio pacemarpnBarh B KayecTBe CBO- eoGpasHLIX MONIEKYJSPHDIX 4aCOB, KOTOPbIE 1103B0- JISHOT YCTAHOBHTh POJCTBEHHMIE OTHOMIEHHA MeEKY BizaMH. O6utne HHcpopMaiuioHHbie NyTH CBAgbIBAIOT YeTOREKA CO BCeMIT HBIHe KHIBYIYIMI Ha Bextte op- TalH3MaMi, @ TAKE CO BCeEMH mpexkite cymlecTBO- bapniuMit Braamn. Hayyenne atux nyteli nomoraer YHEHDIM MPHOTKPHITD 3aHABEC B EPBOM aKTe IbECb!, TOBECTBYIOUEH! 0 BOSHHKHOBEHMH 2K3HH Ha Semute.Cynepexpyumeaxne sxavur pia [JHK Gonsue, vem mpocro canowoii (bakrop; oxo coxpaxaer 6yiiiyio packugucryio HK w yaKwx pankax BKyTPMKneTOX- HBX yenoBHi, — Huxonac Kossapennu, Jlexywn 8 Xapew, 1993 [eHbI M XPOMOCOMbI 24.1, Inementbi xpomocom 7 24.2. Ceepxcnupanu3ayna MHK 15 24.3. Crpyktypa xpomocom 27 aawep JLHK npeanaracr am mtepecnyio 610- AOTHMECKYIO BarAULKY, HOCKOABKY MoseKystbt THK o6biuno snauntesno Kpynnee casux KGIETOK HTH BUIPYCHDIX YacTiM (pc. 24-1). BosnuKa- €T BOTTOC, Kak TAKHe KPYIIIMe MOTeKYsIbI yrlakOBArTD B Knerke. Ur06t OTBETHTD Ha 9TOT BOItpOC, CHEAYET nepeiit oF paccMorpenist BToprHoii cTpykTypot THK (cm. rn. 8) & e€ yansutennoii tpetunoii CTPYKType, sexKauleli B OCHOBe CTpoeHiiA XpoMo- com — xpaniunia renerieckoit midopsauuat. B navase rapt Mot paceMorpHiM OcHosHble auiemen- ‘Tht XPoMOCOM, a saTeM OCTAHOBHINCA Ha OGcyAeHI 1X pasMepa 11 opranusanun. Jjatee Mbi OGpariiMcA K ronowornn HK 1 o6cyquM Bapiairrit ckpyuuBaninst Wt cyllepcKpy4niwanuia’ Monexys. B sakmoyenne Mot oGcyaii Bsamoztelicrans JJHK GesKaxn1, cr1oco6- CTBYIOMUHE KOMHAKTHOI vIETasLKe XxPOMOCOM. 24.1. JnemenTbi xpomocom Knerowias JIHK conepant renin Mexkrenmnte 06- JACTH; W-Te HW APYTHe MOFYT BbITOTHATD 2kTISHEHHO arkHbte dpynrkniit, Bociee catoskibie renioMns, Hartplt= Mep TeHOMDI aYKapHOT, NyKAAIOTCA B Gostee CrOrK- a Ob sane Puc, 24-1. Benkosbiii kancua Gaktepnorpara T2 oxpynKa- eT eAMHCTBeHHy!O AuHelHyIo MoneKyny AHK sroro dara. B peaynsrare nuauca vactuy 6aktepnopara B RMCTMANKpO- paxnon Boze JHK puma u3 xarcuga u pacnpocrpaxunacs To miowepxxoctn Bonk. Yacruna 6axtepmodara T2 cocrour 3 TONOBK M XBOCTa, ¢ TOMOMIBIO KOTOpOTO Gakrepuodar Tpukpernserca K BHEMMeH NoBepxHocTM GaxTepwand- Hoi KnerH. Ber HK, noxasannan Ha 9T0h sneKtpoHHoH muxpodororpadun, obo conepKUTCA BRYTPM FONOBKH ara.[8] Yactb TIT. 24. Tent n xpomocomet HbIx ypopriax opranisanmit xpomocon. Mut niauitest ¢ paccmorpertia pasiminbrx THITOB oceLORATEAD- nocreii HK 1 crpyktypiibix aziementon xpomocom. Tettbi — ato yuactkn monexyn HK, Koaupylowve nonunentuabi M MoneKynbt PHK 3a nocaennee croneTHe Halle npencrarenne o Tenax CyIWETBEHHO HaMeHILIOC. PaHee reHUM Ha- SHIRAI YYACTOK XPOMOCOMLI, KoZMpyIoUULTi HLH onpenermiouutit out mpusnar wm dbenorunnse- cxoe (nimMoe) cnoiicrso, HanpuMep wer ras. B 1940 r. /bkopax Biuut n Sapapa Teiitem npen- AOKIAM MoseKysIApHOe ollpenereHHe rena. 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Conpementioe GuoxumnyecKoe onpexerenne rena eme Goree KouKpeTio. enaxnn aapinarores Bee ywacrkn HK, koampyroune nepsimnyie nucie- AOBATEADHOCT KOHEYHIX POMYKTOB, K KOTOPbIM orHocatcs fomuuenriab wu PHK, o61azatoume ‘Ibxopmx Bunn, 1903-1989 ‘Spsapy Tetirem, 1909-1975 THE mPHK Toannentag Pe bnd® bs |] soe Gc G Arg TOA U Gc G Gac G Gly A T A TOA u Aut A ‘Tyr CG c AoT A CG c Thr TA U TA u THA U Phe TA U Gc G CG c Ala CG c Gc G THA U Val TOA U TA u CG é Ser T » * u 5 IIc KOHeTt ‘TpancxpnOmpyemas nem Puc. 24-2. Coorserctswe Merkay KOAUPyIOWMMM yYacTKaMn AHK, MPHK # annHokcnoTHoi nocnenosaTenbHocTeIo no- AwnentnaHoh yenM. Tpiiers: aykneorupos 8 HK onpe- Rensior aMuHoKMCnoTHyIO TocneROBaTensHocTs Genka mp. nocpenrmvecrse mPHK. Opa ua werrem JJHK urpaer pont anpust Ana cHntTesa MPHK, nykneotwarsie TpAnneTk (KO- owt) KoTOpolt KOMTUIeMeRTapitA TpurLIeTaM JIHK. ¥ KeKo- TOpHx Gakrephit M MHOTHX syKapHoT KoANpyoUMe Mocne- ROBATEMBKOCTH MpepMBAOTCR HEKOAMPYOMMMH yHACTKAMH (rai waaneaemsinn uxmponanu). cTpyktypuoii wan KaTanrriueckoii bynKnueli, Ha- paay c renamu JJHK conepxur 1 apyrie tocaexo- BATEADUOCTH, BHITOAHAOLINE HEKOMUTEABHO pe rysTopnyio ¢bynkuuo, Peryasropubie nocaes0- BATCABHOCTH MoryT OGosHavATh Hayao HIN KOHELL TeHOB, BIMATH Na TPALCKPHILINO WO yKAsbIBATH MecTO HIMHUIAIUIH pelKALUHL WML peKonGrna- ruin (rx, 28). Hexoroppie rent Moryt akenipeccupo- BATBCA PASHBINI HYTAME, IPH TOM O/H HL -TOR Ke yuacrox HK caysxirr marpiueli aa o6pasonannis pasHbix tipoayktos. CourererByioulne MexaHiaMBt ‘TpaHicKPHMIUIH H-TpaNcaAMUN MCaAUL B ra. 26-28.Mbt Moxkem upnGsmgirerbuo paccunrarh sit WiiMaabbiii pasMep retta, KoaUpylouero cpeauttii Genok. B ra. 27 noxpo6Ho pacckasatio 0 TOM, 470 Kark/lal AMMHOKHCIOTA B HOSHNEITHANO! HEN KO- AMPYETCA HOCIELOBATEHOCTHIO 13 TPeX HYKEO- THOR (pukc. 24-2); HOCAENOBATENBUOCTH 9THX TPH tieToR (KOAOHOB) COOTBETCTBYIOT eHOUKe aMH- HOKUCIOT B HOAMUELTHAe, KOTOpLI KOAUpyeTest Aaitbim reiom. Tosmnenrijuias web 13 350 amu HOKHCHOTHMX OCTATKOB (Ienb cpanel sU1tHBI) cootnercrayer nocnejonareabnocti 13 1050 1. 11. Oanako muorne rent aykapnor i HeKoTopnle rena upoKapuor npepsisaiotes cermenranu JIHK, ne Hecynuintit impopsanit 0 Gerke, 1 HosTOMY OKA- IBANOTCS! SHAYUTEUO JUICE, HEM HOKAsDIBAET. npocroii pacuer. Ckonbko reno B oAnoii xpomocome? Xpo- Mocoma mpoxapota Escherichia coli, weit renom ToaHoctmo pacimdporan, mperctarnaet coboit Koabuyenyio Moxexyay JJHK (11a camo seme. 9T0 ne Upawniabnniii Kpyr, a ckopee neta Ges tavana n Konna), cocroamyio H3 4 639 675 1. H. B stot HOCHEAOBATEABHOCTH — CONepANTCH — IPUMepHO 4300 renios Genkos 1 ene 157 renos crabusbnDx Mouekya PHK. B renome yeoseka upiepio 3,1 MAIpA nap HYyKseOTHMIOB, cooTRETCTRYIOLUIX nowrn 29 000 renam, pacnonoxennbim Ha 24 pas- HbIx XPOMOCOMAX. Monexynti [HK ropazqo Kpynuee, YeM KneTONHbIe MN BMPYCHbIe CTPyKTypbI, B KOTOpbie OHM yNaKOBanbi Monexyat xpomocomnoit LHK o6niano ma siro- To MOpAlLKOD JUICE, 4eM KIeTKH HH BupyCHDIe MacTHIUbl, B_KOTOPHIX OME comepaKaten (puc. 24-1; ta6.1. 24-1). 970 oTHOcHTEs! KO BCeM KiTaccaM opra- HSMOB HK BHpycaM. 24.1 Snementei xpomocom [| Bupyest. Bupycot ne Moryr akvTs BHe Apyroro op- Taniiama, pie akNaHecHOCObHOlT KaeTKH, CKopee IIx MOKHO Ha3kaTb BHYTPIKICTOUHDIMH Hapaattraatit, MCHOMP3YIONLIMM PeCypCbl KNeTKH XOaAHHa [UIs paamioxentis, Muiorie sipyciinte sacri CoctosT ‘TombKO 119 reHoa (OBbINIHO o;tHOft MoaeKytst PHK Win JJHK), oxpyakeunoro Geskonbis 4ex:10M Terionot nour Beex supycos pacreiii 1 neKo- TopbIX BUpyCoB GakTepitit H KUBOTHEIX COCTOSIT 113 PHK. Take retions o6s14Hi0 HeGombunoro pasmepa. Hanpimep, reioit perposipycos wzieKomtrato- murx, TaKHX KaK BHU. coxepskar oxo.10 9000 nyk.te- oruon, a Saxrepnodar QB — 4220 nyKacotHa0n. Teriomp1 oGonx Bupycon npestcran.anor coGoil ofmo- netiovesnyio PHK. Teviomor JJHK-conepaanmx siipyco Haxmoro Kpynice (ta6.1. 