r

PALl." An" 0 H H A HE" 0 JI0 r "H. PALl." 0 3 K 0 JI0 r " H

T.36 1996 BblD, 4

,

a!fntk/!f;-fis d ~-t-lCftg flf)/z :sp£ta~1ug pI w;/2:i-ir ¥t.onl-ctmin-ed ..; D!2.- v tv ,+17.. MwLl..'c)atf+r'i/l .. f!A1 IJPP :r:ctJ!lotti

YAK 581.5:57.084.2:575.224.46

© 1996 r. E ..sI.3H6mm;KaH, C.A. repacbKHH, A.A. Y~aJIOBa,E.B. CnHpHH

AHAJI"3 rEHET"qECKUX nOCJIELl.CTBuH3AfPH3HEHUH nOCEBOB03"MOH P}KJf PALl.HOAKTIlBHLIMHBbillALl.EHHHMH

qEPHOEbIJIbCKOH A3C

ITpH H3Y'leHHH reHeTH'IeCKHX nOCJIe)1CTBlifi 3arpSl3HeHHJI noceBOB 03HMOfi P)l(Hpap;HoaKTHBHbIMH Bhma)1eHHSlMH qepHOObIJIhCKOfl A3C B Hall60JIee OCTpylO <Pa3Ypa)1Hal\HOHHoro B03)1eHCTBHJI (BereTal\JWHHblfl nepHoA 1986 r.) YCTaHOBJIeHa3aKoHoMepHaSl H3MeH'IHBOCTh B 3aBHCHMOCTHOT )1031,1 BbIXO)1aaoeppaHTHhIX KJIeTOKH HX HarpY)I(eHHOCTH nOBpe)l(AeHHSlMH. IToKa3aHO, 'ITO CTHMYJ1S1l\IIS1MOp<poMeTpH­'IeCKHX nOKa3aTeJIefi npopOCTKOB, xapaKTepH3YIOll\JlX JlX )l(1I3HeCnoc06HOCTb HapaHHHX 3Tanax OHToreHe3a, HaOJU{))1aeTCSIHa <pOHe )10CTOBepHoro pOCTa l\HTO­reHeTJI'IeCKHX HapymeHHfI ..

CfepHo6blAbCKaR aBapWl, paouoaKmUBHoe 3a2pR3HeHue, a6eppau,uu XpOMOCOM, 03UMaJl

pOJICb, ceAbCKOX03Rucm8eHHbie pacmeHWl.

ABapJUt Ha l..{cpHo6blJIbcKml A3C He HMeeT aHaJIOrOB KaK no I1JIOll\a,[(H pa,[(HoaKTHBHoro

3arpJl3HCmUI, TaK Ii 110 BeJIH'UmC nOrJIOll\eHHhIX 06'beKTaMH )KHBOH nplipO,[(bI A03. Bl.{epHo6bIJIbCKOM Bbl6poce npHCYTCTBOBaJI 60JIee IIlHpOKliH cneKTp paAHOHYKJIliAOB no

cpaBHeHHIO C aBapHeH 1957 r. Ha IO)!(HOM YpaJIe. 3Ha'lHTeJIbHbIH BKJIa,[( B aKTIIBHoCT!,BbmaAcHIiH BHeCJIH 131}, 137CS, 134CS, 90Sr, 238+239pu, 240pU, KOTopbIe, 6JIaro,[(apJl CBoeH

BbICOKOH nO,[(BH)KHOCTH H BblCOK03HeprCTR'l.CCKOMY cneKTPY R3JIYQeHRH npe,!!,CTaBmnOTHall60JIce onaCHOCTb ,[(JIJI paCTeHllH II )!(IlBOTHbIX. CJIO)!(HaJl 3aBllCllMOCTb OT BpCMeHH

paAHOHYKJIR,[(Horo COCTaBa BbI6poca H H3MeH'IHBbIe MeTeOpOJIOrll'l.eCKHe YCJIOBIDI B Te'leH~e10 cYT aKTllBHoU 3MllCCRR npllBemr K 'Ipe3BbI'IaiiHO HeroMoreHHoH KapTllHe Bbma,[(eHHR.

HeO)\HOpOAHoe pacnpeAeJIeHRe paAHoHYKJIH,[(OB 06YCJIOBHJIO 60JIblllOH pa36poc ,[(030BblX

HarpY30K Ha 06'beKTbl )KllBOH npHpo,[(bI, KOTOpbIH B p}l,[(e CJIY'l.aeB ,[(a)!(e Ha He60JIbIllHXY'IaCTKaX BblXO~llJI 3a npe~eJIbI nOpJl~Ka.

HaH601IbwCMY pa,[(llal\HOHHoMY B03AeHcTBHIO nOI(BeprJIllCb npHpo,[(HbIe R arpapHbIe

3KOJIorH'teCKHe CHCTeMbI B 30-KHJIOMeTpOBoH 30He KOHTpOJI}I aBapllH »a l.{A3C, Ha3Ha'HITCJlbHOH 'I.aCTH KOTopoH pa,!!,HoaKTHBHoe 3arpJl3HeHRe B 1986 r. 1(0CTHrJIO HecKolIbKRX

TblCJI'I KWKM2. l.{epHo6bIJlbCKaJl aBapIDI npOH30111JIa B KOHl\e anpeJIJI - ncpno,!!, YCKopewlOro

pOCTa II <pOpMHpOBaHHH penpOAYKTnBHbIX opraHOB, KOrl(a paCTRTeJIbHble co05ll\eCTB3Han60JIee pa~RO'lYBCTB:l!TelIbHbI. MaKCRMaJIbHOe pa~HaL\HOHHoe B03,[(eHCTBHe Ha 06'beKTbl

)!(HBOH npHpOAbl npHlllJIOCb Ha nepno~ oCTporo o61IY'leHRJI (10-20 CYT C MOMeHTa aBapKH),

