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Текст
                    ('

'

II Al:ilffYPHEllTOB

!l.'l.H III KC>:Ihlll ll{OB

H.B. 4e6b1Wee, r.c. ry3"1KOBa, K>.6 . .naaapeea,
C.H . .nap"1Ha

EI10JIOfl1}l
HOBEHillHH

CITPABQqHHK

1lo.t oolltcii pc.w1<1u1cii a1«tW'.\1111rn 11.B. lleo1>1111eBa

!\lon;na
«:\L\XAOII»

2007


Y,Il;I\'. 57 BBK28 q_34 qefihillleB H.B., fy3HKOBa q_34 r.c., .ilaaapesa 10.B., JlapHHa C.H. B:irnJiormi:. HoBeiiumii cnpaBO'-IHMK. - M.: MaxaoH, 2007. (.Il:mi IIIKOJibHHKOB M a611Typ11eHTOB). 512., 11.'J. - ISBN 5-18-001064-0 (978-5-18-001064-3) JTOT conpe;1.1eHHbltt, D0"1HLiii, y1nrnepca:11,HLIH cupaBO'lHl1J( CO.L\CPlKWf llCC pa3,1€!Ibl 6uo.1orm1: o6ll(aH 6uo.101·11H, 1·etteT111rn, 60Ta11uKa, 300"'Ionrn, a11aTOMHH. BLICTpo Hai1n1, CllCTeMaTll3llpOBaTL 11 3aUOMHHTb 60"'Iblll0tt 06'bCM HH<jJOpMal(HJl IJOMoryT MHOr04llC"'lellHLie p11cymo1, Ta6:rnuLI, cxeMbI, 'leTI<aH CTPYKTypa H y,106Hbiii noncic E.'laro,1apH TecToBLIM 38,iJ,aIIHHM 11 HOBOH, HHTepaKTHBHOH cjJOpMe l!O/Ul'IH '.\laTepua"'la, ;JTa Kllttra MO)KCT o6ecrre4HTJ, CaMO!IO,[(rOTOBKY I( ;JJ(31l:vICHllM (Ef;), BCTYIIHTe.'Ib!lbJe B BY:lbl) upn C3Mb!X Bbl- COKHX rpe6osamrnx K a6wrypne1ny. A,;1peconaH nmpoKoMy Kpyry 'IHTaTe:ieii: y'Iall\11:vicH cTapmHx 1c1accos mKoJJ, rn:v111a3Htt, JJH!\Cen c yr.'ly6,,eHHbIM H3y'IeHHCM 6HCmor1111, a61nypHeHTa:vi, CTYilCIITaM cpe,iJ,HHX 11 BLICill!1X yqe6!1hIX 38BCi\€HHH, upeI!O.[\aBaTCmlM. Y.ll,K57 BBK28 © ISBN 5-18-001064-0 (978-5-18-001064-3) 'le6h!llleB H.B., J'y3HKOBa © 3AO Ko:vma1111H r.c., C.H., 2007 «Maxao11», 2007 Jfa3apesa 10.E., J1ap1ma
Co~epmaHHe BBE,UEIUiE YpoBHH opraH11aanm1 >K11Boii :viwrepHH . . . . . . . . . . . . . . . . . . . . . . . . . . . . . • . . . . 10 OcHOBHhie cBoiicTBa nornhIX opraHH3MOB . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 OBIIV\H J-3110.TIOnrn: OCHOBbl I ~I1TOJIOfl1I1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . MeTOJ\bl H3Y'ICH!1H Ko1€T!Of . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 061qce CTpOCHHe K.1€Tl{H . . . . . . . . . . . . . . • . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . XHMW!eCKHiI COCTall T«1CTKH . . . . . . . . . • . . . . . . . . . . . . . • . . . . • . . . . . . • . . . . • . Opranw1ecK1te BeII\CCTBa KJieTKH . . . . . . • . . . . . . . . . • . . . . . . . . • . . . . . • . . . . . . . BeJ1KH . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . • . . . • . • . . . . . . . . • . . . • . . . <lJepMCHThl . • . . . . . . . . . . • . . • . . . . . . . . . . . . . . . • . . . . . . . . . . . . . . . . . . . • . . . . JlHTIH,l\hl . . . . . . . . . . . . . • . . • . . . . . . . . . . . . . . . . • . . . . . . . . . . . . . . . . . . . . . . . . Yr,1eBOJ\hT . . . . . . . . . . . . . . . . . . . . . . . . . . . . • • • . . • . . . . . . . . . . . . • • . . . . . . . . HyKJICHTIOllh!C J{HC.TIOThl . . . . . . . . . . . . . . . . . . . • . . . . . . . . . . . . . . . . . . . . . . . . . . ,ZJJ-IK (,l\eao1<c11p1160HyK,1emroBa.H 1rnc.1orn) . . . . . . . . . . . . . . . . . . . . . • • . . . . . . . . • PIIK (pn6ony1G1c11HOBaR KHCJTorn) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . AT<P (a,l\eTI03HHTpmllOCQ>opna.H 1mc.10Ta) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13 14 16 17 19 19 21 22 24 24 25 27 28 HeK:1cT01 111h1c cpopMbI n<H3HH . . . . . . . . . . . • • . . . . . . . . . . . . . • . . . . . . . . . • . . . . 30 B:upyChl . . . . . . . . . . . . . . . . . . . . . . . . . . . . • • . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30 KJrcTO'lllhIC cpop:vib1 n<H3HH . . . . . . . . . . . . . • . . . . . . . . . . . . . . • . . . . . . . . . . . . . . . 34 Ha,l\I~apcTBO upoKapHOThI. . . . . . . . . . . . . • • . . . . . . . . . . . . . . . . . • . • . . . . . . . . . . 34 Ha,l\napcTBoayKap110Thl . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . • . . . . . . . . . . 36 06MeH llCII\CCTB 11 IIpeBpaII\Cl!He 3Hepr11H . . . . . . . . . . . . . • . . . . • . . • . . . . . . . . . . 48 lPOTOCHTITC3 . . . . . . . . . . • • . . . . . . . . . . . . . . . . . . . . . . . . . . . . • . . • . . . . . . . . . . . 50 XcMOCHHTea . . . . . . . . . • • . . . . . . . . . . . . . . . . • . . . . . . . . . . . . • . . • . . . • . . . • . . . 55 8HepreT11qec1<11n o6:vieH . . . . • . . . . . . . . . . . . • • • . . . . . . . . . . . . . . • . • . . . . . . . . . 55 ,IJ;e.1eH11e KJTeTKH . . . . . . . . . . • . . . . . . . . . . . . . • • . . . . . . . . . . . . . . . . • • . . . . . . . . 58 lIHTcpcpa3a . . . . • . . . . . . . . • . . . . • . . . . . . . . . . • • . . . . . . . . . . . . • . • • . . . . . . . . • 59 MHT03 . . . . . . . . . . . . . . . . . . . . • . • . . . . . . . . . . . . . . . • • . . . . . . . . . . . . . . . . . . . . 59 Mefio3 . . . . . . . . . . . . . . . . . . . . . . . . . • . . . . . . . . . . . . • . . . . . . . . . . . . . . . . . • . . . 61 Pa3MHO>KCH11eopraHH3MOB . . . . . . . . . . . . . . . . . . . . . • . . • . . . . . . . . . . . . . • . . . . 64 Becrronocpa3:viHon<enuc • . . . . . . . . . • . • . . . . • . . . . . . . . • • . • . . . . . . . . • . . . . . . . 64 IlOJTOBOC pa3MHO>K€HHe • . . . . . . . . . . . . . . . • • . . . . . . . . . . . • . . . . . . . . . . . . . . . • 66 faMeTOI'CHC3 . . . . . . . . . . . . . . . . . . . . . . • • • . • . . . . . . • . . • . • . . • . . . • . . . . . . . • 67 I1H,ll.HBH,l\ya,1bHOC pa3BHTHC opraHH3Ma •••.......••..••...•.........•.•. 72 IlOCT3M6p11ona.1bHOe pa3BHTHC . . . . . . . . • • . . . . . . . . • . . • . ..••.••....••.... 79 fEHETJIKA OcHOBHhlC IIOHHTHH :u onpeJJ;e,1en11H reneTHKH. 3aKOHhl r. MenJJ;e,1H .•••.....•. 81 Baa:uMoJ(eAcTB11ere110B . . . . • . . • . • • . . . . . . . . . . • . • . . . . . . . . . . • . . • • . . • . . • . 86 Cu;enJiennoc Hac.1e,l(oBaHHe. feHeT:uqec1we onpeJJ;eJTe11uc no.1a. ll;ttToreneT:uKa •.• 91 MoJieKyJIHpHaHre11cTHKa ..•..•.•..........••............•.•••..••.•.. 97 PerrJI:uKaQHH ,UHK ••.....••..•.•.........••...........••.••...•...•. 98 PeaJittaar~:uH reHeTwICcKofi 11H<)JopMau;m1 B KneTKe. Bttoc11HTC3 6eJTKa ..•..•... 99 TpancKp11rru11H ...•........•.•.••.••.......•••......••.••..••.•... 100
4 CoiJepJ1Canue fipouecc1rnr sy1rnp1wT1P1ec1-rnx H-PHK . . . . . . . . . . . . . . . . . . . . . • . . • . . . . . • . . TpaHc"1HUHH . . • . . . • . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . • . . . . . . . . . . . MenHUHHcKaareHCTHKa . . . . . . . . . . . . . . . . • . . . . . . . . . . • . . . . . • • . . . . . . . . . OcnoBHL!e '.\1.eTO/lbI 11ayqeHH.H renen1K11 qe;10BcKa . . . . . . . . . . . . . . . . . . . . . . . . . feHeanorH'!CCKHft MPTon . . . . . . . . . . • . . . . • . . . • . . . . . . . . • . . . • . . . . • • . . . . . QHTOreHCTH'leCKHH MCTOn . . . . . . . . . • • . . . . . . . . . . . . . . . . . . . . . . . . . . . . • . . . feHeTHKa COMBTH'leCKHX KJICTOK • . . . . . . . • . • . . . . . . . . . . . . . • . . . • . . . . . . . . • MeTO.L\ .u;epMaTor1m<jlnKH . . . • . . . . . . . . . . . . • . . . . . . . • . . . . • . • . . . . . . . . . . . • BJu13Heu.oah1tt MeTon . . . . . . • . • . . . • • . . . • . . . . . . . . . . . . . . • . . . . . . . . . . • . . • BnoxHMH'lecKHe MeTonhr ...•.•..•.•.•..•............•••.......•.•.•. IlonyJl.f!ll,l!OHHO-CTaTHCTl!'!CCKl!ll MeTon . • . . . . • . . • . . . . . . . . . . . . . . . • • • . • . . MeTOALI npenaTaJihHOtt (Bo ape MR 6epeMCHHOCTH) .u;HarHOCTHKH ..•...•....•.. lfaMeH'IHBOCTh. Tmrhr H3MeH'!HBOcTH. Pe11apa1ou1 .IJ:HK •.•.....•.•...•..•.. feHeTH'leC!(HC OCHOBbI CCJ!eK.UHH )fUJBOTHLIX, pacTeHHH, Ml!KpoopraHH3MOB. BnoTexuonorHH •••.........••..........•......•.•.• Ce,1eKU.HH pacTenntt ..••.•..•.•....•.•....•..•...•......•.....••...• CeJieRU.HH mnBOTHhIX ....•..•.•.•...........••......•...••..•.•....• CeJJeRll,IIH MlmpoopraIIH3MOB (B OCHOBHOM 6aRTepttii H rpH6oa) ••••••....•... BttoTexHonorH.H. fenHaH ilIDKeHeptt.H •..•.•.•.••...•.•.•.••........•..• IJ;l1TOIIJI83MaTw1ec1we nacJie)J;OBaHl!e •••.•.•...•..••..•.•.•.••.•.•..•..• 3BOJUOU:ll.H Bo3HHKnosenll.c muanu na 3eMne. Pa3Bl!Tl!e opranuqecKoro M11pa .•....•...• Pa3Bl1Tile mtt3HH na 3eMJie ........••.....•...•.•.•......•••..•.•.•..• OcHoBnLre :JTanbl <iBo,110u.ttu opranll.'!ecKoro Mttpa .•......••...•........•. Kpynnei1:nrne apoMop<jlo3bI B ::iaon10r~Hll opranuqecKoro MHpa .•..•........•. .IJ:oKa3aTeJibCTBa 3BOJII0ll,llll ...•..•.•..•.•...•.•.•.....••.•.•....•.... llaneoHTOJJOrH'ICCKHC,[\OKa3aTeJibCTBa .•......•.•.....•.•..•...•..•...• BuoreorpacpttqecKne ;:i;oKa3aTeJihCTBa . . • . • . . . • . . • . . . . . . . . . . . • • . • . • . . . • . Mop<jlonorll.'lecK11e;:i;oKa3aTeJILCTBa .••.•....•......••.•..••..•..•...•. 3M6puonoruqecK11e noKa3aTeJihCTBa ..••.....•...•.•.•..•....•.....•.... feHeTtt'leCKHe AOKB3aTeJibCTBa •.•...••.•...•.......•.• .- ...•..•.•...•.. BHOXHMl!'!CCKHe l! MOJICRyJIHPH0-6HOJIOI'H'!CCKlle noKa3aTeJihCTBa •.•.•..•..• Ilpoucxomnentte qenoseKa ..•......•.•...••..•..•......•.•.•.....•... llaneoHTOJiorll.'lecK11e noKa3aTeJILCTBa npoucxomnenu.H 'leJIOBeKa. 3BOJIIOI~HH npHMBTOB .•.•.•.•.•..•.•.......•.........•...•.• Pa3Bl1Ttte BLrcrnnx rrpll.MaTOB •.••.•.•..••.•.••.•.•.•.•....•......•.... 0CHOBHhie CTBAHH ::JBOJUOI~llll qeJIOBCKa ...•••.••.•.•.•...••.•......•.... IlttTCKBHTPOTibI (npeBHeiinme JIIO,[\ll, apxaHTpOIILI) •.•....•.•.•.•..•.•..•.• Hean.n;epTaJihll,LI (.upeanue nro;:i;u, naneoanTponLr) .••.....••...•..•.....•• qeJIOBCK COBpeMeHHOI'O THITa, llCROnaeMblll ll cospeMeHHblH (neoaHTpOITbI) .••. CoapeMeHHhIH 'lenoae:K u ero 3BOJIIOIJ:I1H (ueoanTponLr) ...•.••.•...•..•..•. .IJ:o.uapBHHOBCKllll rrepHOA pa3Bl!Tll.II 6HOJIOI'llH ••..•..••.•.•..•...•.•..••• Tpanc<jlopMtt3M ..•..•.•••.••.•••••.•.•.•...•••••.•••.•••.•.••...•• llepsa.11 u.enocTna.11 Kouu.enuu.11 :3Bon10u11u. JK. B. JlaMapK ••...••..•.•..••.• 3soJIIOll,ilounoe yqenll.e qapnaa .IJ:apsuna •...•.••••.•••.•..••.••.•...••• Ml1KP03BOJIIOll,ll.11 ••••.••••••..•.•••••••.••••••••.•.•.••..•••••.••.• llorryJIHll,HH - 3JieMeHT8pHa.H CllCTeMa 3BOJIIOUHH .•.•..••.••.•.•.•.•••••• 102 103 106 106 106 109 110 111 111 112 112 113 114 120 121 122 123 123 124 125 126 129 129 131 132 132 133 134 135 135 135 138 138 138 141 141 142 143 144 144 144 145 146 146
CoiJepJtCanue cf>aKTOpbl 3BOJIIOUH11 . . . . . . . . . . . · · . • . . . . . . . . . . • . . • • · · · · · . . . . . • . . . . . . OcHOBHOH 3Tarr 3BOJIIOUlIOHHOro npOilecca - Bll.ll . . . . . . • . . . • • . . . . . . . . • . . . . MaKp03BOJIIOHllH . . . . . . . . . • . . . • . . . . . . . . • . . . . . . . . . . . . • . . . . . . • . . . • . . . OcHOBHhIC nanpan.11eH11H 3BOJI]{)I~llll . . . . . • • . • . . . . . . . • • . • . • . . . . . . • . . . . . . . OcHOBHhIC rro.11o;irnHHH Cl!HTeTnqecKott Teop1111 3BOJIIOilHH ••......•....•.•... CaJibTailHOHnaH TeopnH ;:momou1111 .•.•........••••.•.......•....•.•..• ConpeMeHHa.H Teopn.H 3no1110uu11 . • . . • . . . . . • . . . . . . • . . • . . • . . • . . . . . . . . . . • • 148 152 154 155 15 7 158 158 3KOJIOfIIH ITonHTHH «cpe.11.a 06J1TaHHH» 11 «3Ko.11or11qecKne cjJaKTOpbI» . . . . . . . • • . . . . . . • . 3K0.110f'HH rrorryJIHUllll . . . . . . . • • . . . . . • • . . . . . . . . . • . • . . • . . . . • • . . . . . . . • . 3K0.110f'J1qecKaH xapaKTepHCTHKa BH,Ua .•.......••••.••......•...•..•... BnoreoileHoa • . . . . . • • • • . . • . . . . . . . • . . . . . . . . • • • . • • . . . . . . • . . . . • . . • . . . CTPYKTypa6noreoueHoaa • . . • . . . . . . • . . . . . . . • • • . . . . . . . . . . . • . . . . . . . . . • . IToKaaaTe.'ln cpyHKUHOHnponan11.H 3KOCHCTCMhI • • . . . . . . . . . . . . • . • . . . . . . . . • • Arpouenoah1 • . . . • • • . . . . . . . . . . . . • . • . . . . . . • . . • . . . . . . • . • • . . . . . . . • . . . . BnoccjJepa • • • . . . • • . . • . . . . . . . • . . • . . . . . • . . . . . . . . . • . . • . . • . . . . . • . • • . . . JK1rnoe BClllCCTBO ll ero cj:>yHKUHH ..••......•.......••.•.••...•.•..••... ,Il;eHTeJihHOCTh 'IeJIOBeI<a. HooccjJepa •......•••.......•.•.•..•.•••..•...• BmrnHHe .11.eHTCJihHOCTH •rn;wneKa Ha cTa6H"'lhHOCTh 6J1occjJeph1 ..•...•...•.•.• OcHOBHhre 31rn110rw1ecKne rrpo6.11eMhI . . . . . . . . . . . . . . . . • . • . . • . . . . . • . . • • • . 159 166 167 168 168 169 1 70 171 172 174 174 174 BOTAHIIKA T1<aHH 11 opraHhI BhIClllHX pacTeH11il . • . • • . . . . • • . . . . • . . . . . • . • . . • . . . . . . . . BereTaTllBHhre opraHhI (Kopen1>, cTe6e"1h, Jll!CThH) . . . • . . . . . . . • . • . . . . • • • . . . MopcjJo.1orw1ecKoe cTpoeH11e Kopn.H. KopHeBa.H c11cTeMa . . . . . . . . . • . . . . • • . • . . AHaTOMJ1qecKoecTpoeH11eKopHH • . . . . • . • . . . . . . • • • • . • • . . . . • . • . . . . . . . . . . CTe6e.11b . • . . . . . . • . . . . . . . . • . . . . . . . . . . . . . . . . • • . . • • . • . . . . • . . . . . . . . . . JlHCT .... ,,, .. ,,,, •...... ,,,, •..... ,,, . . . . . . , .•.•..... , •••.....•• feHepaTHB!Ihie opraIIhI (UBCTOK, rr.110.u, ceMH) .........••..•....•••....••• IJ;BeTOK .. , , , ... , , , , , ...... , , , ..•.. , , , , •....... , •.•.... , ••..••.••. fIJIO.U, , .. , , , ... , , , , , , . , .... , , , , , •.. , , , , , ..... , , , , , , , ... , , , , , ••... CeMH . • . . . . • • . . . • • . . . • • . . . . . . . • . • • . . . • . . . • . . . . • • . . . • . . • . . . • • . . . . . KJ1accncjJ11KaUHH 110Kpb1ToceMeHHhIX pacTeH11tt ••.......•••.•..••.....••. OcHOBHhre ceMelicTBa K.11acca ,Il;ny.uo.11hHbre (KpecTOilBCTHhie, PoaouneTHhie, Bo6onh1e, ITac.11eHOBb1e, C.11omHouneTHhre, MaJihBOBhre, MapeBbie, BHHorpa,nHbre) ••.......••••.•....••.•.....••... CeMelicTno KpecTouneTHhie (KarrycTHhre) ......••••.••...•••.........•.. CeMelicTBO Po30ilBCTHhre •..•••....••......•.•...••...•.••...•......• CeMeil:cTBO Bo6oBbre (MoThIJThIWBhre) • . • • . . . . . . . . . . . . . . • . . • . . . . . . • . . • . • CeMelicTBO ITac.11enoBb1e ..••......•.••••...•.••••.•..•.••.••..••••... CeMelicTno CJ1omnouBeTHhre ..•••....•...•...••••..•....••.•.....•••.. OcHonnwe ceMeiicTBa KJ1acca 0.llHO.UOJihHhie (JI11.11eliHble, 3.11aKOBh1e) ••......• CeMeil:cTBO Jiu.11ei1:1Ible ..•.•••.•......•••..•••...••...••.•.••...•..•. CeMeil:cTno 3"1aKH, 11J111 MHTJIHKOBbie •.•••...•....•....•..••••...••.•.• Ha.uuapcTno ,Il;oH.llep11b1e, HJIH ITpoKapn:oThI . . . . . • • • • . • . . . . . . . . . . . • . • . • . . BaKTepHH •.•...•••........•.•....•••...••.•••.•••.......••.....•• IJ;apCTBO I'ptt6br ...•...•..••.•.•...•..•...••••.•••........•....•... 0TJJ;eJI Jl11lllaHHllKH .•.....••.•••...••••..•••...•..•....•.••..•..••• 1 76 183 184 185 188 192 195 195 200 201 203 204 204 205 205 206 207 208 208 209 209 210 212 216
6 Coilep:Jl(;anue napCTBO PacTemrn. JlO[\UapCTBO HacT05111~11e flOi~OpOCJil1 . . . . . . . . • . . . . . . . . . . 218 OT l\eJI 3eJICHhIC BO,'l;opoc;rn . . . • . . . . . . . . . . • . . . . . . . . . . . . . . . . . • . . . . . • . . • 218 0T[\enBypweBOAOPOCnl1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . • 220 OTJ\C.'l KpacnhIC BOl\Opocmt, 11.'IH Barpmmn . . . . . . . • . • . • • . . . . . . . . . . • . . . . . . 221 II01v~apcTBO Bhicmuc crwpoBbie pacTemrn . . . . . . . . . . . . . . . . . . . . . . . . • . . . . . . 223 OTJ\CJI MoxoBH,UHbie . . . . • . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . • . . . . . . . • . . . 223 K.'!acc JI:ucTOCTe6e.'lbIIhie MXH, IIOAI<Jracc Bc.1b1e MXIL CcjiarttyM ..•.•.......• 225 0TAeJI IlarropOTHHKOBlfl\l!bie . . . . . . . . . . . • . . . . . . . . . . . . . . . . . . . . . . . . . . . . • 225 0Tp;eJI TI;rayHOBHAIIbie . . . . • . . . . . . . . • . . . . . . . . . . . . . . . . . . . . . . . . . . . • . . . . 228 0Tp;en XBou~eBH,ll,HhIC • . . . . . . . . . . . . . . • . . . . . . . . . . . . • . . . . . . . . . . . • • . • . . . 229 CeMCHHbIC pacTcH11n ....•.....•...••.•..••.•.•..•...........•..•..• 230 0Tp;CJI foJ!OCCMeHIIbJe . . . • . . . • . • . . . . . . . . . • . . • . . . . . . . . . . . . . • . . • . . . . . • 230 0Tp;eJ! IloKpbITOCCMCIIHhIC, 11.'IH IJ)eTIWBh!C • . . • . . . . . . . . . . . • . . . . . . . . . • . • . 233 300JIOI'IUI KJ1acc:ucjnmautt.H mttBOTHoro MMpa . . . . . . . • . . . . . . • . . . . . . • . . . . . . . . . . . . . . 237 Ilo;:i;uapcTBo O;:i;HoKJieTo'-IHhIC mHBOTHh1c, uJI:u TipocTeilurne ...•...•.....•.. 237 06rua.H xapaKTepHCTHKa npOCTCHllIHX ..•....•....•.•......•.....••..•.• 238 TttrrcapKoMaCTHrocjJophl .•...•.•.........•..•...•....•.•..•.....•..• 240 Tttn arrHKOMUJieKca •.•.•......•.•..•.•....•..•••....•....•.......•. 248 Tnrr HHcjJy3opHH (pecHH'-!Hb1e) • . . . . • . . . . • . . . . • . . . . . . . . . . . . . . . . • . . . . . • . 249 Tio;:i;u.apcTBO MnoroKJJeTO'-!HhIC . . . . . . • . . . . . . . . . . . . . . . . . • . • . . . . • . . . . . • . 251 Tttn ry6K:u . . . • . . . . • . . . . . . • . . . . • . • • . . . . . . . . . . . . . . . • . • • . . . . • . . • . . . . 251 Tttrr KHUie'IHOIIOJIOCTIIhIC . . . . . . • . . . . • . . . . . . . . . . . . • . • . . . . • . . . . . . . • . . . . 252 KJiacc fttAPOH,ll,Hhie . . . . • . . . . . • . . . . . . . . . • . . . . . . . . . . . . . . . . • . . . . . . . . . . 253 KJiacc Cu.mpOHl\Hhre, :u.'!:u C1~ttcjJOMep;y3hI . • . • . . • . . . . • . . . . . . • . • . . . . . . . . . . . 256 KJiaccRopaJ1noBwerroJittllhl . . . . . . . • . . . . • . . . . . . . . . . . . . • . . . . . . . . . . . . . . 257 Tttrr UJIOCK11e '-!CPBH • . . . • • . . . . . . . . . . . . . . . • . . . . • . • . . . . . . . . . . . . • . . • . . • 258 KJiacc PecHH'-!Hhie '-IepBtt • . . . . . • . . . . . . . . . . . . • . • . • . . . . . . . . . . . . • . . . . • . • 261 Knacc Cocanh~HKH, :u.1n Tpel\rnTOAhI. . . . . . . • . . . . • . • . . . . . . . . • . . . . . . . . . . . 263 Knacc JleHTO'IHhIC qepB11, u.rrn 1~ecTo.n;b1 . . . . . . . . . . . . • . . . . . . • . • . . . . . . . . . . . 265 Tnrr KpyrJihIC, ttJIH ncpBH'-!HonoJiocTHhJe, '-ICpBH . . . . . . . • . • . . . . • . . . . . . . . . • . 268 KJiacc HcMaTOAhI, l1JIH co6cTBCHHO J<pyrJihIC qepBH •..••..•..•.•..•.•...... 270 THrr KOJih'-laTbJC '-IepBH, HJ!lf KOJ'Ib'-ICI~bl .•.•..•..•.........•.••.•.•.....• 273 KJiacc MaJio~eTHHKOBbie qepB:u (o.'!:uroxeThI) •..•.•..•.......•.•....•.... 275 KJiacc MHoro~eTHHKOBhie '-!CpBH (rroJittXeThI) ..•.•..•..•.••••.•.....•... 278 KJiacc II11nBI<H . . . . . . • . . . . . . . . . • . . . . . . . . . . • . . . . . . . . . . . . . . . . . . . . . • • . 279 T11n MOJIJIIOCKH, :u.'!H MHl'KOTe,'lh!e •.•.•.......•...•....•.•...•...•.••. 280 KJiaccBp10xoHorueMoJIJIIOCI<tt ..••.•.•.....•.........•......•.•...•.• 284 KJiacc ):J:BycTBOp'-!aThre, HJIH IlJ1aCTHH'-!BToma6epHbie .•..•...•.....•....•.. 286 KJiacc foJI0Bonor11e MOJIJIJOCKH ......•..•.•.•......••.•.......•.•.••.• 288 T11rr'-!JieH11cTonor11e •...•....••........•..•.•.•..• . •...•......•.•.•• 289 KJiacc PaKoo6pa3Hhie .••.•.••..•.•.••.•....•...•..•.....•.•.•..•..• 292 KJiacc Tiay1<006pa3HbW • . • . . . . . . . . • . • . . . . . . • . . . • . • . . . • . . . • . . . . . . . . . • 296 KJiacc HaceKOMhre ......•.....•..•.•.••.•..•.•.•..•.......•.•..•.•• 298 T11rr xopp;OBhie ..••....••.•.•..••.•.......•....••.....•...•.•..•.•. 309 JlH'-!HHO'-!HOXOp,!J;OBb!e ..•......••..•.......••...•..•........•••.•.•. 310 KJiacc RpyrJiopoTb!C •...•...•..•..•.....•....•.•.......•.••...•...•. 314