24-1). Muorme moxexys JJHK bit PYCOB Kaky10-TO YacTb 2kU9HEHHOTO WHK:IA HaXoHAT- ca B3aMKHYTO!t KoAbIEROIi cpopae. [prt permmnKartt Bupyca B KneTKe XOaAIIHIA MOryT HoaBAATHCA cre uduecke hops supycinix JLHK, raspimaenpie penikarusriban dopmannn; ikanipristep, sHOTHE JN~ neiinbte Moxeky-ub1 JK cranowaten KosbuenbIaM, a oanoncnoyednie JJHK o6pasyior sumepsi, Tums noi LHK-conepaxania Bipycom cpeanero pasmepa sipasierea Gakrepnodar 2, wxbunupyiouuii E. coli. Penuxarupias bop JLHK chara 2. anytpit keToK lipetcrawiena KOUCBOIL AByXIEHONeMHO! critpa- subio. ayxnenoueunian JHK coaepxirr 48 502 1111, a uma ee Kourypa cocrapaser 17,5 mkm, Tenom Gaxcrepuodara 6X 174 Torke conepaxur JHK, no ero paamep nanoro Menbine; B supycuoit yactune HK pencrapztenia osnonenoxesnioli KouIbUEBO!l MosteKy- noii, a ABYXHeHOMeNIas perLIKaTHBHas dopMa c nepaku't 5386 ni. 1. Xora Bupycntse renomb MaLtetth- kne, juunia ux JLHK naxnoro Goxme, 4m pasmep CaMUX BUPYCHBIX 4aCTIIL, COepARALLUX ITH MOEKY- aii JUHK (ra6z1, 24-1). Paamep supycuoli ‘Lamia saipyenoit Lama oupyerioit Bupye AUK, nn TUK, 1 acrmubt, M4 OX 174 5386 1939 25 7 39.936 14377 78 n 48 502 17460 190 TA 168 889 60800 210 Tipmmevanne. Pasmep JUHK ykasan suit pemmxarussoii (uyyxnenovenoit) dpopwnt, JLnny HK onenpann, cunrraa pasmep nappt Hycreornion papi 3.4 A (em. pc. 8-13 #7. 1),[10] Yacts TT 24. Tenet xpomocomes Baxrepun, B onnoii kere E. coli conepxures ups- Meptio & 100 pas Goabine JJHK, ¥en p vactine Gak- Tepworpara 2. Baxepna E. coli uneet onny anyxite- Hlovetttyto KombueByto Momexyaty THI. Ona cocto- ver na 4 639 675 11.1.1 tocTuraer 8 sUINHY upusepHo 1,7 wiv, «70 mpeppimaer Muy caMoit KaterKH E. coli MC PAnoM npmsenena knerKa E. coli (2 mKM). C ee Kaerna E. « TUK E. coli Puc. 24-6. AHK wa nManposannon KnerKw E. coli. Be- IMMA CTpenKaMM OTMeYEN KOMBYeEMe MoNeKynEt mnasmwq. Bente 0 veprure nara — apredarmns, Xpomocoma E. coli (qamHoi 1,7 mm) npeacrasnena 8 nMHeiHon opme; npHGssunTenbuo B 850 pas (puc. 24-3). Tlomumo KPyMIHOii KoAbILERO! XPOMOCOML B CocTaRE HYK:IC0- ida MHorHe GaxTepitt conep»kaT ANY HI HecKOmb- KO MATENbKUX KOAbIeRKIX MoreKyn JK, cpo6on- HO pacrionaraiouunxcs B irosose, TakHe Biex- POMOCOMHDIC SJeMCHTHL HasniaIOT TuaAMILaNt ot 4; em, rawoKe c. 439 8 1. 1). Bomba CTRO TinaAMI COCTONT Beer HA HeCKOADKHX Th CaM Tap HYKACOTHAOB, HeKOTOpbIe comepxaT Gos1ee 10000 nH. Ont necyr reneriteckyto sicpopmanino 1 periiupyiores ¢ o6pasonanuest .04epHHX 1. MIA, KOTOpHIe ToTlaRaIOT B.LOYEpUTE KACTKU RB IIpO- uecce jeslenus posurreascKoii Kerk, Thiasstiabt oGtiapy2KeIlbt He TOMDKO B GakTepiAX, HO TAKE & ApoxoKax 1 apyrinx rpaGax. Bo MHoriix cayuanx raasiytbl ne jawor muKa- KIX TpeUMYLUeCTR KIETKAM-XOBsICHAM, HX eH CTBeHHAs dautatia ~ NesapiIcHMoe Bocpor3BesteHnte. Onnako nexoropsie tuiasMiutbt HecyT noxtesHie Lit xossuttia rents. Hanpnep, comep»kamneca B titan ax reli MOryT MIpiyiaparb kaleTKaM Gaktepusii ycToii- unwocr kanTHGaKrepuaabHbis arertas. Thiassttbt, hecynuie ren B-TaKTaMaanl, oecneuunaior ycroil- SMBOCT K B-TAKTAMIDIM AITHGHOTHKAM, TAKUM KaK neni H aMOKeHULLAI (eM, pite. 6-28 BT. 1). Thiasmivis Moryt nepexosirTs oT KieTOK, yetoiiut- BBIX K ANTHOMOTHKAM, K PYTHIM KALeTKAM TOTO 2Ke HIN apyroro nia Gaxrepuit, B peaysubrate Yero aTH wer Kil TakKoKe CrationsiTea pesuicrenTupisu. Hirrencin- Hoe MpuMencHue aTHGHOTHKOB anAcTeA MOWUIEIM cestextuniisiss cpaxtopos, cilocoGeTBYIOULIM pacttpo- Crpanenino m1asMiut, KOLIPYIONUIX yeToIiMMBOCTH K aHTHGHOTHKAM (a TAIOKE TpalicTIOsoHIOD, KOTOPLAe KO- AUpylor anastornuiste rent!) cpestn GosesnerBopHibIx Gaxrepuii, 1 npuBomur k HostB.TeLIO GakrepmaztbHbtx IITAMMOn C YCTOITITBOCTSIO K NECKOTDKUM atrTHO0- Tukast, Bpauit iauimiaior 1oMMaTb onlacHocts UM PoKOro HcHHOAESORANIA ANTHOHOTHKOR H HastiasaIor HX TosIbKO B Caysae OcTpoti HeoGxoaHMocTH, Ho atia- JOTMEDIM IpHIMAM OrpalliiBaerest LiNpOKOe 11C- TlomsOwAHHe ALTHGHOTUKOR {UIA seYeHNs cembcKO- XoasilicTReMHbIX 2BOTHEIX,DyKapnorn. Bcrerke aporokeli, onHIX M3 CaNDIX MarenbKUX aykapnor, B26 pasa Goabme JLHK, yea B kaerke E, coli (raGzx. 24-2), Kner nr010- Boi Mymikn Drosophila, Kraccueckoro otwerta renlerineckHx HecenoRAKii, coneprKaT B 35 pas Goamine HK, a kneTKH HeaoneKa — nprMmepno B [TET Ait remiponoconinevoropumeprmios ueno: Tipneproe OGuas JIHK, uu. xpomocos —_uutea0 renoe Escherichia coli Gatepus) 1639 675 1 4435 Saccharomyces cerevisiae (npoxxu) 12.080 000 16° 5.860 Caenorhabditis elegans (iwemarona) 90 269 800 12° 23.000 Arabidopsis thaliana (pacvenne) 119 186 200 10 33.000 Drosophila melanogaster (nnononan myuika) 120 367 260 18 20.000 Onyea sativa (puc) 480 000 000 2% 57000 ‘Mus musculus (tatu) 2.634 266 500 40 27000 Homo sapiens (uenovex) 3.070 128 600 AG 29 000 24.1 Snementsi xpomocom [11] 700 pas Goabme HK, vem kaerku E, coli, Muo- rue pacrenua 1 aMiiGun coxepxat ene Gomi AK. Teneriueckitit marepnian KaeroK aykapHor opranigoBant p ie xpomocos. JyuLtoHsAUbIT Ha- Gop xpomocom (2 n) aanncir or pia oprannasa (1a6z. 24-2). Hanpunep, B comarieckoii Knerke Tipumesanne. Hicpopsaniis nocrosinuio ouonnsieres, 1019 no.ary PATHITECH K Caitras, HOCHSLICHIBIM OTAEADIDIM TeHOSILIN. pOeKTAM. * Jb1s ucex aykapnor, KpoMe {poACKeti, mpHHo, Fanionunait naGop. JLuKHe nrraststit ApoxO aGopon TaKix xpomocos. Test ANN OTL ® JL camox ¢ anys X xpomoconantn, ¥ canton ecrs X xpomocoma, no ner Y.,€, neeto Tf xpomocons o oGeHHO HMCIET HoOceMS (oKTANTON AND) HAH GobAE suns Goce cwexeit nuupopsaist 6 1a6op xpomocont ucatoneKa 46 xpomocom (pric. 24-5). Kaxkaas xpo- MocoMa aykapHoTHeckoil KAETKH, Kak TloKasaN0 Ha pic. 24-5, a, ConeprkITT ONY OYeHD KpyILHyIO apyxcrupambuyio MoaeKyay HK. eaquate uermpe xpomocoms! venoReKa (22 napHite xpo- MOCOMBI Ht jIne HoTOBDIE XpoMocomnl X it Y) pas- anmmarotea 110 suiMMe Goace sem B25 pas. Karka XpoMOcoMa ayKapHor conepxUT onpegesenHbtii a6op rerion. Ecant coenmnnits mexxnty coGoit Monexyat HK seoneveckoro renoma (22 xpomocoml it xpomoco- Mot X iY aun X it X), noaysnrrea locaenowateay- Puc. 24-5, Xpomocomel aykapmor. c — mapa CBAIAMHHX H KOHJeNCHPOBAMMBX CECTPHACKUX XPOMATABL Wa xpomocomi Nenopexa. B raxKoit (bopme aykapMorHveckie xpoMOcoMit pebEtBaIoT MOcne perAKa- nH MB MeTadase B mpoecce mnTo3a. 6 — ronneti Ha60p xpomocom W3 nemtKOYWTA OAHOrO H3 aBTOPOS xanrn. B x@xknoit HOpMaNbHOHt