KOfl~a 3Ha'IKTeJIbHbIH BKJIa,[( B nOrJIoI.l\eHHyIO A03Y BHOCllJIH KOPOTKO)KHBYll\He pa,[(HO­HYKJIHAbI. BTopa}l <pa3a BKnIO'IaeT JIeTO H paHHIOIO oceHb 1986 r., B Te'leHRe KOTOpbIX

MOll\HOCTb A03bl Ha nOBepxHocTH nO'IBbl ynaJIa ,[(0 20-25% OT nepBOHa'laJIbHOH BeJIH"IHHbI

[IJ. B pa60Te [2] 6blJIa ~aHa npe,[(BapHTeJIbHaJi llH<popMal\HJI 0 reHeTH'IeCKHX .II 6HOJIO­

rH'IeCKHX nOCJIe,[(CTBJUtX3arpJl3HeHIDI nOCCBOB 03HMOH P)KH pa,[(RoaKTHBHbIMH Bbmal(eH1UI:MH

498

lfepH06blJIbCKOH A3C B 1986 r. Terrepb, Korga C MOMeHTa '-IepHo6blJIbCKm'i KaTaCTpo<pbl

npOlllJIO 60JIee 10 JIeT, oc06eHHO o'-leBllgeH YHHKaJlbHbIH xapaKTep ):\aHHbIX, rrOJIY'-IeHHbIX B

nepBYlO, Hall60JIee OCTpYIO cpa3Y pa):\Ha~llOHHoro B03):\eikTBHH, '-ITO rro6ygHJIO Hac BHOBbBepHYTbCH K aHami3Y rrOJIY'-IeHHOrO 3KcrrepHMeHTaJlbHOrO MaTepHaJla.

MATEPliAJILI Ii METOJJ:liKA

CeMeHa 03llMOH p:>Kll paHOHllpOBaHHoro COpTa BeJITa co6paHbi 4-6 aBrYCTa 1986 r. B 30­

KllJIOMeTpoBoH 30He aBapllll Ha l.J:A3C C Y'-IaCTKOB rrJIoru,agblO 5 M2 rrHTll pa3JIll'lHblX rroMOru,HOCTll 3KCrr03ll~JfOHHOH ):\03bl Y-Jf3JIY'-IeHHH rrOJIeH C 50-100 rJIaBHbIX KOJlOCKOB Ha

Y'-IaCTOK; KOHTpOJleM CJlY:>KllJIll ceMeHa H3 3epHOXpaHHJIllru,a . .D:JlHorrpe):\eJleHHH cYMMapHm1nJIOTHOCTa pagaoaKTllBHoro 3arpH3HeHHH II rrOrJIoru,eHHblx paCTeHJUIMH g03 B Mae, lllOHe H

aBrycTe 1986 r. Ha Ka:>K):\OMll3 rrHTll 3KcrrepllMeHTaJIbHbiX rrOJIeH C Y'-IaCTKOB rrJIoru,a):\blO

5 M2 MeTO):\OM KOHBepTa oT6llpaJIll rrp06bl nO'lBbl C nocJIegYIOru,eH y-crreKTpoMeTplleH

06pa31.\0B nOBepxHocTHoro CJIOH (0-2 CM) B 5-KpaTHOH rrOBepXHOCTll, K03cpcpll~lleHTBapllal.\llll - 30-40%. :

.D:JIHI.\llToreHeTll'-leCKOrO aHaJIll3a ceMeHa rrpoparu,llBaJIH B '-IallIKaX ITeTpll B TepMOCTaTerrpll 25-26° C, cpllKCal.\lllO npopocTKOB ocyru,ecTBJIHJIll B a~eTOaJIKOrOJIe (1 : 3) Ka)!():1ble 2 '-I

C 16-ro rro 24-H '-I, Ha 40-H, 48-H II n-H '-I rrpoparu,HBaHllH; Ha Ka:>KgylO TO'-lKY 6paJIa 10-15

pacTeHllH. OgHOBpeMeHHo ll3MepHJIll gJIllHY rJIaBHbIX KopelliKoB rrpopocTKOB. BCXO:>KeCTb

ceMHH rrepBoro roga xpaHeHHH B Jla60paTopHbiX YCJIOBmIX orrpegeJIHJlll rro YCKopeHHoMY

MeTogy [3]. MllKpOCKOIIJf'-leCKJfft aHaJIJf3 npoBogllJIll Ha arrllKaJIbHblX KopHeBblX MepllCTeMax

Ha gaBJIeHblX rrpenapaTax, oKpallleHHblx al.\eTOOpcellHoM. B Ka:>KgoM rrperrapaTe rrpOCMa­TpllBaJIJf 1000 KJIeTOK gJI5I Ol.\eHKH MllTOTH'-IeCKOrO llHgeKca, Bce aHa- II TeJIocpa3bI gJI5I

OnpegeneHll5I 'lHCna KJleTOK C a6eppal.\IUIMH XpOMOCOM; aHaJIH3 crreKTpa a6eppal.\llH rrpOBO­gHnH no KnaccllcpllKal.\llll [4] C BblgeneHHeM xpoMaTllgHblx (OgHHO'lHbIX) H XpOMOCOMHbIX

(ABOftHbIX) MOCTOB H cpparMeHToB, a TaK:>Ke OTCTaBaHHH XpOMOCOM (reHoMHble MYTaI.\Hll).

ITPJf Ol.\eHKe OTCTaBaHJfH Y'-IHTblBaJlJf XpOMOCOMbl, Jle:>Kall.\He oTgeJIbHO OT pa301lleglllHxcH

aHacpa3HbIx llIarrOK Ha paCCTOHHJfH, He MeHee '-IeM B gBa pa3a npeBbllllalOlI.\eM TOJIlI.\llHYXPOMOCOMbI.