7 CoiJepJ1Canue 317 326 JCwcc Ilpec:vihIKRIOII\HCCH (pCIITH,11111) • • . . . . . . . • . • • . • • . . • . • • . • • • • . • . . • • 333 JC1acc I!THllhI • . . . • • . . . . . . . • . . . . . . . . . . . . • • • • • • • • • . • • • • • • • • . • . • . • . • • 339 KJrace M.1e1rnm1TaIOll\He (3Bep11) . . . . • • . • • • • • • . • • • . • . . • . • . . • . . . • • . • • • . . 351 CpaimttTCJibHaH attaTo:v11rn cncTeM opranoB II03BOHO'!HbIX . • . • • . . . . • • . . • • • . • 364 <PirnoreHe3 Hapym:Hh!X IIO!<pOBOB . . . . . • • . . . . . . . • • • • . • • • • • . . . . • • • • • • • • • . 364 <P11JIOl'CIIe3 Cl{C.'IeTa . • • • • . • • . . . • • . . . . . . . . . . • • • • • • • . • • • • • • • . • • • • . . . • . 369 .I(mpcpepelll\HPOBKa IIOaBOHO'lHHK:J B np01~ecce 3BOJIJO~Ifl1 . . . • . . • . . • . • . . . . • . 369 3BOJI!Olil1H '!Cpena Il03BOHO'f!Ih!X • • • . • • • • • • . • . • . . . . • . . . . • . . . . . • • . . • . . • • 369 3BOJ110lil1H pa3Bl1Tl1H IIllTHIIaJIOH IW!Ie'fl!OCTH • • • . . . • • • . . • • . . . . • • • . . • . • • • 370 <P11JioreHe3 IIHll\eBap11Te.1LIIOH CHCTC:vlbl • . • • • • • . . • • • • • • • • • • • • • • • • . • • . . • • 370 <Ptt.'!Oreuea ,ll,bIXaTe.1hl!OH CliCTC'.ll.LI . . • . . • . • • • • • • • • • • • • • • • • • • • • • . . • • . . • • 372 <PttJIOreuea KpOBeHOCllOH Cl1CTC'.ll.LI. . • . . • • • • . • • • • • • • • • • • • • • • • • • • . • • • . • • • 3 73 3BOJIIOl\IUI COCY,ll,OB . • . . . • • . . • • • . . . . • • • • • • • • • . . • • • • . • • • • . • • • . • • . . • • • 373 Ha,ll,K.'Iacc pb16b1 . • . . • • . • • . • . . • • • • • • • • • . . . . . . • . . . • . . . . . . . • . . • • • • • • . . ICwcc 3eMHOBO/J,IIbIC (a:viqm6m1) • • . • . • • . . . . . . . . . . . . . . . . . • . . . . . . • • • • . • . <PttJiore11e3 nepmrni1 c11cTeMbI . . • • . . • . • • • • • • • • . . • • . • • • • . • • • • • . . • • . . • • • CTpoenne roJ10nnoro M03ra noaBOHO'!HbIX • • • • • • • • . • • • • • . • . • • • • • • . • • • . . • . <Ptt.'IOl'Cl!C3 opral!OB '!YBCTB . . • . • . . • . • • • • • • • • . . • • • . • • . . . • • . • • . . • • • • • • • <PttJIOl'CHC3 Bhl,ll,CJI!1TCJibHOH 11 no.1osoii CHCTeM .••••..•.•.••.••..•••••.•• CBH3L Bhll\CJIHTCJILHOH l1 IIOJIOBOH CHCTeM • • • • • . . • • . . • • . • . • • . • . . . • • • • • . . • AHATOM11H 11 3 74 3 74 376 377 379 <Pl13I10JIOrIUI "l!EJIOBEKA ••••••..•...•...••.••••••.••• 380 385 396 397 400 403 CTpyKTypuo-cpyHKn11onaJibHh1e qacT11 nepm10U: c11cTeMbI . • . • • . • • • • • . • . . . • • • 404 TKaHH . . . . • • . . • . • • • • • • . . . • . . • • . . . . . . • • . • • • • • • • • • • • • • • • • . • . . . • • • . • OnopHO-,l\Bl1raTe.'lbHbIH armapaT . . • . • • . . . • • . • • • • • • • • • • • • • • • • • • • . . • . . . • Mbunel!HaH c11cTeMa •••••••...••.•..•.•••••.•••••••.••••••••....••• CTpoeHHC MbillllibI • . • • • • . • . • • • • • • • • . . • • • • . • . • • • • • . • • • • • • • • • • • • . . • • • HepmiaH c11cTeMa ...••••••••••••••••••••••.•.•••...•..•.••.••••••• Ilepe,llalJa B036yn<,ll,eHl1H nepBHOH IWCTKOH 405 407 <PyHKlil111 OT,ll,eJJOB l'OJ!OBHOf'O :vioara • • • • • • • • • • . • • • • . • • . • . • • . • • . . • • . . • • . 408 IIepmpepn'IeCKaH IICpBHaH CHCTe:.i:a . • • . • • • • • . • . . . • • . • • . • . . • . • . . . • • • • • . . 411 CocTaB pecliJieKTopnoU: ,ll,yr11 . . • . • . . • . • • • • • • • • . • • • • . • • . . • • • • • • . . • • • • • . • 413 Bbrcll!aH HepsHaH ,ll,eHTC.'lbHOCTb • . • • . • • • . . • • • • • • • . • • . • . • • • . • . . • • • • • • • • 416 Mexatttt3M 06paa0Ban11>1 ycJIOBHoro pecjJJieKca • • • • • • . • • . • . • • • • • • • • • • • • • • . 41 7 Bo36ym,ll,eH11e 11 TOpMom:enne yc.10BHbIX pecp.1eKcoB • • • • • • • . • . • • • • • • • . • • • . • 41 7 Y'-IeH11e 0 ,ll,BYX CHrHaJILIIb!X CHCTCMax • . . . • • • • • • • • • • • • . • • • . • • • • • • . . • • • • • 419 T11rrb1 BhICllleii HepBnoU: ,ll,CHTCJibHOCT11 . . . • • • • • • • . • • • . . . • • . • • • • • • . . • • • • • 419 Ilpo~eccLI IIaMHTH • • • • • • . • • . • . • • . • • . • • • • • • • • • • . • . • • • • . • • • • • • . • • . • • • ,121 BH,ll,hI naM>1T11 . . • • • • • • • • • . • • . • • . • . • . • • • • • • • • • • • • • . • • • • • • • • • • • . • ••• 421 Oprauh1 '!YBCTB • • • . • . • • • • • • • . • • • • • • • • • • • • • • • • • • • • • • • • • • . • • • . • . • • • • 422 OpraH apeHttH . • • . . • . . • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • . • . • • • . . • • • • . • 423 BcrroMoraTeJihllbIC opraHhI rJia3a • • • • • • • • • . . • • . • • • • • • • • • . . • . • • • . • • • • • • • 425 3pttTC.'lbHLiii anaJIH3aTop • • • • • • • • • • • • • • • • . • . . • • • • • • • • • . • • • • • • . • • • • . • • 426 Oprau cJiyxa tt paBHOBeCHH • • • • • • • • • • • • • • . . • . • • • • • • • • • • • • • • • • • • • • • • . • 427 3BYKOBOCITpl1HH!\1a!Ollll1H anrrapaT BHyTpemrero yxa • • • • • . . • • • . • • • • • • • • • . • 429 Oprau paBHOBec11>1 (BecT116yJIHPHhIH anrrapaT) . . • • • • . . • . • . • • • . • • . • • • • • . • • 431 Crr11uttoii M03r • • . • . . . . • • • • • • • • . • • • • • • • . • • . . . . • . . • . • . . • . • • • • • • . • • . • foJIOBHOH M03r •.••....•.••.•.••..•••••••....•..•.•.••.•••.....•••
8 CoiJep:H<:anue OpraHhI o6oHmnrn YI BKyca . • . . . . . . • . . . • . . • . . . . • . . . . . . . . . . . . . • . . . . . . . . 432 Koma . . . . . . . . . . . . . . . . . . • . . . . . . . • . . . . . . . . . . . . . . . . . . . . . • . . . . . . . . . . 433 KmKHhIH ana.1Y13aTop . . . . . . . . . . . . . . . . . . . . . . . . . • . . . . . . • . • . . • . . . . . . . . . 435 BHyTpeHHHH ceKpeu,1u1 . . . . . . . . . . . . . . . . • . • . . • . . . . . . . . • . . . . . . . . . . . . • . 436 06Me!I Belll,eCTB H ::rneprirn . . • . . . . . . • . . . • . • . . • . . . . . . . . . • . . . . . . . . . . . . . . 445 06MeH BOAhl H MHHepa.'IbHblX COJierr . . . • . . . . . . . . . . . . . . . . . . . . . . . . . . . . . • . . 446 06'.\feH 6e,1KOB . . . . . . . . . . . • . . . . . • . . . • . • . . . . . . . • . . • . . . . . • . • • • • . . . . . . 44 7 06MeH yr,1eBOl\OB . • . . . . . . • . . . . • . • • . . • . . . . . . . • . . . . . . . . . . . . . • • . . . . . . • 448 06MeH )fU!pOB ••••.•......• · . • . . . . . . . . . • . . • . . . . . . . . . . . . • . . . . . . . . • . 449 BHT3MIU!bi • . • . • . . . . . • . . . . . . . . . . . . . . . . . . . . • . . • . . . . . • . . . . . . . . . . . . . . 449 KpoBoo6pauleHtte . . . . . . . . . . • . • . . • . . . . • . . . . . . . . . . . . • • . . . . . . . . . . . . • . 452 CTpoeHtte RpoBenocHhIX cocyn;os . . . . . . • • . . . . . • . . . . . . . • . . . . • . . . . • . . . . . • 453 KpoBHHoen;aBJietttte ..•.••......•.•..•......•......•...•..••...••••• 456 PeryJI.HU,Yl.H ABHJKCHUH r<posu IIO cocy,ZJ,aM •.•..•.•.•....••......•.•...... 456 CTpoen11e cepAila •...••.•••...•...•.....•..•...•....••..••......•.. 457 BHyTpeHHH.H cpe,ll;a opraatt3Ma .•.•.......•..•..........••.....•.••.•.. 460 CTpoea11e JIUMcPaTII'!ecRoil c11cTeMh1 .•..•.••.•....•.....•......•......• 461 KposeTBopHbleopraHhl ..•............•.••.•....•...•••.....••.•....• 462 ~OpMeHHbleaJieMeHTbIKpOBH ...••.••.•...........•.•..••••.•.•..•.•. 464 MexaHH3M csepThrnaHHH Kposu . . • . . • . . . . . . • . . • . . . . . . • . • • . • • . . . . . . . . . . 467 fpyrrnhI KPOBH 'Ie;IOBeJ<a ....•..•.•.•••...•..•.•.•.....•.•.•....•.••. 468 L\b!X3HllC . . • . . . . . . . . . . . . . . . . . . • . . . . . • . . . . . . . . • . . . . • . . . . . . . • . . . . . • 469 Mexantt3M s11oxa H BhI,noxa •..•...•....•....•.•......••..•....•.....• 4 72 fa3006MeH B JICI'KHX ll Tl<aJIHX . . . . . • . . . . • . . . . . . . . . . • . . • . . • . . . . • . . • . . . . 4 7-4 IlnuleBapeHtte •••.••.•.•......••.•...•.........•.•.......•......•. 4 76 fioJIOCTh pTa .•.••.•....•....•.•........••.•.•.........••••.••...•. 477 Ilrrlll,esapIITe.'Ibllbie meJie.1bl .......•.•..•.•........•....••.••...•.••. 484 PeryJIHU,ll.JI JJHIIleBape1mH ....•.•......•....•.•......••.•.....•...•.• 486 Bhl.n;eJieHIIe .....•..•.•....•.•........•......•.....••..••...•..•.• 48 7 Pa3MHOJKeHll.e .•........•.••...•......•.•......•.•.•.....•..•.•.•. 492 MymcKa.H no,1osaH Cll.CTeMa •.•...••.•.•.•.•...••....•.•.....••••...•. 492 JKeHCKaH IlQjIOBaH Cll.CTeMa ....•.••••.••..••.....•.••.•..•.••.•.•.•.• 493 3M6pll.OHaJibHhlll nep110,ll, pa3Bl1Tll.H '!eJIOBeKa •...•.....•..•.••.•••..•..•• 496 TECTbI 06Ill,aH 6ttOJIOl'll.JI ...•.•••.....•.•.•.....•.....•..... , •.••.....•.•.• 497 il;HTOJIOI'IIH ......•.•......••....•....•.••.•.•••..•••.•••....... . • 497 feHeTHI{a ..•.......•...••.•••...•......••.•.••..•.••.......•.••.. 498 3BOJIIOU:HOHHOe y'!eHll.e .......•.••...••...•....•...•........•.•...•. 499 AHTponoreHe3 ..•..•.......•..•..•..•...•.•.....••...•...•.•.•.•.• 500 3KOJ!OI'HH .•...•..•.•..•.•..••.•••.•........••......•...•...•.•.• 501 BoTaHnKa ....••.••.•.•.........•.....•.•.•..•......•.•.•.......• 502 3ooJiornH ..••••..•....•.....••.••....•..•.....•...•••.•.••....•.. 504 AHaToMHH ....•.•.•..••.•.••...••.•.••..•..••.•. • •...••...•..•... 506 OTBETbl ........................................................ 51 O
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14 06UfllJl 6UO/lOZUJl. 0cH06bl lfUmO/lOZUU cTaBJieHHe 6hrno orrpoBeprttyTo HeMeu;KnM yqeHhIM Py.n;oJih<lJOM BHpxoBoM (1858 r.). OH rrnca.11: «Bouca.H KJtem1w npoucxo()um u.3 rJpyioil KJlf'nrn:u ... » - TO ecTh o6octtoBa.T£ rrpnttu;Hn rrpeeMCTBeHHOCTH J<JieTOK rryTeM .n;e.1eHHJL 3To yTBepm.n;eH11e CTaJIO TpeTbHM IIOJIO:IK€HH€M KJICToquoii: TeopnH. III aTarr - cTaHOBJieHne coBpeMeHHOH KJieToqHoii: TeopHH. Bo3MO:IKHOCTH cBeToBoro MHKpOCKorra XVII-XVIII BB. 6hIJIH orpaHH- qeHHhI. IloaTOMY HaKorr.1eHHe MaTepnaJia 0 KJI€TOqHoM CTpoeHHH pacTeHHH H :IKHBOTHhJX, 0 B 30-x rr. CTPYKType caMHX KJie'l'OK IIIJIO M€,ZlJI€HIIO. lfao6peTeHHC xx B. 3JI€KTpOHHOro MHKpOCKOIIa H ycoBepweHCTBOBa1-rne B CBJI- 3H c 3THM M€TO,ll;OB ncc.1e.n;oBaHHH JI03BOJIH:IH HaMHOrO r.1y6me ll3YlfHTb cTpyKTYPY n cpyHKIJ;HOHnpoBaHne pa3JIHlfHhIX KOMIIOHeHTOB KJieTJHL Ha ocHOBaHHH IIOJiylfeHHhlX pe3yJibTaTOB 6hIJIH ccpopMyJIHpoBaHhl OCH06Hb/e noJlOJICeHUJl co6pe.MeHHOU KJLemo•mou meopuu. Bee JICU6bte opia1tU3Mbt cocmo.Hm u3 KJLemoK. KJtemKa - 3Jte.MeHmapHa.H J1Cu6a.H cucme.Ma, erJuuu4a cmpoe1tu.H, .HCU.311eoe.HmeJtbHOcmu, pa.J.Mlto.HCeHUJl U UllrJU6UOya.'lbHOi?O pa;wumUJl JICU6blX opiaHU.J.M06. Bne KJtemKu 1tem JICU.JHU. KJtemKu 6Cex opianu.JM06 cxodttbt no cmpoeHu10 u xu.MU'lCCICOMy cocmaay, u 6 nux cxodllbt.M o6pa3o.M npome1Ca10m npo4eccbt 06.Mena Bew,ecm6, xpaHenue u peaJtu3a4uJl nacJteocm6enHou un<jJop.Ma4uu. Ho6bte 1CJtem1Cu 60.Jttu1w10m moJtb/CO nymeM deJte1tu.H panee cyw,ecmBy10w,ux 1CJtemo1C. KJtemo'lnoe cmpoeHue Bcex llblHe J/CuByw,ux opiaHU3.M06 - c6uoemeJtbcmBo eduncm6a ux npoucxoJ1CdeHu.H. TaKHM o6pa3oM, B HaqaJie XX B., 6Jiaro.n;apJI co3.n;aHHIO KJieTOlfHOH Teopnn, ccpopMHpOBaJIOCh npe.n;cTaBJI€HHe 0 e.n;HHCTBe opraHnqeCKOH rrpHpO,ll;bl. I. CneTonan MHKpocKODHH: 1. <Pa3060-ICOHmpacmtta.R .MU1Cpoc1conu.H - 3TOT MCTO,ll; II03BOJIJieT pe3KO IIOBhICHTh KOHTpacTHOCTh H3o6pameHHJI o6'beKTa. 2. JlOJt.Hpu3a4uonHaJl .MUICpOCICOllUJl - MeTo.n;, rrpu KOTOpOM B rro.1€ 3pe- HH.H IIOJI.HpH3aIJ;HOHHOro MHKpOCKOIIa B pe3yJihTaTe ,ll;BOHHOro JiyT-IerrpeJIOMJI€HH.H ( «aHH30TPOIIHH») o6'h€KThl OKa3hIBaIOTCJI RPKO CB€TJIIIJ;HMHC.H Ha TeMHOM cpoHe. 3. HHmep<jJepen4uOHHaJl .MUICpOCICOllUJl - MeTO,ll;, npH KOTOpOM CBeT pa3.n;eJIJI€TCJI Ha .n;Ba rryqKa. 0.n;HH H3 HHX rrpoxo.n;HT qepe3 o6'heKT, .n;pyro:H - MHMO Hero. BcJie.n;CTBHe aToro cBeT, rrporne.n;nrnii qepe3 o6'h- eKT, HCIIhIThIBaeT cpa30BhIM C,ll;BHr, H H306pameHH€ CTpOHTCJI TaKHM o6pa30M, qTo yqacTKH KJieTKH, 06Jia.n;a10ru;He pa3HOH IIJIOTHOCTbIO, 6y.n;yT OTJIHlfaThC.H .n;pyr OT .n;pyra no CTerreHH KOHTpaCTHOCTH. 4. Mu1Cpoc1Conu.H B me.MHO.M noJte - rrptteM, rrpn KOTopoM n3o6pametttte nona.n;aeT B o6'beKTHB B pe3yJihTaTe OTpameHHJI CBeTa OT o6'beKTa, 6Jia-
15 MemoiJbi u3y'lenun 1aiemKu ro,napJI lJeMy MeJih'Ia:HUIHC 1.J:UCTHI~bl Bbir JIJ!,[IJ!T CBCTJ!lIJ;HMHCJ! TO'-IKRMH Ha TeMHOM cpoHe. 5. YJ1bmpa<fJuoJ1emoea.H MUICpOCKOllUJl - crroco6, rrpn KOTOpOM KOMITO- HeHTbl ofrheKTa 06J1a,na10T cneu;ttcpHllecKHM rrorJio~eHHeM Y<P-J1yqei1: c orrpe,neJieHIIOH ,ll;JIHHOM BOJ!Hbl, '-ITO II03BOJIJieT BbIJ!BHTb HX 6e3 OKpallIHBaHIIJI. 6. <PJtyopecu,eHml-taJl MUKpOCKOllUJl - MeTO,ll; IICCJie,noBaHHJ! )KlfBbIX K.!e- TOK B YJ1bTpacpHOJ1€TOBOM CBeTe. Il03BOJ1JleT H3YlJIITb co6CTBeHny10 cpJIIOOpecu;eHI~HIO (cBelJeHHe) Be~eCTB, a TaK)Ke HX CTpoeHHe IIOCJIC .ll;06aBJieHIIJ! B HHX crreu;HaJihHbIX KpaCHTeJiett (cpnyopoxpOMOB) • .IJ:aHHhIII MeTO,ZJ; II03BOJIJieT orrpe,ri;eJIHTb MeCTa paCIIOJIO)KeHHJ! HYKJieHHOBbIX KHCJIOT, BHTaMHHOB, mttpOB H .ri;p. 7. JI,ucprpepettu,uaJtb1toe yJ1bmpau,ettmpurpyzupoeattue - MeTo.n; II3ylJe- HH.H COCTaBa KJieTOK, OCHOBaHHbIM Ha pa3,neJieHHll KJ'IeTOlJHbIX opraHeJ'IJI H BKJIIOlJeHHH no llX IIJIOTHOCTH. IlpH 6hICTPOM Bpa~eHHH pacTBopa B crreu;ttaJihHOM rrpII6ope - yJihTpau;eHTpttcpyre - opraHeJIJ!hI KJleTOK Bhma.n;aIOT B oca.n;oK, pacrroJiara.HCb CJIO.HMH B COOTBeTCTBHll co csoeii IIJIOTHOCTbIO. 8TH CJIOH pa3.ll;eJI.HIOT H H3ylJaIOT OT,ZJ;€JlbHO. II. 8JieKTpOHHaH MHKpocKonm1: 1. Tpanc.MuccuOHHa.H :JJleKmpoHHa.H MUKpocKonu.H - 11306pa)KeHHe Ha cpJiyopecu;Hpy10~eM 3KpaHe 3JieKTPOHHOro MHKpOCKOIIa rrpH pacceHBaHHH rryqKa 3JleKTPOHOB o6'beKTOM. qeM 60Jihllle paccellBaIO~a.H crroco6HOCTb Toro HJIH HHOro yqacTKa, TeM 6oJiee TeMHbIM BbirJI.H,ll;HT OH Ha 3KpaHe. 2. CKattupy10w,a.H (pacmpoea.H) 3JteKmpo1t1ta.H MUKpocKonu.H - MeTo.n; H3ylJeHH.H TpexMepHOH IIOBepxHOCTH KJieTKH .ri;ettCTBHeM rryl.J:Ka 3JleKTpOHOB no IIOBepXHOCTH 06'heKTa. III. PettTrettocTpyRTypHLiii attaJIH3 - H3ylJeHHe KOHcpurypau;HH MoJieKyJ1 6eJIKa, HYKJieHHOBhIX KHCJIOT ITO pacrrpe.n;eJieHllIO Ha 06'beKTe peHTreHOBCKOro H3JiylJeHIIJ!. IV. BHTaJILHoe (npmKH3tteunoe) H3yqettHe KJieToK - H3yqentte cBo6o.n;Ho- :anrny~Hx IlpOCTCMllIHX opraHII3MOB, KJieTOK KYJibTYPhl TKaHH, a TaK)Ke KJleTOK KpOBH. lfayIJenue r/JUKCUpo6aHHblX KJlemoK ( c HCIIOJib30BaHHeM crreu;ucpHl!eCKOM o6pa60TKH H rrpuMeHeHHeM pa3JIHlJHbIX KpaCHTeJie:H) rrpOBO,ZJ;HTC.H CJ1e,ny10~H­ MH MeTO,ZJ;aMH: 1. F(umOXU.MU'ieCKUe .MemoiJbl HCCJie,ZJ;OBaHH.H OCHOBaHbl Ha npHHIJ;IIIIe crreu;HcpHlleCKOrO CBJ!3bIBaHHJ! KpaCHTeJie:H c orrpe.n;eJieHHbIMH KOMIIOHeHTaMH HJIH Be~eCTBaMH 2. AemopaiJuozparjJu.H KJieTKH. ocHoBaHa Ha perucTpa1~HH se~eCTB, MelJeHHhIX pa,ZJ;HOHYKJIH,ZI;aMH. 3. HMMyHozucmoxu.Mu'iecKue .MemoiJbt ocHoBaHhI Ha orrpe.n;eJ1eHHH aH- TureHOB B KJleTKax H Ha o6Hapy)KeHHH TKaHeBhIX aHTHTCJI. 4. [_{ umocneKmporjJOmOMempU.H 6a3HpyeTCJ! Ha onpe.n;eJieHHH KOJIIIlJeCT-
06~an 16 6uo1wiun. Oc1to6bl 1.fumo1wzuu Ba sern;ecTB l1 l1X COCTaBHblX ::weMeHTOB B KJleTKe B 3aBIICl1MOCTl1 OT nor Jiorn;eHJUl CBeTOBblX Jlyl.JeH onpe,neJieHHOH ,iJ;JIMHbl BOJIHbl. 5. ll,umo<jJU3UOJl02U'leclWU .lft,emoi) OIIHpaeTCH Ha JI3yl.JeHIIe q:nI3IIOJIOrl1· l.JeCKHX npou,eccoB Ha OT,neJihHbIX II30Jil1pOBaHHblX KJieTKax ll,bl, HepBHbie CTBOJ1bl, raHrJIMU HT • 6. A1emoo ICYJlbmypbt m/CaHeu '.Vlhllll- .n;. npe.n;noJraraeT HCC.'Ie,ll;OBaHIW JKIIBbIX KJieTOK, IIOMern;eHHblX B COOTBCTCTBYIOID;YIO cpe.n;y, B KOTOpoii OHH cnoco6Hbl K aBTOHOMHOMY pocTy. IlpIIMeHHIOTCH TaIOKe MUICpoxupypzua - BblIIOJIHeHIIe c IIOMOID;bIO cneu,H- aJihHblX MaHIIIlYJIHTOpOB pa3JIHl.JHblX orrepau,nii Ha KJieTKe Hee opraHOH,n;ax, l1 npUJICU3HeHIWe OICpawuBanue JKIIBbIX KJieTOK KpaCIITeJiflMII B .n;1rnrra30He KOHU,eHTpau;nH:, He Bbl3hIBaIOrn;nx TOKCII'IeCKOro 3cpcpeKTa. 06m;ee cTpoeuue 1meT1m KJtem/Ca - amo 3JleMenmapnaR cmpyKmypnaa, <jJyn1Cu,uonaJ1blta.R u zeuemu'leCKa.R eiJunuu,a JICUBbtX opzanu3M06, 02panu'leH1ta.R no.llynp<muu,ae.wou MeM6panou, cocmoaw,aa u3 aopa u u,umon.naaMbt, cnoco6naa IC caMopezyJLRu,uu, CUM006H06JleHUIO u CUM060CnpOU36eiJeHUtO. <l>opMa KJieTKH (pnc. 1). Pa3Jin'-laIOT KJieTKn c n3MeH'IHBOtt cpopMott (aMe6hI, JiettKOll,HTbl H .n;p.) H Ha n KJieTKII, cpopMa KOTOpbIX 6oJiee HJIH Mettee IIOCTOHH- cneu,IIcpII'IHa ,ll;JIH Ka:m.n;oro nx TIIrra (ttttcpy3opmf, crrepMaTo30II,ll;hI, 3pIITp01I,IITbl, 3IlIITeJIIIfu'IbHbie H uepBHbIC KJieTKII, a TaKme 6oJibIIIHHCTBO pacTnTeJibHhIX KJieToK). ct>opMa KJieTOK 3aBIICl1.T OT HX cpyHK1I,IIOHaJibHOrO rrpIIcrroco6JieHIIH, IIOBepxHOCTHoro HaTHJKeHMH M BH3KOCTM rrpOTOIIJia3MbI, MeXaHM'IeCKOI'O B03,n:eifcTBHH rrpHJierruorn;nx KJieTOK, pnrH,n:HOCTII (:meCTKOCTH) KJieTO'IHOH MeM6paHbI. Pa3Mephl KJ1eToK. Pa3MephI KJieToK KOJie6JIIOTCH B rrmpOKMX rrpe.n;eJiax. Lt:ttaMeTp caMbIX MeJIKHX JKl1BOTHbIX KJieTOK paBeH 4 MKM. 0.n;tta- KO HeKOTOphie pacTnTeJihHhie M JKHBOTHhie KJieTKH MOJKHO Bn,n;eTh HeBoopymeHHhIM rJia30M, TaK KaK OHH ,ll;OCTMraIOT HeCKOJihKHX caHTHMeTpOB B ,n:waMeTpe (ttarrpnMep, Hifu;eKJieTKM). B a TKattax JKMBOTHhIX o6'beM KJieTOK orrpe.n;e- JieHHoro THila ,ll;OBOJihHO IlOCTOHHeH H He 3aBMCHT OT o6rn;ttx paaMepos TeJla. HarrpnMep, KJieTKM PMC. 1. a - C!>opMa KileTOK: HepsHa11 l<JleTKa; 6- r- IIO'IeK II rre'IeHM y 6h1Ka, JIOIIIa,n;M II MblillH IIMean11- IOT rrpMMepHO 0,ll;IIH H TOT me pa3Mep; pa3JIH'!He Ten11anbHb1e KlleTKL1;,l1-COeAL1HL1- B BeJIH'IMHe opratta o6ycJIOBJIHBaeTCH 'IHCJIOM, TenbHOTKaHHafl KneTKa; e - a He o6'beMOM KJieTOK. 8To npaBMJIO HHOr;:i;a Ha- KlleTKa; JK - 11iiiu,e- Mb1we•iHa11 KneTKa 3hIBaIOT 3a/COHOM nocmORHHOZO o6oeMa.
XuMu'teCKUii cocmaB KJlemKu 17 XHMH'lleCKHii cocTan KJieTKH B COCTUB )KllBblX opraHI:I3MOB BXO).(.HT Te me XHMHqecKne ;)JJC:\1CHThl, KOTOpbie COCTaB."IfffOT H 06'heKTbl HeiKHBOll npHpO.D;bl. Co11epmautte n 1rn:enw xHMH'lec1rnx coe;i:uue1rnii: (6 % 1w Cblpy10 Macey) ~-- L -----· ~~ -B~~e(-;,~a=--- -- --~--1 ____He_opram1_q_ecK11e. _____ _j______ __Opra1_1w1ec:KJ:!~--- __ . Bo.n;a I M11HepaJihHhle coJIH 70--80 1-1,5 L_ __________________ · Bemrn JK11pb! YrJieBOAhI 10--20 1--5 0,2---2 ---~·_HYl{:l!C11H<~!3h~li!1_CJIOThl ____!_-:-_2__ ; B 1rneTKe o6ttapymetto 6oJiee 80 x1:1M1:1qecKnx sJieMeHTOB Ilep1:10.1J;11qecKorr c11cTeMhI IL Mett,n;eJieesa. Ilp1:16JIH3HTeJihHO 40 3J"IeMeHTOB np1:1H1:1MaIOT yqacTHe B npou.eccax o6MeHa semecTB 1:1 o6Jia,n;aroT Bb1pameHttorr 61:10.1or1:1qecKorr aKTHBHOCThIO. 8TH 3JieMeHTbl Ha3bIBaIOTC.H 61:1oreHHbIM1l. Ilo co.n;epmaHHIO HX ,n;eJI.HT Ha TPH rpynnh1: 1. M aKpo;ue.11-ie1-tmbt (98 '%) - 0, C, H, N. 2. Mu1Cp03JteMe1-tmbt (1,9 °/.i) - K, P, S, Mg, Cl, Ca, Na, Fe. 3. Y.llbmpaMuKpo:uteMeHmbt (0,01 'Xi) - I, Cu, Co, Zn, Mo, Br, Mn, B 1:1 .n;p. Boi}a B cpe,n;HeM COCTaBJI.HeT 80 % MaCChl KJieTKH (.n;o 95 % - B KJieTKax MeAY3hl, 90 % - B KJieTKax 3apo.n;hrllla qeJioBeKa, 79 '% - B MhIIIIu,ax cep.n;u,a, 60 %1 - B CTapbIX KJieTKax H 10 % - B KJieTKax 3MaJIH 3y60B). Bo.n;a Haxo,n;HTC.H B KJieTKax B .n;syx cpopMax: CB06o,n;Horr - 95 'Xi H CB.H3aHHOll - 5 %i. IloTep.H 20 % BO,ll;hI cMepTeJihHa ,n;JI.H opraHH3Ma. Taxoe BhICOKoe co.n;epmaHHe BO,n;hr B KJieTKe onpe.n;emieTCH MHoroo6paa1:1eM ee cpyHKIJ,Htt: - cpe.n;a ,ll;Jl.H npoTeKaHHH 6HOXHMHqecKHX peaxu,1:1i1; - YHHBepcaJihHbIH paCTBOpHTeJih; - cpe,n;CTBO TpaHCIIOpTHPOBKH BemecTB B KJieTKax (.n;mpcpy3H.H) H B opraHH3Me (Kposoo6pamettlie ); - o6paaosattMe KJieToqHhlX CTPYKTYP (60Jibllloe co.n;epJI{aHHe B U.HTon.1a3Me); - onpe,n;eJI.HeT o6'beM 1:1 ynpyroCTh KJieTKI:I ( 06ecneq1rnaeT ocMOTHqecKoe H Typropttoe ,n;aBJieHHe); - yqaCTByeT B TepMoperyJI.HIJ,HH (6Jiaro.n;ap.H BblCOKOH TeIIJIOeMKOCTH H TellJIOIIpOBO,/l;HOCTlI); - yqacTsyeT B peaKIJ,H.HX rM,n;poJIH3a lI cpoToCHHTe3a (HcToqHHK HOHOB H-t ); - cpe.n;a ,ll;Jl.H OIIJIO,ll;OTBOpettm1; - y pacTeHHH o6ecrreqHsaeT TpattcnHpa11;tt10 H rrpopacTaHHe ceM.HH. BMoJiorMqecKa.H poJih BO,n;hr orrpe,n;eJI.HeTc.H MaJiblMH paaMepaMH MOJieKyJI, crre11;mpHKOH HX CTpOeHHH (IIOJIHpHOCTb, crroco6HOCTb o6pa30BbIBaTh B0).(0PO.n;Hhie CB.fl3H). Ilo OTHOIIIeHlIIO K BO,n;e Bee semecTBa ,n;eJIHTC.H Ha 2uapo<PU.llb1-tbte (paCTBOplIMhle B Bo,n;e) - MHHepaJihHhle COJIH, KHCJIOThl, meJioqH, .n;.