CoMaTHHECKO! KneTKe NenoBeKa COnepKUTCA 46 xpOMOCOM. res oa eo _ =~acre IIL. 24. Tenet xpomocomel Hocth {LNMOii OKO:IO ouIOrO MeTpa. BosutMeTBO KIIETOK HeAOHEKA ANIAONANDL, OSTOMY O6utaA ATH hia [HK raxnx kneroK oKos102.m. ¥ napocutoro wes10- Beka mpuMepHo 10"! keToK, TAKUM O6pasoM, oGuLast auimna Bcex Mosiexya JUHK cocrapsser 2 - 10'' KM. Jaa cpasnenus, okpyxxnocrs 3eaumt — 4 10! KM, a paccrosnne or Semin no Coma — 1,5 - 10% Ka. Bot kak yAMBHTEADHO KoMHaKTHO yilaKoBana JJHK B HanIMx KyleTKax! B wierkax 3yKapHioT ecrb HM Apyrue opranea- ant, conepaxaumne JJHK, — sto MHToxonpHn 11 xst0- pomacrit. Moxexysu muroxonapuansuoii HK (str/IHK) namnioro Menpme sepuprx xpomocom. Pasmep aByxuenouewix KoabueRbIx MTIHK B KyleTKAX AKUBOTHBIX CocraBiseT MeHee 20 000 n. HL (B MuToxoNApHAX YenoBeKa MoseKysa \r/IHK coctont 3 16 569 11. #.). Kaxnas surroxonapua OGuUNO HeceT OT ABYX AO jeCsTH KONI Mose kya MTJLHK: ix siics10 MoxkeT BospacTHt 110 coTen B HEKOTOPLIX KsIeTKAX, HallpiiMep B ANdxpepentyt- pylouUIxca KxeTKAX aMGpHOHa. Y HEKOTOpHIX Op- TariiaMoB (HampuMep, TpilalocoM) Kakitas Mit TOXOHAPHA CoMepAUT THIEHIN KOMI MTJLHK: Mu TOXOHAPHM OpranHaoRAN ht B CIOKUDIT KOMTUTEKC, hasbinaemmii KHMeToracToM. Paameph MTstHK PACTHTeNDHBIX KeTOK RapbpyioT or 200 000 20 2 500 000 1. nH. Moaexysr JJHK xaoponaacton: (xiJIHK) Toxe zByxuenouesnie 1 KotBHeBDIe, Puc. 24-6, flenaujanca muroxonapun. Hexoropsie suto- XOWApHanbHME Genk Monexymst PHK (Ha bororpadwn He BUAKEI) KOAUPYIOTCA ORKON KOMEN MUTOXOHAPHAMDHOK THK. Muroxoxapyansian HK (xnJHK) penmanapyerca KakAMi pas, KOrRa DEAMTCA MUTOXONDPMA, 4170 MpenMecTBy- er nenexwio KnerKn. ux junta Kore6nerca oF 120 000 x0 160 000m. H. Buigsuranoch MHOKECTBO THMOTES OTHOCHTEDHO upoucxoasienia JJHK mitoxonapuit 1 x0po- quiactos. Ofuenprsnannan ceroaus TouKa 3pe- Hii BAKMONAETCA B TOM, YTO OHI IIpeACTaBsOT co6oii pyAUMeHTb XxpoMocoM j:peBHHX GakTepmtii, KoTOpble MPOHINKAH B LUTOTABMY XOasitCKUX KITe~ TOK W CTA LIPeUNECTBEHHHKAMH ATHX OpranesLt (cm. pic. 1-36 B 1. 1). Muroxonapnasbias JLHK Kojumpyer MHTOXOHApHasbuMte TPHK 1 pPHK, a TaIKKE HECKOALKO MITTOXOMApHATbULIX GeKOB. Bo- nee 95% MUTOXONApHADHEX GeAKOB KOANpyeTes snepnoit JJHK. Mutoxonapnn 1 xtoportacrer 1e- ausitca pMecte ¢ KierkaMn. LHK atix opranesia pe- ILINWHyeTeA epet WeENEHHeM KETKH HB TIpoUec- ce jenenns, a sare novepune MoneKyst Mr/[HK Tolasaior B Opranezutbt AoNEpUNX KIeTOK. Tetibi m xpomocome! 3ykapwor O4Hb CHOKHO OpraHW30BaHbI Y Gakrepuit muorix BtoB Bcero Osa xpomocoa, I MOYTH BO BCeX c:AY4AX B KAXKO! xpoMocoMe IIpII- cyrerayer 110 oni’ KonmH KaxKOFO rena. Jv He- Miorne renbl, Hanpumep renbt pPHK, coneprxatest b HeckOKHX KONHAX. Terbl 1 peryisEropHDle 110- CACHOBATEMbIOCTH COCTABIAOT IIpakTHYECKH Bech reviom poKapnor, Bostee toro, mOuTH Kaxkjtbtit rel crporo coorsercrayer amnnoKHtestoTHOii trocaIejtoBa- TexbHOCTH (II NOCAeOBaTeMOCTH PHK), KoTO- pyto on Koaupyer (puc. 24-2). Crpytypray u ¢byuxumonanaas opransa- IMA renoB aykapnoT ropasio ca1orxKHee. Hecneno- BAH XPpoMOCOM dyKapHOT, a MoOs/Hee ceKBEHt- ponanne HoAnbXx HoCeAORaTeMNOCTel reHOMoR aykapilot UpHuecAO \Woro CHprpHa0B. Muorue, ecan He GombunIcTBO, reHOB ayKapnor o6sanator UnTepeCHOii OCOGENHOCTDIO: HX HYKAOTHAUNBIE O- caeqOBATeABHOCTH COMepAAT OLMH IIH HECKODKO yuacrkos JJHK, 8 korupsix ne Kolupyeres aMuno- KHCIOTHAA HOCHAELOBATEHHOCT HOsMMMEHTHANOO fpoaykta. Takie HeTpalicpyeMble BCTABKH Hapy- WaiOT LIPAMOe COOTBETCTBHE MCKAY HYKCOTHIHON HOCACAOBATEIBUOCTbIO TeHa MH AMHHOKHCHOTHOI HOCHELOBATEMDHOCTLIO KOAMpyeMOro —losHETTeN- Tila. ITH HETpancaupyeMbie cerMenTst B CocTABe Feion HagbinAlOy HUTPOHAMH, 121M BeTPOeHHDIMH HocaeAOBaTeMHOCTAMH, & KOAEIPYIONUNE CerMen- Tht — okaonamn. Y tpoKapnor am HeMnorue reHbl Coep2xaT MUTPOUbI.Ten sustoro A BO annGymmna 1 2 90 n, #. 222 n. 4. 24.1 Snementei xpomocom | 13] Sxon 3 Verpor i 126 nH. B-CySnenmmnna’ A remors1o6una 131 mw. B 851 nH. Puc, 26-7. M¥rpowbl 8 AByx aykapMoTHYeckMx reHax. Tex ausHOrO ANLGYMUNA CORePMUT CeMb wiTpoxoe (or A no G), pasrensiouwx KORMpyIONMe TIOcReROBATEMBHOCTH BOCEAN 3K30HOB (L, of 1 po 7). Tex fi-cy6nequimus remorno6una HeceT va WiETpOHA M TPM SK30Ha, BKMIOYAR ORMH WHITPOH, conepwamnit Gonee NONOBUEE Nap KYKeOTHROB LaHHOrO Teva. Brinnivunpix renax Bpicunx ayKapitor nocsen0- BaTeNbHOCTH HIITPOHOB OGHUNO HaMHOrO AMUHeE, vem TlocsieonaTemBHOCTH aKs0HOB. Hanpumep. 6 rene ofuoit NOTMHENTHANO Her ANMOrE GeaKa onanbOymuna (pic. 24-7) MHtTpoNL HaMMoro AHH Hee 9K3ONOB: CeMb HHTPOHOB BMECTe cocTaBIsIOT 85% JUNK rena. B rene S-cyGbennmint remorno- Ginia Goxee nosoBitM JLHK conepxxutes 6 enutti- crsenHom uuTpone, Ten MBnucunoro Genka THTHHA yeMIOH o KomNYCCTBY HTpoHOB — ux 178. B re- ax rncToHon, Ho-nitzutsrowty. Hrrporton ner. B Goat IMHETRE CTYYALB YUKU HATPOHOB He yeTATOB- aena. B nero, anmb oKo10 1,5% JJHK yestopexa SIBAGHOTCA ), ConpoBoxsaioMux ocratHyo JLHK. Hpocrnie nocteaosarensnocru HK ne Konupyor Genny wit PHK. Ho p otanune or 10jBHKHBIX re- NeTHYeCKUX azeMenToB caTewuTHad JLHK Moker BLIIOJIT B KuleTKAX YeNOBeKA onpestesIEHnEIe PYHKLMH, LoCKObKY MHOTHE IIpocTite nocseEOBa- remuocrit HK stoxassonanet 8 abyx cirensiit- MECKHX yuaCrkaX ayKAPHOTHYECKUX XPoMocoM B IleHTpomepe 1 Te1oMepax.[14] acre TH. 24. Tens n xpomocomes 45% sppaNcnosons, 13% 21% SINE Puc. 24-8. Tunbt nocnegosatenbHocTen B reHome YenoBe- ka, Ha nuarpanne mpencrasnenst Tp rape coctaBnsio- ume reHoNa — rpaNcrosonet (nlonBWKHHe reneTHYecKHe aneMexTw), TeHbt K cMeuIaHNWe MOcneRoBaTenbHOCTH. H3- BeCTHO NeTNpe OCHOBHNX Knacca TpaxcTIosONOB (Ka AMa- rpamme mpencrapnenst pM 13 Hux). [InMMNOZe paccertmnxe nostopst (LINE) pasmepo of 6 n0 8 7. mH. (17. 1. H. = 1000 r. 1.) o6uruo conepaar HecKonsKO reHoB Genkos, Ka- ranvsupyioulwx Tpaxcnosywio. B rexome HacummbBaeTcA mpumepxo 850 000 LINE. J[nuna xoporKnx paccentisx To- topos (SINE) npuxepxo of 100 no 300m. x. B retome yeno- Bexa Hx CORePAMTCR OKONO 1,5 MnK, M3 KOTOPHX Gonee 1 MAK cocrasnaor snemextit Alu, HasBaHHBe TAK Mo To MMMM, To OHM OGHURO COnepMaT ORME Cait PECTPMKYNK AA 3H- Rowyxneass pectpnxnun Alul (cm. puc. 9-2 BT. 1). Brenome raxwe oGxapyxexo 450 000 KonMii TpaxnosoHOB, GnusKHx perposupycam, anHom or 1.5 no 11 7. 