ITpJf aHaJllf3e 3KCrrepJfMeHTaJIbHbiX gaHHblx rrpHMeHHJIJf MeTogllKY CTaTllCTJf'-leCKOrO aHa­

JIH3a 3MnllpJf'-leCKJfX pacnpeAeJIeHJfft [5], rr03BOJIHIOru,ylO, B '-IaCTHOCTll, onpegeJIllTb OnTJf­MaJIbHblH 06"beM Bb16oPKll, HeooxogHMblH gJIH nOJIY'IeHll5l Ol.\eHOK ll3Y'-IaeMblX napaMeTpoB C

¢llKCHpoBaHHoH OTHOCHTeJIbHoft BepoHTHoH norpeIIlHocTblO rrpll 3agaHHoH gOBepHTeJlbHoHBepOJITHOCTH.

PaC'-IeT g030BbiX HarpY30K OCHOBblBaJICH Ha gaHHblx 0 paglloHYKJlll):\HOM COCTaBe Bbma):\e­

HHH H npH6JIIDKeHHft, OrrJfCbIBalOUJ,JfXpOCT 3JIaKOBbiX paCTeHllH H MHrpa1.\H1O pagHoHYKJlll):\OB

rro rrpocpHJIIO nO'-lBbl. Ol.\eHKY nOrJIoru,eHHoH g03bl 'Y-ll3J1Y'-IeHll5I rrOJlY'-IaJlll rrYTeM CYMMHPO­

BaHH5I BKJlagoB OT Bcex JIJfHJfH crreKTpa, HCXOgSi H3 npegnoJlO:>KeHHH 0 paBHoMepHoM

pacnpegeJIeHJfJf pagJfOHYKJIHgoB no rrpocpJfJIIO nO'-lBbl Ha rJIyoHHY 1 CM (TaKoe 3arJIy6J1eHHe

HCrrOJlb30BaJlll, 'lTOOM Y'-IeCTb 3KpaHHpoBKY Y-Jf3JIY'IeHllH MHKpOpeJIbecpoM nO'-lBbl) . .D:JISI

paC'IeTa nOrJIoru,eHHoH g03bl ~-H3JIY'leHHH C'-IHTaJIH, '1TO '-IaCTb aKTllBHOCTll pacnpegeJIeHa

rro nOBepxHoCTH nO'-lBbI, gpyra51- paBHoMepHo B CJIoe nO'lBbl 1 CM. YrJlOBOe pacnpegeJleHHe

~-H3JIY'-IeHH5I nOBepXHOCTHoro H TOJlCTOCJIOHHOrO HCTO'-lHHKOB rrpllHHMamf KOCHHycoHgaJlb­

HbIM: B nepBOM CJIY'-Iae gJIH Y'-IeTa 3KpaHHpOBaHllH ~-H3JIY'-IeHHH MHKpOpeJlbe¢OM nO'-lBbl, BOBTOpOM - H3-3a 6J1H30CTJf KOCHHycongaJlbHOrO 3aKOHa K 3aKOHY gHCPCPY3Horo pacrrpegeJleHIUI

~-H3J1Y'leHIUI TOJlCTOCJlOHHOrO HCTO'lHHKa. K03cp¢llI.\HeHT 3KpaHHpOBaHllH P-ll3J1Y'-IeHHH

paCTHTeJlbHblMll TKaHHMll, nOKpbIBalOII.\HMll KpHTH'IeCKllH opraH, 6bIJI rrpllHHT paBHblM 0,5.

ITorJloru,eHHble g03bl P-H3J1Y'-IeHH5I paCC'lHTbIBaJlH HHTerpllpoBaHHeM g030BOH CPYHKl.\llHOCJlaOJleHHH TO'-le'lHOrO HCTO'-lHHKa gJlH Ka:>Kgoro pagHoHYKJlllga no Y'-IaCTKaM nO'-lBbl

He3KpaHHpoBaHHbIx H ogHoKpaTHo 3KpaHHpoBaHHbix cTe6J1HMH pacTeHllH no cxeMe,

rrogp06Ho H3JIO:>KeHHOH B pa60Te [6] . .a;o30Bble HarpY3KH Ha pacTeHHH, paCC'lllTaHHble no

npegJIO:>KeHHOH cxeMe, MorYT OTJIH'-IaTbCSI OT peaJIbHblX B OCHOBHOM H3-3a OTCYTCTBHH

HHcpopMal.\HH 0 nepepacnpegeJleHHH BO BpeMeHH pagHoHYKJlHgoB B rroBepxHocTHOM CJIoenO'-lBbl.

499

&:"'fL?U)fcftlcrd oIz£vllele/lt'y//t!- fi;r g~NPwz-ftdl tfC?CP..-$.

~ _ 1f,jerC{,~(SUmmaf'im!)r::UnS;t-# o!- soil!- e£J1u-CL11Una-i-M 1/2

iaye.-r- 0-;). c-m ) kit/yon 2/ Dr! Dp; .~D '. ~Y\ [J ArJ d fo . f}' la-Gte -I':J' 4r - aoSOI '"OA/U -oS£. r fV!I.ff (:at jk'Ctftl o;;tJal1~T a 6 n H ~ a~

40r 1I~+aI-(OI2 p-fN Dct) ti;y ([jrLlf)Paa~O/102U'leCKaJ/ xapaKmepucmuKa 3KcnepUMehma/lbllbiX y'wCntKOB

:: ~:J.iI-l/llinM 3KcncpHMcHTanbHbIH I CJI

DrIDpIDr>+r

peCleE.

Y'IaCTOK

v UlftV))KO)f{YIllKH kozhvshkJ

5300,181,151,33M /1L( '--:::Jt.{aMKOB Ciilllnko.'"

15700,332.83,13PaAHH

Rail",16500,583,44,00

EOpll\CBKa vor'SM vi; (L 35001,2 '.6,57,7

KpacHo

kr-ci> ho71301,1510,812,0

flpUMe'lallue. CJ- cYMMapHa!l nnOTHOCTb 3arpSl3HcHHSI nO'lBbI B CJloe 0--2 CM, KH/KM2; D"(> Dp.