06utaJI 6uo1wzuJI. OcnoBbt 1fUmo1wzuu 18 Buo.'loruqec1rnn po:1h HeK0Topb1x xnMH'lec11:ux JJieMeHTOB cf>Il3IlOJIOfl1'!eCKaH pOJlb 8;ieMeHThl Yr.wpo.n (C) Bxo;vrT B COCTaB BO/~bl, opral!ll<ICCKilX BernecTB (6eJ1Kll, 11yK- Bo11opo,1J; (H) ;1e11HOBbJe KllCJIOThr, yn1cno,1J;b1, JI111IIIL\hI). R'.IlCJIOpOl\ (0) Y<IacTsymT B c1rnTe3e opraHH'Iec1<11x uernccTB Il q1yHKl\IIHX, AaoT (N) ocyn1ecTB.1HCMbIX oTlfM11 opraHll'ICCKllMll BeU(eCTBaMII. Y<IacTsyeT B npou:cccax so36yn<J(eH1u1 K;1cT101, B 11011,1J;cpma- HaTpllll (Na) Hllll OCMOTil'ICCKOI'O ,ll.aBJICHilH Il pH cpe,ll.hl, B,1IlHCT na pa6oTy noqeK BXO!lllT B COCTaB KOCTHOII TKaHll, HC06XO,ll;llM npn csepTbIBa- Ra;1bll,Ilii (Ca) Hllll KpOBll, MhIIIIe'IHOM COKpan(eHlll1 Heo6xo11HM 11.1n Bo36ym.neHilH nepBHhJX KJICTOK, rrp0Be11e1111a RaJIIltt (R) llMIIY-1bCOB, COKparneHllH Mbllll1I YqacTnyeT B 11011AepJKaHilll pH JKe,1y11o'IHOI'O COKa, OCMOTil<IC- XJiop (Cl) CKOI'O l(aBJICHl1H IIJia3Mbl KpOBH ct>occiJop (P) CTpyKTYPHblll IWMITOHCHT IWCTeii 11 3y60B, BXOJ(HT B COCTaB JI~e.'IC30 (Fe) CTpyKTYPHh!H KOMIIOHeHT !'CMOr,10611:Ha KPOBH, MHOI'JI06HHa ATct>, HA)l;ct>, cpoccpoJ1mrn11on Mbllll1I, cpepMCIITOB I(errn nepenoca 3,1eKTpOHOB Mo11 (I) Mell.h (Cu) BXOl\HT B COCTaB I'OpMOHOB 11IllTOBil!llIOii JKeJie3hI - - -- - - - -- -· YqacTByeT B npou:eccax KpoBCTBopeHMH 11 c:11nTe3e reMOI'JI06MHa cl>TOp (F) Marm1if (Mg) C1·pyKTypHblll KOMIIOHCIIT 3y6uoii TKaHll BXOJ(HT B COCTaB XJIOpOcp11JIJia, KOcpepMCHTOB, aKTHB11pyeT 0HepreTH'IecK:11iJ: o6MCH :11 c11nTe3 AHR Cepa (S) BXOJ(IlT B COCTaB aMill!OKl'!CJIOT, 6eJIKOB (FrnCyJimI) H BllTaM:rrHOB (B ) 1 1l;IlHK {Zn) K0Mnoue11T cpepMeHTOB, 11eo6xo11:rrMbIX .n;m1 uopM3.1hHOro pocTa Ko6aJibT (Co) Bxo.n;MT B cocTaB BttTaMMHa B 12 MapraHeu (Mn) Heo6xo11ttM .n;Jia oKMCJICHllH m:rrpHbIX KMCJIOT, yqacTByeT B rrpou:eccax l\hixamrn Il <PoTOCilHTc3a MOHOcaxapII,D;hI, 6eJIKll ll .n;p. ll zuiJpo<jJo6ttblC (BO,[(OHepaCTBOpIIMbie) - jKif- pbI, IIOJIIIcaxapII,D;hl ll .n;p. MunepaJlbflbte coJlu cocTaBJIRIOT 1-1,5 'Yo MaCChr KJieTKH. BoJihIIIa.H 'laCTb HeopraHII'leCKIIX Bell(eCTB HaXO,[(.HTCR B KJieTKe B BII.n;e COJieH HOH, COJI.HHOH, cpoccpopHoii:. cep- MttHepaJihHhre COJIII IIrparoT BamHyIO pOJib B pa3BIITMII mIIBhIX opraHII3MOB. IIx He.n;ocTaToK MJIH II36hlTOK MOJKeT rrptt-
Opzanu'!ecKue 6eUfecnrna K.riemKu. £e.ri1<u 19 B€CTH K r116eJIH opraHH3Ma. COJIH :v10ryT Haxo,n:HTbCH B K.1CTKe Jll16o B BHµ:e HOHOB, JI!160 B TBep,n:oM COCTOJIHHH. Munepa.lhHbie COJII1 B BOlWblX paCTBOpax ;:i;11cco1(1111pyKiT ua aHHOHbI 11 KaTHOHhI, y 1rncTBYIOT B no,n:.n:ep:maHnn ocMOTIIqecKoro JJ,aBJTeHHH (3a cqeT pa3HOCTH KOHI(eHTpau,Hii no o6e CTOPOHbl MeM6paHhI KJieTKH), TIO)J,)(epJKHBaIOT c;ia6orue.10lfHYIO pH (7,2-7,4) B TKa- HeBOM Jl\I:!,llKOCTH. Oprattnqec1me Bem.ecTBa KJJeTKH BamueiIIIIIIMII opram1qecKHMn Beru;ecTBaMn KJieTKH RBJIHIOTCR 6e"1KH, mtt- pbr, yrJieBO!l:bl n HYKJieHHOBb!C KHCJIOThI. CTpyK'l'ypubre n ;::i;pyrne oco6euuocTH KJieTKH Tecuo CBJI3aHhI c ,n:JIHHHhIMH MOJieKyJiaMH, COC'l'OJIIIJ;I:!Mll H3 TIOB'l'Opmoru;nxcJI ;)JieMeHTapHbIX C,11,HHHU., coep;HHeHHbIX KOBaJieHTHbIMH CBJl3JIMH. 3TH e,n:HHHUbl Ha3hIBaIOTCJI MOHOMPpa- MU, a o6pa3yIOrn;uec.H MaKpOMoJieKyJihI - no.rtuMepaMu. OprauntJecK11e coe;::i;I:Iueu11a cocTaBJIJIIOT B cpe,n:neM mirnoro opraHH3Ma. B pa3Jrn:qnbre 20-30 '% :vraccbr KJie'l'Kn TI:!IIbI KJI€TOK BXO,ll;HT Heoµ;IIHaKOBOe KOJIHqe- CTBO opraHuqecKnx coe;::i;nHennii. TaK, B pacTHTeJibHhIX KJieTKax rrpeo6Jla)(aJOT CJIOiKHLie yrJieBO)J;hI - noJincaxap11)(hI, B aornoTHhIX - 6eJIKH 11 a<Hphr. Op;HaKo KaJK,l(aJI rpynna opraHI1'IeCKI1X Beru;eCTB B JII060M THrre KJI€TOK BhIITOJIHJieT CXO,l(Hbie QJYH1\IXl1H. Bemrn y pacTeHHM 6eJIKH CHHT€311PYIOTCJI B pu6ocoMaX l13 aMHHOKHCJlOT. y )KH- BOTHbIX 6eJIKH IIOCTyrraJOT B opraHH3M c rrnru;eii, pacru;enJIHIOTCJI AO aMHHOKHCJIOT, KOTOphre HJI;YT na CHHT€3 co6cTB€HHbIX 6eJIKOB B COOTBeTCTBUH c retteTHqecKHM KOJ(OM. Cocma6 6e.ri1Co6 BuonoJlUMepbl. MonoMepaMn JIBJlRJOTCH aMMHOKHCJIOThI - HH3KOMOJ1eKyJIRpH&re coe)(HHenuJI. Pa3JIHqaroT 20 aMn:HOKHCJIOT. 0614aJL <jJop.wy.ria: R I NH 2 - C - COOR, I R r.n;e NR 2 aMunorpynna ( ocHOBHhie cBonc'l'Ba), COOR Rap6oKCHJibHaH rpynna (KHCJIOTHhre cBoncTna), R - pa,n;n:KaJI (20 BapuaHTOB). 8To aMcpoTepHhie coe,n;HHeHHR. <l>H3HqecKue cnoi1:cTBa: 6ecl(neTnh1e, xpucTannnqecKMe, pacTnopnMhI B Bop;e, HepaCTBOpHMbl B opraHH'I€CKHX paCTBOpHTeJIJIX. AMHHOKHCJIO'l'hI, coe;::i;nH.HJICh, 06pa3y10T rrerrTH)(HYIO CBJI3h (xonaJieHTHyIO, npoqHyIO). -C-N- 11 I 0 R
06UfUJl 6uoJtoZuJl. Ocno6bt 1.J,Umo;wzuu 20 KJWCCU<jJUKQL{UJl 6PJlKOB." I. Ilo KOJIII'IeCTBY aMHHOKI1C~IOT: 1. 2. II. 2 ,!l,O 10 a:vi11HoKucJioT); 10 .no ••. ): co6cTBeHHO IIOJJI1IJellTH,ll,bl (OT 10 .n;o 100 aMl1HOKHC.10T) ropMOHbI; 6eJIKl1 (npoTel1Hbl)- OT 100 aMI!HOKHC.10T ,ll,O HeCKOJibKI1X Ml1.1.Jll10HOB. O.:rnronenn1,11,b1 (OT IloJittnenTI1,ll,bI (OT IIo cocTaBy: Ilpocmble 6e.il!CU ( npomPU/lbl) COCTOJIT T0.1bKO H3 OCTaTIWB aMIIHOKUCJIOT: npoTaMHHbl H rHCTOHbl - BXO,ll,JIT B COCTaB HyK.1enpoTeH,ll,OB. MrpaIOT Ba:IKHYIO pOJib B pery.1Hl(HH MeTa6o.1H'feCKOM aKTHBHOCTH reHOMa; npoJiaMHHhI H rmoTe.1n:HhI - 6e.i1KH pacTn:TeJibHoro npo11cxom,n;enn:JI, co- cTaBJIHIOT OCHOBHYIO Macey KJie:UKOBHHhI; aJih6YMI1Hbl n: rJio6yJIUHbl -- 6eJIKH :IKHBOTHOro npmICXQ)l{,ll,eHirn (5111'1HbIM 6eJlOK, Mbllllll;bI, MOJIOKO). CJWJICHbte 6eJtKU ( npomPudbt) pa3.1wrn10T n: na3hIBaIOT no npocTeTn:<IeCKOM (He 6eJIKOBoi1) rpynne: xpoMonpoTen:,11,hI - 6eJIKH, B KOTOpb!X npocTeTH'feCKOM rpynrrofi: CJIY:IKHT ITHrMeHT (reMor JI06HH, Il;HTOXpOMhI, ,ll,bIXaTeJihHhie cpepMeHTbI); HYKJieorrpoTeH,ll,hI - 6eJIKH, CB513aHHhie c HyKJiel1HOBbIMI1 KHCJIOTaMn: (oc- HOBa 51,11,epnoro Bem;ecTBa JIHITOilpOTe:H,ll,bI - xpoMaTHHa); coe,n;r-rneHH51 6eJIKOB n: JIHilll,ll,OB (c]JepMeHTbl ITJia3MaTI1- qecKHX MeM6pa11); cpoccpoupoTen:,n;hI - coe,n;11nemrn 6enKoB 11 cpoccpaToB (B MonoKe, meJITI{e I<ypn:noro RHu;a, ttKpe pbr6, II,HC); rmmorrp0Te11,n;b1 - coe,11,n:nemrn 6eJIKOB H yrJ1eso,11,oB (KoMnoneHT KJieTo<I- noii MeM6panLr); MeTaJIJIOIIpOTeH,ll,hI - III. coe.n;nnenmI 6eJIKOB II MeTam10B (cpepMeHTbI). Ilo CTPYKType: <Pu6pUJlJl.HpHbte 6eJtKU - IIOJIHITeITTI1,ll,Hbie Hll'reBH,ll,Hbie u;errn, ITJIOXO pac- TBOpllMbl B 130,ll,e (KepaTn:H BOJIOC ll poroB, MI103HH MhIITII.J;, KOJIJiareH KOCTe:H, cpn6pn:noren KpoBn:). I'Jl06yJtRpllbte 6eJtKU - TI0.7II1ITeITTH,ll,Hble u;entt, CJIO:IKeHHbie B cpopMe ITiapa, BO,n;opaCTBOpllMbie (npoTeHHhI TIJia3MbI KpOBH, cpepMeHTbI). CTpyKTYPhI 6eJrna (ypoBHH opraHHaau;HH) II epBu'lnaR - (pnc. 2) JIHHeii:na51 nocJie,n;oBaTeJihHOCTh aMHHOKHCJIOT, coe,n;uneHHhIX ,n;pyr c ,n;pyroM nerrTH,ll,HbIMll CB51351MH B ITOJIHI1eIITH,ll,HYIO u.errh (crreu;HcpH'!Ha, 3aKO,ll,HpoBaHa B .IJ;HK, OT Hee aaBMCRT CBOHCTBa ll cpyHKD;Hll 6eJIKOB). BmoputtnaR: IIOJIHrrerrTH,ll,Ha51 u.errh, 3aKpyqeuHa51 B crrHpaJih. a-cnupaJ/,b rrpaB03aKpy<IeHHa51, B03HMKa10m;a51 B pe3yJihTaTe o6pa30BaHH51 BO,ll,OPO,ll,HhIX CB513eii Mem,n;y C=O II NH, Mem,n;y nepBhIM u TpeThHM aMMHOKHCJIOTHhIMH 3BeHh51MH (rJI06HH); - CKJia,n;<IaTa51, {3-cnupaJtb, B03HHKaIOm;aa B peayJihTaTe o6pa3oBaHH51 Bo,n;opo,n;HhIX CB513ell Mem.n;y napaJIJieJihHO opn:eHTHpOBaHHhIMll yqaCTKaMH no-
21 CmpyKmypa 6enKa JI11nenn1;i:no:H Mo.1eKyJrh1 (KepaTHtt BOJIOC); -- Tpexq,1ettttaa cn11paJih - crr11pa.1e- B11.n;tta.H CTPYKTypa, B03HHKaIOIIJa.H B pe3yJihTaTe rreperrJieTett11.H Tpex rro.111rrerrTH.n;tthrx ri:erreii (KOJIJiareH). Tpemu'l1taJt - rroJI11rrerrTH.n;Hhre ri:err11 B pe3yJihTaTe rrpocTpattCTBettttOH yKJia,ll;KH o6pa3yIOT paKTeptta i1to6yJty (KJiy6oK). Otta xa- ,ll;Jl.H 6oJibllIHttCTBa 6eJIKOB H rro.n;.n;epmHBaeTC.H HOHHbIMH, BO,ll;Opo.n;HbIMH CB.ff3.HMH, ,ll;HCYJih<pH,ll;HbIMll MOCTI1KaMH (-S-S- ), r11.n;pocpo6HhIMH B3aHMO- 6enKa: 3 - .n;eii:CTBH.HMH. Bee rJio6yJI.HpHhre 6eJIKH - 4 2 P111c. 2. Pa3nV14HbJe crpyKrypb1 MoneKyn I - nepBV14Hasi; TpernYHasi; 4 - 2 - BTOPV14Hasi: YernepTVIYHasi (Ha cpepMettTbI, attTIITeJia, ropMOttbI. q emeepmU'llWJl - xapaKTeptta ,ll;Jl.H 6eJIKOB co CJIO:?KttbIM CTpoettIIeM, co- CTOHT H3 tteCKOJibKHX IlOJIHileITTH,ll;ttb!X ri:eneii:, y.n;epmHBaeMbIX B MOJieKyJie ,ll;HCYJih<pH,ll;ttbIMH MOCTHKaMH, Bo.n;opo.n;tthIMH CB.ff3HMH H r11.n;pocpo6ttbIMH B3aHMo.n;eHCTBH.HMH. HarrpHMep, MOJ1CKyJia reMOrJI06Htta COCTOHT H3 'ICTbipex rroJIHITerrTH,ll;ttblX l~erreii, Kam.n;a.H 113 KOTOpbIX CB.H3atta c o.n;tto:i1: rpynrroiI reMa, y.n;epm11BaIOIIJero KHCJiopo.n;. HapyllleHHe CTPYKTYPhI 6eJIKa tta3brnaeTc.H iJe1wmypau,ueu. Otta ttacTynaeT rro.n; .n;eHCTBHeM cpH3H'IeCKHX H XHMM'IeCKHX cpaKTOpOB (TeMrrepaTypa, B03.n;eiICTBHe KHCJIOT H .n;p.). BbrnaeT o6paTHMOH (pettaTypari:H.H) H tteo6paTHMOH (eCJIH ttapylllaeTc.H rrepBH'Itta.H cTpyKTypa MOJieKyJihI). <Py1ucu,uu 6eJt1Coe: 1. Cmpy1Cmyp1taJl (cTpOHTeJihtta.H) - BXO,D;.ffT B COCTaB Bcex MeM6pattttbIX CTPYKTyp. 2. <PepMe1tmamU6/taJl (KaTaJIHTH'IeCKa.H) 3. 3aw,um1taJl (HMMYttOJIOrH'IeCKa.H) - - cpepMettTbI opraHH3Ma. aHTIITeJia, HMMyHorJio6yJIHtt, HttTepcpepott. 4. Co1CpamumeJtbltaJl (.n;BHraTeJihtta.H) - .n;BHmettHe pecttM'IeK, mryn1- KOB, COKpaIIJettHe MblllIIJ:. 5. I'opMOHllJlbHllJl (peryJI.HTOptta.H) - ropMOHbI (HttCYJIHH). 6. 3nepiemu'leC/CaJl - ITPH CJKHrattHH 1 r 6eJIKa BbI,D;eJI.HeTC.ff 1 7 ,6 K,Il,JK 3HeprHH. 7. TpancnopmnaJl - reMorJI061:1H. 8. Peu,enmopnaR - po.n;orrcHH rJia3a (peri:errTOphr). 9. 3anaca1ow,aR - .HH'IHhIM aJih6yMHH, Ka3eHH MOJIOKa, ::m.n;ocnepMa ceM.HH. 10. To1Ccu'lec1CaJl - R,n; 3Meii:, rrayKoB, HaceKOMhIX. «l>epMeHTLI <PepMeHmbt (OT JiaT. fermentum - 3aKBaCKa) - 3TO 6eJIKH, rrp11cyTCTBYIO- Ill;He BO BCex »<HBbIX KJieTKax H HrpaIOIIIHe pOJib 61:10Jior1:1qecKHX KaTaJIH3aTOpOB.
06~aJI 22 6uo.1iozuJI. OcnoBbi lfUmonozuu CuHTe3MPYIOTC.H M3 aMMHOKMCJIOT Ha ptt6ocoMax B COOTBeTCTBMM c reneTV!'IeCKHM KO,D;OM. Ilo XHMH'IeCKOll npwpo.n;e ;JTO 6P.!llW (H:MeIOT CJIOJI<HOe CTpoeHH:e, qeTBepTlI'IttP6e.iK06ble KO.Mnoueflmbl (HOHbl MCT3Jlv'IOB, BHTaMHHb!). BeJIKOBa.H qacTh cpepMeHTa Ha3hIBaeTc.H ano<jJep.Me1tm. He6eJIKOBa.H qacTh cpepMeHTa Ko<jJaKmop - HOH MeTaJI.W HJIH HeopraHI1'IeCKOC seru;eCTBO; KOrpep.Meflm (K03H3HM) - opram1qecKoe seru;ecTBO. CBoilcmea rpep.Menmo6: HYIO CTPYKTypy) M 1. BhlcoKa.H crrer~ml:mqHocTh, TO eCTb cpepMeHT H cy6cTpaT rro cTpoemuo COOTBCTCTBYIOT )J.pyr )J.pyry. 2. 3. ,[(ettCTBHC B CTporo onpe)J.eJieHHOH IIOCJI€)J.OBaTeJihHOCTM. YcKopettue orrpe)J.eJieHHhIX peil.KIJ;HH. K.naccurjJuKau,u.R cpep.MeHmoe: Cyru;ecTsyeT rnecTb KJiaccos cpepMeHTOB B 3aBHCHMOCTM OT Tuna KaTaJIH3MpyeMbIX peam:i;nfi:: 1. OKcuDopeDyKmaJbl - KaTaJIH3HpyIOT OKMCJIHTeJibHO-soccTaHOBMTeJib- Hhie peaKIJ;HM. 2. Tpanc<jJepaJbl - KaTaJm3MpyroT uepeHoc <PYHK1IHOHaJibHhIX rpyrrrr OT O)J.HOro seru;ecTBa K )J.pyroMy. 3. I'uopo.na3bl - KaTaJIH3HPYJOT rH)J.pOJIH3 - peaKIJ;HH pacru;ernieHH.H CJIO)l{- HhIX opraHuqecKHX seru;ecTB Ha rrpOCThie IIYT€M rrpncoe)J,HHeHH.H BO,ll;bI. 4. JI ua;Jbl - KaTaJIH3HPYIOT HCI'H)J.pOJIHTHqecKoe rrpncoe)J.HHeHHe HJill OTru;errJieHHe <PYHKIJ;HOHaJihHOH rpyrrrrhI. 5. HJo.Mepaabt - H30Mepu3ar:i;u.H. 6. JI U?a3bl ( cuttmemaabl) - KaTaJIH3HPYIOT peaKIJ;HH CHHTe3a c BaHH€M attepruu AT<l>. <PyHKU,U.Jl rjJep.Meflm06 - HCilOJib30- ysemfqeHMe CKOPOCTM KJieToqHbIX peaK11Hll B MHJI- JIHOHbI pa3. Hu O)J.HH rrpor:i;ecc B KJieTKe He rrpottcxo)J.MT 6e3 yqacTH.H cpepMeHTOB (pe)J.yrrnnKa111rn ,[(HK, CMHTe3 PHK, 6e.JIKOB, AT<l>, cpoTOCMHTe3, )J.hIXaHue H )J.p.). Jlnn~hl y paCT€HHH JIMIIH)J.hl CMHTe3HpyIOTC.H B KaHaJiax rJia)J.KOH 8TIC. y JRHBOT- HhIX IlOCTyrraIOT c rmru;ei1:, pacru;errJI.HIOTC.H )J.O COCTaBHhIX qacTeii (rJIMIJ;epMH M mHpHbie KMCJIOTbl) H BHOBh CHHT€3MPYIOTC.H B co6CTB€HHhie JRHpbl. Co.n;epJRaHHe JRHpa B KJI€TKe COCT3BJI.H€T CocTaB JIMIIH)J.OB. 5-15 'X, MaCChI cyxoro seru;eCTBa. JKupbt ( mpui.nuu,epuobt) - CJIOJRHhie acpnphI TpexaTOMHoro CIIHpTa l'JIH11€PHHa H BhICOKOMOJieKy.TI.HpHbIX JRHpHblX KHCJIOT. cH 2 - o - cp - R1 CH - 0 - CO-R 2 CH 2 - 0 - CO - R 3 ! JKnpbl 6bJBaIOT HaChlIIJ;€HHbie H HeHaCblill€HHhie. 06w,a.R <jJopMy.na:
<Pep;nenmbt. JI unuilbi H acbtuJ,eH llbU' 23 (TBCpJJ;bIC) - cTcapttHOBaa, naJihMHTIIHOBMI, :vrnc;u-IHaJJ KHC.'10Thl (rOBJ:I)KHH, CBHHOH mttp). H e1taCbLUJ,eH Hbte (mtt.n;Ktte) - OjICHHOBaa, JIHHOJICBaa KIICJIOThI (no,r:i;co:r- tteqrroe MacJ10, pb16tti1: )!{ttp). ct>II3WlCCKHe CBOHCTB3 JKIIpOB: JKHpbI nepaCTBOpHMbl B BO,r:i;e; paCTBOPHMbl B opraHIFieCKHX paCTBOPIITCJI.HX (aqrnpe. CITIIpTe, 6eH3IIHe II JJ;p.). JI unowh1 - C.'IO)KHhie scjmphl BbICOKOMOJieKyJIHPHhlX JKIIpllbIX I{IICJ'lOT K IIHM OTHOCJITCJI IrnpOTHHOHJJ;hI (cjJOTOCIIHTCTnqeCKHC IlHI'MCHTbI), II CIIHPTOB. cTepOHJJ;HhIC ropMOHhl (ITOJIOBbie, I'JIIOKOKOpTllIWH,[\hI H .n;p.). KJ1,accurjm1rnu,uJt JtunuooB: 1. IIpocmbte J1,Unu<Jb1 rrpe.n;cTaBJIJ:IIOT co6oi1: scpIIphI crrttpTa 11 :tJ{IIPHhIX KttcJIOT ( mpuz.11 uu,epUObt ): cmepouiht - rrpOH3BO,L\Hhie crrttpTa XOJICCTepoJia (IIOJIOBhie ropMOHhI, xoJieCTepo.'I, KOpTmwcTepoII,nhl, BHTaMHH D); mepneHbt - BCIIJ;CCTBa, B octtoBe CTPYKTYPhI KOTOphrx JiemttT rrJITIIyroJ1hHhIM yrJreBo,r:i;opo,r:i; uaorrpett (xJiopocpttJIJI, KapoTHHOH,L\hI, BHTaMHH J1cupopacmBopu.Mbte Buma.MUHbt (A, LI;, E, K); BOC/CG - C.'IO)KHbie scpttpbI BhICIIIHX JKHpHbIX A); KIICJIOT H BhICOKOMOJieKy- mipHbIX CIIHpTOB. 2. CJLO;HC/lbte J/,UllUObl - :JTO COeJJ;HH8HHe H JJ;pyr11x BCIIJ;eCTB. K HHM OTHOCJITCJI: rjJOcrjJo.11unuobL - ::icpttpOB CITHpTa, JKHPHhIX KHCJIOT coe,r:i;ttHeHIIJI I'JIHI~epmia, JKHpHh!X KHCJ'IOT H OCT3TKOB q>0ccpopHOM KHCJ'lOThI; crpuHzo.rtunuiJbt - coe,n;IIH8HHJI JIHIIH)J,OB H CIIHpTa CcjJHHI'03HHa; Z.!IUKO.!IUnuiJbt - coe,n;11HeHHJI JIHilII)J.OB II yrJieBO,L\OB; J/,UnonpomeuiJbL -- C08)J.HHCHHJI JIHilH)J,OB H 6eJIKOB; nuz.Mellmbt (KapoTIIHOIIJJ;hI, reMOI'JI06HH, XJIOpocjJHJIJI H .n;p.). <Py1-tKu,uu .11unuiJoB: 1. 3Hepzemu 'leCKaR - rrptt OKHCJI8HHH 1 r JIIIIIH)J,OB Bhl,L\CJ'lJICTCH 38, 9 T8IIJI3. JlttIIIIJJ;hl o6ecrreqHB3IOT 25-30 '% K,llJK BceH: sHeprHH, Heo6XOJJ;HMOH op- raHII3My. 2. Cmpoume.flb/iaJl (cTpyKTypHaJI) 3. I'op.MOHaJlb/taJl ll pezyJ[,JtmOpHaR - BXOJJ;JIT B COCT3B 6IIOMeM6paHhl. CTepOH,L\Hhie ropMOHhl peryJ'IllPYIOT o6MeH BeIIJ;eCTB H pa3MHOJKCHHC, BHT3MIIHhl. 4. 3anaca/OUJ,llR B )KHBOTHhIX - B paCTHTCJihHhIX KJICTKax ceMHH IIJIOJJ;OB pacTeHHH, JKHpoBaJI TK3Hb. 5. If cmO'lllUK ;miJoceHHOU 60iJbt 1,1 6. 7. 8. 9. rrpH OKHCJICHHH 1r JKIIpOB o6paayeTCJ:l MJI BO)J.hl. Tep.MOU.JOJ/,RU,UOHHaR - yqacTIIe B TerrJioo6Mette. 3aUJ,UmHaR - OT MexattnqecKHX IlOBpCJK)J.CHHM. Bb1iJe.1umeJ1bliaR - y ttaceKOMhIX - «JKIIposoe TeJIO». Cneu,uaJibllbte rjJynKu,uu - xttMnqecKtte curHaJihI (cpepoMOHhI y HaceKo- Mhrx), o6pa30B3HHe BO)J.OOTT3JIKHB3IOIIJ;HX ITOKphITIIH y pacTeHIIM (BOCKa), OKpacKa IIJIO)J.OB H CCMHH, yqacTHe B cpoTOCHHTC3e (rrHI'MCHThI).
06Ufa1l 6uo;wzu1l. OcnoBw ztumo;wzuu 24 YrJieBO,UbI B KJieTKax pacTeHHH yr;reso.n;hr CHHTe3npyIOTCH B X.i'Ioporv:racTax B npo11,ec- ce cpoToCHHTe3a 113 C0 2 H 20 11 no.n; .n;eii:cTBHeM coJIHe•moro cseTa. B opraHH3M ;1rnsoTHhIX nocTyna:10T c TIIIIll,eii:. Co,n.epmamrn yrJ'Ieso.n;os B KJieTKe (B KpOBll 0, 1--0, 12 'fr,), B paCTII'reJibHOH KJieTKe .n;o 90 'Yr, 1-5 'Yo (1rny6HH KapToc}>eJIJI, ceMeHa). YrJieBO,D;hI - :JTO yrJ1eso.n;opo.n;Hhie coe.n;IIHeHIIH. 06I11,aJI cpopMy.w: Cn(H 20)m. KJtaccu<jJu 1cau,u.R y2JtrnoiJ06: 1. IlpocThie yrJieBOJJ;hl - MOHocaxaptt,n;hl. B 3aBttCHMOCTH OT qIIcJia yrJie- mpuOJbl (rJIHll,epaJih,IJ;erII.n;), mempoJbt (spttTpo3a), nntmo.3bt (p11603a, .n;e30Kcttptt603a, sxo.n;amne B cocTaB HYKJieHHOBhIX KMCJIOT, AT<l>), zeKCOJbl (rn10xo3a, cppyKT03a). BOJJ;HhIX aTOMOB B HX MOJieKyJie pa3JIH'IaIOT: 2. CJIO)KHhie yr.'IeBOJJ;hI - 3TO ITOJIMMepbI, COCTOJ!Ill,He H3 MOHocaxapH,ll;OB, coe,n;HHeHHhIX rJIMK03HJJ;HOI'i: CBH3hIO. Pa3JIIFrnroT: • O.llUZOCaxapuiJbl - yrJieBO.D;hI, IlOCTpOeHHhie H3 tte60JihIIIOro qucJia (2-10) MOHocaxapH,D;HhIX OCTaTKOB, paCTBOpHMbie, HMeIOT CJia,ll;KHH BKyc; • iJucaxapuiJbl BOH caxap - yrJieBO,D;hl, 06'he,D;HHHIOill,IIe ,D;Ba MOHOCaXaptt.n;a (rmw;e- caxapo3a, COCTOMT H3 O.D;HOH MOJICKYJihl rmoK03hl H O,D;HOH + rJIIOK03a; JiaKT03a = + cppyKT03a); MOJieKyJihI raJiaKT03hr; MaJihT03a = rmoK03a K03a + raJiaKT03a; • noJtucaxapuiJbt - caxapo3a = rJIIOKo3a rmo- cJIO)KHhie BhICOKOMoJieKyJiapHhie yrJieso.n;hl, o6pa3o- BaHHhie COTHHMH H ThIC.HqaMH MOJieKyJI MOHocaxap11,n.oB, HepaCTBOpIIMhle, He HMeIOT C.1'Ia,D;Koro BKyca. ::ho JIHHCHHhie 11 pa3BeTBJieHHhie TIOJIHMephl, MOHOMepbl KOTOphlX coe.n;ttHeHhl l'JIIIK03MJJ;HOH CBJI3hl0; • ZOMono.riucaxapuiJbl - COCTOHT II3 MHO)KCCTBa OJJ;HHaKOBhIX MOHOcaxa- pH.!l;HhIX OCTaTKOB (KpaxMa.1'1, l'JIHKOl'eH, u;eJIJIIOJI03a); • zemeponoJtucaxapuiJbl - COCTOHT II3 MOHOCaxapH,D;OB pa3HhIX BIIJJ;OB (re- nap1rn). <PyHKU,UU yZ./l€60006.' 1. CTpyKTypHaa (cTpOHTeJihHa.H) - BXO)J;HT B COCTaB KJieToqHOH CTeHKll pacTeHlIH, 6aKTepHH, rp1160B. 2. 3HepreTnqecKaa 1 7 ,6 3. 4. 5. - IIPII OKIICJieHHII 1 r yrJieBO)J;OB BhI)J;eJIReTC.H K,Um :meprIIn. 3anacarow;aa - KpaxMaJI y pacTeHHH, rJIHKOl'eH y )KHBOTHhIX. CocTaBHOtt KOMnotteHT ,UHK, PHK, AT<I>, <I>A,U 2 -, HA,U', HA,U<I>·. 3alll,HTHa.H - pell,eIITOphl TKaHeBOll COBMeCTHMOCTH, XHTHHOBhlll IIO- KPOB qJieHHCTottorttx. HyKJieHHOBhie KHCJIOThI HyKJteuno6bte KUCJtOmbt (oT JiaT. nukleus- a.n;po, snepBhre 6hIJIH o6HapymeHhl B a.n;pe) - cJio:arnhle npHpo,n;Hhie BhICOKoMoJieKyJIHpHhie coep;irneHtta, 06ecnequsa10I11,ne xpatteHHe II nepe.n;aqy Hacnep;cTBeHHon nHcpopMau;mr B mnBhIX opraHH3Max. CocTaBJIHIOT 1-5 % HhI MOHO- H TIOJIHHYKJieOTH)J;aMH. OT cyxoiI Macchi KJieTKH H npe.n;cTaBJie-
Y i1te60dbt. II yK1teu1to6ble KUCJWmbt. )_( / / K 25 A3or~1cToe OCHOBaH11te aAeHlltH, "1/1111 ryaHlltH, 11tmt 4ll'IT03111H, ll'IJlH OcrnTOK Tll'IM"1H cpoc<f>0pHoll'I Ki.1CJ10Tbl P111c. 3. CTpoeHVle HyKneorngci II 0Jtutty101eomuab1 Maccoi1. JIHHettHhre 6ttorroJIHMepb1 c orpoMHOH MOJieKym:i:pttoii Motto.111.epa.l1tu HYKJteu1toBbLX KucJ1om m;tte H3 a30THCToro OCHOBaHH.H, caxapa JIOThI (pHC. .HBJI.HIOTc.a HyKJ1eomuabt, cocToB- rreHT03hI H OCTaTKa cpoccpopHOH KHC- 3). A.3omucmbte octtoBaHUR rrpe,n;cTaBJieHhr npoH3BO.L\HhIMH rrypn:Ha, nn:pHMH- .n;n:tta. IlypHHOBbie OCHOBaHH.H: OCHOBaHH.H: u; - IJ;HT03HH; T - A - a,n;eHHH; THMHH; y - r - ryaHHH. I1HpHMH,ll;HHOBhIC ypau;HJI. TIIMHH OTJIHqaeTC.H OT ypaIJ;HJia HaJIHqHeM MeTHJihHOH rpyrrrrhI (-CH;)· B o6pa3oBaHHH HYKJieoTn:,n;oB rrpHHHMaIOT yqacTn:e ,n;Ba MOHocaxapu.n;a Jbt. 3To pn:603a - B PHK H ,n;e3oKcn:pn:6o3a - JieOTH,ll;OB .HB.'l.HeTc.a ne1tmo- B .IJ:HK. TpeThHM KoMrrotteHTOM HYK- ocmamoK rpocrpopHou KUCJLombt - rpocrpam. B 3aBHCIIMOCTn OT qn:cJia cpoccpaTHhIX rpyrrrr pa3JIH'18IOT MOHO-, ,L\H- H TpHcpoccpaTbl HYKJieOTH,11,0B. Coe,n;HHeHH.H a30THCToro octtosattn:.a H caxapa Ha3hmaIOTCH a c rrpn:coe,n;n:HeHHhIM cpoccpaTOM - 1tyKJteomuoa.Mu. HyK.rteoJuaa.111.u, Ha3BaHn:e HYKJieOTM,11,0B o6yCJIOBJieHO BXO,ll;HIIJ;HMH B HX COCTaB a30THCThIMH OCHOBaHHHMH H o603HalJ:aeTC.H 3arJiaBHhIMH 6yKBaMH. B rrpn:po.n;e cym;ecTBYIOT ,n;Ba Tttrra HYKJieHHOBhrx KHCJIOT, OTJin::qaJOIIVIXCB no COCTaBy, CTpoeHHIO H cpyHKIJ;lIHM: .IJ:HK JIOTa, PHK - .n;e30KCHptt60HYKJieHHOBa.H KHC- pn:6ottyKJieHHOBa.a KHCJIOTa • .ll:HK (,z:i;e30KCHpH60HYKJieHHOBaH KHCJIOTa) TpexMeptta.a Mo,z:i;eJih CTpoeHHH .IJ:HK 6hIJia rrpe,nJiometta ,nm. YoTCOHOM II <I>. KpHKOM B 1953 r. MoJieKy JI a ,[:t;HK cocTOHT H3 ,n;Byx rroJIMHYKJieoTH.n;- Hhix u;errei1, 3aKpyqeHHhIX B rrpaBOCTOpOHHIOIO CIIHpaJib (pHC. CIIHpaJIH paBeH 2 3,4 HM, mar CTIHpaJIH - 4). ,[:t;n:aMeTp HM. 0.n;HH BlITOK HMeeT 10 nap HYKJieOTH,ll;OB. HarrpaBJieHHe u;enei1 ,IJ;HK B3aHMHO rrpoTHBorroJiomHo. Ha KOHIJ;e BCer.n;a pacrrOJIOJKeH OCTaTOK cpoccpopHOH KHCJIOThl, Ha 60,n;Ha.H OH-rpyrrrra. 3' KOHIJ;e - 5· CB0- OcTOB ,[:t;HK cocTaBJIHIOT MOJieKyJihI ,ne30KCMpH6o3hI H cpoccpaTOB. Ilapbl a30TlICThIX OCHOBaHHH HYKJieOTH,ll;OB pacrroJioJKeHhl BHYT- PII cnHpaJIH. Mem.n;y ocHoBaHH.HMH o6pa3yIOTCH Bo,n;opo,n;Hhre CBH3H. OcTOB 06pa3yeTCH 3a cqeT KOBaJieHTHhIX CBH3e:H (pn:c. Ba npHHIJ;Hrrax Ko.MnJLe.MeHmapttocmu 11 5). CTpOeHHe ,IJ;HK OCHOBaHO anmunapaJLJLeJtb1wcmu. ,[:t;JIHHa MO- JieKyJihI MomeT .n;ocTHraTh HeCKOJihKHX caHTHMeTpoB. MoJieKyJrn:pHhIM Bee .n;eCHTKH H COTHH MHJIJIHOHOB KJieTKH qeJioBeKa - oKoJio d (,n;aJibTOH). CyMMapHa.H ,ZJ;JIHHa ,[:t;HK .a,n;pa 2 M. B ayKap110T11qecKHX KJieTKax ,[:t;HK 06pa3yeT