1. x, Xora OHM «a- xBaveHM» eHOMOM 4 Me MOTYT Mepemenjarsca U3 KNETKH B knerky, 380m0yKONHO OK GnuaKH perposupycam (rm. 26), x KoTopHm o7HOCHTCR BAY. Enje owe Knacc Tpaxcno30HOR (cocrapnnounii <3%, 3nech He Moxa3ax) Mpenctasnex pas- AMMAIOUMMUCA NO AAMKe d)parmertaMu TpacnosoHOB. Tipumepiio 30% renoma cocrasnaior rocnegosarensHo- crm, conepmanyne rexst BenkoB, HO TONBKO ManaA YacTb 3T0; HK waxonurcr 8 sxsonax (KonMpyIOMMx rocneRoBatens- voctax). K cMewlaHHBIM MocnenoBaTenLHOCTAM OTHOCATCA mpocrize xoporxwe mosropit (SSR) « nosropst xpymatunx cer- mestos (SD); nocnenue scrpevaiorca Gonee vem B OKO KOMMM pasnwynesx monomeHviax. Kak orMevaeTcA BTN. 26, NOTH BEC) TEXOM B PesynBTaTe TPANCKPUNUMH Npespama- eta 8 Mlocnenosarensuocrs PHK mpuvem MHOrMe yuacr- kat PHK enje xe oxapaxrepusonai. Kpome Toro, 8 rexome conepmarca OCTarKH TpaxCnoz0KOB, KOTOpNe ® Mpouecce SBONIONMK WSMEHMNMCh HACTOMBKO CHNBHO, NTO WX TPYAKO upextuduyuporars. Lerrpomepa (pic. 24-9) npeacranaser coGoit n0- caejovateabnocrs JJHK, k Kotopoli p mponecce AeNCHH KIETKH TIpHKpermoTeA Genk, chaabima- Joule xpoMocomy ¢ MiITOTHTeCKHM BepeTcHOM. ITO naaniMoztelierpite naxkHO si panlioMepHioro it Tou Horo pasyjeernia naGopow xpoMocoM Tio odepHTNt Kaetkam, Bbrvit pbieneHib 1 Hay'4eHtbI LlerrTpOMepBt Saccharomyces cerevisiae, Baxneiiune ana dynnunt- Onlmponaitis eH TpoMep HocAeAORaTesDHOCTH. HMe- lor umtiy oko 130 n. H. 1 coRepKat GostbIMIOe KO- aimcerno nap A= T. Lerrposeps nbicurix aykapi- OT HaMHoro JUIMHHICe H, BOTANIC GT APOXKEDEIX, o6brMO comepxKaT Mpocty!o MocstenonaTeTbHOCT JAK, coctoamyto 13 Toca Tartentinx Kom O7- Holi HIN HeCKOMBKHX KOPOTKHX NOCTCORATENIDHO- creii 13 5-10 1. n. p onitHakopoii opmerrrann. Pos Mpocrix nonropon npit ¢byHKIUOKTPoRANITH eH ‘Tpomep Tioka ToUNO He yeranon.teHta. Testomepsi (or rpeu. felos — Kone) — nocsienona- TeAMIOCTH Ha KONIIAX xpoMocom aykapHioT. cra6ut- JINSMpylouUste XpoMocomb. TeoMepnt saanunpa- lores MHOTOKpAaTHO HOBTOpsIOUUIMHEs HOCAEORA- TembHOCTAMH BIA G)T.Gy), BYVAL)» rae x Uy O6MUNO UMeIOT sHaueHHA OT 1 20 4 (ra6.1, 24-3), Uneao teromepnbix nostopon ny Goabunmicrea oAHOKTeTOMX aykapHor cocTaD- aaet or 20 20 100, ay NureKonMTaiomutx, Kak npa- BILTO, mpestamact 1500. Kort nmelinoit moneKy- at JIHK we soryt nomnoctsto penatutponarsca O6sNDIN CHOCOSOM. KeTOUTBIM. perLANKaIUOH- HIM armnapatom (noaMo2kHO, 9TO OANA US MPHIL Kouuenoii crpyktypm JLHK y Gaxrepuii). Tero- MeplbIe HOBTOpHI tIpHcoeAHUsOTES K KOI dy KapHOTHYECKHX XPOMOCOM B OCHOBIIOM € MOMONUEIO ehepmerrra resiomepasnl (cm. pic. 26-39). ‘Tenomepa Lexrpoxepa EE eee Yuuxaspunie nocnexosateuoct (rex), PacceMMEBIe HOBTOPH, MHoxKeCT HEHE TOWKH Hayaia penunKALn Puc. 26-9. Bamtuble oco6eHHocTH crpoeHina xpomocoMes ApoxoKeii.(CEE terse noc Hocaexonarennnocns. ‘eaostepintx nowtopon ‘Homo sapiens (aen08eK) (TTAGGG), Tetrahymena thermophila (TTGGGG), (peciuruaroe npocteiiaee) Saccharomyces cerevisiae ((TG);-(TG)p.3)n (aporoxn) Arabidopsis thaliana (pacrenne) — (TTTAGGG), Jlaa nayuenna dyixnmonanbuoii pont erpyk- TYpHDIX QleMeHTOR DyKAPHOTHYECKHX XpoMOCOM 6pm CKOHCTPyHpoBaHDt HeKyCeTBeHHDIe Xpomo- comis (11. 9 Bt. 1). JLaa bynknonponania 20- crarouo craGuannoii snneiinoii HeKyecTREHHOii XPOMOCOMBI HYKHbI JHU ‘TPH KOMIIONEHTA: Wen ‘Tpomepa, TeOMEpHI Ha KAKOM KONIC HocTeLOBA- TEADHOCTH H Y4ACTOK, B KOTOPOM MPOHCXOAUT HLH nalts periuKkanit. UekyeerRenible XpOMOCOMbI apoxoxeii (YAC; em. pic. 9-7) 6biin cosnanbl Kak MUCTpyMeHT Ant GuoTexHonorHuccKHX Uecseno- sani, Jaa acuenua renernseckux sa6onenannii ¢ MOMOMHIO CoMaTHeCKO!t reHHo‘i Tepamun GLH paspaGoranm HeKyecrseHiue xpomocomn YeAOBe- xa (HAC; em. on. 9-2). Kpatkoe conepkanue paapena 24.1 SNEMEHTbI XPOMOCOM Ten — yuactox xpomocomns, KoTopulii neceT mt dopmauino o bymeunoiaibioM HomMeNTIe win mouexyae PEK. Hapauty ¢ rewast xpomo- coMbI coepxxar pasnoo6pasiiiie perysisTopHble NocsieaonareabHOCTH, YuacrByIONEe B perv Kalliti, TpatickpHAINH H ApyTHX Mponeceax. = Tenosian HK 1 retomias PHK supycos OGbIMNO Ha HECKOALKO NOpsKOR jee CO- AUPIKALLUUX HX ICTETOK HI BUPYCHIX YacTHL, & — Mntorne revs aykapiior (io auth HeKoTopHIe rent Gaxrepnit 1 apxeli) mpepsmnaiores rieKo- AMpy!OUUIMM NOCTeTORATeABHOCTAM — He ‘porta, Koanpyionute yuacren, pasnenenipre HiTTporann, HasbIBatoT aksOHAMH. & Mence tperu renosnoii HK uenopexa cocran- syuor reubi. B ocraabuoit HK suioro nosrops- 24.2 Caepxcnmpanwzayun DHK | 15] JOWIMXCA NOCHELOBATEANOCTCH Pad AMYIIX TH nos. Tlapasiirupylone HyKseHHOBIAe KHCIO- Thi, HSBCCTHDIC Kak TpalcHosonDt, CocTABsAIOT TIpHMeptio oatoniiny Teoma YenoneKa. m® Xpomocombi aykapuor cojepxkaT Awe BwKHDIE nowropsionuiecs nlocuexovarensnocrn JJHK co cnemmpruccKoii dbysKuneli: erTpomepst (yuacriat upnxpeuteuiot surroriueckoro wepe- Tella) 1 TEMOMEPHI, PACHOTOXKCHHHE Ha KOHNAX Xpomocom. 24.2. Caepxcnupann3zayua HK Kaerouran JJHK, kak mpl puaenn, ypessptuaiiio KOMIAKTHa M, CIeLORATEDHO, O6s1aRaeT BbICOKOIL crenienuio erpystypnoit opramisanuit. Mexanusst YANK CHYKUT HC TOKO JUL yNaKOnDIBALLIA Se Cynepexpysennan ermpamm Puc. 24-10. Ceepxcnupannsayna. OG TenedorHEit yp 3axpyyex 6 crupans, vax JJHK, u cam Moxer o6pasonn- Barb AOMlOMHMTeNbHWe NeTNK CBepxcnMpanu. Ira aHanorHA YMecTHa eule Mo Toit MPMNMHe, ¥TO WMEHHO TeneporRD amp nomor /jKepomy Buxorpany M ero KonneraMm TORATD, To MHOrwe cBOlicrBa MaNeNLKHX KONBUEBEEX MoneKyn HK Moryt o6AacKaTbca cynepcxpyanaxnem. Bnepstie oxM 06- Mapyauinn aBnenwe ceepxcrmpanuseyn [HK 6 1965 1. 8 obpasax ManenbKHX Monexyn BUpycHO; [JHK.[16] Yacte TIT. 24. feats u xpomocomer HK, no 1 ans oGecnevennst aoctyna « conepaxa- ielica B Heli mKbopmaiin. FpeaKite Yem pacemo- ‘per paGory aTOrO MEXAIIIKIMa B TAKHX KsIeTONHBIX lponeceax, Kak PeruIHKAIIa HTpancKpHTIUAA, MbL AODKHML O6cyANTh BAKHYIO CTPyKTYPHYIO OCcObeH- nocth JK ~ enepxenupamsanuo. Crepxenmpamusanust — expyunpaume yoxe cy- mectayionteli cnupasi. Tpamepom taxoii erpyk- Typhi sinasieres ckpysennmii TeaccbounBtit unyp. Ha nya Mexay TpyOKoit 1 annapatom 4acto 06- pasyeres oa HIN HecKOAbKO crepxenupasteli (ue. 24-10). Monexyaia JUHK expyvena 6 2noii- Hylo clinpanb, B KoTOpO!l oGe He JLHK aaxp: yep BoKpyr esol! oH. Jlanbuelimee saxpyan- pantie TAKOii OCH BOKPyT camoii ceGx (pic. 24-11) Ippo K OGpasoBalliuo cBepxcHupacHsoBANHoii JIHK. Kak mot yeiunst aasiee, chepxenmpasiuaanist OGMUNO sBAKETCA peay.LtaTOM CTpyKTYpHOrO Ha- nipskenns, Eoa JLHK ne 3axpyyena Boxpyr cvocii OcH, Ce Hashinaior peraKenponannoii Jlorisno upeanonoxirh, #To UpH YnakoBbIna- nut JUHK posimxator pasmiunute sapnanrit cnepx- crmpamisanunt. Mevee oxesnno, ro pemisKanutst i tpantcxpamuts LHK toxKe wansior ta eepxerit- Jleotnan cupam HK Oce Cynepexpyvennas: THK & wv Puc. 24-11. Caepxcnapanwsayun HK. Korga qeotinian cru- pans HK saxpysneaetca Boxpyr ceoeh ocu, o6paayerca Ropaa cmmpans (caepxcmmpant). Cyrepexpyerryio [HK o6unnito KasbiBaloT ceepxcrupanio. . ma" , Seay «ft E e f es 7 Puc. 26-12. MIpH paaenenuu witTei cnapanbHoi crpykrypet TpOMcKoAMT CBepxcnHpanHsayMA. 