Dl>+r - nornOll\CHHbIe KpHTH'ICCKHMH opraHaMH pacTeHHH A03bI 3a BereTal.\HOHHblH nepHoA, fp.

PE3YJIbTATLI H OECYJKAEHHE

B Ta6JI. 1 AaHa paAlIOJIOrHqeCKaSi xapaKTepHcTHKa :3KCnepHMCHTaJIbHbIX YlIacTKoa Ha

MOMCHT c60pa CCMSlH. BRAHO, liTO TaKHC 6a30BhIC xapaKTcpHCTRKH, KaK CYMMapHaSi

nJIOTHOCTb paAHoaKTHBHoro 3arpH3HCHHSI nOBepXHOCTHoro CJIOSInOllBbl H nOrJIO~eHHaSi 3a

BereTal\HOHHbIH nepHoA KpHTHqeCKHMH opraHaMH paCTeHHH A03a, pa3JIHlIalOTCH nO'lTH ua

nopHAOK. COrJIaCHO KJIaCCHCpHKal\HH 30H XpOHH'leCKOrO AeHCTBHH HOHH3HPYIOlUero

R3JIY'leHlli! Ha npHpOAHble nonYJIHl\HH [7], no KpaHHcH Mepe Ha ABYX 3KcnepHMeHTanbHbix

Y'IaCTKaX, xapaKTepH3YlOIl1,HXCH HaH60JIbUlHMH ypOBHHMI\ paAHoaKTliBHoro 3arpH3HeHlil!,

HHAYl\HpOBaHHhle H3JIY'IeHHeM 6HOJIOrH'IeCKlIe 3¢¢eKTbl AOJDKHbl AOMHHHpOBaTb HaA

AeiicTBHeM APyrHx 3KOJIOrH'leCKHX ¢aKTopoB. BKJIaA ~-H3JIY'IeHHSI B cYMMapHYlO

nomo~cHHYIO A03Y pa3JIH'IeH AJIH Ka)f{AOro Y'IaCTKa, HO BO Bcex CJIY'IaHX B 6-9 pa3 BblIlle

BKJIaAa Y-H3JIY'leHlli!, '1TO XOpOlllO corJIacyeTcSI C pe3YJIbTaTaMH paC'IeTa A03 B onblTax no

3arpH3HeHHIO noceBOB CeJIbCKOX03S1ikTBeHHbIX KYJlbTYP npOAYKTaMH SI/J,cpHoro /J,eJIeHHH [8]

H nOATBep)I<J~aeT CAeJIaHHblH B XOAC aHaJIH3a nOCJIeACTBHH KbllllTblMCKOlI aBapnH BblBOA 0

TOM, '1TO CYMMapHble nOrJIO~eHHble paCTeHHHMH A03bI npH 3arpSl3HCHHH cpeAbl pa/J,HO­

aKTHBHblMH Bhma)J,CHHSlMH MorYT B 10 H 60JIee pa3 npeBbiUlaTb A03Y BHCIllHero ''I-06JIY­'1CHHH [9].

B Ta6JI. 2 npeACTaBJIeHbl PC3YJlbTaTbll\HTOreHeTH'leCKOrO aHaJI1I3a KOpHCBblX MepHCTCM

npopocTKOB CCMSlH 03HMOH p)f{H, c06paHHblx Ha Y'IaCTKaX C pa3HbiM ypOBHCM

paAHoaKTHBHoro 3arpH3HcHHH. COnOCTaBJICHHe '1aCTOTbl BblXOAa a6eppal\HH Ha 16-24-ft,40-48-H H 72-H 'I npopall\HBaHHH CBHACTCJIbCTBYCT 0 BblCOKOH 3¢¢eKTHBHoCTH npol\cCCOB

nOCTpaAHal\HOHHoro BOCCTaHOBJIeHHH, nOCKOJlbKY /J,OCToBcpHble reHeTH'leCKHe 3qJ¢eKTbl

paAHal\HOHHOro B03)J,eHCTBHH Ha6mo/J,alOTCJI rJlaBHblM 06pa30M npH npoxo~eHHH KJICTOK

'1epe3 nepBblH MHT03 (16-24 'I npopa~HBaHnH). nOJIY'IeHHble pe3YJIbTaTbi OAH03Ha'!1I0cBH/J,eTCJIbCTBYK>T B nOJIb3Y paAHal.\HoHHoH npHpo)J,bl Ha6JIK>AaCMblx a nOKOJleHHH M2

H3MeHeHHH. CTaTHCTH'IeCKHH aHaJtH3 npcACTaBJICHHhlX B Ta6JI. 2 /J,aHHbIX cBHAeTCJIbCTBycT

o TOM, '1TO Y)f{e nOrJlOll\eHHaH paCTCHHHMH ]l,03a 3,1 rp npHBoAHT K p,OCTOBCpHOMYYBeJIH'ICHHIO BblXOAa a6eppaHTHbix KJIeTOK. YMecTHo HanOMHHTb, <ITO B HalllHX