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35 K1temo'tnble <jJopMbl J1Cu3nu. HaiJq,apcm6o npoKapuombt JKryTHKH He orpaHHqeHhl MeM6paHOll, HMCIOT BOJIHHCTYIO cpopMy H COCTO.HT H 3 ccpepHqecKHX cy6'he,ll,HHH11, 6eJIKa cpJiareJI.iIHHa. 3TH cy6'he.n;HHH11,h1 pacnoJIO;I<eHhI 110 c1rnpaJIH H o6pa3yIOT IIOJihlll 11,HJIIIH,ll,p .n;HaMeTpOM 10-20 HM. KoJIIIqecTBO II pacnoJioJKeHHe mryTIIKOB MOJKeT 6hlTh pa3JIHqHhIM. Ha KJieTOt!HOtt CTeHKe neKOTOphIX 6aKTepHll: BH,ll,Hhl TOHKIIe BhlpOCThl (naJIO'IKOBH,ll,HhlC 6eJIKOBhie BhICTYTihI), KOTophie Ha3hrnaroTc.H nuJtu, IIJIH cpHM6p1rn. 0HII KOPO'le H TOHhllle mryTHKOB H npe.n;cTaBJI.HIOT co6oi1: KOpOTKHe IIOJlhie 11,IIJIHH,ll,Phl II3 6e"'rna nHJIHHa. CJiymaT ,ll,JI.H npHKpenJieHH.H 6aKTepHll: K cy6cTpaTy HJIH .n;pyr K .n;pyry. f{btXanue y 6aKTepHll: ocy111,eCTBJI.HeTC.H B Me30COMaX, y CIIHe3eJieHhIX BO,ll,0pocJiell - s UIITOrrJia3MaTH'leCKHX MeM6paHax. XJioponJiaCTOB II .n;pyrHx KJie- TO'IHhIX opraHeJIJI, OKpymeHHhIX MeM6paHOM, HeT. Pa3MHOJICG/Omc.H npOKapHOThl nyTeM 6uuapuoio aeJteHUJl c o6pa3oBaHHeM .n;syx ,ll,O'lepHHX KJieTOK. Ilepexo.n; K .n;eJieHHIO orrpe.n;eJI.HeTC.H OTHOIIleHHeM o6'heMa a.n;pa K 06'heMy ll,HTOIIJia3Mhl. Ilepe.n; KJieTO'IHhlM .n;eJieHHeM npOIICXO,n;HT peIIJIHKall,H.H .[(HK, BO speM.H KOTOpoll: Me30C0Mhl y,n;epJKIIBaIOT reHOM s orrpe,n;eJieHHOM rroJioJKeHHH. Me30COMhI MoryT npHKperrJI.HThC.H K rreperopo,n;KaM Mem,n;y ,n;oqepHHMH KJieTKaMH H yqaCTBOBaTb B CIIHTe3e se111,eCTB KJieTO'lHOll CTeHKH. J{eJtenue np01wpuom (pIIc. 12). CHaqa.rra ,n;eJIHTC.H HYKJieoH.n;, a 3aTeM UHTO- rrJia3Ma. CKopocTh pa3MHomeHH.H BhlCOKa. TaK, 6aKTepH.H KHIIle'IHoll: rra.rro'IKH KaJK,n;hle y 20 MHHYT y,n;saIIBaeT CBOIO 'IHCJieHHOCTh. 6aKTepHH Ha6JIIO,n;aeTC.H H ll0Jl060e pa3MHOJ/Cenue, HOB caMOll rrpHMHTHB- HOll cpopMe. IlpII 3TOM HeT o6pa30BaHmI raMeT H CJIH.HHH.H KJieTOK, a rrpOHCXO,ll;HT 1 2~ 4 [CQJ51} II [~] III 4 PMc. 11. CTpoeH11e 6aKrnp11anbHOVi Kf1eTKl1: 1 - lKf)'TlllK; 2 - Kancyna; 3 - 4>0TOClllHTeT111YeCKa11 MeM6paHa; 4- n11n111 (4>111M6p111111); 5- M830CO- P111c. 12. CxeMa AeneH11R np0Kap110T: KJ18TOYHall o60J10YKa; MeM6paHa: 6 - KOllb~eBall MOJ1eKy11a )lHK; 7 - ~111TOn11a3Ma; 8 - 3anacHb1e n111TaTel1bHb1e se~ecTBa; 9 PH6oc0Mb1; 10- nna3MarnYecKaR MeM6paHa; paHbl; 11 - Kf18TOYHa11 CTeHKa MeM6paHbl Ma; 3 - ~111rnnna3Ma. I - II - 2 - 1- nna3MaTlllY8CKall MOJ18KYJ1a )lHK; CTa./:111111 penn111Ka~11111 4 )lHK; CTa./:11111 Cl1HTe3a nna3MaTW-18CKOVi MeM6- Ill - ¢>opM111posaH11e nonepeYHOVi
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06Uj-aJt 6uo1wiuJt. 46 Oc1W6bl 1.fUmo.ioiuu XpoMamuHOBbte cmpyKmypbt: XpoMamuH u xpoMOCOMbL - ,n;se cpopMhI cym;ecTBOBamrn: O)J,Horo MaTepuaJJa: B 51p;pax He,n;eJIRIIJ;llXCR K.JleTOK M€H030M - xpoMaTHH, B p;eJUIIIJ;HXCR MIIT030M MJIH xpoMOCOMbI. XMMMqecKMH cocTas: ,D;HR (B cpopMe .11:e30Kcup1160HyKJ1eonpoTeM;:i;a); rttcTOHOBhie, nerMCTOHOBhie 6emrn:; cpepMeHTbI (,Il;HK-no.1MMepa3a M )J,p.); M-PHK, T-PHK, MHHepaJihHbie KOMilOHeHThl. B 33BHCHMOCTM OT cpyHKUHOH3JihHOro COCTORHMJI xpoMaTHHa pa3JinqaJOT reTepoxpoMaTMH u ayxpoMaTMH. I'emepoxpoMamU1-i - reneTMqeCKH tteaKTMB- Hhie yqaCTKH xpoMaTMHa. Ilo,n; CBeTOBhIM MHKpOCKOilOM OH MMeeT BH,ll; rJihI60K HJIH rpaHyJI, HHTeHCMBHO OKpaUIMBaeTCJI H npe,n;cTaBJJReT co6oil KOH,n;eHcn:po- B3HHbie (cnupaJIH30BaHHbie) yqaCTKM xpoMaTHHa. 3yxpoMamun - reHeTMqe- cxn: aKTHBHhie yqaCTKM xpoMaTMHa, npM CBeTOBOH MMKpOCKOTIHH Hepa3JIHq:MM, CJrn6o oKpaUin:saeTCJI M rrpe,n;cTaBJJReT co6oi1 ,n;eKoH,n;eHcMpoBaHHhie Cn;ec1mpaJIM30BaHHhie, pacKpyqeHHhie) yqacTKH xpoMaTnHa. XpoMocOMbL (oT rpe'L chroma rrpe,n;JiomeH B 1888 r. ll,BeT, KpacKa, n soma - TeJJo); TepMMH HeMeu;KMM yqeHhIM Bn:JJhreJJhMOM BaJih,n;eilepoM) - rrpe,n;cTaBJIJilOT co6oii KOH,n;eHCMpOB3HHhIH xpoMaTMH BO speMJI MMT033 HJIH MeH033. OcHOBY xpOMOCOMhI COCTaBJJ.HeT ,n;syxu;erroqeq:HaH MOJieKyJJa ,D;HK, KOTOpaa no,n;sepraeTCH yirna,n;Ke, rrpno6peTaJI orrpe,n;eJJeHHYIO TpexMeptty10 CTPYKTYPY, HJJH KOHcpopMaun:10. Bh1,n;eJ1HIOT HecKOJihKO yposHeii rrpocTpaHcTBeHHoii yKJJa,n;KM - KoMnaKmu3au,uu /l,HK (cM. pa3,n;eJI «I'eHeTnKa» ). XpoMocoMhI u3yqa10TcJI B MeTacpa3e MMT03a. MeTacpa3HaR xpoMocoMa coCTOI:IT n:3 ,n;syx xpoMamua (pMc. 15). HMeeTcJI nepeu'lna.R nepemRJKKa, HJIH u,ettmpoMepa, KoTopaa ,n;eJIMT xpoMocoMy Ha nJte'lu. HeKOTOphle xpoMocoMhl HM€lOT 6mOpU'l1-tYIO nepem.RJKICY H cnym- HUIC (yqacTOK KOpOTKOPO IIJJetia, OT,n;eJIHeMbIH BTOpntIHOii rrepeTHJKKoii). KoHUbI xpoMocoM H33hIB3lOTCH me.1wMepaMu. <f.>yHICU,UU R8pa: xpaHeHn:e H3CJI€,Zl;CTBeHHOH MHcpopMaUMM H nepe,n;aqa ee )l;ffqepHHM KJleTKaM B npou;ecce ,n;eJieHHH, peryJIHIJ:HH n: KOHTpOJih npoueccos JKM3He,n;eHTeJlhHOCTH KJieTKH, yqacTHe B CHHTe3e 6eJIK3 (TpaHCKPHIIUHJI), M€CTO o6pa30BaHMH cy6'he,ZJ;MHHU pH60COM. PMc. 15. XpoMocOMbl: 1- MeTaL1,eHTJ)11YecKasi; 2- cy6Mernu,eHTp11yecKasi; 3, 4- aKpou,eHTp11YecK11e. CTpoeH11e PaaJIHqHhJe THIIhI ay1mpu0Tuqec1mx .KJieTo.K 3yKapMOTHtieCKHe opraHH3MhI ,n;eJJHT Ha TpH uapCTBa: pacTeHI:IH, :IKHBOTHhlX H rpn:6os. PaccMoTpes o6m;Hii 1rnaH cTpoeHMJI KJieTKH 16, 17), XPOMOCOMbl: 5 - LJ,eHTPOMepa; (pnc. 6 - BTOpwmasi nepeTSJ)KKa; 7 CnYTH11K; 8 - xpoMaT11Abl; 9 - Hble oco6eHHOCTH pacTHTeJihHOH H :IKHBOTHOM TeflOMepbl B pa3,n;eJie «BOT3HMK3» ). KJieTKI:I MOJKHO Bhl)l;eJIHTh OTJIMtIMTeJih- (KJI€TKM rpM60B paccMaTpnBalOTC.H
47 K11,emo'lnbie </JopMw JtCUJnu. HaiJu,,apcmBo 3y1wpuomw CpaBueuHe cTpoeHHH pacTHTeJibHOH H ;KHBOTnoii KJieTOK PacTHTeJihHaH KJieTKa IlpH3HaKH IIJiaCTH/:lbI XJiopornracTbI, :lKHBOTHaH KJJeTKa 0TCYTCTBYK>T xpOMOTIJiaCTbI, JieHKOTIJiaCTbl reTepoTpOcpHbIH Cnoco6 ABTOTpOcpHbIH nuTaHHH ( cpoTOTpocpHbIH' ( canpoTpOcpHbIH, xeMOTpocpHbIH) napa3HTH'leCKHH) B xJioponJiacTax, B MHTOXOH,llpHHX CuuTe3 AT<I> MHTOXOH,llpH.HX Pac~enJieuHe B xJioponJiacTax H Bcex Bo Bcex 'laCTHX KJieTKH, r,11e AT<I> qacT.HX KJieTKH, ueo6xo,11HMbI 3aTpaTbI r,11e Heo6XO,!IHMbl 3HeprHH 3aTpaTbl 3HeprHH KJieTO'IHhIH y HH3IUHX pacTeHHH Bo Bcex KJieTKax n;eHTp ll;eJIJIK>JI03HMI PacnoJiomeua cuapymH OT KJieTO'IHa.H KJieTO'IHOH MeM6paHbl 0TCYTCTByeT CTeHKa BKJIK>'leHH.H 3anaCHble TIHTaTeJibHbie Be~eCTBa B BH):le 3epeu 3anaCHbie TIHTaTeJibHbie Be~eCTBa B BH):le 3epeH H KpaxMaJia,6eJIKa,KaneJih KaneJih (mHpbI, yrJieBo,11 M8CJia; BaKyOJIH c KJieTO'IHbIM rJIHKOreu); KOHe'IHbie COKOM;KpHCTaJIJibICOJieH npo,11yKTbI o6Meua, KPHCTaJIJibl COJieli; nHrMeHTbl BaKyoJIH Kpynubre noJiocTH, CoKpaTHTeJibHble, 3aTIOJIHeHHbie KJieTO'IHblM TIH~eaapHTeJibHbie, COKOM - BO,!IHbIM paCTBOpOM pa3JIH'IHbIX Be~eCTB, .HBJI.HK>~HXC.H Bbl):leJIHTeJibHble B8KYOJIH. 06bl'IHO MeJIKHe 3anaCHbIMH HJIH KOHe'IHbIMH npo,11yKT8MH.0CMOTH'leCKHe pe3epayapbI KJieTRH CB.H3b Mem,11y I1Jia3MO):leCMbl KJieTKaMH Ilepeuoc qepe3 MeM6paHhI, KJieTO'IHble KOHTaKTbl ,lJ;eJieHHe KJieTO'IHa.H neperopo,11Ra IlepeT.amRa 06pa3yeTC.H B Il;HTOTIJI83Mbl npH CTpOHTC.H B n;eHTpo6emHOM n;eHTPOCTpeMHTeJibHOM MHT03e H MeH03e uanpaBJieHHH uanpaBJieHHH rJIHKOKaJIHKC Ha 0TCYTCTByeT llMeeTC.H aapymuoii: MeM6paue
06~aJt 6uonozuJt. Ocn06bL 1fUmonozuu 48 J - -·- \ ~ft=~~..-- - --- 22 -21 2 20 -J9 18 3 .. 17 16 5 6 PMc. 16. CrpoeHi.-ie pacrnTeJTbHOiii KJTeTKi.-i : PMc. 17. CrpoeH11e :>Kl1BOTHOiii KJTerKi.-i: I -3KJou,111- 1- roa; 2- ceKperopHbli1 nyab1peK; 3 KJTeTOYHafl CTeHKa; 2 - flAepHafl o60JTOY- 3 - flASPHafl nopa; 4 - flAPblWKO; 5 flAPO; 6- xpoMarnH ; 7- rflaAK"1iii 3HAOnna3MaT"1Y9CK"1ill peTi.-iKynyM ; 8 - nnaJMantYeCKafl M9M6paHa ; 9 - U,"1TOnna3Ma; 10 - Ml1 KKa; porpy604Kl1 ; 11 - M11KPOt:fmnaMeHTbl; 12 - annapar ronbA)f(11 ; 13 - n113oc0Ma ; 14 - Honnacr; 15 - KfleTOYHbli1 COK ; 16 - XOHAP1111 ; 17 - rpaHa; 18 - ponnacra; ro- Ml1TO- MeM6paHa xno- 19 - xnoponnacr; 20 - p116oc0Mb1 - n11U-1esapi.-irenb4 - annapar ronbA)f(11 ; 5 - M11Kpo<f>11naMeHTb1; 6 - MmoxoH.Qp1111 ; 7 - wepoxosarb1i1 3HAonna3Marw-iecK11i1 pernKynyM; 8 - p116ocOMb1, Ha11 aaKyonb; CBS'l3aHHb18 c 3HAOnna3MaTll1Y9CKl1M per11KynyMOM ; 9- nna3M8Tl1Y9CKaR MeM6paHa; 10 - U.11TO- nna3Ma ; 11 - 13 - 3yxpoMarnH ; 12 - flAPblWKO; 14 - o6onOYKa; 16 - r erepoxpoMarnH ; flA8PH8fl n o pa; 15 - flAepHafl AB9 U,9HTPl10!111 , pacnonmKeHHble nOA npl'IMblM yrnOM Apyr K Apyry; 17 - CB060AHble c 3H.QOnna 3MarnYeCK11M pernKynyMOM; 21 - p1160COMbl; 18 - wepoxosarb1iii 3HAonna3MarnYecK11ill per11Ky- 20 - rnaAK11i1 3HAOnna 3M8Tl1Y9CKl1iii per11KynyM; 21 - JTl-1HOU,11T03Hbliii ny3blpeK ; 22 - n11HOU,11T03; 22 - CB060AHbl9 p1160COMbl; 23 - KJTeT04Hafl CTeHKa COCeAH11X KJTeTOK; 24 - nnaJ- nyM; MO.QeCMbl J11130COMa; 19 - M11KPOTpy6o4K11; 23 - Ml1KPOBOPCl1HKa; 24 - nornou_\eH11e 11!111 ceKpeU,11R y OCHOB8Hl1R M11KPOBOpCl1HOK; 25 - RAPQ 06MEH BEW.ECTB M nPEBPAW.EHME 3HEPrMM Bee JRHBhie opraHH3MhI, o6HTaroru;He Ha 3eMJie, .HBJUIIOTCH OTKphIThIMH CHCTeMaMH, ,ll;Jl.H KOTOpblX xapaKTepeH HerrpephIBHO rrpoTeKarorn;Hii o6MeH Bern;ecTB H aHepruu c 01<pyJ«a10rn;eif cpe,n;oii. 06MeH Bern;ecTB H 3HeprHH - OCHOBHOe CBOHCTBO JRHBoro. Bee rrpou;ecCbl JRH3He,n;enTeJihHOCTH (pocT, pa3BHTHe, pa3MHOiKeHHe H ,n;p.) HeB03MQ;KHhl 6ea IlOJiy'leHH$l H3 OKpyJ«a10rn;ell: cpe,n;bl B npou;ecce IIHTaHH.H Bem;eCTB ( CTpOHTeJihHoro MaTepHaJia) H 3HeprHH, Heo6xo,n;HMhIX ,ll;JIH. nocTpoeHMH. XMMHqecKHX coe,n;HHeHHM opraHH3Ma, a TaKJ«e ,ll;JI.H COBeprneHMJf pa3JIM'IHOH pa60Thl. CoBoKynHoCTh pea1<u;uif o6MeHa, npoTeKa10w;ttx B opraHH3Me, Ha3blBaeTcn Me ma60JIU3MO.M.
06Men Beiq,ecmB u npe6paiq,enue :mepzuu 49 MeTa6oJIH3M BK.moqaeT B cefor ,[(Ba rrpou;ecca: 1. A11a60JtU3M (accn::vrn:m1u;ttH, rrJiacTn:qecm1ii o6MeH) - coBoRynHocTh 3H.L\O- TepMIIqecRttx peaKu;Hii 6IIOCHHTe3a, KOTOpble npHBOl\flT K o6pa3oBaHHIO CJIO)KHhlX Beru;ecTB H3 60J1ee npOCTbIX ( CHHTC3 6eJIKa, rJIHKOf'eHa; cpOTOCHHTe3, xeMOCHHTe3). 2. Kama60JtU3M (,n;IICCHMIIJIHD;IIH, ;mepreTIIqecKIIH o6MeH) - COROI{yn- HOCTh 3K30TepMIIqeCKHX peaKD;HH pacna,[(a II OKHCJieHirn BhlCOKOMo.1eKYJIHpHbIX Beru;ecTB .n;o 6oJiee npocTbIX. 8TM rrpou;eCChl B3aMMOCBH3aHhl BO BpeMeHM M rrpocTpaHCTBe, flB.1flIOTCfl 3BeHbHMH MeTa6oJIM3Ma. Pea.KD;IIM 6MOCMHTe3a HeB03MOIBHhI 6e3 3HeprMM, KOTOpaa BhI.n;em:i:eTCH B peaKD;llHX 3HepreTIIqecKoro o6MeHa. PeaKD;HH ,n;IIcc11M11Jinu;nn ne MYT 6e3 <t>epMeHTOB, 06pa3y10ru;nxcH B peaKu;Max rrJiaCTMqecKoro o6MeHa. Ilo THilY IlHT3HHH BCe )KHBhie opraHM3Mhl rro.n;pa3,n;eJIHIOT Ha: 1. Aemompo<jJ11bte - crroco6Hhre CMHTe3MpoBaTh opraHMqecRne Beru;ecTBa H3 HeopraHnqecKnx. <l>OTOCHHTeTllKll (cpOTOaBTOTpO<t>hI) - K HMM OTHOCHTC.H 3eJieHhie pacTe- HllH ll <t>oTOCHHTe3npyroru;ne 6aKTepnII, KOTOPhie MCII0Jlh3YIOT 3HeprMIO COJIHeqHoro CBeTa. Bee OCTaJihHble )KllBhie cyru;ecTBa MCTIOJih3YIOT 3HepI'HIO, 3aKJIIOqeHHYIO B XHMnqeCKllX CB.H3HX. XeMOCMHTeTMKM (xeMoaBTOTPO<l>hI) - K HMM OTHOC.HTC.H HeKOTOphie 6aKTepnn, KOTOphie llCIIOJih3YIOT 3Heprnro, BhI,ll;eJI.HIOiu;YIOC.H npM OKllCJieHllll HeopraHnqecRnx coe,n;nHeHnii ( ceposo.n;opo,n;a, aMMMa.Ka, IBeJie3a M ,n;p.). 2. I'emepompo<jJHbte IIIHHCTBO 6aKTepnii, IBllBOTHhie, rpn6h1, He3eJieHhre pacTeHIIH, 6oJih- He cnoco6HhIX caMOCTOHTeJihHO CMHTe3MpoBaTh opraHM- qecKHe Beru;ecTBa ll MCIIOJih3YIOiu;MX ::meprnIO XMMlfqeCKHX CBH3eii I'OTOBhIX oprrurnqecKHX coe,n;nHeHnii. Carrpo<t>nThI (carrpoTpo<t>Hhre opraHn3MbI) - rrttTaIOTCH opraHnqecRnMM Beru;eCTBaMn MepTBbIX TeJI (6aRTepnn, rpn6hI). Ilapa3llTbl (napa3llTMqecKMe opraHH3MbI) - IIHTaIOTCH opraHnqecKIIMll Beru;eCTBaMn )KHBbIX opraHH3MOB (6oJie3HeTBOPHhie 6aKTepHM, rrapa3HTnqecKne pacTeHn.a, rptt6hI, IBHBOTHble). 3. Mun:compo<jJHbte opzanu3Mbl HHH) - (3BrJieHa 3eJieHa.a, HaceKOMO.H,D;Hhie pacTe- MOI'YT IIMTaThCH KaK aBTOTpo<t>Ho, TaK ll reTepoTpo<J:>Ho. Ilo cnoco6y nocTynJieHHH IlHTaTeJibHhIX Bern;ecTB opraHH3MhI Ha ZOJW30U1tble - 3aXBaTbIBaIOru;He rrnru;eBble qacTHI.l;hI M pa3,n;eJIHIOT ZOJLO<jJUmltble - BCa- CbIBaroru;tte pacTBopeHHble aeru;ecTBa. Ilo OTHomeHHIO K KHCJIOpO,ll;y pa3JIMqaroT: Aapo6nbte opraHH3MhI (Ty6epKyJie3HaH rraJIO'IKa) - ,n;hIIIIaT Kttcnopo- ,n;oM, Heo6xo,n;HMhIM ,ll;JIH OKllCJIMTeJihHO-BOCCTaHOBHTeJihHbIX peaKD;llll npH TKaHeBOM ,ll;bIXaHllH. Anaapo6Hbte OKMCJinTeJin. H cmpozue opraHM3Mhl - HCTIOJih3YIOT BMeCTO KHCJiopo,n;a ,n;pyrMe 0Hn .n;ennTca Ha necmpozue (,n;M<t>TepMiiHaH rranoq.Ka) (B036y,n;nTeJIH CTOJI6HHKa, raHrpeHhI; noqBeHHhie 6aKTepMH).
06UfaH 6uo.1wzuH. Ocno6&i lfUmo.nozuu 50 TaKHM o6pa30M, rroTOK Bem;ecTB H 3Heprww o6ecneqwBaeT caMoo6ttoBJieHHe H caMOBOCI1pOH3Be):(eHHe opraHH3MOB. «l>oTOCllHTe3 <Pomocunme3 - 3TO rrpou,ecc npeo6pa3oBaHHH coJIHeqHorr 3HeprHH B rroTeHu,HaJihHYIO 3HeprHIO XHMHqecKHX CB.H3e:H: opraHHlfeCKHX se~eCTB. 8To TaIC>Ke: rrpou;ecc o6pa3oBaHHH opraHMlfeCKHX sem;ecTB npw yqacTHH 3HeprHH cseTa. OH cBorrcTseHeH KJJeTKaM, co,D;epmam;HM cneu;waJihHhie cpoTOCHHTe3npyrom;ne nMrMeHThI, rJJaBHhIM H3 KOTOpbIX HBmieTC.H XJIOpocpMJIJI; e):(MHCTBeHHhIM rrpou,ecc B 6noccpepe, Be,D;ym;nii: K 3anacaHnIO 3HeprnH 3a cqeT ee BHernHero MCTOqHHKa; 3BeHO 6HoreoXHMHqecKHX ll;HKJIOB Ha 3eMJie M OCHOBa Bcex u;erreli IlHTaHMH. 3arraceHHaH B rrpo):(yKTax 3HeprH.H - OCHOBHOH HCTOqHnK 3Heprwn )l;JIH qeJIOBe'IeCTBa. EmerO)l;HO B pe3yJihTaTe cpOTOCHHTe3a Ha 3eMJie 06pa3yeTCH 150 MJIPA TOHH opraHH'IeCKOro Bem;ecTBa (rrepsnqHaH rrpo)l;yKu;nH) M BhI):(eJIHeTCH 200 MJIPA TOHH CB060):(HOro KHCJIOpO):(a. KpyrosopoT KHCJIOPO)l;a, yrJJepo):(a M )l;pyrHX 3JieMeHTOB, BOBJieKaeMhIX B cpOTOCHHTe3, ITO)l;):(epmHBaeT cospeMeHHhIH coCTaB aTMOCcpephI, He06XO):(HMhIH )l;Jl.H cym;eCTBOBaHH.H Ha 3eMJie COBpeMeHHhIX cpopM mH3HH. <l>oTocnHTe3 rrpen.aTCTByeT yBeJinqeHHIO KOHu;eHTpau;un C0 2 , rrpe):(OTBpam;a.a rreperpeB 3eMJIH BCJie)l;CTBHe TaK Ha3hrnaeMoro « rrapHHKOBoro 3cpcpeKTa». <l>oTOCHHTe3 B pacTeHHHX ocy~ecTBJI.HeTcH B xJioporrJiacTax. OH BKJIIOqaeT ,ZJ;Ba 3Tarra, IIOCJie,ll;OBaTeJibHO CBH3aHHhie Mem)l;y co6oii: I. 8'rarr 110rJiom;eHnH H rrpeo6pa3oBamrn 3Heprwn ( ceemoeou npou,ecc ). II. 8Tarr rrpeBpam;emrn Bem;ecTB ( me.Mttoeou npou,ecc). ,[(Ba rrpou;ecca cpOTOCHHTe3a BbipamaIOTC.H OT ):(eJihHhIMM o6o6~eHHhlMM ypaBHeHHHMH. I. 12H2 0 + 3Hepm.a cBeTa ~ 12H2 + 60 2 i + 3Heprn.a AT<l> (cBeTOBOH rrpou;ecc). II. 6C02 + 12H2 + 3Heprn.a AT<l> ~ C6 H 12 0 6 + 60 2 i (TeMHOBoii: rrpou;ecc). 06o6m;eHHOe ypaBHeHne <PoTOCHHTe3a: 6C0 2 + 6H 20 :JHeprHfl CBeTa --x-JI-op_o_<jl_HJI_JI__,,. C 6H 120 6 + 60 2 i Ceemoeou npou,ecc rrponcxo.zi;nT B muJta1CouiJax xJtoponJtacmoe, me.r.rnoeou - e cmpo;ne, To ecTh 3TH rrpou;ecchl rrpocTpaHCTBeHHO pa306m;eHh1. IlJiaCTH.D;hI xapaKTepHhI TOJihKO .zi;JI.a pacTeHHM. 0Hn He Hali):(eHhI y rpu6oB My IKHBOTHhIX, 3a HCKJIIOqeHHeM <fJoTOCMHTe3upyrom;Mx rrpocTeii:rnnx. B 3aBMCHMOCTH OT OKpaCKM, CB.H3aHHOH c HaJIHlfHeM HJIM OTCYTCTBMeM rrnrMeHTOB, pa3JIHqaIOT TpH OCHOBHhIX THIIa IIJiaCTH)l;: XJtOponJtaCmbt (3eJieHoro u;BeTa), xpo;nonJtacmbt (JKeJIToro, opaHJKeBoro, KpacHoro u;BeTa), JteuKonJtacmbt (6ecu;BeTHhre). 0)l;HH rrJiaCTH)l;hI MoryT rrpeBpam;aThCH B )l;pyrue. CTpoeHHe rrJiacTH,n; MO/KHO paCCMOTpeTh Ha rrpHMepe XJIOPOIIJiaCTOB. XJtoponJtacmbt - 3TO )l;ByMeM6paHHhie opraHOM)l;hI, BHyTpeHHHH MeM6paHa KOTOpbIX 06pa3yeT muJtalCOUObl, qiopMnpyrom;ue epattbl, Mem,n;y KOTOpbIMM Ha-
06MeH. 6€Ufecm6 u npe6paUfenue :Jnepzuu. <Pomocunme3 51 xo,LIMTCJI cmpwna. B MeM6pattax TIVJaKOI1,LIOB Haxo,L\aTea rr11r:vieHThI (x:iopoqmJIJI, KapOTI1HOI1,L\hl 11 .11,p.). B CTpOMe co.n,epiKaTeJI cpepMeHThl H p1160COMhl. retteTWieeKHH arrrrapaT rrpe.n,eTaBJICH UHKJIHqeeKHMH MOJieKyJiaMH .II:HK. 3To UOJiyaBTOHOMHhle opratteJIJlhl, erroeo6Hhie eIIHTe3IIpOBaTb co6eTBCHHhie 6CJ1KOBhle MoJieKyJihI. OcttoBttaa cPYHKI~Ha xJioporrJiaeToB - cPOTOeIIHTe3, a TaKJKe e11ttTe3 AT<t> II A.II:<P (c1Joe¢op11JI11posattIIe), eHHTe3 11 rII.n,poJIH3 JIHIIIIJ(OB, aeeIIMHJIJIUHOHHoro KpaxMaJra II 6eJIKOB, OTKJia,D;bIBaIO~Hxea B CTpo:vie. B oenoBe eTpoeHIIJI XJIOPOcPHJIJIOB JieJKHT Mg-nop<jJupu1w6btu CKeJtem. Ero eoeTaBJrnIOT rrop¢11pnttosa.a «rOJIOBKa» e aTOMaMH Mg B uettTpe (rII,11,poclJ11J1bHa, JieJ'KHT Ha IIOBepxHoeTH MCM6paHbl) II cPIITOJihHhIH «XBOeT» (rIIJJ;poq>o6HhIH, y,n;epJKHBaCT MOJieKyJiy XJIOpocpHJIJ1a B MeM6paHe). Cy~ecTByeT HeeKOJibKO MO.L\HcPHRau11i1 xJiop01p11JIJIOB (a, b, e, d), OTJmqa10rn;nxca erreKTpaMH rrorJio~eHHJI. Bee paeTeHHJI B KaqeeTBe oeHOBHOI'O IIHrMeHTa eo,n;epJKaT eHHe-3eJieHhIH XJlOpO<jJUJlJl a, a B KaqeeTBe ,11,0IIOJIHIITeJihHhIX 3eJieHO-J'KeJIThlH XJlOpO<jJUJlJl b. B xJioporrJiaeTax xJiopocpnJIJI n ,n;pyrne rrnrMeHThI, rrorpyJKeHHhie B TmrnKoH.L\hI, eo6paHhI B <l>YHKUHOHaJihHbie e,n;HHHUhl (rro 250-400 MOJieKyJI), Ha3hIBaeMhie <jJomocucmeM.a.Mu. HMe10Tea ,n;sa Tnrra cpoToe11eTeM: <PC-I n <PC-II. Kam.n,a.a clJoToencTeMa eo,n;epmnT 250-400 MOJieKyJI rrnrMeHTOB. Bee n11rMeHThI clJoToeneTeMhI MoryT rrorno~aTb qaeTHUhI eBeTOBOH aHeprnH (cPOTOHhl, KBaHThI eBeTa), HO TOJlhKO 0,11,Ha MOJieKyJia XJiopocpnJIJia aTOH eneTCMhI MOJ'KeT HeIIOJlh30BaTh rrorJio~eH­ HYIO aHeprHIO B q>OTOXHMnqeeKHX peaKUHJIX. 8Ta MOJieKyJia Ha3hIBaeTeJI peaKU,UOliliblM. u,eHmpoM., a ,n;pyrne MOJieKyJihl IIHrMeHTOB Ha3bIBaIOTC.H aHmeHllblMU, TaK KaK yJiaBJIHBaIOT aHeprnIO eBeTa, IIO,LI06HO aJITeHHaM, ,D;JIH IIOeJ1C,n;y10~ei1 rrepe,n;aqn pea1<unoHHOMY ueHTpy. B <PC-I pea1<unoHHhiii ueHTP o6pa3oBaH MoJieKyJioii xJiopocpnJIJia a, o6o3HaqaeMoi1 P 700 , r,L1e 700 - onTnMyM rrorJI01uettna B HM. B <PC-II peaKUHOHHhIH ueHTP TaKme o6pa3oBaH MOJieKyJIOM XJIOPOcPHJIJia a, o603HaqaeMOH p 680' r,n;e orrTHMyM norJio~ettna JremnT B pai1oHe 680 HM. <t>oToeneTeMhI I II II pa6oTaIOT eHHXPOHHO ll HerrpephIBHO, HO <PC-I MOJ'KeT <l>YHKUHOHllpOBaTh OT,D;eJihHO. CpaBHHTeJILHaJJ xapaKTepncTnKa pa60Tb1 ,11;Byx l}>oTocncTeM npu l}>oTocuHTeae <l>OTO~OC~OpHJillpOBaHue He11uKJmtiecKoe ll;ttKJllltJCCKOe °YqaCTBYIO~tte $OTOCHCTeMhl <PoTocucTeMhI I, II <PoTocucTeMa I IlyTb :rneKTpOHOB HeIIttKJitttiecKuii littKJilltJeCKllH IlePBhlll ,D;OHOP aJieKTpOHOB Bo,n;a HA,U<P. <PoTocucTeMa I IlocJie,n;ttuii aK11enTop Ilpo,n;yKThl peaKuuu AT<P, HA,U<PH, KllCJIOPO,L\ AT<P -- -- <PoTocncTeMa I I. CeemoBaR <jJaaa rrpoTeKaeT Ha eBeTy Ha MeM6paHax THJiaKOH,L\OB npH yqaCTHH XJiopocpmrna, 6eJIKOB - rrepeHoeqHKOB aJreKTPOHOB n clJepMeHTa - AT<PCllHTeTa3hI (pne. 18).
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06;n,en Beiq,ecmB u npe6paiq,enue ;mepzuu. <l>omocunme3 53 HbIH aK11,enTOp P 430 (6eJIOK, co.n;epmaI11,11ii: JKeJie3o 11 cepy). P 4a0 nepe11awr cB011 :3JieKTpOHbI Ha .n;pyroil: JKeJie30Co.n;epJKaI11,MH 6eJioK - cpeppe.n;OKCHH, a IIOCJieJJ;HMM Ha KocpepMeHT HA,ll.<t> , KOTOpbIM BOCcTaHaBmrnaeTc.H /IO HA.D:<t>H 2 . MoJieKyJia P700 B xo.n;e rrpo11,ecca OKHCJI.HeTC.H, HO 3aTeM BOCCTaHaBJIHBaeT IIOTep.HHHbie :3JieKTpOHbl 3a cqeT :3JieKTpOHOB, IIOCTynaIOill,MX TIO :3JieKTPOHHO-TpaHcnopTHOM 11,enu OT <PC-II. 0,n;HOHarrpaBJieHHhIM IIOTOK Ha3hIBaeTC.H fteU,UICJlU'leC/CUM nomo/CO.llt 3Jle/Cmpo1t06, a o6pa3oBaHtte AT<t> - 1teu,u1CJtU'leC1CUM <Pomo</_Joc<PopuJtupoea1tueM. <t>C-I MOJKeT pa6oTaTb He3aBMCMMO OT <t>C-II. 3TOT rrpo11,ecc Ha3hIBaIOT 11,l:IKJIRqecKRM IIOTOKOM 3JieKTpOHOB. B xo.n;e Hero He rrpoMCXO,ll;MT cp0TOJIM3 BO,ll;hl, BbI,n;eJieHRe 0 2 M o6pa3oBamrn HA,ll.<t>H 2, o.n;HaKo 06pa3yeTC.H AT<t>. TaK KaK cnHTe3 AT<t> CB.H3aH c U,UICJlU'leCICU.M nomo/Co.M 3Jte1Cmponoe, ero Ha3hIBaIOT u,u1CJtU'leC1CU.M <Pomo<Poc<PopuJtupoea1tue.M. Y ayKapttoTuqecKttx KJieToK oH ocyill,eCTBJI.HeTC.H pe,n;Ko, o,n;HaKO y cpOTOCMHTe3I:IpyIOI11,l:IX 6aKTepttii: JieJKMT B ocHOBe rrpo11,eccoB aHaapo6Horo cpoTOCMHTe3a. TaKttM o6pa3oM, 1Co1te'l1tbt.Mu coeaune1tuJl.Mu ceemoeou <Pa3bt .HBJI.HIOTc.H: AT<t>, BoccTaHOBJieHHhIM HA,ll.<t>H2' 0 2• CBo6o,n;HhIM KMCJiopo.n; rrocTyrraeT B aTMoccpepy, AT<t> M HA,ll.<t>H 2 TpaHcnop™PYIOTC.H B CTPOMY XJIOpOIIJiaCTa M yqacTBYIOT B npo11,eccax TeMHOBOM cpa3hl. IL Te.M1toea.R <Pa3a rrpoTeKaeT B cTpoMe xJioponJiacTa n MOJKeT nponcxo,n;HTh He TOJibKO Ha cBeTy, HO u B TeMHOTe. XnMnqecKa.H aHeprn.H (B Btt.n;e AT<t>), 3anaceHHa.H B xo.n;e CBeTOBOM peaKll,HR, HCIIOJih3yeTC.H JJ;JIP, BOCCTaHOBJieHH.H yrJiepo.n;a. PeaK11,nn TeMHOBo:H cpa3hI rrpe,n;cTaBJI.HIOT co6oii: 11,erroqKy rrocJie.n;oBaTeJibHhIX npeo6pa30BaHMH yrJieKMCJioro ra3a, IIPHB0,11;.Hill,YIO K o6pa30BaHHIO rJIIOK03bl H ,11;pyrHX opraHttqeCKMX BeI11,eCTB. B cocTaBe xJioporrnacToB HMeeTc.H cpepMeHT (pH6yJio3o-l,5-6ucpoccpaTKap6oKcttJia3a), KOTOphIH KaTaJIH3HpyeT coe.n;I:IHeHtte MOJieKyJihI yrJieKHCJIOro ra3a c rrpott3BO,ll;HbIM II.HTHyrJiepo.n;Horo caxapa - pu6o3hI c pu6yJto.3o6u<Poc<Pamo.M (PB<P). 0.n;Ha MOJieKyJia PB<t> coe,11;HH.HeTc.H c MOJieKyJio:H C0 2 c o6pa3oBamrnM wecmuyiJtepoa1tou .MoJte/CyJtbt [C 6 ], KOTopaa HerrpoqHa H 6hlcTpo pacrra,11;aeTc.H Ha ,11;Be TpexyrJiepo,11;Hhie MoJieKyJihI <Poc<PoiJtu4epu1weou ICUCJlOmbt, <PI'K (C3)· 3aTeM rrpOHCXO.D;MT U.HKJI peaKll,MM, B KOTOpbIX ttepe3 pa,11; npoMeJKyToqHhIX rrpo.n;yKTOB cpoccporJIH11,epI:IHOBa.H KHCJIOTa npeo6pa3yeTC.H B ,11;Be MOJieKyJihI <Poc<PoiJtuu,epoaJtbaeiuaa (<PI'A), 3aTeM o,11;Ha MOJieKyJia <t>fA tt,11;eT Ha pereHepa11,n10 PB<t>, a BTopaa Ha o6pa3oBaHtte l'JIIOK03hl. B :3THX peaK11,H.Hx MCIIOJih3YIOTCH 3HeprmI AT<t> H HA.D:<t>H, o6pa30BaHHhIX B CBeTOBOH cpa3e. lJ;MKJI 3TRX peaKll,MH IIOJiyqHJI Ha3BaHne «11,MKJI KaJibBHHa» (pttc. 19). CyMMapttoe ypaBHeHtte CHHTe3a l'JIIOK03hI: 6C0 2 + 12HA,D;H + 18AT<t> -i- C 6 H 12 0 6 + 12HA,ll.+ + 18A,ll.<t> + 18<t> + 6H20 KpoMe rJIIOK03hI, B rrpo11,ecce cpoToCHHTe3a H3 <t>fK o6pa3yIOTC.H .n;pyrne MOllOMephI CJIOJKHhIX opraHnqeCKHX COe.D;HHeHHH - aMHHOKHCJIOTbI, rJIH1J,epHH, 'KHPHhle KBCJIOTbl, HyKJieOTM,ll;bl.