3axpyTATe [Be PeIMHKie B Mpasosaxpyvenxyio ABoiMyI CmApaMs, KaK MOKasAHO Ha pepxiem pucysixe. Tompocure xoro-mM6ymb nonepwars ont Koren, u nonpoGylire paanenurs HiTe C npyroro KoHNA. Bu Gynere HaGmonan> cymepcxpyymparne. Pamsarno HW camH saBHest oF Hee. O6a mponecca cBaganinl ¢ pasmenenuem mei JLHK, cocoéersy- fom OGpasouanino AottomMUTTeMbUDIX BITTKOB cH pas (kak NoKasaHo Ha pric. 24-12). To, 410 1IOTHO YHaKoBaNad KAeTOUHaN JJHK aaKpysiinaerca BoKpyr caNoii ce6a, upHoGperast CBEPXCLINPAIBUY IO CYPYKTYPY, KAKETCA JOTHAHBIM 1 ARE TPURIAIDIbIM, COM Gbt He OfNO HaGmOACHHE: Mure KAeTOUTIUe KOTDIeBLIC MoIeKyztbI JLHK ocra- 1OTCH B BbICOKO! CTEHCHH CyHepcKPYYCHHbIMH JAKE HOHE TOLO, Kak 1X DKCTPArUPYIOT Ht OMUMUOT, OCHO- Goran or GeRKOB 1 JIPYTHX KaeTOUHBAX KOMMOHEH- ‘ror. H3 arore clenyer, 470 cBepxcHupaZiauuis — HeOTbeMIeMOC CBOICTHO YeTBEprHSNO!L CTpyKTYpHE ALK. Ono xapaxtepno .y1a Bex Monexya Kerounoli AK 1 erporo perysmpyeres kaxkitoi4 kerk. Crepxenupamt xapakrepnayiores ecko. MH TapaMeTpaMut, KOTOPbIE MOXKHO H3MeputTD; Ha octowe aTHx janix Guun rayGxe nomATD CTPYK- rypy mt bynniuim JUHK, Hecesienonaiis Bo MHOrOM. Gasupylorca Ha KOHLEMILAX ToMOTOrM — ojutOrO M3 paSAeNOB MaTeMaTHKH; ToNONOrHs HCeHeAyeT cpoiicrna oGbeKra pH HerpepuBHoli AccbopManuin. Beayuae JUHK nenpeppipias fecbopmalist BONA er HgMenteriis KondopMalUitst 13-3 TeHIOBOTO ABN 2KCIIA 1 BaAHMOAeTICTBNA ¢ GesKAMHL HUH sIpyrHMit MonleKysiaMu; NpepprBiictas jecbopMatta npiROT K pasppiny uenei JLHK. Jaa Konpuenbix Mosexy.1 JIHK rononornueckue caoiicrna — 910 Te cnoii- cTBa, KOTOPhIe He MeHsIOTCA TPH AecpopMannit We- neit JLHK no Tex nop, oka He nosHuKaloT paspHipb.Tonoxoriueckne cnoiicrna HaMCHAIOTCA TONDKO B pesyaibraTe paspbina H rlocaeAyIoULero cocaMHCHHA ontioii win oGeHx nenteit THK. Tenepb pacemorpum pynaamenranpipie cnoii- cTBa H (pusH4eCKyIO OCHOBY CBepxCNHpamHsanHn, Bonbuimicreo KneTounbix HK packpyyexsi Yro6pi NonATh cyTb cRepxerpaMsatHH, Cocpeno- TOWMMCA CHayasa Ha cBoiicrBax NeGOsUDUIX KOs nenbix Monekya JJHK, raxnx kak iraaMiast 1 He: Gonbume JLHK-cozepxaune supyce. Ecam raxue Monekyibt JIHK ne umeror paspbinos Hn B onHO! Hs Leneii, OHM HasbIBAlOTes BAMKHYTHIME KONBIEBHIMA Mosexyasamn JIHK. Eean copma samxnytoit Kom nesoit JLHK 6anaka no crpyktype B-chopme HK (ctpyxrype Yorcoua~ Kpinka; em. pic. 8-13 BT. 1) C ogi oGoporoM jpoiinoii enupann va 10,5 1. 1. ola ckopee peKcHpoBanla, YeM cynlepeKpyyena (pue. 24-19). Cynepexpysnpanne nponexozurr npit HekOTopoM HanpskeHHM CTpyKTypbt. Ounuentiast gamkuyTas Kobucran JJHK pero Guipacr pezak- cuposannoli, Hesasncnmo oF ee GHowortecKoro npontcxorxtenst. Bostce Toro, Mosieky.it JLHK, 110- AYYCHHLIE 13 ONPEMEICHHOFO KICTOUHOFO HCTOUHH- ka, HMCIOT XapakTepHylO HMCHHO JUIA HHX CTeICHb cpepxenupaszannn. Caenosaresbno, HK nanpa- 2keHA B-TAKO!i CrelteHH, YLOGbI B Heli BOSHUKAIH Cy- TCpBITKH, 1 9TO CocTosHHe peryaNpyercs KIETKO! Tpaktisteckst B s1o6oi Moment mpesemt ta npaxenua swotnolt cumpan JIHK sosniKa lor H3-3a ce YacTHMNOrO packpyunBanna. Vina- ue ropopa, 8 JIHK okaspivaetes Menowe vittKon 24.2 Ceepxcnipannsayua HK |i cpa, set B Beopwe. Bausnne yacrirato- TO packpy4nparinis OGo6meCHO Ha pie. 24-14, Yuacrox KombneBoii JLHK paamepom 84 11.11. B pestak- CupoBaNIHO‘i popMe MOrKET CO;EpAKATH BOCEM BATKOB atwoiinoii crupaam, sur oywit BHToK Ha 10,5 11. u. Ferm OsMH #13 TAKUX BHTKOB yrtasien, Nomyaaercs 84 11H. = 12,0 1. #. na BuToK, 7. ¢. 6ombmte, vem 10,5 — Bem uuma, xapaxrepnas sia B-IHK (pute. 24-14, 6). Ito orkronenne of HanGoace cra6wHoii opt JK IPUBORUT K TepMOsUHAMMNECKOMY HaNps>KeHIHO B Mosiexy.1e, 7. €. k neycroliunsoii dpopme JIHK. Ora- CTH tanpaxeune Moker GbiTb custTo Mp ckpyHiBa- iin JUHK noxpyr coGersentioii oct ¢ oGpasonarinies cpepxcrmpartt (pc. 24-14, 6; HeKoTopas sacTh Hanps- OEHHA Ha YuacTKe 13 84 11. H. MOXKET MPOCTO paciipey JlstTse8 10 packpy4entioii crpyitype Gotee Kpyitiioro ébparmenra JUHK). B mpiuigine itanpsxere Moxer GbiTh KOMIeHCHPOBAHO Ip pasfeenHN ABYX ere JUHK 11a paccrosut Koso 10 1. i. (pic. 24-14, 2). B nsornposannoii samKuyroii Konpucsoit JJHK na- TIp/PKEHNC, BHISBANNOE YACTHMHDIM: packpy4sBautH- em, 06b14uN0 B Gonbieit cTeleHH KoMneHcHpyerea cRepxcHupataamuieli, 4em upH pasjenenunt yeneli, nockobKy ckpyynpanne ocu JUTK, Kak npasisio, ‘peGyer Meubille SHeprii, 4eM paspaB BOROPOAIIEIX casei MeKY Hapamit HyKeoriytoB. Onnako creny- eT OTMETHTD, YTO TOCAe YacTHAHORY packpy4HBaHHA ALK in vivo ce wiru serve pasnensuorcs, 470 10380- JACT CUMTbIBATS COMCPAKALLLY ICH B MX HACPOpMalLitto. Kaxkaaa Klerka akriBHo ocyuecTBAReT pac- Kpyuntwanne cBoci JUHK c nomonmio cpepmeutatie- MbIx Uponeccon (onstcaHo HIDKe), H BOSHMKAIOINCe B pesysiTare HalIpsPKEHHOe COCTOAIME CAY KUT [UA Puc, 24-13, Penakemposatian 4 ceepxcnmpanuaosanHan nnasmmanan AHK. Ha xpaiineri newoit anexrpouvor muxpodororpadun rovasaa penakcuporakvaa Monexyna; crertetb Caepxcrupa- smgayjvu Bospactaer cnewa Kanpaso.[08] Yacte TEL 24. fenei xpomocomes 6 Cynepexpywennas JJHK 2 Pacnnerere neneit uc. 26-14, Peayntraret packpyymeanun [HK. a — ywacrox BK samnnyrolt Komyesor monexynit AMMO 84 nL H. B enaxcupozaxnoii dbopme ¢ Bocembio oGopoTanm criMpamnt. 6 — ynanewine onHoro BNTNA BMSbIEAeT HATPAXEHNe CTPYK- Typét. ¢ — O6nHO HaTpaXeHNe CrnanMBaeTcA MPH Cynep- cpympaninn. 2 — vacrwvirioe pacxpysupante [JHK neckonb- xo o6neraer pazsenuvewne uertel Toxasaxo, wr0 Teope- TMMeCKH packpysteaxne Ha ORME OGopoT MoNeT obnerwvAT® pasfenenue yerielt Ha yuacrKe ANMNOM OKoNO 10 n. H. OAKAKO oGsranio paanenentiio KTei a TaKOM KOpOTKOM yuacrKe Mpe- TATCTEVIOT BOROPORNME CRAIN MeMY NapaNn HYKNeOTHAO, wapbexr cravonutca svaNHMHIM TOMSKO AMA Gonee AMMAEENK yvacrxos JJHK # Gonee cunsioro pacxpysneaxin. s3anacania aneprin, Kaerkn nogutepxunaror JLHK B YACTHYHO packpy4eHHOM COCTOAHHH sua Tro, YTO- 601 o6xcrurTp ce KoMMaKTHYI0 yHakoBKy. Packpyyut- panne JJHK tarxe nario jut (byHkuonnpopartits cbepmentor, yyacrayrouyix 8 MeraGomame JIHK, KOTOpHIe JLIA BLITOANEHHs CBOE Zara LONKUBL pasaeaaTb ane natn JLHK. B rakom packpyyerttiom cocrosuu JLHK moxker CYIecTHOBATH JHUND B TOM CALyHAC, CCIM Ola 3aMKHY- Ta B KOLO HIN cBsgata 1 craGHaHpoBalta Gest Kant TaKHM O6pasom, TO er He MoryT CBOGORHO Bpaitarbea oTHocuTeNbIIO ApyT Apyra. Ecant osuta 13 nleneit 6 HsOAUpoBAHHOii, cBoGOAHON OT GenKa Korb nesoii JLHK pasopnatia, 8 stoit TouKe