3KcnepHMeHTax no H3yqeHHIO HHAYKI.\HH l\HTOrCHeTnqeCKHX 3¢¢eKTOB B KOpHCBOH

MCpHCTCMC APyrHx CCJlbCKOX03j/iicTBeHHhIX KYJIbryp [10, 11] AOCTOBcpHoe YBeJIHqCHHe'laCTOTbl a6cppaHTHbix KJICTOK Ha6JIIO/J,aJIOCb npH conOCTaBHMblX A03ax oCTporo "1­

~~~re~ ..Ha pHcYHKe npCACTaBJleHa 3aBHCHMOCTb BblXOAa l\HTorCHeTH'IeCKHX nOBpe)f{)J,cHHH B

nepBoM MHT03e OT nOfJIOll\eHHoH 3a BereTal\HlO A03hl 06JlY'IeHIDI. OT'!eTJIHBO npeACTaBJlSISI

06YCJIOBJICHHYIO o6DeKTHBHbIMH npH'IHHaMH B03MO)KH0CTb cy~eCTBeHHmi norpcllIHOCTH npHOl\CHKC nOrJIOIl\CHHblX paCTeHHHMH A03, CJIeAycT Bce )f{e OTMeTHTb HcmlHeHHblH xapaKTep

nOJIyqCHHOH 3aBHCHMOCTH H HaJIH'IHC ABYX A030BbiX ]l,Hana30HOB, B npeACJIaX KOTOpblX

BblXOA a6eppaHTHblX KlICTOK He YBeJIH'IHBaeTCSI. AHaJIOI'H'IHaH no <PoPMC ]l,030BaH 3aBHCH-

500

·-rr ...A j,~ p _ 0 ~/~, .,ik ~c£vICJl u't v V·U{JL rf ¥offM1-fi(:()Ifr VUl.17VJ~ . (iL ('ODf- ,

fnf)'t(l si-un uptg .cifr e(;- cJ. '1- hour s) /AI j M-ft ft.1 e ~S-P-P- d fj ns s ~ dwelvp I rtf­CIk \ th/'DYllL!4£ irmdiQ)/,of'2) tf'Orf}. a.ocSof\-B.uL do.s.-e du,!'-t)w tJ.e.ff-iWCJ>

p~'ool .if

{ '1.••...••..

.••....•..

). .••.../ ,...

T-v/ -rGaL2

-fNJ( l££ V1~ D~

{1U':t rcuvi­u.lli .j)i/'cW

/f

/}/!'D 16

12

8

1" 618 10

G~ OX dos~ £jf1... I I12

••••

3aBilCHMOCTb BblXOAa wrroreHe1"H'!eCKHX nOBpe)!(]:\eHllli B KopHeBoli MepllCTeMe 16--24-'IacoBbIX npOpOCTKOB ceMJUJ

OJlIMoli P)!(Jj, cct>opMHpOBaHHbIX B YCilOBIIJIXxpoHII'leCKOrO 0611Y'1eHIIJI, OT nOrnOll\eHHOli 3a BereT3l1.II0HHbni nepllonA(Db1

no OCII a5cL\Hcc - nor1l0ll\eHHaJI A03a, rp; no OCII 0p/lllHaT - 'IaCTOTa a6eppaHTHblX KJIeTOK, %

BeHHbIMH H CJIO)!(HbIMH "OBpe~eHWlMH. B AaJIbHeHllIeM aHaJIH3HpYIOTC~ TOJIbKO Te a6ep-

paL\HH, KOTopble YAaJIOCb HAeHTH<pHL\HpOBaTb, KJIeTKH co CJIo>i<HbIMH, He nOMalOll.\HMHCjI &.l 'pacn03HaBaHHIO nOBpe>KAeHWlMH, a TaK)!(e C MHKpOHApaMH H3 aHaJIH3a 6blJIH HCKnlOqeHbl.~ lis. W,I~5ICHO, '1TO "PH 3TOM OL\eHKa '1HCJ1anOBpe)!(AeHHB Ha KJIeTKY SlBJUleTCSI3aHH)!(eHHoH. AHaJIR3 e()rr~ C'.£\-I-U,;"t..­

H3MeHeHWI HarpY>KeHHOCTH B 3aBHCHMOCTH OT nOrJIOll.\eHHoH A03bl CBHAeTeJIbCTByeT 0 ct.Wrc~;hons and.---

AOCTOBepHOM pOCTe nOBpe)!'1~eHHOCTH KJIeTOK C YBeJIH'IeHHeM CTeneHH paAHaI.\flO/lHOrO Wt,\(:.MJYUIUe)

B03AeHcTBWI. 06pal.L\aeT Ha ce6S1 BHHMaHHe CYll.\eCTBeHHoe (HO CTaTHcTH'IeCKH HeAOCTO- lNer.e N.YYWV.t.oL-

BepHoe) CHH)!(eHHe nOBpe~eHHOCTH KJIeTOK "PH A03e 1,3 [p. B L\eJIOM >Ke 3HaqeHWI :noro IAV--''''' ~,'.,.I v' '1 Ct-t1 C!,.().J.-Y/ ~ .nOKa3aTeJIH He BblXOASlT 3a npeAeJIbl, KOTopble Mbl Ha6nlOAaJUi B 3KcnepHMeHTax C OCTPbIM \ Q ~.

Y-06JIyqeHHeM CeMjlH cenbcKOX03j1HCTBeHHbIX KYJIbTyp [to, I I].AHaJIH3 HHAYL\HpyeMoro B YCJIOBWIX paAHoaKTHBHoro 3arp~3HeHWI MeCTHOCTH cneKTPa

a6eppaL\HH H 3aKoHoMepHocTeif ero H3MeHeHIDI B 3aBHCHMOCTH OT nornoll.\eHHOH: KPHTH­

QeCKHMH opraHaMH paCTeHHH A03bl "MeeT Ba>KHOe 3HaQeHHe An~ nOHHMaHIDI MoneKynSlpHblX

MexaHH3MOB cpOpMHpOBaHlli! paAHaL\HOHHbIX nOBpe>KAemni H OL\eHKH BKlIaAa HOHH3HpYIOJ.l.\lfX