06UfaH 6uo;wzun. OcnoBbt lfUmoaozuu 54 HecTa6ttJihHhI: rrpoMemyToqHhiii rrpo,n;yKT co P:u6yJio3o-l,56mpocct>aT (PB<I>) <Pocct>orJim:i;ep:uHoBaR KllCJIOTa (<l>I'R) 2C3 pereHepau;11JI 2AT<I> 42AT<I> _ ___, <I>ocq>0rJI11u;epoaJib,n;ertt,n; (<I>I'A) 2C3 I'JIIOK03a + 2<I> r 2HA,ll;<I>H2 '-+2HA,l:J;<fl+ ~-___,<flpyKT03a Pitc. 19. l.J,111KJ1 KanbBlllHa <l>aKTOpbl, BJIIUIIOITI;He Ha cpoTOCHHTe3 Ceem - cKopocTh cpoTOCHHTe3a npHMO nponopu;noHaJihHa HHTeHCHBHOCTH CBeTa; npH ITOBbiilleHHH HHTeHCHBHOCTH CKOpOCTb cpOTOCHHTe3a 3aMe,n;JIHeTCH, TaK KaK pa3pyrnaroTCH MOJieKyJihI XJiopocpHJIJia. BpeMR: qeM 6oJibrne nepno,n; OCBern;eHHOCTH, TeM ,ZJ;OJibIITe npo,n;oJimaeTCH cpOTOCHHTe3. Ko11u,e11mpau,uR yzJLelCUCJlOZO za3a 0,03-0,04 % ); npn KOHD;eHTpau;HH (B HOpMe B aTMoccpepe co,n;epmHTCH C02 ,n;o 0,1 °/c, ITOBbIIITaeTCH CKOpOCTb TeM- HOBhIX peaKD;HH cpoTOCHHTe3a. BofJa - cy6cTpaT cpoTOCHHTe3a; npn yMeHhIIIeHHH co,n;epmairnH BO.D;hI 3a- C0 2 He MomeT ,n;ncpcpyH,n;npoBaTb. onTttMaJibHaH TeMnepaTypa ,n;JIH cpoToCHHTe3a KphIBaroTCH ycThHD;a, H Te.Mnepamypa - +25 °C (,n;JIH pacTeHHH cpe,n;Heii ITOJIOCbI); CHHmeHne npHBO,ll;HT K 3aMe,n;JieHHIO pa60Tbl cpepMeHTOB. ,UocTaToqHaH ICOHU,enmpau,uR XJlOpO<jJUJlJlU. Heo6xo,n;HMo ,n;ocTaTOqHoe MHHepaJihHOe nnTaHne Koc.Mu1leC1CaR pOJlb pacme11uu CH B TOM, qTo pacTeHHH - (onncaHa K. A. (Mn, Fe, Mg, NH ,n;p.). THMttpH3eBhIM) 3aKJIJOqaeT- e,n;HHCTBeHHbie opraHH3Mhl, ycBaHBarorn;ne COJIHeq- HYIO ::rneprHIO H aKKYMYJIHpyrorn;ne ee B Btt,n;e ITOTeHu;HaJihHOH XHMfiqecKOH 0 2 no,n;,n;epmnBaeT mtt3lfa KHCJiopo,n;a o6paayeTCH 030H, ::rneprHH opraHHqecKHX coe,n;nHeHHH. Bhl,n;eJIHrorn;niicH He,n;eHTeJihHOCTh Bcex aapo6HhlX opraHH3MOB. KOTOphlii aarn;Hrn;aeT Bee mHBoe OT YJihTpacpHoJieTOBhIX Jiyqe:H. PacTeHHH nc- C0 2, tt36b1ToK KOToporo co3,ZJ;aBaJI «napHHKOBhlH acpcpeKT»' H TeMnepaTypa nJiaHeThl ITOHH3HJiaCh ,n;o Hhl- ITOJih30BaJIH H3 aTMoccpepbI rpoMa,n;Hoe KOJIHqecTBO HeIIIHHX 3HaqeHHH.
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06~all 56 6uo1wiull. Oc1W6bl 1tumo1wiuu conpoBom.n:a10:w;11xcH Bhrp;eJieHHeM ::rneprttu. Kone'-IHhIMH npo.n:yKTaMH pacna.n:a HBJUIIOTCH co~, H20, ATcD. 3Hepl'HH, Bhr.n;eJIHIOIIJ;aflCH IIpH paapyrueHHH XllMHqecKHX cBH3eH:, 3anacaeTcH B MaKpoapruqeCKHX CBH3HX MOJieKyJ1 ATcD " .n;pyrwx BhICOK03HepreTttqecKHx coe.n;HHeHHtt. ATcD - yuwBepcaJihHhrtt ucToqHHK aHeproo6ecneqeHHH KJieTKH. CttHTe3 A Tell nponcxo.n;HT B KJieTKax Bcex opraHH3MOB B npouecce ¢ocipopuJzupo6a1tuR - npncoe.n;ttHeHr-ur HeopraHttqecKoro cpoccparn K AJJ:cD. IlpeBpa:w;eHHe Be:ru;eCTB l1 3HeprHH B npouecce .ZJ:HCCHMHJIHUHH BKJIIOqaeT B cefoc cJie,n;y10:w;11e 3Tarrhr: I aTan - no11.roTOBHTeJ1bHhIH. CJIOJRHhie opram1qecKtte Be:w;ecTBa no.n: ,n;ettCTBHeM cpepMeHTOB pacna,n;aIOTCH Ha npoCTbie (MOHOMepb1): 6eJl}(U iJo aMUIWKUCJlom; JICUpbt 00 ?Jluu,epul-la u JICUpllblX }(UCJlOm; yiJLeBOiJbt (KpaxMQJl) oo .M011ocaxapuoo6 ( ?JllOK03bt ); nyKJteunoBbte KUCJtontbt iJo 1-lYKJLeomuiJoB Um. 0. Ilp11 3TOM BhI,!l;em1eTCH 3Heprl1H B BH,n;e TeIIJia. II aTarr - rJ1HKOJ1H3 (6eCKHCJiopo,n;HhIH, aHaapo6Hbrtt) - MHorocTyrreHqaThitt <lJepMeHTaTHBHhIH npouecc pac:w;enJieHmI I'JIIOK03hI ,n;o .ZJ:BYX MOJieKyJI IIHpOBHHOrpa,n;HOH KHCJIOThI (IIBK, IIHpyBaT), ,n;Byx MOJieKyJI ATcD 11 BOCCTaHOBJieHHOH <lJopMhI nepettocq11Ka Bo.n;opo,n;a HA.UH, c o6pa3oBantteM npoMeJRyToqHhIX rrpo,n;yKTOB. Ilpouecc ocy:w;eCTBJIHeTCH B UHTOIIJia3Me 6e3 yqacTIUI KllCJIOpo,n;a, c MeM6paHaM11 He CBJI3aH. ITBK I' ..'1101{0:13 Ilp11 3TOM 40 'Xi ::rneprtttt aanacaeTcH B Bn,n;e ATcD, a 60 'Y,, 3Heprnn paccenBaeTcH B BH,n;e TenJia. III. llpu omcymcmBuu KUCJLOpooa: y ,n;pommeH: 11 pacTeHHH rrpo11cxo,n;HT CIIl1pTOBOe 6pomeHHe, nptt KOTOPOM CHaqaJia o6pa3yeTCH yKCyCHhIH aJib,!l;erH,n;, a 3aTeM 3Tl1JIOBhIH CIIHpT: nnK Yl\.CyCHhIH a.1h;J,CfltJJ. JTHJIORbii1: CTIHPT y H€KOTOpbIX a<HBOTHhIX 11 6aKTepnH: npn He.n;ocTaTKe K11CJIOpo,n;a npo11cxo,ll;HT MOJIOqHOKHCJIOe 6pomeH11e B MhlllieqHhIX KJieTKaX c o6pa30BaHHeM MOJIOqHOR Kl1CJIOTbI: ITBK :\IIO.TJOllIHtH li.H.C.110Ta B peayJihTaTe rJI11KoJIH3a o,n;HOH MOJieKyJihI rJIIOK03hI BhICBo6om,n;aeTcR 200 K.D;m, 113 KOTOphrx 120 K,llm pacce11BaeTCR B Bn,n;e TerrJia, a 80 K)lm pacxo,n;yeTcR Ha CHHTea ATcD.
06Men 6PUfeCm6 u npe11paUfenue anepzuu. :J11epzemu'leCK1tii o6Men 57 !Ipu llU.'lU'lUU KUC.rtopoiJa rII)l,pOm13 (KliCJIOpO.ll,HhIH, a::ipo6Hoe /lbIXaH11e) noJinoe cpepMeHTaTHBHoe pacn~err.TJettne IIBK ;no yrJieKMcJioro ra3a H BO.D;bI, c o6pa30Ban11eM 36 MOJICKYJI AT<f>: .n;Be Mo.'IeKyJlhI o6pa3yIOTCR B pemni;ttRx u;irn.11a Kpe6ca (B MaTpttKce MttTOXOH.ll,pttil) H 34 :vro.'IeKyJihI - B rrpouecce OKHC.'lHTCo'IhHOrO cpoccpopH.'"!llpOBaHttR (Ha BHYTPCHHett MeM6paue MHTOXOH.ll,· pn:ii, B KOTopyIO BcTpoeui,1 cpepMeHThI .ll,bIXaTeJihHOll ILerrtt). Ha6.'lIOJJ;aeTcH U,UKJl mpuKap601-w6blX Kuc.1wm ( u,uKJL Kpe6ca ); 3aKJIIOq11TeJibHYIO cTa.n;n:IO OKHCJieHna - ;J.'teKmpo11Ho-mpancnopm1ty10 u,enb. TOJlbKO y aspo6Hb!X opraHH3MOB. 8TOT npouecc BR.JIJOqaeT: IIttpyBaT IIOCTyrraeT B MllTOXOH.n;pnIO n B3aHMO.ll,ellCTByeT c cpepMeHTaMn: Ma· TpttKca. 3.11,ech nponcxo.11,MT .n;eru.11,pn:poBaHMe (oTru;emieHtte BO.ll,opo.n;a), .n;eKap6oKCMJittpoBantte ( OTru;errJieHtte C0 2) n: npttcoe,n;ttHeHtte KocpepMeHTa A (HS KoA) c o6pa30BaHHCM aueTMJIKocpepMenTa A (CH;iCO - S-KoA), 06Jia,n;a10w;ero BhICOKosuepreTttqecKoil CBH3hIO. 8To coe.n;n:Hentte BOBJieKaeTca B umrn Kpe6ca uenh nocJie,1i;oBaTeJihHhIX peaKUHll. AueTHJI-KoA coe,n;ttHHeTca c MOJieKyJioil w;aBeJieBoyKcycnoil KMCJIOThl (mYK), rrpu 3TOM 06pa3yeTCH TPMKap6oHOBaH JIHMOHHaH KHCJIOTa. B xo,n;e IIOCJie.n;yJOw;MX cpepMeHTaTMBHhIX peaKUHll JIHMOHHaH KHCJIOTa OKMCJIHCTCH, rrptt 3TOM BOCCTaHaBJIHBaJOTCH 3 MOJieKyJihI HA,U · p;o HA.II:· H, o.n;Ha MOJieKyJia <f>A,U B <f>A,U • H 2 H 06pa3yeTcH MOJieKyJia ryaH03MHTpncpoccpaTa (fT<f>) c BhICOK03HepreTttqecKoil cpoccpaTHOll CBH3hIO. 3HeprttR IT<f> ttcnoJih3yeTca ,n;JIH cpoccpopmmpoBaHn:H A,U<f> H o6pa30BfilIHH AT<f>. JlttMOHHaH KHCJIOTa TCpJJ.eT .n;Ba yrJiepO.ll,HhIX aTOMa, 3a cqeT KOTOpb!X o6pa3YJOTCH .n;Be MOJICKYJlhl C0 2 • B cyMMe, B pe3yJihTaTe ceMM nocJie.n;oBaTeJihHhIX cpepMeHTaTHBHhIX peaKUHll, JIMMOHHaH KHCJIOTa rrpeBpaw;aeTCH B mYK. 06pa30BaBlllaRCH MOJICKyJia mYK COC.ll,HHHCTCH C HOBOM MO.'lCKy.1oll aI~eTHJI-KoA, H UHKJI IIOBTOpHCTCH. B COCTaBe JIHMOHHOM KHCJIOThl croplleT rrpHCOC.ll,MHMBlllHMCH OCTaTOK aueTHJIKoA. IlpM 3TOM o6pa3yIOTCH yrJieKHCJIL!H ra3, aTOMhI Bop;opo.n;a; sJieKTPOHhI rrepeHOCHTCH Ha aKUCIITOphI HA,U- H cf>A.II:".; 3HeprmI XHMMqecKHX CBH3Cll yrJICBO- .ll,OB, mttpoB, 6eJIKOB HaKarrJin:BaeTCR B MOJieKyJiax HA.II: · H 2, <f>A,U • H 2 M AT<f>. n: <f>A,UH 2 rrpottcxop;uT B .n;hIXaTeJihHOM uerru MHToxoH.11,pn::H, Ha BHyTpeHHei-i: MeM6paHe. OHM nepe.n;aIOT aTOMhI Bo.n;opo.n;a no ,UaJihHeiirnee OKHCJieHtte HA.II: · H uerru BCTpOCHHhIX B MeM6paHy 6eJIKOB. Ilptt 3TOM rrpOTOHhl OKa3bIBaIOTCH Ha BHCUIHell CTOpOHe BHyTpeHHell MCM6paHhl H HaKaITJIHBaIOTCH B MemMeM6paHHOM rrpocTpaHCTBe (cHapymn: MeM6paHbI B03pacTaeT ITOJIO)KHTCJibHhlll 3apa,11;), a 3JICKTP0Hbl OCTaIOTCH c BHyTpeHHCH CTOpOHhl MeM6paHbI, r.n;e coe.n;HHHIOTCH c 02. Memp;y BHelllHeii tt BHyTpeHHeiI cTopoHaMH MeM6paHhI B03HHKaeT pa3HOCTh mpa1tcMe.w6paHHbtu nome1tu,uaJL. B MeM6paHy BCTpoeH cpepAT<P-cu1tmema3a. Bero MOJieKyJie OTKphrnaeTcH KaHaJI, qepe3 KOTOphIM ITOTCHIJ;HaJIOB MeHT MoryT rrpoxo.n;nTh rrpoTOHhI rrptt pa3HOCTH noTeHUIWJIOB 200 MB. IIpoTOHhI Ha- 'IHHaIOT rrpoxo.n;HTh KaHa.11, .ll,BHJKHMbie CHJIOM 3JICKTpHqecKoro IIOJIH (TaKHM 06pa30M, BHyTpH B0306HOBJ1RCTCH co,n;epJKaHHe npOTOHOB, H3pacxo.n;oBaHHhIX Ha BOCCTaHOBJICHHC Q2-), H COC.ll,HHHJOTCH c rrpOTOHaMH.
0614aR 6uo.rwzuR. 58 Oc1t06bl ~umo.rwzuu 3a c'-!eT ::meprtttt ;~smKylll;ttXCH npoTOHOB B Mo:1eKy:1e AT<P-cttHTeTa3hI npottcxo,n;ttT CMHTe3 AT<P tt3 A)J;<P tt <P. 06lll;ee IWJII1'-!8CTBO :meprttM, BhI,n;eJrnIOlll;8HCH Ha 8TOM 8Tane, --- 55 '/\, 2600 K)J;m. ::iuepr1rn 3anacaeTCH B <}:>opMe AT<P. 06lll;aH <}:>opMyJrn atta::ipofo1oro 11 a::ipo6ttoro sTarrns: --1 cf>OTOCl1HTe3 ~ Aspo6noe ,n;b1xaH11e -----------------· AHa60JI1P1eCKMH rrpou:ecc, B Ilpou:ecc ,ll;HCCHMHJIHU:HH, B pe3yJihTaTe KOTOporo H3 rrpoCTbIX pe3yJibTaTe KOToporo l\IOJICKYJlbl HeopraHM'IeCKHX coe,n;HHeHHM yrJieBO,ll;OB pacru:errJIHIOTCH CHHTe3HpyIOTCJl MOJieKy Jibl npOCTb!X HeopraHH'leCKHX yrJieBO,Zl;OB coe,n;HHeHHH 3HeprMH AT<I> HaKaIIJIHBaeTCH H 110 3neprHH 3arracaeTcH B Btt,n;e I j AT<I> 3arracaeTcH B yr JieBo,n;ax I RttcJiopo,n; pacxo,n;yeTCH j YrJieKHCJihIM ra3 H BO,n;a YrJieKHCJihIM ra3 H no,n;a I IIOTpeOJIHIOTCJl Bb!,!\eJIJIIOTCH RttcJiopo,n; Bbl,!\eJIHeTCH --------- IlpOHCXO,!\HT yBeJIH'leHHe IlpOHCXO,ll;HT yMeHb111e1rne opraHH'leCKOH MaCCb! opraHH'leCKOH MaCCbI Y syKap110T rrpou:ecc rrpoTeKaeT B Y syKapttoT rrpou:ecc rrpoTeKaeT B XJIOPOIIJiaCTaX MHTOXOH,n;pm:ix IlpoHCXO;IHT TOJibKO B KJieTKax, IlpOHCXO,ll;HT BO BCex KJieTKax B co,n;epmaru;ttx XJiopocpHJIJI, Ha cseTy Te'leHHe :lKH3HH nenpepb!BHO J AEJ1EHHE KJlETKH KJLemo'lllbtit u,uKJL - sTo rrep:tto.n; :JKM3He,n;eaTeJihHOCTH KJieTKM OT MOMeH- Ta ee IIOHBJI8Hl1H .n;o TIOCJie,n;yIOlll;ero ,n;eJieHtt.H HJil1 CMepni:. ll;:ttKJI ttMeeT pa3HYIO rrpo,n;OJI:JKl1T8JlhHOCTb B 3aBHCHMOCTl1 OT TKaHeBOM rrp:ttna,n;Jie:>KHOCTl1 H BHeIIIHHX <}:>aKTOpoB: TeMrrepaTyphI, IIHTaTeJihHhIX Belll;eCTB, KHCJiopo.n;a (y 8p:ttTpOIJ;HTOB 120 CYTOK; y 3IIHT8JIHH KO:JKH 20 cyToK; y KJl8TOK KOCTHOI'O Mo3ra 8-12 qacoB). KoMnoHeHTOM KJieToqttoro u;:ttKJia HBJIHeTCH HOCTh 6-12 qacos). .Mumomu'lecKuu U,UKJl, KOTOpbIM BKJilOqaeT rrep:tto.n; IIO,Zl;I'OTOBKH KJieTKH K /18JieHHlO ( uHmep<jJa3y) H co6cTBeHHO ,n;eJieHHe - Mumoa ( cpe,n;HHH rrpo.n;omK11TeJih-
penenue x:nemx:u. Hnmep</Ja3a. Mumo3 59 IIHTepcI>aaa H ttmep<jJa3a - rrep1rn,n; Jini::me,n;eJITeJibHOCTH KJienrn Mem.ny .D;C.'IemrnM 11 . 3aHHMaeT 90 <;(, BCero BpeMeHH KJieTO'-IHOro IJ;HKJia. IIepttO.D;hl HHTepcpa3bI: 1. Ilpecw1memu'lec1Cuit (nocm;.wmomu'lecKuit}, HJIH G 1-nepuoiJ, B KOTOpoM rrpoHCXO,[\HT rro,n;rOTOBKa K penJIHKaIJ;HH (caMoy,n;BOeHHIO) .D:HK. OH BKmoqaeT: CHHTe3 6eJIKOB, cpepMeHTOB, tteo6xo,n:ttMhIX .D;JIH o6pa 30 _ BaHHH PHK, H Beru;ecTB, Bxo,n;Hru;ttx B cocTaB .D:HK; CHHTe3 PHK; o6pa30BaHtte ptt6ocoM; CHHTe3 AT<P; o6pa3oBatttte opraHeJIJI - MHTOXOH,n;pttii, XJIOponJiaCTOB (y pacTeHHM), 8IIC, JIH30COM, annapaTa fOJib,[\JKl1, BaKyoJieii, IlY3blpbKOB; pocT KJieTKH ,n;o «KPHTH:qecKOM MaCCbI ll;HTOIlJia3MbI»; o6pa30BaHtte Beru;eCTB, no,n;aBJIHIOIIl;HX HJIH CTHMYJIH:py10ru;ttx HaqaJio cJie,n;y10ru;eii: cpa3hI. XpoMOCOMa rrpecttHTeTHqecKoro rreptto,n;a - oottoxpoMamuiJtta.R, c retteTttqecKHM Ha6opoM 2n2c, r,n;e n - qHCJIO xpOMOCOM, c - KOJIHqecTBO .D:HK. 2. Cuttmemu'leCKUil, uJtu S-nepuoiJ, B KOTopoM npottcxo,n;ttT perrJIHKau;ttH ,n:HK IIOJIYKOHCepBaTHBHbIM cnoco6oM (CM. pa3,n;eJI «feHeTHKa» ). B HaqaJie rreptto,n;a xpoMoCOMhl - oiJttoxpoMamuiJttbte (o6pa3oBaRbI o,n;ROM xpoMaTtt,n;oii, 2n2c). IIocJie perrJittKau;tttt .D:HK - iJByxpoMamuiJHbte (cOCTOJIT H3 .D;BYX O,[\HHaKOBhIX xpoMaTH,[\, 2n4c). CttHTe3 6eJIKOB (rttCTOHOB), yqacTBy10ru;ttx B cpopMttpoBaRHH CTPYKTYpbI xpoMOCOM. 3. Ilocmcuttmemu'leCKuit (npeiJ.Mumomu'leCKuit), HJIH G2 -nepuoiJ, B KOTOpoM rrpOHCXO,[\HT rro,n;rOTOBKa KJieTKH K ,n;eJieHHlO. BKJIIOqaeT B ce6JI: CHHTe3 6eJIKa; o6pa3oBatttte MHTOTttqecKoro anrrapaTa; CHRTe3 AT<P; CHHTe3 PHK; CHHTe3 6eJIKOB, orrpe,n;eJIJIIOru;ttx ,n;eJieHtte KJieTKH; yBeJittqeHHe MaCCbI IJ;HTOIIJia3MhI; pe3KOe B03pacTaHtte o6'heMa H,n;pa. XpoMocoMa - ,n;ByxpoMaTtt,n;HaJI, c reHeTttqecKHM tta6opoM 2n4c. MHT03 M umo3 - aTo tterrpHMOe ,n;eJieHtte KJieTKH, rrptt KOTopoM npottcxo,n;ttT paBRoMepaoe pacrrpe,n;eJieatte ttacJie,n;CTBeHHoro MaTepttana Me:>K,n;y ,n;oqepHHMH KJieTKaMH (pRc. 201. Oco6ettttocmu .Mumo3a: ocHOBHOM crroco6 ,n;eJiemrn ayKapttoTHqecKoii: KJieT1m; BepeTeRo ,n;eJieRttH 06pa3yeTcH; y,n;Boett11e tt paBHOMepttoe pacrrpe,n;eJiemrn reaeTHqecKoro MaTep11ana; paBHOMepttoe ,n;eJieHHe u;RTOIIJia3MhI. BuoJtoZu'lecKoe 3Ha'lettue .Mumoaa - o6ecrreqeHHe Toqttoii rrepe,n;aqH ttacne.n;cTBeHHOii HHQJOpMaIJ;HH Ka:>K,[\OMY H3 ,n;oqepRHX JI,n;ep. qHCJIO xpOMOCOM B 06e11x ,n;oqepHHX KJieTKax paBHO qHcJiy xpoMOCOM MaTepRHCKOM KJieTKH. MRT03 BKJIIOqaeT qeThipe c]la3hI: npo<jJa3a, .Mema<jJa3a, attarjJa3a, meJto<jJa3a. fio BpeMeHH 3aHHMaeT B cpe,n;HeM 10 °lo OT BpeMeHH MHTOTHqecKOrO I.J;HKJia. 1. llpo<jJa3a ( 2n4c) - CaMaJI rrpo,n;OJIJKHTeJibHaJI c]la3a MHT03a, B KOTOpOH rrpoHCXO,[\HT KOR,n;eHcau;m1 xpOMOCOM H c]lopMRpoBaHHe MHTOTHqecKoro arrrrapaTa.
06~aR 60 6uo.1wzuR. OcnOBbt 1.j.Umo.1wzuu Te.w(\1aail )H'.110on1<rn K.:ienrn A Pfll.:THTe:1bHaa K.1Pih'.ll 6 P111c. 20. A - 06Ll\a5l cxeMa M"1T03a: I - MaTep"1HCK"1e xpOMOCOMbl 2n2c; II - XpOMOCOMbl, COCT05lL1\"1e a-13 AByx xpoMarnA nocne G2-nepa-10Aa 2n4c; Ill - A04epHa-1e xpoMoCOMbl 2n2c. 5 - cxeMa M"1T03a B )1("1BOTHoi;i "' pacrnrnnbHoi;i KneTKax: 1 - nepeT5l)l(Ka; 2 - KneT04Ha5l neperopOAKa Bo BpeM.H rrpocpa3hI rrpottcxo,n;MT: crrttparrtt3a1IM.H xpoMocoM; pa3pymemi:e .H,n;epHoif o60JIOqKM M .H,ll;pbIIIIeK; cpopMHpOBaHMe MMTOTMqecKoro arrrrapaTa: pacxom,n;eHMe 11eHTpMOJieH K rrpOTMBOilOJIOJKHbIM CTOpoHaM (nomocaM KJieTKH), MeJK,n;y KOTOpbIMM 06pa3yeTC.H BepeTeHO ,n;eJieHM.H. 2. M ema</Ja3a ( 2n4c) - BhlCTpaMBaHMe ,n;ByxpoMaTM,ll;HhIX xpoMocoM B 3KBaTopnarrhHOH rrJIOCKOCTM (MeTacpa3Ha.H rrJiaCTHHKa). Bo BpeM.H MeTacpa3bI rrpottcxo,n;MT: ,ll;BHJKeHHe xpOMOCOM K BepeTeHy, rrpnKpenJieHne HX KMHeTOXOPOB (11eHTpHOJieif) K HHT.HM MHTOTMqecKoro arrrrapaTa; BbICTpanBaHMe xpOMOCOM ITO 3KBaTopy BepeTeHa ,n;eJieHH.H KJieTKH. 3. Atta</Ja3a ( 4n4c) - cTa,n;H.H pacxom.n;ettn.H xpoMaTM,n; K rromocaM KJieTKM. Ilpn 3TOM rrponcxo,n;MT: .n;eneHne 11eHTpOMep; o6pa30BaHHe O,ll;HOXpOMaTH,ll;HblX ,n;oqepHHX xpOMOCOM, KOTOphle ,ll;BHJKYTC.H K rrpOTHBOIIOJIOJKHblM nomocaM MMTOTMqecKoro anrrapaTa; B KJieTKe HaXO,ll;HTC.H ,D;Ba ,ll;HIIJIOH,ll;HblX Ha6opa. 4. TeJw<jJa3a (4n4c - 2n2c B 1caJ1Ci)ou KJtemKe). B Heft rrponcxo,n;nT: pa3pyweHne MHTOTHqecKoro arrrrapaTa; pa3,n;eJieHne KJieTKM (llHTOKHHe3); ,n;oqepHHe xpOMOCOMbl .n;ecrrMpaJIH3YIOTC.H; o6pa30BaHHe .H,n;epHblX MeM6paH; BOCCTaHOBJieHHe .H,ll;pbIIIIeK. ,UeJieHne llHTOrrJia3MhI ( u,umo1Cu11e3) B J/CueomttblX 1CJtem1Cax npottcxo,n;nT 3a cqeT Bn.HqHBaHH.H MeM6paHhI (oT rrepncpep1:11:1 K 11eHTpy) H o6pa3oBaHHSI 6opo3,D;hI ,n;eJiemrn. B pacmumeJtbHblX 1CJtem1Cax - 3a cqeT KJieToqHoif rreperopo,n;KH (oT 11ettTpa K rrepncpepnn) B 3KBaTopHaJibHOH IIJIOCKOCTH. B pe3yJihTaTe H3 KJieTKH c Ha6opoM 4n4c 06pa3y10TC.H ,n;Be ,n;oqepttne KJieTKH c Ha6opoM reHeTnqecKoro MaTepnarra 2n2c Kam,n;a.H. 3ttaqeHne MHT03a. MnT03 o6ecrreqnBaeT: 1. IlOCTOSIHCTBO xpoMOCOMHOro Ha6opa B p.H,n;y IIOKOJieHHH KJieTOK. 2. ToqHyro rrepe,n;aqy ttacJie,n;cTBeHHOH nttcpopMa11nn.