cnoHTaHHO Bos Hukaer coGosHOe BpaleHue H packpyscrniast JJHK HepexomuT B peaakcupowannoe cocrosuue. B_Mo- aieKyae samKHyToii Korbuenott JK sco nirrKon crmipas neaKas HaMenitTD Ges Xora 661 BpeMenttoro paspnina onnoii us neni JTHK. Ho aroit npisnie unics10 BIITKOB crupasm B MoaeKyzIe JLHK spaserea XxapakrepucTitkoii cpepxermpamaargtit Crenene ckpyymeanna AHK onpegenserca Tononormyeckhm napamMeTpom — NOpAAKOM 3ayenneHna ‘Tononorns nperaraer HeKOTOpHIe KOHILCTIIHH, KOTO pole TOMOrYT aM B OGcyAuteNIH cBepxenpacTsaUTH JLHK, 8 sactrocru Kormenniio nopsika sanertennst, Topanor sanenacia — TonoxoriecKoe cBojicTBo siwyxienoveunoit JK, nlockomby oo He HaMenseT- ca pH crnGamut nt sedpopmatut JLHK 20 Tex nop, noKa Yen o6e Her. Uvieno sanentennit o6ositava- wor kak Lk — or aura. linking number (pre. 24-15). Jlapaiite pacemorpum niponece pasnenenna ue- neii AByxttenouewHoii KonbuCROl HK, Eca jise UCM cHagAIIDL TAK, KaK MOKAgAHO Ha pute. 24-15, a, HX UPOYHYIO CBS MOXKHO HasBaTb TOrOOrNNe- cKoii cBasbio. JlaxKe ecm paspyuiaiores BCE BORO- PONE CBAIH HM CTOKHUTOBHIC BaANMOneiicTBHA M@KAY OCHORATIHAMH, TAK YTO Hapymtaerea «usu UeCKUii KONTAKT MEXKILY HETAMH, OLN NO-TIpExHeMy 6 Lk=6 Puc. 24-15. Mopanok sayennenia (Lk). 3nece, Kak o6%KO, xaxnas rony6aa neta o6oatavaer opty 13 nenek nByxue- noweuxoi IK. faa monexynu Ha pic. a Lk = 1, png mone- xynit Ha pwc. 6 Lk-= 6, Onia wa HuTei Ka Puc, 6 uso6panena HesaKpyeHHoi, 4T06m AcKee NoKazaTb rpaniuy Boobpana- eno rosepxnocru (6nenio-rony6oro yeera). Yucno nepe- cevenuii jaHHoii movepxHocTH crupanbHoit HUTS cooTBeT- crayer nopanxy sayennenna.OcTaOTes CBASAIHDIMEL TONOSIOTHMECKHM OGpasoM. Tlpexcrasum ceGe, 4ro oaHa M3 KOMbICBBIX 1e- nei orparuunBacr HeKylo MonepxHocT (Hampi Mep, TaKyl0, Kak MOBepXHOCTh MbUIbHOM MuIcHKH BHYTpHt KOubIA Hepes TEM, KaK 13 Hero BbYLyBAlOT MbuIbuDIi HyabIpb). HopatoK aantennenia MoxkHO onperenttD Kak HCO Tepeceyenstii ITO! NOBEpX- Hocrit Bropoli yeribio. B Momexyse Ha pic. 24-15, a Lk = 1; 8 Monexysie Ha puc, 24-15, 6 Lk = 6. Jas samKnytoit KombueRoli JJHK nopanox sanewieniia Berna oMlcbiBacTes UeTbIM HNeOM. IpuusTo, 40 B ripanoaakpyyentioli cnupain nopanoK aarerite- Hat — HlonoxKiTeNbHOe YHCHO (+), a B JeBOAAKpy- ueioii cpa — oTpnutarebnioe (~). B JLHK or- piutarezpnbtit wopsi0K sauertciuis| we BeTpesactca. Tpumenuat arm paccyayienna « Mosieky.1e 3aM- xnytoii Koapuesoit JIHK, cocrosueii us 2100 n. u. (pc. 24-16, @). Kora Monekyaa penaxcuposatia, HOpalOK sauerseHHa Nairn Mpocto: ITO oTHOMTe- Hino Yer Acex Map ocHOBaNUii K SHeAY Hap OCHO- patti, IpHXOAAUIXcA Ha ON OGopor crtMpatt (apumepuo 10,5 11); cexoBarenbiio, B Aalto cayyae Lk = 200. Uro6b1 kombitevast Monexy.ta JLHK Morsia ObITh XapakrepHsopata orIperteNcHiBIM HO PAKOM 3alleruieHiis, Hit OHA 113 ce Lereli He AOJDKHA cosepxate paspbipon. Ect xorb B ouoii ent ects PASPHIB, CHPA MOXHO HOTMOCTHIO pagent Wa ane erin. B aTOM culy4ae TorlosIOrHMECKHe CBsIaH OT- cyrersyior, 1 Lk nenbsa onperennr (pic. 24-16, 6). Tenleph Mut Moxkem onucath ckpyuennocts TLHK, oucpupysi ismenennes nopsuica saericuita. 3a TOUKY OTCUETA IPHHVIMaloT MlopsoK saNeTCHHA b penaxcnponarnoii JHK, Lky, Jaa Monexysib1 11a pic, 24-16, a, LRo = 200; ect 13 MosteKy.104 YratntTb jpa Burka, Lk = 198. ALK = Lk — Lho ALk = 198 — 200 = -2 (24-1) O6miuno yno6n0 BEIparkaTh HMeHenHe HopsyiKa sa- Henuienit Yepes Mapamerp, He sannicaugit or JUIN unt Moseky.ttt JLHK. Dror napamerp naspinaiot ILIOTHOCTHIO cBepxcHUpamisamun (0), 11H ce (puyeckHM H3MeHeHHeM OpsKa 3allemienns; ILIOTHOCTH CBEpXCTpAUIsABI Panta OTHOMEHILO HSMEHCHIA HCA BUTKOB CHUAN K HX NERY B e- siakeuporannoii JLHK: o-ALk/Lky (24-2) B upuepe nia pic, 24-16, ¢ 0 = 0,01; ato osnta- uacr, wro ynaten 1% (2 us 200) sirrkos crupasiut 24.2 Ceepxcnapanusayuia HK | 1:)] ‘al = went 6 Lk weonperesrenno a De 6 Lk=198 Puc. 24-16. Mlopagok 3ayennenua Ha npwmepe moneKyn aamKHyTon Konbyenoh HK. Konsyecar [JK pasmepom 2100 n. H. noxasania ® Tpex dbopmax: a — penaKcuponarran, Lk = 200; 6 — penaxcuposaxnan c onoueroveuHHin pas- potbon (HnKOM), Lk Kemban onpeneniTs; @ — YaCrno, Ka pa Burka, packpyventian JJHK, Lk = 198. Yacruuno pacxpy- ‘enNan Monexyna OGu'HO cyulecrEyer B cynepcKpyeHHOM cocromiMn, Ho YacruuHoe packpyyMBaHne TAKE MoxeT 06- neruuts pasbenunenna yerteli HK. b JHK (6 B-opme). O6tiun0 crenen» packpysn- Banna Kerounoit JLHK coctasaser 57%; T. ©. 6 or ~0,05 a0 -0,07. Orprnatenpioe snayerine no- KasbinaeT, YTO HSMeHeHHe Mopsa/Ka salenmeHns cBasano ¢ packpyunpainem JUHK. Taxus oGpasom, cBepXCHMpassaltiis, BRISBANE YACTHYHBIM pac- Kpywinaniiem, — 970 orpitiaresbitas cBepxcnnipa- ausaiuta. H, nao6opor, np HeKoToppix O6cTOATEh- crnax JJHK moxker Gurr lepeKpyyenta, 470 BhIparKa- eTCA NONOAUTENBUDIM 3HAYCHBEM CBEpXCHMpAitsa- tun, Orwerim, #70 8 caysae, Kora JK yactHano pactuterena (orpuuatespiias cRepxcrpatsanna), saKpyyeitiad BoKpyr ocH Yacrb cumpaa JLHK sB- AAKCTCA SepKAIBHDIM OTPAKEHHEM. HepeKpy4eHHoii chupan JLHK (nosoxiresibtaa cBepxcnmpam- sauna) (pic. 24-17). Caepxcrmpamsanus He c: uaiivibit mpouece; xapakrep cnepxcnmpactiaanuti 10 Gonbmreit 4actt onmcbipacres nedbopmannueii Kpy4e- 1H, BosnUKaOMelt BH ups yMenpmenn nant YBCAMMCHI HOpsAKa 3aleHIeHHs LO CpaBHEHIIO C B-popmoii LHK.[20] Yacte TIL. 24. Tett w xpomocomet Pesaxenposannas JHK Lk = 200 ame J \ Lk = +2 « tmnt y Y Tlonoxrempnoe -ynrepexpystha ») eynepexpysnmmase Lk = 198 \NY/ Le = 202 Puc, 26-17, Orpnyatensian 4 nonokuTenbHaR cBepx- cnwpanwsausn. [Ina penaxcuposannoit monexyms /IHK, ToKasakHoit Na pwc. 24-16, a, wacrMuHoe pacpyuMaxne wa miepexpyumeaine Ha Ava oGopota crupann (Lk = 198 wax 202) MpMBORMT x OTpHuaTeNbHOm WnM TlonoMMTEMBsOR CBEPXCTIMpANHsayMH, COOTBETCTEeHHO. 3ameTBTe, ¥TO Och cnupann [JHK w 8 Tom 4 8 Apyrom cnyyaax saxpyyMBaeTca 8 MpoTMBononomHwx KampaBnennax. Tlopauox saneiienna moxer usMeusTbca Ha © 1 mpit paspniue oanoii nei JLHK, nonopora on- Horo 43 KOMLIOB Ha 360° BoKpyr Bropoit Her 1 co- cqumenis pasopparnnntx KoroR. Takoe Hancrenite Ne BANSET Ha HiICO Nap HYKACOTHNAOR HLTH HCO aTomon & KobIEBO!T MOaeKy-Ie /LHK. Jkwe «bopatet kompnenoii JIHK, ormvaionmees TosKo TaKuIM ‘TOMOMOTHYCCKHM CBOIICTBOM, Kak HOpAVLOK 3alten.1e- uv, HasbipaloTeA TonONsOMepaMH. ‘Mle [pumep 24-1 ONPEMENEHME NAOTHOCTH CBEPXCNMPANU3ZAUMK Kakowa 1ioriocth cBepxerupasiisailin © aamkty- Toil KosbUCBOI MoreKystbt JLHK samuoii 4200 1. H., Aga KOTOpOit NHOpsZOK saitenstenusn Lk = 374? Ka- Kowa 1mioTHocrb cBepxenupanusaiun JLHK Toit xe guint mp Lk = 412? Coepxnurkn atHx Monexy TONOKITTENTbB HIN OTPHNATEABHLI? Pemenne. Hatinem Lk. Hua toro amuuy Komb- ucsoit HK (8 u. 1) aenus na 10,5 1. 