H3JIYQeHHH B Ha6JIIOAaeMYlO peaKL\H1O reHeTHQeCKHX TeCT-CHCTeM, B Ta6n. 4 npeACTaBlIeHbl

AaHHbie 0 QHCJ1eCTPYKT)'pHbIX MYTaL\HH onpeAeJIellHblX THnOB H HX OTHOCHTeJIbHOH '1aCTOTe

D' '., B ~6ll.\eM cneKTPe MYTaL\HH B 3aBHCHMOCTH OT A03bl. CTaTHCTH'IeCKHH aHaJIH3 nOKa3aJI, '!TOI >-fN tUA-IOYl. 0t Cia-mC/"&!6 ( aU-erct-h'Dl15 -: ) p-eo C.e1-t cvn..oL. VCL.UleS Tct eft ~.

~ vLQ)J)~ for CLUtrcfA'l;f- c.ell3 i oLe-fL-e. n dA ~ fr'o m T a 6 JI H U a 3I rCtcllct-+\'O\'1 ctOg..e_ c ( Ib - :)llp +1)( er-no Y1 ')

PacnpeOe.lleHUe '/uc.lla n06pe:JKOeHUUHa K./IemKY U 311ll'/eHUR Tl06pe:JKOeHHOCmua6eppaHmHbix 6-f'd.e1ZK./IemOK 6 3a6UCUMocmu om 003bl o6.11y'/eHUR (16-24 '/ IjJUKCaI~UU) ..__..--... ~ II l-t I L'" I /J1Zll:,/ r·e-l.ll.n I- e. u.. S /

1,34 ± 0,11 32,51,15 ± 0,10 12,11,35 ± 0,07 36,61,57 ± 0,19 39,21,40 ± 0,13 40,81,59 ± 0,12 54,7)

npUMe'jaH,u~, MTI,,- }tOlIJi KneTOK C MHO)l(eCTBeHHblMH H CJJO)l(HblMH nOBpe)i(,!\eHHJlMH'I!/

~1,fi.1n, . L (!"~ fJ:/J.r. I. j502 0J~l MP - g0b- V.f- utU w;'ih ru1 Jne.flOUS af1-o . vu' -u.tA.-t-.L.tAL-cUI rna~~ aJ-tu"'a- fron s)

'-lHCJIO atSeppal.\HH Ha KneTKY

--A moUnJ of c.fM-e fClA1\On~ {)-Q('- uUDDS..t A03a,

f rp

~~ _/• ~o

A~ ;- '-QJ110itil -/;- 1,3,\ 3,1

vI cr Uaa tvt- 4,0

utf!.!> 7.712,0

._'\IA .( K

~

4033

1015]4995

2729643129

43

51

23

119

30

4o3245

oo1

1

24

ooIIo2

43955

11

HarpY>KeH­HOCTb,

OTH.e}t •

Uh-i£!1;- )(.uu..,-

MTI,%

kP

~Yct'P -4.J;'/J-ftk 01 s-tl'll roJural1lUlfrUJ ons

\Jv~ ~m rcwlial-/oYl d{)S-fL,~rY-

()(xp.end i YLf

Tu/.-w t; .Ta6nlfl1a 4

XapaKmepucmuKu cneKmpa cmpYKmypHblX Myma4uu 13 3aeUCUMocmu om 003bl 06J1Y'JeHuR

,[J,03a,rp06~eeqlfcno a6eppaL\lfll pa3HbIx TlfnOB

DOS2, fi#

'1HCnOfimotervt c::iCiJP.rf..t eel ion sa6ep- &1 r;{i~· -bfp.€ s./ paL\lfll

i i i

~

gf'

Im"IgIf m' Ifnm"

jerklt 0

49527 13I310,255,126,52,06,1

wmoUil{..

1.332419 90 0.. 12,559,428,10,00,0

t13,1127962415107,148,832,33,97,9

4,0671331180 519,145,626,50,07,4

(t,~t-em+I'o liS7,766734220310,450,732,80,04,5

12,01441959554 713,241,038,22,84,9

llpuMe'JaHue. g - OTCTaBaHHSI XpOMOCOM (reHoMHbIC MYTaL\HIf); f, m' - xpoMaTIfAHbI~

(OAHHO'lHbIe) <pparMcHTbI H MOCTbI C conYTcTBYIO~HMH <pparMeHTaMH; f", m" - XpOMOcoMHbIe(ABollHbIe) <pparMeHTbI H MOCTbI C conYTCTBYIO~IfMH <pparMeHTaMIf.

npaKTll'leCKll He Ha6nlOAaeTC~ ll3MeHeHll~ OTHOCHTenbHoro crreKTpa MYTal(HH B 3aBHCH­

MOCTH OT A03bI. Bll,f\HO, 'ITO rrpe06naAalOIl.\HM THrrOM l(llTOreHeTll'leCKllX HapyweHllH y

ll3Y'IeHHOrO copTa P)Kll B HopMe II rrpH XpOHll'leCKOM 06JIY'leHHH BO BceM AHarra30He A03

HBJI~lOTC~ xpOMaTllAHble a6eppaQHll, 3aTeM CJIeAYIOT OTCTaBaHH~ XpOMOCOM H XpOMO­

COMHbIe a6eppaQllH. HH3KHH Y,f\eJIbHbIH BKJIaA XpOMOCOMHblX a6eppal\HH, ~BJI~IOIl.\HXCH

MapKepaMH pa].\Hal.\HOHHOro B03J1eHCTBIHI. Ha cpOHe CraTHCTH'IeCKH J10CTOBepHoro YBeJIH­

'1eHll~ 06~ero Bblxo].\a l.\IlTOreHeTll<leCKHX rrOBpe)KAeHllH C pOCTOM nOrJIOll.\eHHOH paCTe­

HIDIMH ,f\03bl 06YCJIOBJIeH, no HaweMY MHeHHlO, oc06eHHOCT~MH ll3Y'IaeMOrO 06'beKTa.