,aenenue Knem1eu. ftfeii.o.J 3. 4. 5. 61 Yne.'IH'IeHue 'IHc.ria I<JieToK, T. e. MexaHH3M pocTa u pa3BHTHH opram13Ma. Crroco6 pereHepau:i111 KJieTOK. .flBJ1HeTC5i ocHoBoii 6ecno.11oro pa3MHomemrn. IlaTOJIOrH5i MHT03a. HapyrneHue HOpMa.'lbHOro Te'IeHH5i II HenpaBHJlbHOe pacrrpe,neJieHIIe xpOMOCOM Mem.ny .ll;O'-!CpHIIMH KJieTKaMH rrpMBO.ll:HT K B03HMKHOBeHMIO KJieTOK c uec6aJiaHCHpOBaHHbIMH KapHOTHII8MH (MYT81J;H5IM). <t>opMbI rraTOJIOrlrn:: 1. IloBpem.neHHe xpOMOCOM ITO,[\ ,[\eiicTBHeM 5i,ll;OB (MeTaHOJI, KO.'IXHI~HH). Ilp11 3TOM ~ee B03HHKaeT HapyrneHHe U:CJIOCTHOCTII xpOMOCOM, rrpHB0,[\5i- K HerrpaBHJibHOMY paCXO:>K,[\eHHIO HX K IIOJIIOCaM. 2. IloBpem,L\emre MHTOTH'IecKoro arrnapaTa npwBo,nMT K HepaBHOMepHo- 3. HapyrneHHe U:HTOKl1He3a - MY pacrrpe,L\eJieHHIO xpoMocoM Mem,L\y ,L\O'IepHHMH KJieTKaMn. B03Hl1KHOBeHne rrpem,[\eBpeMeHHoro 11JJtt rro3,L\Hero o6pa3oBaHM5i 6op03.L\ ,neJieHna. A.Mumo3 - 3TO npaMoe ,[\eJiettne KJieTKH tta,L\BOe, rryTeM rrepeTamKn. Oco6eHHOCTl1 aMl1T03a: H,L\pO H8XO,L\11TCf! B HHTep<Pa3HOM COCTOf!H11H; xpoMOCOMbI He Bblf!BJif!IOTCH; BepeTeHo ,neJieHMf! He o6pa3yeTC5i; paBHOMepHoro pacrrpe,neJieHMf! reHeTWieCKoro MaTepMaJia He rrpOl1CXO,L\MT (ll3 0.D;HOH KJieTKll o6pa3yIOTCf! ,n;Be He 11,neHTM'-IHbie .npyr ,npyry); B03HMK8IOT .ll:BY- M MHOroa.n;epHbie KJieTKH; o6pa30B8BllIMeC5i KJieTK11 ,n;eJIHTbCf! MllTOTff'IeCKl1 He MOryT. B HOpMe y qeJioBeKa aMHT03 BCTpeqaeTca B KJieTKax crreu:ttamI3ttpoBaHHhrx TKaHeii (3apo,nbIIIIeBbie o60JIO'IKH, cpOJIJil1KYJif!PHbie KJieTKl1 5il1'1HMKa), rrpM HC06XO,ll;MMOC- 1 TM 6hICTporo BOCCTaHOBJieHllf! TKatteii (rroc.'le orrepau:nii, TpaBM MT. IO~MX ,n.), B OT:>KMBUIHX CTape- 2 KJieTKax 11 ,np. Ilpu namoJtoiuu y qeJIOBeKa BCTpeqaeTca B «II8TOJIOr11qecKH M3MeHeHHbIX» M o6peqeH- 3 HhlX Ha rtt6eJib KJieTKax, He crroco6Hhrx B ,na.'lhHeiiUieM ,n;aTh KJieTKM IIOJIHOU:eHHbie (BOCIIaJieHMfl, 3JIOKa'IeCTBeHHbIH pOCT rrpM orryxOJif!X). 4 Me:Ho3 M euo3 - 3TO pe;:i:yI<U:MOHHOe ,neJieHMe, B pe3yJJhTaTe KOToporo rrpoucxo,nnT pe,n;yKu;tta 'IllCJia XpOMOCOM c .D;llIIJIOll,[(HOrO (2n) .no rarr- JIOll,[(HOro (n). KOToporo CBR3aHO c IIOf!BJieHneM IIOJIOBOrO pa3- PMc. 21. CxeMa Mel1o3a: 1 - MaTePHHCKHe xpOMOCOMbt; 2 - xpOMOCO- MHOateHua (C03peBaHMe raMeT), II03TOMY ero Mbl, COCT05Hl.IH8 H3 ABVX xpoMaTHA no- Ha3hlBaIOT ,D;eJieHneM co3peBattua. OH rrpoucxo- cne G2-nep1-10Aa: 3 - KOHb10ra1..11-1S'I; 4 -· Kpocc1-1Hrosep; 5 - nepsoe Mel10rn4ecKoe AeneH1-1e; 6 - sTopoe Mel1o- Meiio3 - cnoco6 ,neJieHna, B03HllKHOBeHwe .n;HT IlpH o6pa30BaHMM raMeT y )Kl1BOTHbIX H cnop y pacTeHHH, rptt60B 11 BKJIJO'IaeT ,ll;Ba IIOCJie,D;oBaTeJihHhIX .ne.11eHMH a,npa. B Kam,n;oM TH48CKoe AeneH1-1e
62 06UfaH 6uo.1wzuH. OcnoBbl l{Umo1wzuu ,n;e.rremu1 BhI.n;e;:r.HIOT <IeThipe cTa,z:urn:: rrpo<l_:>a3a, MeTacpa3a, attacpa3a, Te.11oc}ia3a c 1rn.n;eKcaM11I11.1111 II (pnc. 21). Hep Boe .n;eJieHHe MeH:oaa (pe;:i;y1m;Ho1111oe) H Hmeprpaaa I ( 2n2c----+ 2n4c) - rrpo11cxo,n;11T yBeJ1ttt1ett11e 'lHc.w opratte.ru1, CHHTe3 BeliICCTB H HaKOIIJieHHe 3HeprHH, Heo6xo,n;HMhIX ,ll;Jl.ff ,[(CJICHIUI, yBeJIH'leHHe pa3MepoB KJieTKH, y,n;BoeHHe I!eHTpHoJiett, penJIHKanmi .D:HK. KaJK,[(a.H xpoMocoMa npe,n;cTaBJieHa napoii: xpoMaTH,n;, coe.n;11tteHHhIX u;ettTpoMepoi1. IlporjJa.Ja I ( 2n4c) - caMa.H rrpol(OJIJKHTeJihHa.a <l_:>a3a, BKJIIO'laronwa IIHTh CTa,[(HH: 1. Jlenmome1-w - Ha'lHHaeTCH CIIHpa.'lH3aI!Hff xpOMOCOM. XpoMOCOMhl CTa- HOBHTCH BH,[(HMhIMH B MHKPOCKOII, KaK ,[(JIHHHhie TOHKHe HHTH. Kam,n;aa xpOMOCOMa COCTOHT H3 ,n;Byx xpoMaTH,[(. 2. 3uiomena - B a,n;pe BH,D;eH ,[(HIIJIOH,[(HhIH Ha6op xpoMOCOM. rrpo,n;oJimaeTca crrupaJIH3aIJ,MH 1(0/lo/O?aU,U.Fl roMOJIOrH'lHhIX I'O.MOJWZU'lHblMU H33hIBaIOTCH xpoMOCO- xpOMOCOM H rrpoHCXO,[(HT xpOMOCOM. Mhl, HMeIOliIHe O,[(HH3KOBYIO cpopMy H pa3Mep: O,n;Ha H3 HHX IIOJiy'leHa OT MaTepH, ,n;pyraa - OT OTLI,a. foMOJIO- rH'lHhie xpOMOCOMhl rrpHTHrHBaIOTCH H rrpHKJia,D;bIBaIOTCH ,n;pyr K ,n;pyry ITO BCCH ,[(JIHHe. U:eHTpOMepa 0,[(HOH H3 rrapHbIX xpOMOCOM TO'IHO rrpMJieraeT K IJ,eHTpOMepe 61AeaneHT ,n;pyroii:, H KaJK,n;aa xpoMOMepa rrpm:reraeT K roMOJIO- np011CXOA11T KOHbtora41-1si rH'lHOH xpoMOMepe ,n;pyroil. llapa KOH'hIOrlipOB3BIIIHX roMOJIOrH'lHhIX xpOMOCOM Ha3hrnaeTCH 6ueaJleHmO.M. Kam,n;hIH 6HBaJieHT cocTOHT H3 'leTBepKH xpoMaTH,n; - TeTpa,n;hI. "llttcJio 6ttBaJieH- TOB paBHO raIIJIOH,[(HOMY Ha6opy xpOMOCOM. 3. Ilaxumena - rrpottcxo,n;RT .D;aJihtteii:IIIaa crr11paJIH3aI!MH. TectthIH KOHTaKT Mem,n;y xpoMaTH,[(3MH ,n;aeT B03MOJKHOCTh o6MeHMBaThCH H,[(CHTH'lHbIMH Y'JaCTKaMH roMOJIOrH'lHhIX xpOMOCOM. 8To HBJICHMe Ha3bIBaeTCH 4. ,l(unJlomena - KpOCCUH206epo.M. xapaKTeptt3yeTCH B03HI1KHOBCHHeM CMJI OTTaJIKHBam·m. XpOMOCOMhl, COCT3BJIHIOliIHC 6MBaJieHT, Ha'lHHalOT OTXO,[(HTh ,n;pyr OT ,n;pyra B 06JiaCTH I!CHTpOMep. llpH pacxom,n;eHMH IIJIC'l xpoMOCOM B HCKOTOPhIX MCCTax 06HapyJKMBaeTCH HBJICHHe rrepeKpecTa I1 cu;errJieHHH - XHa3MhI, KOTOpbie CJiyJKaT CTPYKTYPHhIM BhipameHtteM KpoccttHroBepa. napa cecn>HHCKllX xpoMaTl1,!I X11a3Mbl ~a ceen>HHCKllX ><pOMaTl1,!I npo11CXOAl1T KpOCC11HrDBep nax11neHa ,lJ,11nnornHa
,ae;ieuue K1£em1m. M euo3 5. l(ua1wHe3 - 63 xapaKTeptt3yeTCJI MaKCHMaJihHOM crm:paJitt3au;ttei1, yKopo- qeHneM H yTOJilll;eHHeM xpOMOCOM. 0TTaJIKHBaHtte xpOMOCOM rrpo,n;OJIJKaeTCJI, HO OHH OCTalOTCJI coe,n;HHeHHhlMH B 6ttBaJieHThl CBOHMH KOHIJ;aMH. 51.n;phlIIIKO H a.z:i;epHaJI o6oJioqKa paCTBOPJIIOTCJI. IJ;eHTPHOJIH pacxo,n;JITCJI K IIOJIIOCaM, 06paayIOTCJl HHTH BepeTeHa ,n;eJieHHJI. Mema<jJa3a I (2n4c). BttBaJieHThr xpoMocoM pacnoJiararoTcn no 3KBaTopy KJieTKH. K HHM rrpttKpenJI5IIOTCJI HHTH BepeTeHa .n;eneHH51. AHa<jJa3a I ( 2n4c). HttTH BepeTeHa ,n;eJieHHJI coKpaw;aroTcn, HK rro.JirocaM KJieTKH xpoMocoMbt, a He xpoMaTH,D;hI, 1mK nptt MHT03e. B .z:i;oqep- pacxo.n;nTcn roMo.JiorttqHhre mrn KJieTKH nona.n;aroT TOJihKO no O,D;HOH H3 rraphl rOMO.JIOrHqHbIX xpoMOCOM. TeJto<jJa3a I ( 2n2c - n2c B 1caJ1CiJou KJtemKe ). qHCJIO xpoMOCOM B Kam,z:i;oil: K.JieT- Ke cTaHOBHTCJI rarrJIOH,IJ;HhIM, HO KaJK,n;an xpOMOCOMa COCTOHT H3 ,D;BYX xpoMaTH,IJ;. Ha KOpOTKOe BpeMJI o6paayeTC.H H,n;epHan o6onoqKa, rrpottcxo.n;ttT ,n;eJieHtte u;ttTOIIJiaaMbI. Y MHOrHx pacTeHHH KJieTKa H3 atta<Pa3hI I cpaay me nepexo.n;ttT B MeTallJaay II. Merncpa3a I (2n4c) AHa(j)a3a I (2n4c) Te11ocpa3a I (2n2c - n2c a Ka)l(,ll,oC1 KilSTKe) BTopoe .n;eneuHe Me:Hoaa (a1rnau;Houuoe) HHmepfjJa.Ja JI, uJtu uHmepKuHe3 ( n2c) - xapaKTepHa ,n;n.H JKHBOTHhIX KJie- TOK. IIpe,n;cTaBJI.HeT co6oii KOPOTKHH rrepephIB Mem.n;y nepBhIM H BTOphIM MeiioTH'IecKHMH .n;eneHH.HMH, BO BpeM.H KOToporo He rrpOHCXO,D;HT perrmrnau;tt.H ,n:HK. IIpofjJa3a II ( n2c ). XpoMocoMhI yToJiw;aroTc.H H yKopaqttBaroTC.H. 51,n;pbIIllKO H SI.n;epHa.H o6oJio•ma paapyillaIOTC.H. 06paayeTc.H BepeTeHo .n;eneHH.H. M emafjJa3a I I ( n2c). ,n:ByxpoMaTH.D;Hhre xpoMOCOMhI BhICTpaHBaIOTCJJ B aK- BaTopttaJihHOH IIJIOCKOCTH KJieTKH (MeTa<PaaHaJI IIJiaCTHHKa), K HMM rrpttKperrJISllOTCSI HHTH BepeTeHa ,n;eJieHH.H. AHafjJa3a II (2n2c ). ,n:ByxpoMaTH,D;Hhre xpoMoCOMhI .n;enSITCJI Ha ,z:i;oqepHtte XPOMOCOMbI (6bIBillHe xpoMaTH,D;bI), KOTOpbre pacxo,n;.HTCJJ K rrpOTHBOIIOJIOJKHbIM IIOJIIOCaM KJieTKH. TeJtofjJa3a II (2n2c - nc B KaJ1CiJou K.nemKe). XpoMOCOMhI .n;ecrrttpamrny- IOTc.H, CTaHoB.HTC.H HeBH.D;HMhIMH. 06paayeTC.H .H,n;epHa.H o6oJIO'IKa. Kam.n;oe JJ,n;po co.n;epmttT rarrJiott,n;Hoe qttcJio xpoMocoM. IIpottcxo,n;ttT ,n;eJieHtte u;ttTorrnaaMhl, Ma ttcxo.n;Hoii KJieTKH o6paayroTC.H 'lembtpe ian.nouOHbte KJieTKH.
06~aJt 64 npoc)la3a II (n2c) AHac)la3a II (2n2c) 6uo1wzuJt. Ocno6bL ifumo,11,ozuu Merac)la3a II (n2c) Te11oc)la3a II (2n2c - nc s Ka>KAO~ KlleTKe) 3Ha'!eHHe Mei1:03a 1. IIpe,1:i;oTBpa:u.~aeT y,n;BoeHtte qttcna xpoMOCOM B KaJK,n;OM HOBOM noKoneHHH. 2. 06ecneqHBaeT retteTttqecKoe pa3Hoo6pa3He raMeT, cmrnaHHoe c KpocCHHroBepoM H pacxom,n;eHHeM B attacpa3e I pa3HhIX Ha6opoB HeroMoJiorttqHhIX xpoMOCOM. 3. 5!BmieTCH OCHOBOH KOM6HHaTHBHOH H3MeHqlfBOCTH. PA3MHO>KEHHE OPrAHH3MOB Pa3.M1t0J1Cenue - CBOMCTBO JKHBhIX opraHH3MOB npoH3Bo,n;HTh ce6e no,n;o6HhIX. Cyru;ecTByeT ,n;Ba ocHOBHhIX cnoco6a pa3MHOJKeHHH - 6ecnonoe H nonoBoe. BecnoJioe paaMuoarnuHe BecnoJtoe pa3.M1-toJ1Ce1-tue ocyru;ecTBJIHeTcH npH yqacnm o,n;Hoii po,n;ttTeJibCKOH oco6u H npottcxo,n;HT 6e3 o6pa3oBaHHH raMeT. Oco6H o6pa3yIOTCH H3 o,n;tto:i1 HJIH HeCKOJlbKHX COMaTnqecKHX KJieTOK po,n;HTeJI.H ITYTeM MHTOTHqecKoro ,n;eneHHH. IloTOMCTBO HBJIHeTCH reHeTHqecKHMH KOITHHMH po,n;HTeJibCRoro opraHH3Ma. Hatt6onee urnpoKo pacupocTpatteHo cpe,n;H npo:KapttoT, rpu6oB H pacTeHHH. Bttonortt'!ec:Koe 3HaqeHtte 6ecnonoro pa3MHOJKeHHH: ycttJIHBaeT pOJih cTa6ttJIH3ttpyroru;ero eCTeCTBeHHOro OT6opa, cnoco6cTBYeT coxpaHeHHIO npHCIIOC06JIHeMOCTH K nocToHHHhIM ycnoBHHM cpe,n;hl o6HTaHHH. Crroco6h1 6ecnoJioro pa3MHOJKeHHH: 1. ,l{eJtenue - cnoco6 6ecnonoro pa3MHOJKeHHH, xapaKTepHhIH ,D;JIH o,n;HoKJieToqHhIX opraHH3MOB, rrptt RoTopoM MaTepttHCKaH oco6h ,n;eJIHTCH Ha ,D;Be HJIH 6oJibIIIee KOJIHqecTBO ,n;oqepHHX KJieTOK. Bhl,n;eJIHIOT: rrpocToe 6nttapttoe ,n;eneHHe (rrp0Kaptt0Tb1); MHTOTHqecKoe 6HHapttoe ,n;eJieHHe (npocTeHIIIHe, 0,D;HOKJieToqHbie BO,ZJ;OpOCJIH); MHO-
pa 3 MnoJtCenue opzanu.1M06. Becno1we pa3M1toJ1Ce1tue 65 PaaMHO»WHHC - .u.eJ1eHHe (6H11apnoe, MHTOTlfliecKoe, m113oron11H), - II0l!KOB8HJIC, - cjlparMe11rn111rn, c OOJ10)10'1'BOpemieM: - KOH'bJOraLU1fl' - - - KOIIY~1.HIUU1 IIOJIH3M6pHOHH.H, H30raMH.H, - cnopoo6paaona1me, - aereTaTHBHOe, - reTeporaMIUI, - Bea onJio.a,oTRopenHH: - ooraMH.H. KJ"I0HHpOB8HHC 118pTCHOfCHC3 .meCTBeHHOe .n;eJieHJfe, MJIM IUH30J'OHM.H (MaJI.HpMMHhIM IIJia3MO,Zl;MM). fl.n;po .n;eJIMTC.H c o6pa30BaHJfeM ,ZJ;BYX HJIM 60Jihlllero 'IMCJia .a.n;ep, KOTOphle OKpy.maIOTC.H CJIOeM II;HTOIIJia3Mhl, If B pe3yJihTaTe <jlopMnpyeTC.H caMOCTO.HTeJibHhlH opraHM3M. 2. IlO'llWBattue - cnoco6 6ecrrOJIOI'O pa3MHOJKeHM.H, rrpM KOTOPOM HOBhie oco6M o6paayIOTC.ff B BM,ZJ;e BhlpOCTOB Ha TeJie pO,ZJ;MTeJihCKOM oco6M • .n:oqepHMe oco6tt, OT,ZJ;eJI.rr.rrch, rrepexo.n;.rrT K caMOCTO.HTeJihHOMy o6paay .IKM3HM (rtt.n;pa, .n;po.mmtt) MJIM MOI'YT OCTaBaTbC.H rrpnKpernieHHhIMH, o6paay.rr KOJIOHMM (KopaJIJIOBbie IlOJIMilhI). 3. <PpazMettmau,u.H - crroco6 6ecrrOJIOI'O pa3MHOJKeHM.ff, rrptt KOTOpOM HOBbie oco6M o6paayIOTC.H M3 <jlparMeHTOB, Ha KOTOpbie pacrra.n;aeTC.ff MaTepHHCKa.H oco6b (peCHM'IHhJe, KOJih'IaThJe qepBM, MOpCKMe 3Be3,ZJ;hl, CIIMpOrMpa). 4. IIOJtU3M6puottu.H - crroco6 6ecrroJioro pa3MHomeHtt.rr, rrptt KOTopoM HO- Bh1e oco6tt o6paayIOTC.H M3 qacTeii:, Ha KOTOpb1e pacrra.n;aeTc.a 3M6pttoH (6poHeHOCII;hl, MOH03MI'OTHhie 6JIM3HeII;hl qeJIOBeKa). 5. BezemamuBttoe pa3MHOJ1Ce1-tue - crroco6 6ecnoJioro paaMHomeHn.H, nptt KOTOpOM HOBhie oco6tt o6paayIOTC.H M3 qacTeii: BereTaTMBHOI'O TeJia MJIH H3 oco6bIX CTpyKTYP (KOpHeBmIJ;e, KJiy6eHb, JIYKOBHII;hI, OTBO,ZJ;KH, ychl, JIMCTh.H M .n;p.). MomeT 6bITh ecTeCTBeHHhlM H ncKyccTBeHHhlM. XapaKTepHo ,ZJ;JI.rr pacTeHHM H HCIIOJih3yeTC.H B ceJieKII;HH (HcKyccTBeHHoe BereTaTHBHOe pa3MHOJKeHHe). 6. Cnopoo6pa3oBattue - pa3MHomeHHe aa C'IeT o6paaoBaHn.rr crrop. Xa- paKTepHo ,ZJ;JI.H pacTeHMM (BO,Zl;OpOCJIH, MXH, rrarropOTHHKH), rpH60B (neHHII;HJIJiyM, IUJI.HilO'IHhJe rpH6hI), rrpocTeMIUHX (crropOBHKH). 7. KJto1-tupoBa1-tue - ttcKyccTBeHHhIM MeTo.n; pa3MHomeHH.H, KOTOphIH 110- .HBMJic.a B Ha'IaJie 60-x rr. XX B. OcHOBaH Ha noJiyqeHMH HOBoro opraHH3Ma H3 O,ZJ;HOM KJieTKH HCXO,ZJ;HOI'O. B OCHOBe o6pa30BaHH.ff KJIOHa Jie.IKHT MH- T03 • .Il:JI.rr KJIOHHPOBaHH.H pacTeHHH KJieTKH o6pa30BaTeJihHOM TKaHM BhlP~HBaIOT Ha cnerr;HaJihHhlX IIHTaTeJihHhlX cpe.n;ax (MeTO,Zl; HCIIOJih3YeTC.ff ,Zl;JI.H noJiyqeHn.rr HOBhIX copTOB pacTeHnii:). B 1997 r. KJIOHnpoBaHneM 6hlJia IlOJiyqeHa OBe'!Ka .Il:oJIJIH, KOTOpa.H He OTJIH'IaJiaCh OT ,ZJ;OHOpa .rr.n;pa, BO CHJihBO OTJIH'IaJiaCb OT perr;nrrneHTa • .D:oHopoM 6bIJIH KJieTKH MOJIOqHOM
)f<CJie3hl 0.L(HOH rropO.L(hI, a peI.1,I:IITI:IeHTOM - .HMIJ,eKJieTKI:I c npe.nBapI:ITeJib- HO pa3pymeHHhIMH a.npaMM OBIJ.hI .npyroii: rropo.nhr. B HacTomu;ee BpeM.H Be.L(yTC.H pa60Thl no KJIOHI:IpOBaHI:IIO IIeJIOBeKa, 'ITO Bhl3hIBaeT 6ypHbie crropbl He TOJihKO cpe.L(I:I YlJeHhIX, HO II y pa3JII:IlIHhIX rpyrrrr HaceJieHI:Ifl. lloJIOBOe pa3MHO~eHHe IT oJwBoe paaM1wJ1CeHue ocyru;ecTBJI.HeTc.H rrpM yIIaCTI:IH .ll:BYX po.nMTeJil>CKHX oco6eii, conpOBOJK.L(aeTC.H IIOJIOBhIM rrpou;eccoM, rrpH KOTOpOM rrpOHCXO.L(HT CJIH.HHHe .L\BYX IlOJIOBhIX KJieTOK (raMeT). y HOBhIX oco6eii COIIeTaIOTCfl rrpH3HaKH .L\BYX pO.L(HTeJieii. IlO.HBHBIIrnec.H IIOTOMKH OTJII:IIIaIOTC.H reHeTH'IeCKH .npyr OT .npyra. 3TO rrpHBO.L(HT K IlO.HBJieHHIO HOBhIX KOM6HHaIJ,Hll reHeTHIIeCKOro MaTepHaJia. IIoJIOBoe pa3MHomeHHe ,ZJ;eJiaeT opraHH3M 6oJiee a.narrTHpoBaHHhIM K H3MeH.HIOIIJ,HMC.H ycJIOBI:IHM 0Kpyma10ru;eii cpe.L(hI. B npou;ecce 3BOJIIOIJ,HH rroJioBoe pa3MHOJKeHHe 6oJiee rrporpeccHBHO. He.nocTaTKH noJioBoro pa3MHOJKeHH.H: Heo6xo.nnMo yqacTne .L(Byx oco6eii (KpoMe napTeHoreHe3a) H Tpe6yeTC.H 6oJihIIIHM npoMeJKyTOK BpeMeHH, IIeM npH BereTaTHBHOM pa3MHOJKeHHH, He06XO.L(HMhlll B .ZJ;JI.H noJIHoro cpopMnpoBaHH.H opraHH3Ma. 3aBHCHMOCTH OT oco6eHHOCTH CTpoeHH.H raMeT BhI,ZJ;eJI.HIOT CJie.L(yIOIIJ,He cpopMhl IlOJIOBOro pa3MHOJKeHH.H: 1. 1130ZaMUH (OT rpeII. isos - paBHhIM H ga'mos - 6paK) - rrpHMHTHBHa.H cpopMa IIOJIOBOro npou.ecca, rrpH KOTOpOM CJIHBaIOTC.H .[(Be O.ZJ;HHaKOBhie ITO BeJIH'IHHe MaJihie IlO.L(BHJKHhie raMeThl (paBHOJKryTHKOBhie, B0.[(0pOCJIH, XHTpH.L(HeBhie rpH6hI). 2. I'emepozaMuH (oT rpeII. he'teros - HHOii, .npyroii, H gamos - 6paK) - npHMHTHBHa.H cpopMa IlOJIOBoro rrpou.ecca, npH KOTOpOM CJIHBaIOTC.H ,ZJ;Be IIO.ZJ;BHJKHhie co mryTHKaMH KJieTKH pa3Horo pa3Mepa (BO.L(OpOCJIH, XHTpH,ZJ;HeBhie rpH6hl). 3. OozaMUH (oT rpeq. oo'n - .Hiiu;o) - Han6oJiee pacnpocTpaHeHHa.H cpopMa IIOJIOBOro npou;ecca, npH KOTOpOM JKeHCKHe raMeThl - .HMIJ,eKJieTKH - He no,ZJ;BHJKHhI H 6oJiee KpynHhie no cpaBHeHHIO c MY.IKCKHMH raMeTaMH crrepMaT030H,ZJ;aMH (MHOrOKJieTO'IHhie JKHBOTHhie, rpH6hI, BO.D;OpOCJIH, BCe BhlCIIIHe pacTeHH.H). 4. KoH010za4ua (OT JiaT. conjugatie - coe,ZJ;HHeHHe) - npou;ecc coe,ZJ;HHe- HHH .ZJ;BYX oco6eii, rrpH KOTOpOM npOHCXO.ZJ;HT o6MeH HaCJie,ZJ;CTBeHHhlM MaTepHaJIOM, B pe3yJihTaTe qero IlO.HBJI.HIOTC.H oco6H, reHeTH'IeCKH OTJIH'IHhie OT pO,ZJ;HTeJihCKHX opraHH3MOB (HHcpy3opH.H-TycpeJihKa, KHIIIelIHafl naJIOIIKa). B pe3yJihTaTe KOH'hIOrau;HH He nponcxo.nttT yBeJIH'IeHH.H 'IHCJia oco6eii. 5. KonyJta4ua (oT JiaT. copulatio - CJIH.HHHe) - noJioBoii npou;ecc y O.ZJ;HO- KJieTO'IHhIX opraHH3MOB, npH KOTOpOM IlOJIHOCThIO CJIHBaIOTC.H KOilYJIHpyIOIIJ,He oco6H, BhIIlOJIH.HIOIIJ,He cpyHKIJ,HH IlOJIOBhIX KJieTOK (raMeT). IIpn aToM rrpoHCXO.ZJ;HT peKoM6HHaIJ,H.H (o6MeH) HacJie,ZJ;cTBeHHoro MaTepnaJia (MaJI.HpHMHhIM IIJia3MO.ZJ;Hll). 6. IlapmeHozeHe3 .neHne) - (oT rpeq. parthe'nos - .neBCTBeHHHIJ,a H genesis - pom- cnoco6 ynporu;eHHoro noJioBoro paaMHOJKeHH.H, npn KOTopoM
no1wBoe pa.1MHOJ1CeHue. I'aMemoiene3 67 aapO.ll;billl pa3BlIBaeTCH 113 HeOil.'IO)J;OTBOpemwii HllrJ,eKJieTKII. I1CTO'IHlfKOM HaCJie;:J;CTBeHHOro MaTepHaJrn }l,JIH pa3BHTllH HBJ'IReTCR ):t:HK H:ihi;eKJ'IeTKll (iw-wielie3). IIapTettorettea npe;:i;cTaBJIReT co6oii Mo;:i;mp1maumo IIOJIOBOrO pa3MHOiKeHHR B npouecce 3BOmOIJ,HH onpe;:i;eJ'IeHHbIX BH)J;OB iKllBOTHbIX. Illupmw pacnpocTpaHeH y 6ecn03BOHO'IHbIX (TJ'IH, OCbI, Il'IeJihI, .n.aqmttH) Hy II03BOHO'IHbIX iKllBOTHblX (npeCMhlKaIOIL\HecR, IlTHU.bI). PaaJrn'IaIOT ucKyccmBeflliblU tt ecmecmee1tlibtU (TJIM, oc1~r, nqeJibI) nap- Tettorettea. IIapTeHorettea BKJIIOqeH B iKll3HeHHhre UMKJihr :'.\l:HorHx napa3IITOB. Pa3MHOiKeHne, npn KoTopoM paaBHTHe Riiua npottcxo,n;HT aa c'IeT reHeTu:qecKoro MaTep11UJi:a cnepMaToao11;:i;a ( TaK KaK H;:J;po HH1J.eK"'IeTKI1 rror116aeT), HaahrnaeTcR pe,n;Ko (y iKI1BOTHb!X - I'epMa</JpoiJumbt - aniJpo2e1tea. B np11po;:i;e BCTpeqaeTCH Kpalitte Hae3)J;HllKll, y pacTeHllH - BII.D;hI, y K0Topb1x o.n;tta M Ta KyKypyaa). me oco6b cnoco6tta npoHa- Bo,n;nTh n iKeHCKHe, M MYiKCKtte raMeThl. fepMa<lJpo,n;tt:TH3M CtJMTaeTcR caMo:i1: rrpttMllTllBHOH <lJopMOH IlOJIOBoro pa3MHOiKeHMR H CBOHCTBeHeH rrpHMHTHBHhlM opraHII3MaM (npocTeiillltt:e, nJiocKne, JieHTO'IHhre tJepBM, yJIHTKH H .n;p.). IIpe.n;cTaBJIHeT co6oH: rrpncnoco6JieHHe K Cll,UR'IeMy, MaJIOilO,ll;BHJKHOMy, rrapa3HTHqeCKOMY 06pa3y iKll3Hll. I'aMeToreHe3 I'aMemoie1tea - 3TO npouecc <lJopM11poBaHHR raMeT, OCHOBHbIM 3TaIIOM KO- TOporo RBJIReTCR MeH03. Pa3Bl1Tlle raMeT y MHOrOKJieTO'lHbIX iKHBOTHhIX rrpo- cnepMamo3ouiJbt) o6pa3yIOTCR B CeMeHHMKax, meHCKHe IlOJIOBhie KJieTKll ( JlUl{eKJtemKU) - B HH'lHHKax. I'aMembl - 3TO BbICOKO,n;mp<lJepeHU.HPOBaHHbie KJieTKH, HCXO,ll;HT B noJIOBhlX meJieaax (rotta,n;ax). MymcKHe noJIOBhie KJieTKH ( HMeIOru;He ranJIOH,ll;HhlM tta6op xpOMOCOM. Ilpou.ecc o6pa30BaHHR MyiKCKHX IIO- cnepMamoieHe3, a meHCKHX - 060ZeHe3. CnepMamoienea no,n;paa,n;eJIReTcR Ha qeThrpe <lJaahr: 1. B </JaJe pa3MHoJ/CeHuH tt:3 KJieToK cnepMaToreHHOH TKaHH (roHOU.HTOB) B pe3yJibTaTe MHT03a o6pa3yIOTCR ,ll;HilJIOH,UHbie cnepMamoioHUU (2n2c). 2. <Pa3a pocma COOTBeTCTByeT HHTep<lJa3e I Merroaa (no.n;roTOBKa KJieTOK JIOBhlX KJleTOK Ha3bJBaeTCR K .n;eJieHHIO), B KOTOpoii npOHCXO,D;HT penJIHKaUHH ):t:HK. CnepMaToroHHll yBeJIHl{HBaIOTCH B paaMepax, H 113 Kam,n;orr KJieTKH pa3BHBaeTCH cnepMamo4um I nopRiJ1ca (2n4c). <jJaabt co3peeaHuJl KJieTKH .n;eJinTcn Mett030M. B pe3yJihTaTe I ,n;eJ1eHtt:H Meiloaa o6pa3yIOTCH .n;Be KJieTKH cnep.1namo4umbt II nopRiJKa (n2c), a IIOCJie BTOporo ,n;eJieHHH pa3BHBaIOTCR qeTbJpe O,ll;HHaKOBble no BeJIH'lHHe KJieTKH cnepMamuiJbt c ranJIOH,UHhlM aa6opoM XpOMOCOM (nc). 4. B </JaJe </Jop..MupoeaHUJl npoHcxo,n;tt:T KJieTOtJHa.ff ,n;n<lJ<lJepeauHpOBKa H 06pa3yroTCH c nep.1namo3ouiJbt c rarrJIOH.D;HhlM aa6opoM xpoMOCOM ( nc) · ropMoH, 06ecne'lHBa10ru;Hii crrepMaToreHe3, - mecmocmepoH. Kam,n;hre 2-3 Meca~ cnepMaT030H,Uhl o6HOBJIHIOTCH, H cnepMaToreHe3 npoMCXO,UHT HenpepbIBHO. 3. Bo BpeMn