1./snroK: (4200 n. 1.) / (10,5 11. / prox) = 400. Tenepb 10 dhopmy.ne 24-1 moxko air ALE: ALR = Lk — Ly = = 374 — 400 = -26. Tloacrapum anaycins ALR u Lhy B chopmysty 24-2: 0 = ALR/Lky = -26/400 = -0,065. Tlockoasky cpepxcnupamisaiuia orpimarenpas, Mosteysia JHK wecer orpuiarestbttbie cBepxBHTKIL Ecan aaa toi %e Monexyapt JJHK Lk = 41 ALk=412—400= 12, cnenonatespHo,a= 12/400 = 0,03. Caepxcripammsaniia TosloxkKHTeMbHas, —MONIeKy.1a AUK necer nonoxxirrestbubie cnepxpurrict, Lopsiox sauerienis umeer Be etpystypHbie cocraBisiouuie — Kpyyenne (Koadepulen saKpy vennocti; Te — or aura. feist) u paizsmur (usr6; Wr — or aura, writhe) (puc. 24-18). Vix ropaano ‘Tpyaiice onuicaTD, Hem MOpALOK sarterIents, uo kpyveuie Moxkiio paceMarpuBaTb Kak Mepy cKpyaH- Batinst (HCHO BHTKOB) OCH cHpasH, a paiisunr KaK Mepy sloKaIbHOFO HarHGA WIN HpoerparcrBentiolt coasit COceAUMX Hap OcHORATH (4NCIO CuepxmHtT- kon). Ip namenermt nopsaiKa sauennenia Yacrb BostKa‘oMero HtanpaxcrtiiA OGLISHO KoMMeHCHPY- crea MIpH saKpy4unannn (chepxenmpaciisarutt) 1 Narube, 410 abiparkaeres ypantientest: Lk = Tee + Wr Hapamerpur Tie 1 Wr ne oGssarebio nenoune aenipie, Kpyserine 11 paiianir umeior cKopee reo- MeTpHYeckYI0, eM ToHOTOTUMHecKyIO PUpoxy, 110- CKOAbKY H3MensOTeR pu AedpopMatint gamKnyToit Konntesoii Monexyint JUHK. Yactininoe packpyunanne LHK enocaberny- er cnepxcnmpamisanitit 1B oNpeneaeHHo’ mepe DAWA Fpmwas serra (penaxcnponarmas J(HK) Bommuoit murrox. HesHAMITTeTbHBIe HIMeHEHHA CKPYWHBARHS lS SE) Birrox nyzenoro pamuyca, SHAUINTEJIBHBIC HAMeHeHIs B CKPYUHRAHITH Puc. 24-18. Kpy4eHwe w paiisMHr Ha MoqenH pesnHosoro Hypa. Posositi pesmxonisi umryp na0Gpaxaer oc penak- cuporannor monexyms JJHK. Hanpaxexne, nosmuxawmee TPH saKpyyupaun peswrKn (Kak MPH YacTHYOM pacKpy- wma HK), mporenaercr B ape Kpy¥enur WM pait- amura. Tononommeckue WaMeHeHMA TOpAAKa saNenneKHA oGsrHHO COMPOBOMMaIOTCR UIMEHEHWEM KaK KPYYEHUA, TAK upaiisnara.Penaxcmponannas JIHK TINK __ AWW “actso pacxpyseran JIHK KpectooSpasnas erpysrypa JIHK Puc. 2649, O6nervenne o6pasonanna KpecroopasHtix CTpykTyp npH 4actHunom packpy4HeaHnn AHK. Bripusnue, xpectoo6paskwe crpykryps Moryr (popmuposarsca Ha ma- nuNpOMHETX TlocnenoBaTeMBHOCTAX (CM. PMC. 8-19 8 1), HO OxM penko BosHuKaior 8 penaKcupoBaNHoi JJHK, nocKonbKy B nueiuioi [HK pasmemaerca Gonsure nap ocHosanni, ¥em 8 Kpectoo6pasiux crpyxtypax. Yacruuoe pacxpysmeanne TAK o6nersaer pasnenerwe neneit w (boprmposanne xpe- croo6pasvoii crpyxrypst ta coorsercreyouyux yuacrKax 110- cneposarensHocri. ympomact paspenuucune yeueli, a rake oGer~ Yaer paul CYPYETYPHDIX HsMeHeHItit BMOUIEKy.te usnoaornues CTO BEAKO, HO ON HOMOraoT MpoILLMOcTpI- popars HlocaeqeTBHs| YactHuHOrO packpyuitBaniis. Hariomium. sto Kpecroo6pastian erpyKtypa (cM. pic, 8-19 pt. 1) oGnraiTo comepaUT HecKOMTbKO Te- chapeniux ocHonannii, 1 UMeHHO HexocTaToNHaR ckpyueniocrs HK nomoraer coxpansrr pasite- sienue Huei (pie. 24-19). YacruuHoe packpyan- banne upavosaxpysennoll caupam JLHK raroKe oGacriaer OGpasowanne KOpOTKHX OTPEaKOB .eDO- sakpyucnuoii Z-JHK p Tex Mecrax, re tocsten0- BaTeAbHOCTD HYKTCOTHsOB NOBBOAACT peamusonaTh aty cbopmy (ra. 8). Ke sHauelie 9fux HaMeHeHuli ne 24.2 Ceepxcnupannaayua HK [21] Tonouzomepa3bi KaTanu3upytoT v3ameHeHue nopagka 3ayennenna 6 HK Coepxcnmpangaiuia JIHK — crporo peryaupye- Mbtit npoutece, KoTopbiii BAHACT Ha pastible acticKTbI MetaGomaMa JUHIK. B kaokitoii Kaierke cert (pepsert- Thy, (PYUIKIUIA KOTOPLIX 3aKMOYAcTER HCKMOUNITEND- No B YacTHHOM packpysuBaNuIH H/iUI peaaKcaniiH 1 2 3 Csnpxernte L Puc, 24-20. O6HapyMeHne TonoMsoMepos. B stom axcre- pumeure sce Monexynix JJHK conepaar opwHaKosoe weno Tap OCHOBAHUi, HO OTAMYAIOTCR To CTeTIeH CBepxcMMpa- nusayuu. TockonsKy Monexyas cBepxcrupanusosanion THK xonnaxriee, vem penaxcuposanrue moneKyms, OM RBMKYTCA Guicrpee Mp snexrpoopese B rene. Ha mpen- CrapneHkex Penax pasjeneHs ToTOWsOMepH C GANSKMMK aMaNeNMANM TIOTHOCTH cBepxcnMpannaalun (MONeKYRIE npwxyTea caepxy BHu3). Ha RopomKe 1 cHnbKO cBepxcMH- panvaobarnan HK newxetca 8 EME ExMNOM TOnOcH, XOTR, BosMOMNO, B JHK mpucytcreywot pase TonoWsoMepH. Ha nopomvax 2 1 3 suaxet pesynsrams o6pa6orKn cBepx- crupanwaovamion JHK tonousomepazoi tuna I; HK wa nopomxe 3 rionbepranace Bosnelicrsmo hepmenta pombe, vem HK a opoxxe 2. Koraa nnorwocth crepxcnupan- 3auMH CHWOKAETCA NO SHAYEHMA, HANWHA C KOTOPOTO Ten criocobex pasnenat® oTenbHMe ToMOMsOMepHl, CTaHOBATCR BUBNB ANCKPETHME MonocH. B xaxNOM MoNoce, BAENeH- Holi ckoGxoMt crpasa or NOpomKM 3, ConepmaTcR KOBIeBMe monexynst JHK c onumanossm nopaRKOM sanernennn; AIR ABYx COceMHMX Mone MopaAOK sanjenneHua pasnwsaeTcr Ha enunnuy. Penaxeu ponarias — AUHK Cynep- expyuennan JUHK[22] Yacte TIL 24. fenut x xpomocome: (Ocrionannie] 0 Denionarme [Octionaxite] Puc. 24-21. MEXAHM3M PEAKIIMM. Bakrepnanbibie Tononzomepaabi Tana I wameHsior nopa- ok 3ayennenua. Moxasana nocnenosarensiiocrs peaxnit TonoMsoMepaswt Tuma I. CepNeRT MOMET HaXOAUTECA B 3aKDHITOM U oTKpHITO! KONOpMaNHH, a — Nonexyna JJHK caasutBaeTcA C epmenrom B 3axpuror KoN>opmauwu, v onxa yer HK paspesaerca. 6 — depmext nepexonur BoTKpHrTy¥o KONpKauMo, K BTOpaR eM [HK mporunaer Yepes paspere 8 epsom yer. @ — dbepment Haxonwrca 8 3aKpetroli KonopMannu, yer HK poccravasnusaerca.AK. ®epmenr, nonmarongre et nonpKaK0- mute crenenb vacrimmoro pacpywnpanna HK, HagbiBaiores| TONONsOMepagaMtH; OHI SMCHAOT HO- paiaor sanervieniia B JLHK, Oco6enio paxxttyr0 po.tb atu (pepMeHTbr Hrpaior B pouteccax peruuiKaliit ynakonkit JJHK. Cymectayer ana kacca Tou0M30- Mepas. Tononsomepaant I Tina tia Koporkoe nent paspbiaior ony us apyx weneli HK, sare upo- NYCKAIOT LCAYIO Heth CKBOSb PASPLIB MH COCMISOT pasopramnipte Kony; oH HamcnsioT Lk c wraroMt 6 1. Tonongomepasia I tina paspuinaior o6¢ nirrn JLHK Wutamensior Le marom n 2. Teiierane avinx dbepmenton Moxkno mporemont- CTpuponart ¢ HomomtbIo aa1eKTpochopesa B arapostiom ree (pe. 24-20). Fpynma imentianans massive ue 7 ON Seo \ / e Om en Puc. 24-22, BOSMOMHbIA MexAHaM HBMEHEHMA nopAAKa 33- WenneHnA nog AelicTBMeM ayKapHOTHYeckMx TonoMZOMepas tuna IIA. (?) @epmext, cocronmnit 3 Mworux cyGxenneny, cessuipaet ony monexyny HK (creme monocs). Pacps- ‘Tule Monocra Bue H HIOKE CBRaaRHOM /IHK wasereaorca NW C-noporan. ® Bropoii cermer Toit xe monexyms [JAK (xpac- nie ronocts) cananieaercs 8 N-Boporax W oxasiBaerca sartep- ‘THM @, O6e yer repsoro cermenra [JAK pacuerviaiorca (me- X@HNGM TOR peak axanonWieH MexaxMaMy, Ha06panen- HOMY Ha pic. 24-20, 6), u @ sropoit cermextt HK mpoxonnt soa, paspus. ©) Pasopsannaa [HK soccraniasnusaerca, 4 ~ropoii cermenrt JHK stcsoGoxqaerca vepes C-nopota. B xone naHHOTO YMA CBASBIBAIOTCA M TMAPOMMSYIOTCA ABE MOReKYNEE ATP; ont monexyna ATP, nepontHo, rynponvayercn Ka cranvt o6pasonanua omminexca na aTane @. Ocranbine nonpoonocTH rugponusa ATP B annoy peaxuynn enye mpenctomr ycranoBvT. ®@ 24.2 Ceepxcnupanwaayna HK 23) ¢ OfMHAKOBDIM NOpsAAKOM sallerieHiA ABINKETCA B resle B BILe ANcKpeTHBIX HOC. Mero; nosBonseT paavlemer» Tononsomepnl co aitasicmminat Lk, past YaIOUMUAMHICA cero Ha 1, H OGHApYIKUTH HaMeHeHI opayika 3aLlerL: TCH, BHABANTHDTe TOHOMZOMepazaMtt. Kuerk E.coli seit ne metice ueTbipex pa: unix Tontoiomepas (or [210 TV). Tortonaomepaata rina I (ronontaomepasit 11 TH) o6erito penakcn- pylor JIHK, yaasis orpunateautnie enepxpirrait Gionnimaior Le). Mexamsst, ¢ noMomio KoToporo Gaxrepnaninie Tononsomepasi tia I Hamensi- (OT HopaAOK sanenIenus, HoKasan ita pire, 24-21. Baxepuiaztbinnait cbepweirr Tana TT Crononsomepasa tina I, vont JEEK-rupasa) Moxker ppiapatb orprita- Tenbttoe cynepexpyaitnanue (yenbutact Lk), Dt0T chepmenr jut ocymuectmteHnist peak HeHOTDayeT aneprino ATP. Jlasi namencinia nopsiaKa sanenste- unis ZUHK Tononsomepaait Tuna I paspumavr o6¢ nen Moxey JUHK 1 npontyeKaior 8 oGpasonan- tnytoca Spe» apyroii aynaiexe, Crenen enepxcmt- pasianuns Gaxtepranbuoii JLHK nouepaxunacrea Tem peryaaiuint cyssapHoit akrumnocrit Tonon- somepas [it I, B aykapmoriueckux KueTKax TOXKe ecTh TO nonsomepassi THB 1 IL ‘Tononsomepasnt Tn HI — qbepsenro rina I, a cqumersenusiii epwenr Tuna I y noaponounux npucyteTRYeT B ABYX 130- qopmax — Ha 1 TIB. Bomimiicreo Tononsomepas tuna 1, nkuovas JHK-rupasy apxeli, noxoxn 1 upiuayicaar « onuomy cexelicray — x viny TA. Y apxeli, kpome toro, ect neoGurMMe ToHOHAONC- past LV, coctapasuoumte cemelicrso Tha IB. Dy- kapstoTitieckite TouonsoMepaaia THna Hue MoryT wactitinio packpysnnarb JIHK (chopauiponare orpit- MaATeMPUDIe CREPXBUTKH), HO MOTYT pesakenponath kak nlonoxirrenbttog, Tak W OTpiMaTemLtoe cynep- ckpyunpanne (pie. 24-22). Kak Mb YBIUUIM B CzIe;yIONUIX TsraBax, TOTO HGOMepasbI HTpaloT KHOYeRyIO por B MeTAGO-T: we JLHK, novtomy o1it Baxkible MuIUeHH JleKapcre, Tpentiasnauennx LI GopaGn ¢ SaKrepratostoiNt undbexiuiisns 1 Hopoobpasonaniastn (ston, 24-1), Ana komnaktHoi ynakosKu HK tyKHa oco6an (bopma cBepxcnupanuzaynn Y moaexya caepxcmpaanaonannioit HK ects HeeKoNbKO OMX NpHstaKoR, B MoneKyae ¢ OT- putarenbioit cpepxerupaausauueli cBepxeutKu saKpy4entst Bipano (pic. 24-17) st nerue oGpasyior[24] Macro TH 24. Tere « xpomocomet He KOMIAKTHBIE, & TOHKHE H JUIMHHBIe HeTIH, kacTO ¢ miloxecTBOM orReTR:TeHHi (pric. 24-23). Tpw xa- pakrepHoli 11 KIETKH ILIOTHOCTH cBepxcHHpati- SalUHH JUIMLHa CYMePCKPYYCHHBIX Y4ACTKOB C YNETOM ornernnennit coctaniser npumepto 40% suum HK. Taxoii tin crepxermpanisanun Hasiinaercs IuiextoHemuueckuM (oT rpeu. plektos — saxpyyeH- bili 1 nema — wrth). Deus TepMILHOM MOxKHO 060- aHaUntb 2uobyto Crpyktypy ¢ HuTaMU, epexpy¥eH- HBIMH HeEKIM TIpOCTbIM H pery.tipHtbIMt O6pasom, HOH Xopoitio oGtachsier o6uylo eTpyiTypy chepxcrmpa- aunsovannoii JLHK a pactnope. Tlrektonemuueckas crepxcnupamuaanuis, a Mento B TaKoii dopMe BbIjesseTes! HK Bata Goparopunx yoroBUsIX, HeNOCTATOUNO KoMIAKTHA auit yuaxouki [HK 8 kaerke. B apyroii cnepx- Aonomenne 24-1 AISTTTUTEY MevewnesaGonesavni nyren waraGuposanmnronowsowepas Tonoxorns Kxcrounofi JJHK nenocpenctnentio cnstsana ec chynkuneli, Bes nomonu: Tonomomepas KaerEH He Moryr perLtituIporarh H ynakonnwars cHoio JLHK 1 aKe~ peceuporatt reribi 1 ostomy noruGaiwr. Cresionaresth- Ho, MHTHGHTOpEA ToHOHIOMepS MOTyT CTaTb maXKHUINIT hapsanemrimeckusm cperernansn su Gops6o © mi upyson B kavectue armGuotiikos ticuombaytorest awa Kuna ca mirGuropos Gaxrepianinx Tononsomepas. Kysa- HHA, CPEs KOTOPHIX MOXKHO HasmaTh HOHOGHONU H Ky Mepiiui AL, npenctasamor coboit npupoanute mponyn- ‘Th, nbuzezsteMBre 113 cTperrroMuueroR, Ou airnGupywor cosabinanne ATP Gaxtepnansisisnt Touonomepasasit vania I, JUHK-rupasoii 1 rononsomepasoii IV, Ot aura GHOTHKH TONOAKHO PeRKO HeHOMIVIOTCA JWI see muhexiuionnex saGonenaniit enone! acter ripiiax anTnGnorikon Takoro porta jut KaMIME cKOro UpuNeieHs npoxo.vkaeTes. AnrriGuorikn ximonowovoro pasta, KeTopbie TaroKe uncn6upyior Gastepuatsuyio JHK-rnpaay 1 rononso- sepaay TV, nueppite nonuant m MesumuttIcKylo UpakcTHky B 1962 r, Kora Havas NpHMeUsITECA HaMIUKCOBAA KIC ora, 3To weuecrHo oGuaaer orpanuennioit ixpxpexrin- noctoo nn CLIA yake He mpumenscres 8 KntmvecKort Tpaxtinke, OnHAKO AKTHBHOe HKsyHeNHe jtaNMOrO KwrACcA BeMCCT NpHBesIO K HHOsRWIENTIIO pTOpxMONOHOB, MIpHMC- POM KOTOPBIX Moker CoyxTITb UUETIporpOKCaLUM (IUHpO). Aelicroue xmuoronon ocuonano 1a uiruGupowaniit uo- FCHTAMH HSIOKAMCCTHEHI DIME, KICTKANE, dy O/NAKO HOUCK F H,C~ “N7 Ni ay N HC 3 ux) A Lyenpodoxeanuns Hammauxconas xucnora ceqneli craw Tono1omepastioli peakiumt, sakuo¥aIo- meiica n yerpancumt pasptivon p seta JK. Llanpod HOKCAIUH — aNTHGNOTHE mMpoKorO cHeKrpa jeitcraNs ‘Dro oft Ha HeMMOrHx penapatoR, KoTOpElii adxperTHHeH NpoTHe pos6y/urTenel CHONpCKOH s9BLI H paccMarpHBAeT- ca 8 KavecTHe MOMOFO epenteTRa samUITD OF GuonorHC cxoro opyxtis. Xinosionn ACHeTBYKIT HEKHOMETTENDNO. Gaxrepnanbunte TouongoMepaALs, & MTHGUpowanNe yk phoriieckux ¢pepMeirTon Hanuactea upH KoMNeHTpaLutit 1a HeCKOABKO Mops/tKOB HUMIC TepancesTHecKOli NOAH Cent nanGostee Monumx xusMoTepancETiaceKnx fapaTon JU eeHHs O1LKODIOHMECKHX GobIDIX ecb up nurH6uTopl ToHONOMCpas KiexOBeKa. B_onyxo.tcBLAx KuleTKaX OGIO HaGmOAACTeH MOBUAMICHAS KOMUCITTpA- ia ToOMIOMepad, HosTOMy HIpestaparbs, MUFHGUPyIOTUe CH, Mpnorexar CH, ° y7CHs | cH, ‘Tonorexaxcrtupaonoit dbopse, Tak Hasbimaemoii coaenomA- Holi bopme (pic. 24-24), Moker cymecTRoRa’ uacrano packpyvennas HK. B orsumue or Bb Tanytoli mpaposakpysennoli mtextonesumecKoii cope cnepxcnupami su18 comenonsnoii opm XapaKTeplisI IWIOTHBIE sleBOsAKpyweHMDIe HET, ra ctpyktypa Hanomnnaer 10 cope cazonnii uLtanr, akkypaTuo Hamoraumulii a KatyunKy. Xora 24.2 Ceepxcnmpanwzayna QHK [25] TleKToHeMMHECKaA H coneHOHNAS opaut cREpX- CUMNpasH MPHIMTHAABHO OTAMMHAIOTeA NO CTpYK- ‘Type, 06e atH (bopmbt OrpimtarerbHOit cBepxcIt- PAJIIBAIUIN MOrYT BOSHIKHYTH Ha OOHOM U MON IE yuacrke Hactitirio packpyyentoi HK 1 soryr nepexoantb apyr p apyra. 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