nOCKOJIbKY B npOBe].\eHHbiX paHee [10, 11] HCCJIe].\OBaHll~X cneKTpa CTpYKTYPHblX MYTal.\HH B

KopHeBoH MepHcTeMe 06JIY'IeHHbIX ceMSIH nWeHHl.\bl H SI'IMeHSI BKJIa].\ XpOMOCOMHblXa6eppal.\HH TaK)Ke 6blJI BeCbMa He60JIbWIfM. TaKoH HeO)KHAaHHblH pe3ynbTaT 06YCJIOBJIeH

B03MO)KHOCTblO HHJ1YKl.\llH B cpa3e G1 ,f\JIIfTeJIbHO )!(IfBYIl.\llX nOTeHl.\lfaJIbHblX nOBpe)!(].\eHHH,

peaJIH3YlOll.\HXC~ 3aTeM B cpa3ax S II G2, '1TO If orrpeAeJI~eT rrpe06JIa].\aHlfe xpOMaTIfAHblX

MYTal.\HH HaJ1XPOMOCOMHblMH. T.e. B03AeHCTBlle llOHH3HpYlOll.\erO H3JIY'leHIDI Ha XpOMOCOMY

Aa)Ke B COCTOSlHHH OAHOH 3cpcpeKTHBHOH HHTIf (cpa3a G I) B 3aBllClfMoCTIf OT YCJIOBIflI, B

KOTopble rrorra,f\aeT rrorrymll.\lf~ 3apOAbiweBbIX KJIeTOK B MOMeHT 06JIY'leHll~, cnoc06HO

BbI3bIBaTb He TOJIbKO O)KIf].\aeMble rrepeCTpOHKH XpOMOCOMHoro THna, HO H onpeAeJIeHHOe

'lHCJIO xpOMaTHAHblX MYTal.\IfH. AHaJIOrlf'lHble pe3YJIbTaTbi nOJIY'IaJIH If APyrHe HCCJIeAO­

BaTeJIH [17, 18] rrpH aHamf3e crreKTpa a6eppaQllH B KopHeBoH MepHcTeMe rrpopocTKOB06JIY'leHHbIX ceMSIH CeJIbCKOX03~HcTBeHHblx pacTeHHH. TaKHM 06pa30M, B nepBblH.

HaH60JIee OCTpbIH rreplloA ~epH06blJIbCKOH KaTaCTpocpbI. KOrJ~a rrOrJIOll.\eHHa~ AOJa cpOpMH­pOBaJIaCb B OCHOBHOM 3a C'IeT BHelilHero ~- If Y-06JIY'leHIf~, pa,f\HaQHoHHoe nOpa)!(eHHe

rroceBOB CeJIbCKOX03~HcTBeHHblx KynbTYP rro OCHOBHbIM l.{HTOreHeTH'IeCKHM TeCTaM

HanOMIfHaJIO 3cpcpeKT, Bbl3bIBacMblH OCTpblM Y-06JIY'lCHIfCM B corrOCTaBIfMbIX A03ax.

B Ta6JI. 5 rrpe].\cTaBJICHbl AaHHbie 06 H3MeHeHHll B 3aBlfCHMOCTH OT rrOrJIOllleHHOH

pacTeHH~MIf ,f\03bl HCKOTOpblX rroKa3aTCJIeH, xapaKTepH3YlOlllHX )!(H3HCCrroco6HoCTb ceM~H.Ol.\eHKa MHTOTH'IeCKOrO HH,f\eKCa rro BOCbMIf BpeMeHHblM TO'lKaM (16-72 'l rrpopalllHBallHSI)

rrOKa3aJIa, 'lTO cYll.\eCTByeT HeAocToBepHaSi TeHAeHl.\H~ CHH)KeHH~ MHTOTH'leCKOH aKTHB­

IIOCTH C pOCTOM rrOrJIOllleHHOH pacTcHH~MH ].\03bl. J],ocToBepHaSi paAHal.\HOHHaSi AenpcccHSIMHTOTH'IeCKOH aKTIfBHOCTll Ha6JIlOAaJIaCb TOJIbKO Y 40-48-'lacoBblx rrpopocTKOB, OAHaKO B

3TOH BpeMeHHoH TO'lKe aHaJIH3HpyeMblH rrOKa3aTeJIb He rrpOSlBJISIeT 3aKOHOMepHOH

H3MeH'IHBOCTll C A030H. TIp If aHaJIlf3e BCXO)KCCTH ceMSIH Bbl5lBJICHbi CTaTHCTlf'leCKH

AOCTOBepHble 3cpcpeKTbi CTHM)'JISll.{llH rrpll A03ax 1,3-4.0 fp. TIpH A03e 12 fp lIa6111OAaJlOCb

503

B - (;,{UYYr11)) S Orru..'5,

dthY11( » .vvv</' "/ J .

/tUnO'Yf'L[ inu+aJiof/<.,l(f .

4i) In iMrl{) /fflA:ik-r" cLe- IS

(~ i nftU ) ~~r:1tntiUsClrtV[ .~ fLIf!if- g

tv i-M. ,/" ton cPIni.­

fall-1:- JflrCUl?mrL!',(I

J 1/. 1/ j.,f )m - (!"1( j1{)J1W-

) ()·m f 3 frio ut.& )

""tfl-tw 17'U?I&t f S1 -(/

([110) bridgfsw ;H~ (Wr? UNYU· -

.fcvrtJ "" frcrJmel1.!5

'1k o114~SmL~I

4- 5~ Ln'vI1c£~ I whic-h c.kar-a~.kNv.. 4- vietU-i'tify o;t

rMf~4WlinJ fro111 ~d-e.d Lio u '1 p-f!CUlU ~

'Tt~ STa6J1I1l\a5

a1,33,14,07,7

12,0

3,83 ± 0,262,64 ± 0,23 •2,29 ± 0,17 •2,54 ± 0,18'2,75 ± 0,16'2,61 ± 0,27 •

34,21 ± 3,4234,46 ± 2,1737,97 ± 2,9345,29 ± 3,3336,06 ± 3,2125,48 ± 3,49

62,0 ± 2,487,S ± 1,7'74,5 ± 2,2 •88,8 ± 1,6'70,8 ± 2,331,2 ± 2,3 •