68 06UfaR 6uo1tozu1i. Ocn06bL 4umo1tozuu reueT11- ' 3011bl 'IeCKHii CnepMaTore11e3 ; ( cjia:-ia )!{!13HeHHOro 0Bore11e3 ua6op IJ,HK"'la) cnepMaToro111111 Pa3MHO:iKfHmH (MHT03) i II • I ! I- III l 2n2c - --· - \. ····-· i-cnepMaTori.uT I nopH,n;Ka I PocTa (HHTep<lJaaa) i • ." e 'e f> .: c{ ·t OBOrOHl111 ·-- ·-- ,--- - ·-r· --·+ OBOlJ,HT lnopH,D;Ka 2n4c crrepMaT,OlJ,HT II'nopR,n;Ra I Co3peBaHHH . (Meii:oa) I OBOlJ,HT, pe):(.¥Kll,HOHHOe II rropRJ(Ka teJJbu,e /" I ...... ...... ""0 ·=::s • Q) [ __ f----~ IV -- • ) \,j n2c . .. HHIJ,eKJJe'i\Ka : 'peJJ,yKU,HOlll!b~e •• • nc crrep1:'4aT03oH,n;hl I ' ( ,,--'- , \___) '_ TeJJbll,3 I --- ---· --- -- - <t>opMHpOBaHHH L__ -- i i nc '-· Cmpoenue cnep.Mamo.10uiJa (pttc. 22, 23) 3peJiblH crrepMaT030H,ll; HMeeT rOJIOBKY' rneii:Ky H XBOCT. B ZOJl06Ke HaXO,ll;HTCH RiJpo (c rarrJIOH,ll;HblM Ha6opoM xpOMOCOM) H aKpOCO.Ma (BH,ll;OH3MeHeHHhlH KOMIIJieKC roJih,ll;:>KH, co.n;epmam;nii: <PepMeHThl, pacm;ern1mom;ne o60JIO'!KY HMIJ;eKJieTKll ). B rneii:Ke Haxo.n;HTCH u;eHTPHOJIH (coe.n;HH.mom;tte oceByro qacTh c H.n;poM roJioBKtt), B rrpoMe:>KyTO'!HOH (cpe.n;Heii:) 'laCTH - MHTOXOH,ll;pHll (CHHTeaupyrom;ue ATcf>). Xeocm rrpe.n;cTaBJieH o.n;HHM, y HeKoTophIX su.n;os .n;syMH H 6oJiee J1aymuKa.Mu (o6paaoBaHHbIMH MHKpOTpy60'!KaMH) • .Il:JIH ,ll;BH:>KeHllH :>KrYTHKOB llCIIOJib3yeTCH :meprHH MaKposprtt'!eCKHX CBHaeii: ATcf>. UuTOIIJia3Ma crrepMaToaott.n;a rrpe.n;cTaBJieHa TOHKOH o60JIO'!KOH. Oeoiene3 rro.n;paa.n;emreTc.H Ha Tpu ipaabl: 1. B </Ja3e pa3.MHOJtCenuR H3 KJieToK osoreHHoii: TKaHH .HH'!HHKOB (roaou;ttTOB) B pe3yJibTaTe MllT03a o6pa3yJOTC.H ,ll;HIIJIOll,ll;Hbie 060ZOHUU (2n2c ). 2. <Pa3a pocma cooTBeTCTByeT HHTep<Paae I Meii:oaa (rro.n;roTOBKa KJieTOK K .n;eJieHuro), B KOTopoii: rrpoucxo.n;HT perrJiuKan;HH .IJ:HK. 0Boroattu yBeJill'IHBaJOTC.H B pa3Mepax, BCJie.n;CTBHe HaKOrrJieHll.H IIllTaTeJibHbIX Be-
pa 3 .Mnom:enue opzanu3M06. I'aMemozeneJ 69 B P111c. 22. CnepMaTOreHe3 y MOPCKOiii CBHH- P111c. 23. CrpoeH11e cnepMarn3011Aa: A -06- KH. Y4aCTOK nonepe4HOro pa3pe3a 113B11roro t11Hiil BHA; 5 - KaHaJlbU,a ceMeHHHKa: 1 - uieiilKa; 3 - 2- cnepMaTOu,HT I nopsiAKa; 1..!HT II nopsiAKa; 4 - cnepMaTOIOHHS1; 3- cnepMaro- cnepMarnAa; 5 - coMa); 5 - cxeMa crpoeHHS1. 1 xBocr; 4 - fOllOBKa; 2 - annapar ronbAJKH (aKpo- S1APO; 6 - MHTOXOHAPHH; 7 - HHTb (>KryrnK); ocesasi 8 - L!eHTPHOll11; 9- npoMe>KyT04- HaS1 4aCTb cnepMaro30HA w;ecTB B Btt.n;e ace.TJTKa, H H3 Kam.n;oiI KJieTKH pa3BHBaeTcsr oBou,um 3. p.Rihca (2n4c). Bo BpeMsr </Ja3bt co3pe6anu.R nepe.n; oByJimi;neiI OBOD;HT JIHTCH nyTeM Metto3a. B pe3yJihTaTe I I I no· nopsr.n;Ka .n;e- .n;eJieHHH Metto3a 06pa3y10Tcsr: 60- Jiee KpyrrHaH K.JieTKa c IIHTaTeJihHblMH Bern;ecTBaMH H opraHeJIJiaMH - oBou,um II nop.Ra1ca (n2c) H o.n;Ha MeJIKasr KJieTKa - nep6oe noJt.Rpttoe B KOTopoiI ttMeeTCH (peay1Cu,uottttoe, ttanpa6umeJtb1toe} meJtbU,e, TOJihKO sr.n;po. BTopoe .n;eJieHtte Meilo3a ,n;oxo,n;HT .n;o cTa.n;tttt MeTacpa3hI, HO He rrpo.n;oJimaeTCH ,n;aJihllle .n;o Tex rrop, rroKa He rrpott3oiI.n;eT orrJio.n;oTBopeHtte, IIOCJie KOTOporo OBOD;HT pa3yeTCH KpyrrHasr K.JieTKa - II rropSI.n;Ka 3aBeplllaeT ,n;eJieHHe H 06- .Ritu,e1CJtem1Ca (nc), a TaKme Bmopoe no- Jl.Rpnoe meJtbU,e. Ifa rrepBoro rroJIHpHoro TeJihn;a 06pa3y10TCH .n;Ba MeJIKHX BTOpHqHbIX. Bee OHH B .n;aJibHettllJeM pa3pylllaJOTCH. 0BoreHe3 rrpoHCXO,ll;HT IIO,IJ; KOHTpOJieM ropMOHa npoiecmepotta. B OTJIHqtte OT o6pa3oBaHH.H crrepMaT030H,IJ;OB rrpon;ecc o6pa3oBaHH.H .Hiin;eKJieTOK y qeJioBeKa liaq1rnaeTCH B aM6pHOHaJibHOM rreptto,n;e I:l TeqeT rrpepbIBHCTO. y 3apO,ll;billJa llOJilIOCTbJO ocyw;ecTB.JI.HJOTC.H cpa3bl pa3MHO:lKeHHH, pocTa H HaqHHaeTC.H cpa3a coapeBaHHH. R MOMeHTY poac.n;eHH.H .n;eBoqKI:l B ee .HHqHHKaX HaXO,ll;.HTC.H COTHH TLicaq oson;ttTOB I rropsr.n;Ka, ocTaHOBHBIIIHXCH Ha cTa.zi;HH au1Cmuome1tbt (Bo Bpel\rn KOTOpoiI xpOMOCOMbI HMeJOT BH,IJ; «Jla.Mn06blX w,emolC») npo</Ja3bl 3a. B I .111.eito- nepwo.n; no.Jiosoro co3peBaHHH Metto3 B0306HoB.JI.HeTc.H, H Ka:lK,ll;hIH Mee.Hu; no,n; ,n;eiicTBHeM IIO.JIOBbIX ropMOHOB O,ll;HH H3 OBOD;HTOB ,IJ;OXO,ll;HT ,IJ;O MeTacpa3bI II
06UfaH 6uo.rwzun. Ocno6&t lfUmo.rwzuu 70 Meiio3a. B .n;aJihHeiiweM Meiio3 MomeT H.D;TYI .n:o KOHu;a npn ycJIOBllH OilJIO,ll;OTBOpeHHH, ec.1n onJIO.D;OTBope:f!ne He nponcxo.n;ttT, OBOUHT II nopH.n;Ka norn6aeT H BbIBO,ll;HTCH H3 opraHH3Ma. Cmpomue .HUU,eJC.llemJCu (pttc. 24). ct>opMa OKpyrJiaH, pa3Mepb1 OT HeCKOJibKHX MHKpOMeTpOB .n;o HeCKOJibKMX caHTMMeTpoB. KJieTKa Heno.n;BumHa.a, co.n;ep.m11T BCe K.JieTOtfHble opraHe.JI.JibI, MOpcpo.Jior11treCKH ocpopM.JieHHoe .a.n;po, H.n;epHO-UHTOnJia3MaTUtfeCKOe COOTHOllleHne C.ll;BHHYTO B CTOPOHy I.J;HTOn.Jia3Mbl 38 CtfeT PMC. I - 4- 24. 'rlillu,eKJleTKa MJl8KOm1Tat0Ll.\ero: 2 - 5lApo; 3 - 060110YKa; uvirnnna3Ma; 3anaca nuTaTe.JihHbIX Bem;ecTB - PHK, ¢01lllVIKYll5lpHbl8 Kll8TKVI me.JIT- Ka. JRe.JITOK co.n;epmnT mnpbI, yrJieBO,ll;bl, MUHepaJibHbie Bem;ecTBa, 6eJIKH (JinnonpoTen.n;hI, rJIHKonpoTen.n;hI). KoJIHtfeCTBO nnTaTeJibHbIX Bem;ecTB 38BHCHT OT ycJIOBHM, B KOTOpbIX npoHCXO.ll;HT pa3BHTHe 3apO.lJ;blllla. IIpn pa3BHTHH .aiiu;eKJieTKH BHe oprattn3Ma MaTepn (nTn- 95 '% OT ee o6'heMa, a BHYTPH opram-135 'X,' TaK KaK nnTaTeJibHhie Bem;eCTBa, Heo6xo.n;n- IJ;hI, perrTHJIHH) meJITOK COCTaBJI.HeT 6oJiee Ma (MJieKonnTaIOm;He)- MeHee Mbie .D;JIH pa3BIITMH, 3M6ptt0HbI no.JiyqaIOT OT MaTepn. Ha.n; n.Jia3MaTntrecKoii MeM6paHoii nMeIOTCH .n;onoJIHHTeJihHhie o6oJIOtfKH, pa3JIHtraIOm;necH no nponcxom.n;eHnIO: nep8U'lllbte ( JJCe.TLmO'lltbte) - o6pa3yIOTCH H3 noBepXHOCTHOro CJIOH He- 3peJIOM noJIOBOM KJieTKH (OBOI.J;HTa); npoHH38Hbl MHKpOBOpCHHK8MH H 0Tp0CTK8MH cpOJIJIHKYJIHPHhIX KJieTOK, no KOTOpbIM nocTynaIOT nMTaTeJibHhle Bem;ecTBa. BhlnOJIHHIOT 3am;HTHYIO cpyHKI.J;HIO, o6ecnetrHBa.a BH.ll;OBYIO cneu;mpwrnoCTb npOHHKHOBeHHH cnepMaT030IJ,ll;a. y MJieKO- nnTaIOIIJ;HX 3Ta 060.JIOtfIW Ha3hIBaeTCJI 6.JieCTJim;eii; BmopU'lllbte ( xopuon) - COCTOJIT H3 cpoJIJIHKYJI.HpHbIX KJieTOK HJIH Bbl- .n;eJIHeMbIX HMH ceKpeToB. BTop:ntrHa.a o6oJIOtfKa H:nu; HaceKOMbIX co.n;ep.m:nT KaHa.JI - MHKponn.Jie, trepe3 KOTOphIM cnepMaT030H.D; npomrna- eT B Hiiu;eKJieTKy; mpemu'l.nbte - cpopMnpyIDTCH BO BpeM.a npoxoa<,n;eH:nH Hiiu;eKJieTKH no .aiiu;eBo.n;aM H3 Bem;ecTB, CeRpeTnpyeMhlX meJie3aMH CTeHOK .aiiu;eBO,ll;OB. Xopolllo pa3BHTLI y penTHJIHM n nT:nu;, Hanpn.Mep, TpeTWIHhlMH o6oJiotrKaM:n IlTHI.l; HB.JIHIOTCH 6eJIKOBaJI, no.n;cKOpJiyno:aaJI lI CKOp.JiyrroBaJI 060JIOtfKH. Riiu;eKJieTKH MJieKon:nTa10m;:nx TpeT:ntrHoii o6oJIOtfKH He nMeIOT. Oce.Menenue - 3TO npou;ecc, 06ycJiaBJIHBa10m;:nii BCTpetry meHcK:nx n Mym- CK:nx raMeT y muBOTHblx. PaaJiutraIOT ecTecTBeHHOe u 11cKyccTBeHHoe oceMeHeH:ne (lll:npoKo 11cnoJIL3yeTca B pL160Bo,n;cTBe n m:nBOTHOBo.n;cTBe). HapyJJCnoe oce.Menenue BCTpetraeTCH y MHor:nx muBOTHLIX, 06uTa10m;ux B Bo,n;e. B 3TOM cJiyqae .aiiu;a ll cnepMaT030H,ll;hl Bbl,ll;eJI.HIOTCH B OKpyma10m;y10 cpe.n;y. r.n;e npOHCXO,ll;HT
pa3MnoJ1Cenue opzanu.1M06. I'aMemoiene3 HX cJIMJIHMe. BHympe111-tee oce.11te11e/-lue 71 xapaKTepHO /1.•1R o6I1TaTeJ1eH cywn, r.ne OTCYTCTBYIO'f yc.'IOBIUI ,ll;JIR. coxpaHeHMJI H BCTpeqlI raMeT BO BHenrnei1: cpe,11;e. ITplI 3TOH cpopMe oceMeHeHHR cnepMaT030H,ll;hI BBO.D;RTCR B rro.10Bbie nyT11 caMKH. OceMeHeHMe npe/1.rneCTByeT c.1e)l,y10~eMy npou,eccy - On1wrJomeope1tue - OIIJI0/1.0TBopeHHIO. 3TO npou,ecc c.111JIHHR MymcKoll: 11 meucKoii no.10Bhix J{JieTOK (raMeT), B pe3yJibTaTe KOTOporo 06pa3yeTCR. OIIJIO,ll;OTBOpeHHOe HMU,O, HJIH :mioma, .na10~aJI HaqaJio HOBOMY opraHI13MY. ITpou;ecc OIIJIO,D;OTBOpeHHR MO/KHO pa3,D;CJII1Th Ha HCCKO.'lhKO cpa3 (p11c. 1. <Paaa c6.iuJ1CeHUR. BcTpeqe raMeT cnoco6cTBYIOT raMOHhI - 25): x11M11qe- CKMe Be~ecTBa, BhI,ll;eJIJieMhie IIOJ'IOBhIMH K.'leTKaMM. CnepMaT030M,ll;hI BhI,D;eJIRIOT aH,n;poraMOHhl, a J:lMU,eKJieTKH - I'MHOraMOHbl. 8TH Be~eCT­ Ba RKTHBH3HpyIOT /J.BM/KeHHe H peryJIHPYIOT npOHHKHOBeHHe cnepMaT030H,ll;OB B .aiiu,eKJieTKY. 2. <Pa3a aKmueau,uu. IIpottcxo,n;HT KOHTaKT cnepMaT030tt,n;a H JIHll,eKJieTKH. B o6oJioqKe Jittll,eKJieTKH p.a.n;a mttBOTHhIX cy~ecTByeT OTBepcT11e - MHKponnJie, qepe3 KOTopoe npOHHKaeT cnepMaT030H,ll;. y 60JihIIIHHCTBa BH,ll;OB MHKPOilHJie OTCYTCTByeT H npOHMKHOBeHne cnepMaT030H,D;a eTCJ:l 3a cqeT aKpoco.M1toit peaKu,uu. ocy~eCTBJIJ:l­ IIo.n; .n;eiicTBHeM cpepMeHTa (rnaJiypo- HH,n;a3LI) H .n;pyrHX 6HOJIOI'HqecKH aKTMBHhIX Be1u;ecTB, OCB06om.n;aIO~HX­ CJ:l H3 aKpOCOMhl crrepMaT030M,ll;a, pa3pyrnaIOTCJ:l o60JIOqKJ1 J:llill,eKJieTKM. AKpocoMHaJI HHTh rrpom,rnaeT qepe3 pacTsopeHHYIO 30HY JIHll,eBLix CrrnJLrnat111~ di. cnep~IJTOJOH~!a c nepm1•11w11 / 060.10'1KOl1 \ flpmnn.:.uone1mc '1CpC3 ncpnn'lHYIO IlrraJMJHl 1fCCKaR -- MeM6paHa si:HucKJJeTJG.f' Ilepe11•-mafl - - - - o6oJ101..fKa • Jlapo H'1ltCK.rJCTKl1 """T' Pr.tc. 25. CxeMa npm.iecca on110,nornopeH111H: 1-4- nocne.11oearenbHb1e crn.11111111 npol\ecca
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06utall 6uo;wiull. I'euemu1w 90 y ' ,, 3Ill1- ,[\OMl1IIaHTHhIH. Ilpu.Mep. 0KpacKa orrepeinrn: y Kyp (pHC. . x 00.MUHGHlnllblU 3Ill1CTa3 - a) CTaT11qecK11H reH - 35) . 6) CUI benb."1 peu,ecCU6/l blU 3IlHCTa3 - TJ1qecKlIH reH - 3011CTa- pe11,eCCHBHbIH, npOHBJIHeT ,n;ettCTBlfe B reH I'OM0311I'OTHOM pe11,ecCHBHOM COCTOHHHlI H no,n;aBJUICT 3KCrrpeCCHIO HeaJIJICJibHOro retta. AMep11KaHcK11ii reHeTHK B. MaK- KhJOCHK orrttcaJI cJiyqali B Bo:vi6ee, Kor,n;a y :memu;HHhI c rrepsoii: rpynrroii: KpoBlf po,n;lIJICH pe6eHOK c qeTBepTOH rpyn- rroli. Ilpu.Mep. (Ifa,n;1rn, 1959 BoM6ei1cKHH q>ettoMeH r.) ~(Iolo) P: G: ¥ IBJBhh x IAIAHH cf' I 8 h IAH F 1 : JAIBHh - IV P11c. 35. HacnegosaHvie oKpacK~Y KYP npvi B3aVIMOgeC1CTBVIVI ABYX nap reHOB (rro cl:>eHOTHilY (AB) IOJO (IBIBhh), JBIB). a ITO rCHOTH- rry apOHBJIHeTCH KaK 3. IIoJlu.Mepu.H - s3a11ruop;eii:cTB11e HeaJIJie.'IbHhrx reHOB, rrptt KoTopoM 3a 0,[\HH rrpl13HaK B opraHl13Me OTBeqaIOT HeCKOJibKO nap HeaJIJieJihHbIX reHOB; rrp11 3TOM ct:>eHOTlIIllfqecKoe rrpO.HBJieHHe rrpH3HaKa 3aBHClIT OT KOJIHqecTBa ,n;oMlIHaHTHbIX lIJIH pe11,eCCl1BHblX aJIJieJieiL Ilpu.Mep. ll;BeT KO:>KH y tie;rJOBeKa (3a 11,BeT KO:>KH OTBeqaIOT 2-5 reHOB, Ka:>K.L\YIO rrapy 06oaHaqa10T IIH,n;eKCOM - P- ecTb rrnrMeHT P 1P1P 2P 2 - p - HeT rrttrMeHTa p 1p 1p 2p 2 - P 1p1P 2 p 2 - ¥ P 1P 1P 2P 2). qepH0Ko:m11:H: 6eJihIH MyJiaT P 1p 1P 2p 2 x P 1p 1P 2p 2 cf'=> B03MO:>KHhI BapttaHThr OT P 1P 1P 2P 2 ,n;o p 1p 1p 2p 2 (BepoHTHOCTh 1 16 ). Ilo TIIITY IlOJIIIMepIIII HacJie,n;yIOTCH MHOrIIe KOJI11qeCTBeHHbie nplI3HaKl1: I pocT, Bee, cnoco6HocTb K ceKpeu;1111 rpy,n;Horo MOJIOKa y 4. TleHempaHmHoCmb 9 11 ,n;p. -11acTOTa IIJilI BepoHTHOCTh cpeHOTl1Il11'IeCKOro npo.HBJie- HH.H .n;attHoro npH3HaKa cpe,n;n ecex oco6eii: reHeT11qec1rnx HOCHTeJieii: sToro npH3HaKa. 5. 31Ccnpeccu61wcmb -creneHh npoaBJieHlI.H rrp113HaKa OT MaKCIIMaJibHOH BbipameHHOCTH ,n;o MlIHHMa,1bHOH. Ilpu.Mep. Ilp11 IlOJIH,[\a:KTllJIHlI IIICCTOH rraJieu; MO:>KeT OKa3aThCH HOpMaJihHhIM, a MOateT rrpttHHTb Blill!lJaJiaHrH.
Cq,en.nennoe nac.nedoBanue. l'enemu'tec1we onpedenenue no.na. ~umozenemu1w 91 IletteTpaHTHOCTb H :3KCnpeccHBHOCTb HBJIJUOTCH pe3y.'IhTaTOM BJIHHHMH BCero reHOTttna H cpe,n;h1 Ha onpe,n;e.'JeHHhllI reH. C:u;emieHHoe HacJie.zi;onaHHe. feHeTH"1Jec1me orrpe.zi;eJieHHe rroJia. ll;HToreHeTHKa Bb1,n;eJIHIOT TpH ocHOBHhIX ypoBHH H3y<IeHHH retteTHKH: 1. OpiaHU3.MeHHblU - OCHOBhl 3aJIOiKeHbl r. Mett,n;eJieM B 1865 r. 2. KJtemo'lHbtu, mrn u,umoienemu'lecKuu - ocHOBhI 3aJiomeHhI B ;rn6opaTOpHH T. Mopratta B 1910 r. 3. MoJteKyJt.H,PHbtu - Ha'!aJI pa3BHBaThCH c OTKphITHeM CTPYKTYPhI MOJieKYJihI .JJ:HK (1953 r.). K Ha'IaJIY xx B. c pa3BHTHeM MHKpOCKOilH'IeCKOH TeXHHKH HaKOilHJIOCb MHOmeCTBO ,n;aHHhIX o noBe,n;eHHH xpoMOCOM. B 1902 r. Y. CeTTOH conocTaBHJI noBe.n:eHHe xpOMOCOM H nplrnHaKOB npH MHT03e H Meiio3e H o6HapyiKHJI, 'ITO XpOMOCOMbl H npH3HaKH «BeJIH ce6H» aHaJIOrH'IHO. BnocJie,n;cTBHH yqeHhiii npe,n;noJIOiKHJI, "llTO reHhI Haxo,n;HTCH B xpoMocoMax. K 1910 r. B Jia6opaTopnn aMepHKaHcKoro reHeTHKa T. MopraHa B pe3yJihTaTe onhITOB Ha nJio,n;oBoii MYIIIKe ,n;po3ocpHJie 6h1Jia ccpopMyJittpoBaHa xpo.11toco.11tna.H meopu.H nacJte8cm6eHHocmu. OcHOBHLie noJio:meHHH xpoMOCOMHOH TeopHH uacJie,n;cTBeHHOCTH 1. HacJie,n;cTBeHHhie e,n;HHHllhI, HJIH reHhI, JIOKaJIH3YIOTCH B xpo¥OCOMax. 2. Kam,n;hlii reH B xpoMocoMe 3aHHMaeT onpe,n;eJieHHOe noJiomeHHe - JtoKyc. feHbl B xpOMOCOMe pacnOJIOiKeHhl JIHHeHHO. 3. feHbl, Haxo,n;mu;neCH B O,D;HOH xpOMOCOMe, HacJie,n;yIOTCH c11enJieHHO H o6pa3yIOT rpynny c11erIJieHHH. qHcJio rpynn c11enJieHHf! paBHO ranJIOH,n;HOMY Ha6opy xpoMocoM. 4. Mem,n;y roMOJiorH'IHhIMH xpoMocot MaMH B03MOiKeH o6MeH yqacTKaMH - 1CpOCCUHZ06ep (pHC. 36). 5. PaccTOf!HHe Mem,n;y reHaMH H3MepHeTcH B MOpraHH.n;ax HJIH npo11eHTax KpoccnHroBepa: 1M=1 % . 6. PaccTOf!HHe Mem,n;y reHaMH nponop11ttoHaJihHO npo11eHTaM KpocI \ I \ CHHroBepa, a CHJia c11enJieHHf! Mem,n;y reHaMH o6paTHO nponop11nOHaJII:.Ha paCCTOf!HHIO Mem,n;y HHMH. B Jia6opaTopnH MopraHa 61:.1JIH npoBePi.tc. 36. Kpocci-1Hrosep: 1 - ABe roMono.Zl;eHhl pa3JIH"l1Hble :3KcnepHMeHThl. rw1Hb1e xpOMOCOMbl; 2 - l'IX nepeKp8CT BO OnhIThl MopraHa, ,n;oKa31:.rna10I11He BP8M5l KOHb10rau.i-1i-1; 3 - AB8 H0Bbl8 KOM61'1cn;enJieaaoe aacJie,n;oBaHHe (pttc. 37): HaLJ,l'!H '® 2JW~ QD 00 <DCD CDCD
06UfaR 6uo1wzu11.. I'enemuica 92 !'•. I I h• • b t> I I ., • • v ' •• v "i ib ~· • v 6 r, h •• h BA •• v •• " vl:j I I :tfr . ' -1-~ ..,, !v ~ 50'7" HeHpoccoeepbl 83 '-----v--Hpoccoaepbl 17 % % a PMC. 37. CL1en11eHHOe Hac11eAOBaH11e reHOB 0KpaCKl1 rnna 11 COCTOS1Hl1Sl KpblllbeB y nllOAOBOiil Myx11 (Drosophila melanogaster): a coM y caMuoe AP030¢lL111bl: pp KOKpblllblM caMuOM KpblllOiil caMKOiil TOMKOB: 6 (BBW) CKP01J.10Ha c YepHblM KOpOT- (bbw); Fi - cepbliil caMeu c AJ111HHblMl1 Kpb11lb5lMl1 (BbVv) CKP01J.10H c YepHOiil KOPOTKO- (bbw); F2 - 50 % - no11Hoe cuen11eH11e reHOB scneACTe11e orcyrcre115l nepeKpecra xpoMocaMKa cepa5l c AJ111HHblM11 Kpb11lb5lMl1 nOCKOJlbKY y caMua He npo11CXOAl1T KpOCC11HrDBepa, n05lB5lTC5l ABa B11Aa no- YepHblX KOpOTKOKPblJlblX 11 50 % - c AJll1HHblMl1 KpblJlb5lMl1 AJ111HHblM11 KPb11lb5lM11 6 - HenOJlHOe y caMOK AP030¢>11Jlbl: pp - CaMKa cepblX c HOPMallbHblMl1 KPblJlb5lMl1; (YaCrn'IHOe) cuen11eH11e npl13HaKOB BCJ10ACTBL1e nepeKpecra xpOMOCOM (BBW) CKpelJ.leHa c yepHblM KOPOTKOKPblJlblM caMuOM (bbw); F1 - cepa5l caMKa c (BbVv) CKpelJ.leHa c YepHblM KOpOTKOKPblllblM caMUOM (bbw); F2 - noCKOJlbKY y caM- K11 npo11CXOAl1T KPOCC11HrDBep rDMOJ10rl1'1HblX xpOMOCOM, o6pa3ylOTC5l YeTblpe rnna raMeT 11 n05lBSlTC5l YeTblpe B11Aa nOTOMKOB: HeKpoccosepbl - cepble c AJ111HHblMl1 KPblJlb5lMl1 (BbVv) 11 YepHble KOpOTKOKPblJlble (bbw), Kpoccosepbl - YepHble c AJll1HHblMl1 KPblJlb5lMl1 (bbVv), cepble KOPOTKOKpblJlble (Bbw) rea B b (black) v v (vestigial) II waaaK cepoe TeJio qepaoe TeJio HOpMaJibHbie KpbIJib.H aaqaToqab1e (KopoTKHe) B H V aaxo,n;HTCH Bn 7 V '% (M) B I aTan P: G: o.n;aoii xpoMocoMe :.,B+tB x b t v-~ -t-v ·tb/ v T i-v c:v ~ B++b F1: v_l I -t_ IV cepoe TeJio, HOpMaJihHble KPhlJiblI
93 Clfen11,ennoe nac11,ea0Banue. I'enemu'lec1we onpeae11,euue no11,a. l{umozeuemu1w II aTaII Peu;u11pmrnoe c11:pem;uBa11ue - CKpern;:rrnamrn caMu;OB (a) n caMoK (6) c peu;ecCHBHOH rOM03HrOTHOH oco6bIO. a) ____;__ b B_ I P: G: I vr- I 1 b ) ~b x v . I --v ·-,.I 6) . B +---! P: V ~ (5) ~ G: B+ j-b b-t +b vl -t v v~- lv v+-1 v -r-· v1 'lepnoe TeJio, KOpOTKHe KpbIJib.H 50 % 50 'X, II v y cf ,n;po30!l>HJibl IIOJIHOe cu;e11Jie11ne (HeT KpOCCHHrOBepa) => reHbl HacJie,n;yIOTCH cu;e11JieHHO. B pe3yJibTaTe aToro rroJiyqHJIHCh TOJibKO po,n;nTeJihCKHe coqeTaHHH reHOB. iV Q) ~ CD 83 '% neKpoccoaepttwe - (!D w 17 % KpoccoeepHbie --cepoe TeJio, HOPMBJibtthle Kpb!Jlb.H rb - - F2: b bl x F2: BT r ht v+ . v cepoe TeJio, HOpMBJlbHbJe Kpb!Jlb.H j-b vi- +v 'lepttoe TeJJo, KOPOTKHe Kpb!Jib.H HeKpoccoeepw - 83 % bl VT cepoe TeJio, KOpOTKHe Kpb!Jib.H qepttoc TeJJo, HOPMBJibHble Kpb!Jib.H Kpoccoaepw -- 17 % Y <.J? ,n;po3o!l>HJibI cu;errJieHne 11e110JI11oe. IIpou;eHT 11:poccn11r0Bepa 3aBHCHT OT paCCTOHHHH Mem,n;y reHaMH.