8,99 ± 0,056,71 ± 0,05 •6,87 ± 0,06 •9,75 ± 0,05 •8,43 ± 0,06 •7,91 ± 0,Q3'

m~0+ .1000 :;-Pf!f)l~;gra/m

(9) <

• Pa3J1H'IIISI AOCTOBCpHbl lIpll P < 0,05.

cyw.eCTBCHHoe II CTaTIiCTII'-IeCKII BhICOKOp,OCTOBepHOC yrHcTeHlle pOCTa. Y 40--48-'1aCOBbIX

npopOCTKOB 3acpllKCllpOBaHa AOCTOBepHaSi 3aAep)!{Ka pOCTa rJIaBHO,O KOpBSI BO BceM

p,llana30He IICCJIep,OBaHHhIX A03. Op,HaKO y)!{e K 72-MY '-I 3TII pa3J1II'-IIISI IIlmeJIlipylOTCSI.

EOJIee TOro, npll nOrJIow.CHHhIX gOJax 1,3-7,7 fp Ha6JIIOp,aCTCSI cyw.ecTBeHBaSi (HO

CTaTllCTll'-leCKII HegocToBepHaSl) CTllMYJISIlIlHl pOCTa rJIaBHOrO KOpllSl, a nplI g03e 12 fp ­

yrHeTeHlle. 06paw.aeT Ha ce6S1 BHllMaHlle COnpSl)!{eIlHOCTb ll3MClleHllSi C p,030H 3THX

MOpcpOMcTpll'-leCKIiX nOKa3aTeJIeH, xapaKTcpll3YIQW.IlX )!{H3HeCnoc06HoCTh npopocTKOB Ha

paHHllx 3Tanax OHToreHe3a. CJIeAycT TaK)!{C OTMeTIITh TOT cpaKT, '11'0 cTHMymU.(HSI

BCXO)!{CCTll npopOCTKOB Ha6JIlOgacTcSI Ha cpOHC cyw.eCTBeHHO'O (B 2,6 pa3a) II BbICOKO­

gOCToBcpHoro npllpoCTa lIIiTorCHCTH'-IeCKHX HapywcHi-ili. TIpO.QCMOHc.:rpHpoBaHllhIH B HaIlICM

HCCJICAOBaHlIll cpcHoMeH CJIY)!{llT llJIJIIQCTpallUCH OTHOCHTCJIhHOCTH nOHSlTllSI "CTUMYJISlL\IUi" "

CBHACTCJIbCTBYCT 0 TOM, '11'0 31'01' TCpMHH HCJIb3S1 C'IUTaTb CHHOHIIMOM nonC3HOCTH IIJIlJ

ga)!{c 6C3BpCp,HOCTIi B03p,CHCTBIISI, nOCKOJIhKY BocnpHSlTlie cro PC3YJIbTaTOB C TO'-lKH 3peHIDlX03S1HCTBCHHOH gCSlTCJIbHOCTIi MO)!{CT OKa3aThCSI BeCbMa p,aJICKIiM 01' 6HOJIOrll'ICCKOH

cyw.Hocnl SlBJICHIiSi.

TIplI aHaJIll3C 113MCHCHIiSiMaCChI 1000 3CpCH 06Hapy)!{eHbl gocToBepHaSi CTllMYJISlL\HSInpn

06JIY'ICHHH B g03C 4 fp II p,OCTOBCpHOC YrHCTCHHe npll OCTaJIhHhlX A03ax. TlOCKOJIbKY

1I3MCHCHIIC MaCChi 1000 3epCH He KOppeJIlipYCT HH C nOrJIOll\CHHOH paCTeHllSiMIi p,030H, HII C

U3MeHCHHCM APyrHx KOJIH'-IeCTBCHHhIX npII3HaKOB, CJICgYCT Cp,CJIaTb BhIBOp" 'JTO Ha

BaphlipOBaHHC 31'0,0 nOKa3aTCJISI onpcp,CJISlIOw.ec BJIHSlHliC OKa3hlBalOT cpaKTopbl

HCpap,Ual\llOHHOH npUpOp,bl.

B lICJIOM npCp,CTaBJICHHble B HacToSlw.cH pa60TC pe3YJIhTaTbi CBIIp,CTCJIbCTBYIOT 0 TOM,

'-11'0 pap,UaL\liOHHOC nOpa)!{CHIiC a,pOL\CH030B Ha Hall60JIcc 3arpSl3HCHHbiX Y'laCTKaX'LepH06hlJIhCKOrO CJIcp,a JICTOM - OCCHhlO 1986 I. HOCllJIO KOMnJICKCHblH xapaKTcp II

npOSlBJISlJIOCh Ha pa3Hhix ypOBIISIX 6HOJIOrH'ICCKOH opraHll3al\lill - 01' MOJICKYJISIpHO-rellcTll­'1CCKQfO p,o op,aHU3McHHOro.

ABTOphl Bhlpa)!{alQT 6JIarOp,apHOCTh H.I1. CaH)!{apOBOH 3a nOMOU(h B opraHII3aUlIU II

npOBcp,clIIIII 3KcncplIMeliTOB B 30-KllJIOMCTpOBOH 30HC YA3C.

. Pa60Ta BhInOJIHCHa npH '-IaCTU'IHOII cpUHaHCOBOH nOp,p,ep)!{KC cpoHAa L!:)!{OHa L!:. II K3TpUHT. MaKapTypOB.

cnHCOK JH1TEPATYPbI

I. KOJy6oB r.M .. TaclCaeB A.H. PaAH06HOJIOrll'lCCKUC U pa)lH03KOJIOrH'ICCKHC IJCCJlC)10BaHIISI

ApCBCCHblX paCTCHIJH. Cn6.: HaYKa, 1994. 256 c.2.3R6/l1ll{KaR £.51., Cnupwi £.B .. CallJICapOBa Ii.H., A/leICCaXWI P.M. II PaAH06HOJlOrHSI. 1990.

T. 30. Bbln. 3. C. 291-295.

3. Cmenal/08 B.H. II DpaKTlJKYM lIO OCHOBaMalp0HoMIIII. M.: KOJIoc, 1969, C. 119-122.

504