06~a11, 94 6uo1wzu11,. I'enemu1w f eHeTHqec1we orrpe.IJ;eJieHHe rroJia c;IO)KHhIH rrpH3Ha:K, HacJie,D,OBaHHC KOToporo orrpe,D,CJUieTCH pa3BI1- [fOJl - THeM 20/WO. Bhr,D,eJrnJOT HeCKOJihKO THITOB orrpe,D,eJieHHH rroJia B 3aBHCHMOCTH OT tIHCJia 11 cocTaBa 110.'IOBbIX xpoMOCOM (pHc. 1. 2. 3. 38): 1IeJJOBeK, MJieKOilHTarow;tte H ,D,p. XX Bo,D,ano:i1: KJIOII Protenor HeKOTOphre 6a6otIKH, IITHIIhl, pbI6bI, 3eMHOBO)J,Hhie 4. XY XX IlepenoH'IaTOKphIJihie XO zo zw zw zz 2n n (32 xp.) (16 xp.) Il'Ie,'Ihl foMoraMeTHhIH IlOJI - TpyTHH O,ll,HHaKOBhie xpoMOCOMbI. feTeporaMeTHhlH IIOJI - pa3Hhle XpOMOCOMhI. Y qeJioBeKa orrpe,D,eJiarow;ttM B pa3BHTHH rroJia HBJIHeTcH Ha.'IHtIHe Y-xpoMocoMhl. B Hett, rro-BH,D,HMOMy, Haxo,D,HTCH KJIIO'lesoii: reH, ttrparou~n:i1: pOJih peryJIHTopa. Y MyTaHTOB rrptt JIJ06oM KOJIHtieCTBe X-xpoMocoM ,D,06aBJie1rne Y-xpoMOCOMhI IIPHBO,D,HT K cpopMHPOBaHHIO MymcKoro cpeHOTHrra. XXY XO - CHH,D,pOM KJiaiiHcpeJihTepa. CHH.D.POM IIJepuiescKoro-TepHepa ( XXXoY~ XYY J 2 ). 71 0 Cu;em1eHHoe c IIOJIOM HacJie;i:i;oBaHHe IlpU31W1C cu,enJLell c nOJLO.M - reH, orrpe,D,CJIHIOUJ;HH pa3BHTHC oTOro rrpH3Ha- Ka, HaXO)J,HTCH B IIOJIOBOH xpoMOCOMe. x - cu;errJieHHhie npll3HaKH. y- cu;err- JICHHhie rrpH3HaKH (roJiaH,D,pnqec- x x x y 11 I1 ,~~"CD ~ r!f ii~.~ x x x y Koe HaCJie,D,OBaHMe OT OTIJ;a K ChlHy). BoJihIIIMHCTBO 6oJie3HeH - rrpH3HaKOB peIJCCCHBHbie. l.J;HTOreHeTHKa Kapuomun - aTo Ha6op xpo- MOCOM B COMaTH'leCKOH KJieTKe opraHH3Ma ,D,aHHoro BH,D,a. XapaKTepn3yeTC.H rrpaBMJiaMH XpOMO- COM: P111c. 38. XpoMoCOMHbli'i MexaH1<13M o6pa- P: Cf' 44 ayrocOMbl +XX x 44 ayrocOMbl + XY d' G: 22 ayTOCOMbl rrpaBHJIO IIOCTO.HHCTBa 'lllCJia xpoMOCOM; 30BaH1-tfl nona: '/>< ;; :::~~~~~: + X v~+Y F 1: 44 ayrocOMbl + XX 44 ayrocoMbl + XY rrpaBHJIO rrapHOCTH roMOJIOrMqHhIX xpoMOCOM (y )J,HilJIOH,ll,HblX BH,ll,OB; 0,D,Ha xpOMOCOMa IIOJiyqeHa OT OTIJ;a, ,D,pyraH - OT MaTeptt);
Cq,en;iennoe nac;ieiJoBanue. I'enemu"'ec1we onpeiJe;ienue no;ia. L{umozenemu1w 95 TipaBirnO IIH,D;IIBII,D;yaJibHOeTII - OTipe,n;eJieHHOe eTpOeHIIe XPOMOeOM (pa3Mep, cpopMa, co,n;epmaHIIe reHoB); TipaBIIJIO HeTipepbIBHOeTII - Tiepe,n; ,n;eJieHIIeM KJieTKII .IJ:HK y,n;BaIIBaeTeJI TIO npIIHD;IITIY KOMnJieMeHTapHOeTII. Bee xpoMoeoMhI Tio,n;pa3,n;eJIJIIOT Ha ayToeoMhI II TIOJIOBhie xpoMoeOMhI. II OJL06bte - aTo xpoMoeoMhI, OTipe,n;eJIHIOID;IIe cpopMIIpoBamrn MyJKeKoro Ii meHeKoro TIOJIOB. AymocOMbt - Bee xpoMoeoMhI B KJieTKax, 3a HeKJIIOqeHIIeM TIOJlOBhIX. IlpuMep: 1) KapIIOTIITihl qeJIOBeKa - 46 xpoMoeoM, 23 napbl xpOMOeOM 22 TiaphI ayToeoM +XX IIJIII XY; 22 A+ XX; 22 A+ XY; 2) KapIIOTHTI ,n;p030cpIIJihl - 8 xpOMOeoM; 3) KapIIOTIITI aeKapH,D;hl - 2 xpoMoeOMhI; 4) KapIIOTIITI eo6aKII - 78 xpoMoeoM. B 3aBHeIIMOeTII OT paeTioJiomeHIIH u;eHTpoMephl xpoMoeoMhI 6hrnaroT: 1. Memau,eHmpu'lecKue - paBHOTIJieqHe. 2. Cy6Memau,eHmpu'lecKue - HepaBHOTIJieqIIe. 3. AKpou,eHmpu'lecKue - TianoqKoBII,D;Hhie. 4. TeJtou,eHmpu'lecKue - u;eHTpoMepa paeTioJiomeHa Ha KOHu;e xpoMOeOMhI. PaaBIITIIe MeTo,n;oB oKpaeKli II MIIKpoeKoII nIIpOBaHIIH TipliBeJIO K TIOJIBJieHIIIO MeTo,n;a au<P<PepeHu,UaJlbliOZO OKpautu6aHUH. Kam,n;aJI xpoMoeoMa IIMeeT eBoii: IIH,D;HBli,n;yaJihHhIH pIIeyHOK paeTionomeHIIH ayxpoMaTIIHOBhIX II reTepoxpoMaTIIHOBhIX yqaeTKOB. B 1968 r. mBe,n;eKIIii: D;IITOJior T. KacTiepeeoH IV e KOJIJieraMII, a B 1971 r. IIapIImCKaJI KOHcpepeH~H KJiaCCIIcpIIu;IIpOBaJIII MeTO,ZJ;hl CTaH,n;apTII3a~H xpOMOeOM (paa6HBKa TIO rpyTITiaM): BCero Ce.Mb zpynn - no BeJiffqHHe II TIOJIO;KeHHIO u;eHTs poMepbI (pHe. 39). Ilpwnep. 1. fpyTina A - N2 1, 2, 3 - KpynHhle MeTaD;eHTpHqeeKIIe. 2. X-xpoMoeoMa - rpyTina C, epe,n;HHH cy6MeTau;eHTpIIqeeKaJI. PMc. 39. <t>opMbl xpoMocoM: 3. Y-xpoMoeoMa - rpyTITia G, MeJIKaJI aKI - TenoL1eHTpw-1ecKafl; II - aKpOpou;eHTpHqeeKa.H. Yponuu KOMIIaKTHaau;uu (y1tm1,a:KH) xpoMocoM B PMHhie cpaahl KJieToqHoro D;HKJia (HHTepct>aaa HJIH MHT03 (Meii:oa) xpOMOeOMhl HMeIOT Pa3Hoe CTpoeHHe. L1eHTp1114ecKafl; Ill - cy6MernL1eHT- p1114ecKafl; IV - MernL1eHTp1114ecKafl. 1 - L\eHTPOMepa; 2 - cnyrHlllK; 3 KOPOTKOe nne40; 4 - AJJlllHHOe nne40; 5-xpoMaTlllAbl
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97 Ha xpoMOCOMa MO:>KeT COCTOHTh H3 6oJiee ,UHIC 1 ThIC . .11,Byxr~eIIO"Y:e'!HbIX M0.1eKy.1 TaKtte xpoMOCOMhr BCTpe'IaIOTCH B KJICT·Kax CJIIOHHhIX me.1e3 ttaceirn- MbIX, B KJI€TKax HX MaJibIII1I'I1€BhlX cocy,n:on. IloJittTeHHbie xpOMOCOMhI xopowo Bll.11,Hhl B CB€TOBOH MttKpOCKOil, Korn;a KJieTKa HaXO,ll;ttTCH B HHTepcpa3e. feHeTHKH I1CIIOJih3YlOT 3TI1 XpOMOCOMbl ,ll;.'1.H cocTaBJieHHH reHeTH'l€CKHX KapT H, CJie.11,0RaTe.1hHO, .11,JIH onpe,n:eJieHHH TIOJIO;KeHHH reHOB B xpoMOCOMax. MOJl EKYJlSI PHASI rEH ETll1 KA MaTeptta.'lhHhIM cy6cTpaToM HacJie.11,cTBeHHOCTH H tt3MeH-Y:HBOCTH HBJIHIOTca HYKJiettHOBhre KttCJIOThI. PaaJittY:aIOT ,n;sa Btt,n,a HyK.1ettHOBhrx KttcJioT: llHK (,n,eaoKcttptt6oHyKJieHHOBaH KHCJioTa) H PHK (ptt6oHyKJieHHOBaH KHCJioTa). XnMtt-Y:eCKH Hatt6oJiee ycTOM-Y:HBa .IJ:HK. MoJieKyJia .UHK HBJIHeTCH cy6cTpaTOM HaCJie,D,CTBeHHOCTH H PHK - H3M€H-Y:HBOCTH B 60JihillHHCTBe :>KHBhIX CHCTeM. ocHOBa HacJie,n,cTBeHHOCTH y MHornx Bttpycos (snpyc Ta6a-Y:HOM Moaa- HKH, peTpOBttpychI II03BOHO-Y:HhIX). ,r:(HK H TH,D,hI (pHC. PHK 40). 3TO noJIHMephI, MOHOMepaMH KOTOphIX HBJIHIOTCH HYKJieo- Coe,n;ttHeHHe HYKJieoTtt,n,oB B u;errtt rrponcxo,n;ttT aa cqeT o6paaoBamrn cpoccpo3cpHpHoil: CBH3H Mem,n,y OH-rpyrrnoil: (*) tt cpoccpaTHOM rpyrrrroil: cne,n,y:ioru;ero HYKJieOTH,n,a. CxeMy cTpoeHHH .IJ:HK CM. B paa,n;ene « 11;ttTOJIOrHH » • Ha rrpHHIJ;HIIe KOMIIJieMeHTapHOCTH OCHOBaHhI MeXaHU3Mbl MampU'lliOZO cu11.me3a: 1) PenJLu1cau,uR .UHK (y,n;soeHHe). 2) TpattCl{:punu,uJl - CHHTea PHK Ha MaTpttu;e .UHK. 3) TpaH.CMlU,UJl - CHHT€3 6eJIKa Ha ptt6ocoMe. - aaOTHCTbie OCHOBaHHH (A,n;eHHH,THMHH,fyaHHH IJ;HT03HH) *OH IieHT03a - ,11;e30KCHptt603a (HJIH p11603a)
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105 Pezy.1uHfUR pa6ombt zeno6 - JiaKT03a CBH3hrnaeTCH c 6eJIKOM-penpeccopo!'v1; - penpeccop yxo.n.wr co csoero MecTa Ha MOJieKyJie ,[{HK (orrepaTope): - oTKphrnaeTcH «ITYTh» .n;JI.H pa6oThr PHK-rroJinMepa3hr; - CHHTe3HPYIOTC.H M-PHK H COOTBeTCTBYIOrnHe cpepMCHThl ,D;JLH paCIIJ,CII.TICHHH JiaKT03hI; - JiaKT03a pacrnerrJI.HeTc.a; - c pacrnerrJieHneM rrocJie.n;He:il: MOJieKyJihI JiaKT03hI 6eJIOK-penpeccop BHOBh CTaHOBHTC.H CB06o.n;HhIM, «Ca,Z:VITC.H» Ha )lHK-onepaTop, 6JIOI\'.HpyeT TpaHCKPHTIIl,HIO. IIpn .n.o6asneHHM "i:aKT03hI xo.n. co6hrTuM CHOBa rroBTOpHeTc.H. Be"10K-penpeccop Ko.n;wpyeT oT.n;eJihHhIM reH peryJI.HTop. 8Ta cxeMa pa6oThI reHa y 6aKTepttn rro"1yqnJia Ha3saune ::mcnpeccttn. Onepon - 3TO rpynna CTPYKTYPHhIX reHoB (Il,HCTpOHOB), uaxo.n.HrnnxcH rro.n. KOHTPOJieM o6rnnx peryJI.HTOpHblX 3JieMeHTOB (rrocJie,n,oBaTeJihHOCTe:il: ,[{HK H 6eJIKOB-peryJI.HTOpOB). CuuTe3npyeMa.H M-PHK y rrpoKapnoT rroJIHil,MCTpOHHa, T. e. HMeeT tteCKOJihKO CTpyKTYPHhIX reHOB. 9y1Capuomb1. CnHTe3upyeMhre M-PHK .HBJI.HIOTC.H MaTpwu;aMn ,Zl,Jl:.H oT,n,e.TJ:hHhIX neIITH,D;OB, T. e. OHH MOHOil,HCTPOHHbie. KaJR,D;hlM reH B orrepoHe MMeCT csoIO cucTeMy peryJI.Hil,HH. K peryJinpyIOIIVIM 3"i:eMeHTaM rrpou;ecca 3I<crrpeccun OTHOC.HT )lHK-rrocne.n.osaTeJibHOCTH H 6eJIKH-peryJI.HTOpbI. )lHK-rrocne.n.osaTCJibHOCTH: npo;;wmopbt - yCHJIHBaIOT TpaHCKpHIIIl,HIO B nec.HTKH H COTHH pa3; 3HXancepbt - HaXO,D;HTCH Ha 60JiblliOM paCCTO.HHHH OT rena; CllUJl€HC€pbt ( «rJiyIIIMTeJIH») - HaXO,D;.ffTC.H Ha 3HaqnTeJibHOM paCCTO.HHHM, CHHJRaIOT TpancKpHTIIl,HIO. BenKn-peryJI.HTOphr - 3TO 6eJIKOBhre q>aKTOphr TpaHcKpunu;ttn, KOTOphre MoryT 6bITh o6rnMMH ,D;JI.H pa3HhIX rpyrrrr reHOB. AKTMBHOCTh reHOB MOJReT TaKJRe perynnpoBaTbC.H rryTeM H3MeHeHM.H cTerreHH KOMrraKTu3au;nu xpoMaTuHa (crrnpaJIH3au;uu). Penpeccop CrpyKTVPHble reHbl npaMOlOp .. oneparop ,--------__,.,_----~ f-1oc1+P-0-+- /1eZ-+4cY+/acA-£ ~~. M-PHK lac- M·PHK /ac-penpeccopa lac- penpeccop fJ· ranlKTO:tHA33a f3· ralUIKTOlMAnepMea3a (J. ranaKT03M.ATP8HC81,leTMna~a PMc. 46. naKT03Hb1~ onepoH
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Meau1fu1tc1wR. zenemmw. I'enea1wzu'LeCKuii MemoiJ 107 Cyw;HoCTb MeTo.n:a: COCTaBJ'IeHIIe pO,ll,OCJIOBHOH (rryTeM orrpoca) c lICIIOJib30BaHIIeM CHMBO- J'IOB. Ilpo6aHO - qeJ'IOBeK, pO,ll,OCJ'JOBHYIO KOTOporo H3yqamT; reHeaJIOrlfqecKHH aHaJIH3. Il03BOJI.HeT BbIHCHHTb THII HaCJie,ll,OBaillf.H MHOrHX rrpH3HaKOB. OcHOBHhre THIIhI HacJie,D,OBaHH.H rrpH3HaKoB: 1. Aymoco.M1w-oo.MuHaHmHbLU mun Hac.rieooea1wJ1 (reH Haxo,nHTC.H B ayTo- coMe, .HBJI.HeTC.H JJ;OMHHaHTHbIM). XapaKTepHhre rrpH3HaKH ayTocoMHo-,n;oMHHaHTHoro Tttrra HacJie,n;oBaHH.H: npH3HaK rrpO.HBJIReTC.H B KaJKJJ;OM IIOKOJ'JeHHII (rro BepTHKam:1); rrepe,ll,aqa rrpttanaKa (aa6oJieBaHH.H) OT po,ll,HTeJieil: ,n;eT.HM; o6a IIOJia rropaJKaIOTC.H c O,ll,HHaKOBOH qacTOTOH. Ilpu.Mepbl HacJie,ll,OBaHH.H no ayTOCOMHO-,ll,OMHHaHTHOMY THrry: CHH,ll,pOM Mapq:iaHa ( «rrayqhlf» rraJ'IhIIhI, aHeBpttaMa aopThI, ITO,ll,BbIBHX xpycTaJIHKa); TIOJIII,ll,aKTllJIH.H (rneCTb rraJihl!eB); 6paXH,D,aKTHJIHa (IWpOTKHe naJibIIbI). lfHor,ll,a rrptt auaJittae po,D,OCJIOBHOH o6ttapyJKHBaeTc.H oco6a.H CHTYaIIH.H ,ll,OMHHaHTHbIH reH He rrpo.HBJI.HeTC.H, T. e. He rreHeTpHpyeT. IlpH 3TOM BHJJ; po,n;oCJIOBHOH MeH.HeTC.H. IleHempaHmuocmb - aTo qacToTa rrpo.HBJieHH.H reHa cpe,D,H HOCHTeJ'Jeil: ,n;aHHOro reHa, eCJIH reH rrpO.HBJI.HeTC.H, TO OH rreHeTpaHTeH. IleHeTpaHTHOCTb npe,ll,CTaBJI.HeT co6oH: OTHOlIIeHIIe oco6eH:, HMeIOru;Hx ,ll,aHHbIH npH3HaK, K oco6.HM, HMeIOru;HM ,n;aHHbIH reH. BhrpaJKaeTc.H B npoIIeHTax. HarrpHMep, neHeTpaHTHOCTb ctttt,n;poMa Mapcpana CeTqaTKH) - 30 'X,; peTI·rn06JiaCTOMhI (orryxoJih 80 '%. ihccnpeccueHocmb - 3TO cTerreHb BbrpaJKeHHOCTH retta. CTeneHh BbrpaJKeH- HOCTH IIPH3HaKa TaKJKe yqHThIBaIOT rrpH COCTaBJieHHH po,n;OCJIOBHOH. 2. Aymoco.MHo-peu,eccueHbtu mun Hac.rieooeauuJl (ren ttaxo,n;HTC.H B ayToco- Me, .HBJI.HeTC.H peIIeCCHBHbIM). XapaKTepnbre rrpH3HaKH ayTocoMHo-peIIeCcMBHoro TMrra ttacJie,n;oBaHM.H: npM3HaK rrpO.HBJI.HeTC.H He B KaJK,ll,OM ITOKOJieHMll (qaw;e npll 6JIM3KOpo,n;CTBeHHhIX 6paKax); 60JibHhie ,ll,eTH poJK,n;aIOTC.H OT cpeHOTllIIMqecKH 3,ll,OpOBbIX pO,ll,MTeJieH (HOCMTeJieH reHa); o6a IIOJia HaCJie,ll,yIOT rrpH3HaK c O,ll,MHaKOBOH qacTOTOH. Ilpu.Mepbt HaCJie,n;oBaHM.H ITO ayTOCOMHO-peIIeCCMBHOMY THrry: aJib6MHll3M (HapyrneHMe o6MeHa aMMHOKMCJIOT ll CllHTeaa rrMrMeHTa MeJiaHMHa); cpeHMJIKeTOHypH.H (HapyrneH o6MeH cpeHMJiaJiaHMHa :::> rrpOHCXO,ll,llT Ha- KOITJieHHe B Moqe rrpo,ll,yKTOB ero OKHCJieHH.H, ,ll,eHCTByrom;Hx Ha IIeHTpaJibHYIO HepBHYIO CllCTeMy). 3. X-cu,enJleHHbtu oo.MunanmHblU mun uac.Tleooeanua B X-xpoMocoMe, aBJI.HeTca ,ll,OMMHaHTHhIM). (reH Haxo,n;HTC.H
06Ufa1l 6uo;wzun . I'e1temu1w 108 I3 X-xpoMocoMe ttaxo11,wren 60,1 ee 200 1·ettoll. Y meHII.urn aHOMa: cbHbIH re11 MOJKeT H8XO.ZU1ThC.H B 011,HOii (reTepo:rnroTa) IL111 B 06e11x (rOM03IffOTa) X-xpOMOcoMax; y Mymq11H TOJihKO B 011,noii: X-xp0Moco.:-.1e. Y MY:>K'll1H ucer.ua uptteyTeTBYeT TOJibKO o.n;MH a;:uieJib reHa (reMH3J.ffOTHOeTb), cuenJieHHoro c X-xpoMocoMoii:. I'e.wu.wzomHOCmb - aTo npueyTCTB11e B opnrn11aMc TOJihH:o o,r.uroro a.1.rteJIH 1wKoro-.::iu60 rena. Ilpu.¥Lep: rCHhI B X-xpoMoeoMc y MYJK'Il1H - XY; reHhI B Y-xpoMoeoMe y My:>Kq11H - XY; reHhI B X-xpoMoeoMe y memu:mr - XO (euHp;poM IllepemeBeKoro---TepHepa). XapaKTep11hre npM3HaKH X-euern1eHHoro 11,o:vi:1111a11Tttoro nma 11acJie,[\0Bam.u1: 3a60.1eBaHMe npoeJieJKHBaeTe.H B Kam.n;oM IlOKOJieHMH; ee,1u 60JieH OTeu, TO see .n;oqcpu 6y11,yT 60.'JhHhIMH, a ebIHOBb.fl - 3.llOPOBhIMM; eeJiu 6oJihHa ~vrnTh, TO sepo.HTHOeTh pomp;etnrn 6oJihHoro pe6eHKa paBHa 50 •y., He3aBHeHMO OT no.'Ia; 60,1e10T KaK MyiKqHHhI, T8K l1 .mem:.u;uHbI (6oJihHhlX mem.u;11H B eeMLe B ,n:sa pa3a 60JibUie, qeM 6oJihHhIX My.mq11H). IlpuMPpbt X-c.u:enJieHHoro ,n:oMHHaHTHoro Hae.1e,n:osaHu.H: KOpuqHeBa.fl OKpaeKa :JM8JIH 3y60B; BHTaMMH D-pe3MCTCHTHbIH paxH'r. 4. X-cu,enJLPltH.btu peu,eccuettbtii mun 11ac. iedo6anuR (reH Haxo,n:MTea s X-xpoMoeoMe, HB.'IRCTC.fl peueceHBHh!M). XapaKTepHhie nptt3HaKw X-c1~enJieHH01·0 peueccttBnoro Tnna HacJie,n:oBaHHR: 6oJieIOT npeMMYlllCCTBCHHO My)KqHHhr; 60JihHLie ,n:eTH pom,naIOTCJl OT QleHOTHilHqeci-m 3,l(OpOBLIX po,n:HTeJiei1, HO M8Tb 6oJihHOro .flBJIHeTCH reTep03HI'OTHOH HOCHTeJihHl1UCH COOTBCTeTBYIOU~ero reHa; 6oJihHhre Mymq11Hb1 He nepe,n:aI-OT 3a6oJiesaHne ChIHOBhJ:IM, HO nee ,n:oqepn - HOCI1TeJihHHU:hI COOTBeTeTBYIOlllero reHa. IlpuMPpbt X-euenJieHHoro pe~eeeMBHoro HaeJie.n;oBann11: reMOQlMJIHJJ A (HeCBepThrnaeMoeTh KpoB11) (p11e. 48); MHOnanrn )lrorneirna (aTpOQlH.H Mhlllle<JHhlX BOJIOKOH); ~8JihTOHH3M. 5. Y·cu,enJteunoe 1-tacJLeao6aHue (reH Haxo,n:MTCJJ s Y-xpoMocoMe; Bcer,n:a B I'CMH3HPOTHOM COCTOJ:IHHH, Il03TOMY seer.n:a npO.HBJIJ:leTeH). XapaKTeptthle nptt3HaKn Y-cuerrJieHHoro HaeJie,n:osaHMH: reHhI, JIOKaJIH3oBaHHbie B Y-xpoMoeoMe, nepe,n:aIOTCH TOJihKO ebIHOBhHM (roJiaH~pnqeeKoe HacJie.n;oBarrne); ,n:oqepu Bcer,n:a a.n:opOBhI, T. K. mrnor~a He noJiyqa10T Y-xpoMOCOMy. IlpuMep Y-euenJieHHoro HaeJie,n:oBaHm1: rnnepTpMX03 MoqKM yxa ( «MOXHaThie yrnn »). 6. I'oJLa1tapu'lec1we HacJLeaoeaHue - 3TO HacJief\OBaHMe rrpM3HaKa TOjihKO no MymcKOH JIHHl1H (nepe.n;aqa OT OT~a K ChIHy).
L1,umozenemu•1ec1wu 1tzem0<J 109 II 6 III 6 P111c. 48. PoAocnosHasi ceMblll c reMocjm11111ew A MeTo,n; BKJIIoqaeT ;mcrrpecc-.n;mi.rttocT1rny rroJia (onpe,11,eJrentte X- 11 Y-xpoMaTHHa) H 1Capuomunupoea1tue - orrpe,n;eJieHHe KOJiuqecTBa If KatieCTBa xpo- MOCOM c IJ,eJihIO ,11,HarHOCTHKH xpOMOCOMHhlX 6oJie3Heii (reHOMllhlX MyTaIJ,HH, xpOMOCOMHhlX a6eppaI.1,llH). MaTepua.'loM ,11,JIH I.l,llToreneTHtfecKoro HCC"'le,n;oBaHmI MoryT 6hITh KJieTKH rrepmpepttqecKoii KpOBll (JIHMcpOI.1,HThI); KJieTKH, noJiyqeHI!hie npn aMHHOII;eHTe3e HJIH 6HOHCHH xopHoHa. :J1Ccnpecc·iJua21-wcmu1Ca - ::>TO ttccJie,11,oBaHtte noJioBoro X-reTepoxpoMaTH- Ha (TeJihI.1,3 Bappa) B H,11,pax KJICTOK CJIH3HCTOM o60JI0'!I{H pOTOBOH IIOJIOCTH HJIH B .n;pyrHX KJieTKax. B ttopMe y mettru;HH, MMeIOru;ttx ,n;Be X-xpoMOCOMhI, o.n;na MHaKTHBI1pyeTc.H, o6paay.H reTepoxpoMaTMHOBYIO rJihI6Ky B .H.n;pe - TeJihIJ,e Bappa. Y MYJKLJHH (XY) ee HeT. MymcKoii Y-rro.'!oBoii: xpoMaTIIH (F-TeJihUe) orrpe.n;eJI.HIOT rryTeM cneIJ,HaJihHoro OKpallIMBaHIUI H rrpoCMOTpa B JIIOMHHecueHTHOM MHKPOCKO!le . Y-xpoMaTHH BhI,!l;eJI.HeTC.H KaK CHJihHO CBeT.Hlll;8.HCJI TOqKa. qHCJIO Y-TeJreu COOTBCTCTBYeT q11cJiy Y-xpoMocoM B KapHOTHne. Kapuomunupoeanue - 3TO anaJIH3 cpoToKapttorpaMMhI. ,n:JI.H anaJittaa Ka- PHOTHrra xpOMOCOMhl pacrroJiaraIOT IIO rpymraM B BH,n;e H,!l;HOrpaMMhl (pHC. HiJuozpa.M.Ma - 49). aTo cttcTeMaTH3ttpoBaHHhiii KapHOTHrr. CorJiacno ,n:ettBepcKoii KJiaccmpttKaUHH (,n:ettBep, CIIIA, 1960 r.) xpoMoco- Mhl pacIIoJiaraIOT IIOIIapHO IIO Mepe y6hIBaHHH HX BeJIHqHHhl, c yqeTOM pacrroJlO:i«eHHH ueHTpOMephl, HaJIHqllfl BTOPH'IHhIX rrepeTH:i«eK l1 CIIYTHHKOB. IloJIOBhle XPOMOCOMhl BhI,D;e.'!JIIOTCJI OT,D;eJihHO. IlapII:i«CKa.H KJiaCCHcpl1K8I.l,H.H aJihHOH OKpacKH xpoMocoM. B (1971 I'.) OCHOBaHa Ha MeTo,n;e .n;mpc}lepeHI.I;H- Kam,uoii: nape xpoMOCOM IIpOHB.'!JieTcH xapaK- TeptthIH TOJlhKO ,!l;JIJI Hee YHHKaJihHhIH rrop.H,!l;OK qepe,n;oBaHH.H TeMHhlX H CBeTJlhIX IIOJIOC reTepo- H ayxpOM8THHa.
06utaJL 6uo:wzun. I'euernuKa 110 lJ A 6-12 c f)"Al>Ac')I\ 13-15 XX IA IA 1J F: !6 17 :i:: '----v------' AA AA .t ~'r' F G P11c. 49. Kap~ornn 4enoseKa XpoMoCOMhI qeJIOBeiw pa3,n;eJIHIOT Ha ceMh rpyrrrr: 1. A - KpyrrHhie MeTaueHTpIIqecKIIe (1, 2, 3). 2. B - Kpyrrtthie cy6MeTauettTpIIqecKtte (4, 5). 3. C - cpe,n;ttIIe cy6MeTaueHTpttqecKtte (6-12). 4. D - cpe,n;HIIe aKpoue11TpIIqecKIIe c BTopwrnoii rrepeTJDI<Koii (13, 14, 15). 5. E - 6oJiee MeJIKHe cy6MeTaueHTpw.recKIIe (16, 17, 18). 6. F - Memrne MeTauettTpw1ecKIIe (19, 20). 7. G - MeJIKIIe aKpouettTpttqecKtte c BTOpttqttoil rrepenrnrnoil (21, 22). IlOJIOBaH X-xpoMOCOMa OTHOCHTCJI K rpynrre CKaJI; Y-xpoMocoMa - c, cpe,n;HJIJI MeTaueHTpll'!e- MeJIKIIH aKpouettTpIIK. reHen1Ka COM3TM"l0CKMX KneTOK 8Ta ,ll;HCD;IIIlJIHHa II3yqaeT HaCJie,n;CTBeHHOCTb H H3MeHqHBOCTb COMaTHqeCKHX KJieTOK, HCilOJib3Y.H KyJihTypy KJieTOK pa3JIIIqHbIX TKaHeii H opraHOB. Cyr.u;ecTBYIOT cJie,n;y10ru;IIe MeTo,n;hI reHeTHKH coMaTttqecKIIX KJJeTOK qeJioBeKa: 1. Ilpocmoe KYJtbmuBupoeattue. 2. I'u6pu{}u3au,uH - CJIIIJIHIIe KJieTOK pa3HbIX TIIITOB (HanpIIMep, KJleTKH «qeJIOBeK-Mbllllb» IlOCTeneHHO TepHIOT xpOMOCOMbI; MOiKHO ycTaHaBJIIIBaTb rpyIIIIbI cuerrJieHH.H no O,ll;HOBpeMeHHO ucqe3aIOru;HM rrpH3HaKaM). 3. KJtOHUpoeattue PH,ll;OM. I'u6puaoMa - IIOJiyl.J:eHIIe IIOTOMKOB O,ll;HOH KJleTKH, narrpHMep rtt6:3TO KJieToqHbIH rtt6ptt.a;, IIOJiyqeHHbIH rryTeM CJIH.HHIIJI HOpMaJibHOro JIHMc}>OUHTa H orryxoJJeBOH KJieTKII; rrpOH3BO,ll;HT MOHOKJIOHaJibHbie aHTIITeJia O,n;Horo TJIIIa. 4. CeJteKu,uH - oT6op KJieTOK c 3apaHee 3a,n;aHHbIMH cBoiicTBaMH.
MeiJuijunc1wK zenemuica. E1iu3neqo6btU ;nemoiJ 111 MeTOA AepMaTormu:f>1r1K1r1 8ToT MeTo.z:i; rrpe,n;cTaB;uieT co6o:i1 i-13yqeHtte narrttJI.·urpHhrx y3opoB rraJihr~eB, Jia,L(OHett, CTOil • .IJ:epMaTOrJittqmqecKtte Y30pbl llH,!1;11Bl1,!J;YaJibHhl H ue MemnOTCH B Tel!eHtte Bce:i1 )Kll3HH • .IJ:epMaTOrJittqml!eCKHH amL1H3 ll03B0.1.HCT orrpe,n;eJIHTb 3MI'OTHOCTb 6J1H3HeUOB (MOHO- HJIH 1(11-), HeKOTOphIX rCHOMHhIX l1 xpOMOCOMHhlX MYTaUHH (60Jre3HH .IJ:ayHa, IlaTay ll .z:i;p.), npoBeCTH tt,z:i;eIITHq:mKar~HIO Jllll!HOCTH B KPHMHHaJIHCTHKe. 6nlr13He~OBblH M0TOA MeTo.z:i; rro3Bo.1JieT oueHHTh po.Jih Hac.'le,z:i;cTBeHHOCTM H cpe.z:i;hr B pa3BMTHH rrptt3HaKa. BJJH3HeUhI 6hrnaroT MOH03HrOTHhIMH H .n;tt3ttrOTHhIMH. Motto3ttrOTHhie 6JJtt3HeI~hI pa3BttBalOTCJI H3 o.z:i;Hott orrJio,n;oTBopeHtto:i1 RiiueKJieTKH (3ttrOTbl) B pe3yJibTaTe ee pa3.z:i;eJleHHJI Ha,!l;BOe (Ha CTa,!l;Hl1 .n;po6JJeHttB:) c o6pa30Bam1eM ,n;Byx sM6pnottoB. MoH03ttroTHhie 6.'IH3Heuhr HMeroT o.z:i;ttHaKoBhie reHOTHilhI, Bcer.z:i;a 0,ll;HOro IlOJJa; HMelOT BhICOKYIO CTeneHh CXOJJ:CTBa (KOHKOp,z:i;aHTHOCTb) no MHOrHM rrptt3HaKaM. Pa3JJttqne ttx rrpH3HaKoB 3aBttCHT TOJJhKO OT <})aKTOPOB BH€IIJHeil cpe,n:hI • .IJ:n3ttroTHhre 6JJH3Heuhr pom.n;aroTCJI H3 ,z:i;Byx HJ'IH 60.JJee o,n:HoBpeMeHHO oByJittpoBaBlllttX H OIIJJO,!J;OTBOpeHHhIX pa3HhIMH crrepMaT030HJJ:aMH JIMU€KJJ€TOK. 0HH HM€IOT pa3Hhie reHOTHTihI, MoryT 6bITh 0,!l;HOro H.'IH pa3HOro IIOJJa. 0HH cxo,n:HhI Mem.z:i;y co6oH: He 60.JihWe, qeM 6paThJI H cecTphr, pom,z:i;eHHhie nopo3Hh. XapaKTeptt3yIDTC.H .z:i;ncKop.z:i;attTHOCThlO - Hecxo,n:cTBOM rro :viHorttM rrpH3Ha- KaM. BJJaro.z:i;apB: O,!l;HOBpeMeHHOMy pom,n;eHHIO H BOCTIHTaHHIO HM€IOT o6n~He cpe,z:i;OBhie <})aKTOPhI. Pa3JIHqne npH3HaKOB B OCHOBHOM CBJI3aHO c reHOTHIIaMH • .I1:JJJI Jl:OKa3aTeJJbCTBa po.JIM Hac.1e,z:i;CTBCHHOCTH cpaBHHBalOT ,!J;O.'IIO ('%) KOH- KOp,n:aHTHhIX nap (OJJ:HHaKOBhIX no KOHKpeTHOMY npH3HaKy) cpe,n:H MOHO- H ,!l;H3HrOTHhIX 6JIH3HeuoB • .r:i:aHHhre 0 KOHKOp,n;aHTHOCTH 6JIH3H€UOB no H€KOTOpbIM rrpH3HaKaM np1rne,!1;€Hbl B Ta6Jittue. Ilptt3HaKH I I KoHKOp,11aHTHOCTh (cxo,11cTBO), MoH03IffOTHhie .IJ:tt3IffOTHhie 100 46 99,5 97 28 23 KocoJiarrocTh 32 3 3rr11Jierrcm1 67 3 llil130l}lpeHIUI 70 3 Ty6epKyJie3 fpyrrrra KpOBll ll;BeT rJia3 IJ;BeT BOJIOC I 'Xi 37 15 ICopb 98 94 CaxapHhlii: ,1111a6eT 65 18
06UfaR 6uo1wzuR. I'enemu1w 112 BJJaro,n;apH 6JJtt3HenoBoMy MCTo,n;y 6b1.rrn BhIHCHeua nac.'le,n;cTsemrnH 11pe1~­ pacrroJ10:t1<enHocTh qeJJOBeKa K p.H,n;y 3a6oJJeBaHm'i: lllH30(l)peHIIH, 3IlllJICIICHII, caxapHOMY ,LJ;Ha6eTy H ,n;p. 6MOXMMM'teCKllle MeTOAbl 3TH MeTO,ll,hI OCHOBaHhI Ha u3yqeHHII o6M~Ha Bern,eCTB (qlepMeHTOB, aMHHOKHCJIOT II T. ,n;.) MCTO,n;aMH 6HOXHMHH. 0HH l103BO.'l5HOT o6HapyiRHTb HapyllleHHH B o6Mene BCll~CCTB, BhI3BaHHhre H3MCHeHlUIMH renoB, II, KaK CJ!C.D;CTBHC, aKTHBHOCTH pa3JIIIqHhIX qlepMeHTOB. KpoMe Toro, no3BOJJJIIOT ycTaHOBHTh reTepo311roTHoe HOCIITe.'lhCTBO HCKOTOpbIX 3a6oJJeBaHHH. HacJJe.n;cTBCHHhre 6oJJe3HII o6Mena Bern,ecTB rro.n;pa3,n;eJJJIIOTCJI Ha 6o.'le3HH yrJJeBO,ll;HOro o6MeHa (caxapHhlll .n,11a6eT), o6Mena aMHHOKHC.'IOT, JIHTIH,ll;OB II T. ,n;. IlpuMep. ct>ennJJKeTOHypnJI -- 60J'Ie3Hh aMnHOKHCJIOTHoro o6MeHa. BJJOKH- pyeTcJI npeBparn,eHIIe HC3aMCHHMOH aMIIHOKHC.'IOThI tjleHIIJJa.'laHnHa B Tnpo3IIH, npII 3TOM tjleHHJJaJJaHHH OKHCJIJICTCJI B tjlCHIIJITIHpOBHHOrpa,n;HyIO KHCJJOTy, KOTOpaJI BhIBO,ZJ;HTCJI c Moqoii:. 3a6oJJeBaHJie npHBO,ll;HT K pa3BHTHIO CJJa6oYMH51 y ,n;eTeii. PaHHee BhrnBJieHIIe n ncKJIIO'-IeHne tjleHIIJiaJiaHnna II3 nnTaHnH II03BOJIJIIOT npMOCTaHOBHTh pa3BHTHC 3a60.1eBaHH5l. XHMH'lec1me npo11ecch1 B opraHH3Me npu cI>eHHJIKeTOHypuu tjlepMCHT tjleHtt.1aJJaHIIHrn.n;poKCHJia3a (,n;etjleKTHhrii) I )( •I tjleHnJiaJiaHIIH THp03IIH I X •I MeJiaHnH O=C-CH-CH- 1 I OH NH 2 2 -OH tjleHnJiyKcycnaJI KHCJIOTa ¢ennJIMOJIO'-IHaH K-Ta ¢ettnmmpoB1rnorpa,n;Haa K-Ta nonymll.,\lllOHHO·CT3TlllCTlll'teCKlllM MeTOA 3To MCTO,ll, lI3Y'-ICHHH pacnpocTpaHeHHJI nacJie,D;CTBCHHblX npn3HaKOB (naCJIC,D;CTBCHHblX aa6oJieBaHnii) B nonyJJHl1lIHX. Cyrn,eCTBeHHbIM MOMCHTOM npn HCTIOJib30BaHHH 3Toro MeTo,n;a HBJIJICTCH CTaTHCTlI'-ICCKaa o6pa60TKa ,Zl;8HHbIX.
M eiJu~uncKax zenemuKa. 1Wemodbl npenama1tbnoii iJuaznocmu1m 113 IlollyJUIUUOHHO-CTaTIICTtttieCKlIH MeTO,ll; IT03BOJIJ!eT pacctrnTaTh B nonyJrn:UlIIf ll3CTOTY HOpMaJihHhIX lI ITaTOJIOrIItieCKlIX reHOB lI reHOTlIITOB: reTepo3HroT, roM031IrOT ,ll;OMIIHaHTHhIX lI peuecCHBHhIX. "lJacToTa rettOTIInos II cJlettoTttnoB pacctIHThrnaeTcJI ITO <jJopMyJte XapauBauH6epia: p2 + 2pq + q2 = (p + q)2 = 1, r,n;e p -- qacTOTa .n;oMHHaHTHoro retta A, q - qacTOTa peueccttBHoro reHa a, q2 -- t.IaCTOTa rOM0311l'OT ITO pe1~eCClIBHOMY reHy aa. Bamtto, tITo no 3TOtt cpopMyJie Momno pacctIIITaTh qacTOTY reTepo311roT no naTOJiornqecKIIM peuecCIIBHhlM reHaM, KOTOphlC HaX0,11;.HTC.H B CKphlTOM COCTO.HHIIH. JipuMep. "llaCTOTa cpeHIIJIKeTOHypIIII (aa) B norryJI.Hl[llll 1: 10000, 3HatIIIT, q 2Ha = 0,0001; q = 0,01 p+q = 1, Tor,n;a p = 1 - q = 1 - 0,01 = 0,99 2pqAo = 2 x 0,99 x 0,01 = 0,0198 - 0,02 m:rn 2 'Xi. 3HatIIIT, qacTOTa reTep031IrOT ITO reny cpeHIIJIKeTOHypIIH B II3ytiaeMOM IIOITyJI.HUlilI COCTaBJI.HeT 2 '% . MeTOAblnpeHaTanbHOM (BO epeMSI 6epeMeHHOCTlll) AlllarHOCTlllKlll AMHuou,eHme3 - OT6op (rrpo6a) aMHHOTMtiecKott mu,n;KOCTH c ueJihIO UHTOreHeTHtiecKoro HJilI 6IIOXHMHlleCKOro HCCJie,n;oBaHH.H ,ll;JI.H ,n;11arHOCTMKH HaCJie,n;cTBeHHhIX 3a6oJieBaHHM. Il03BOJI.HeT BhUICHHTh reHHhie 6oJie3HH o6MeHa Bell[eCTB lI xpOMOCOMHhle 6oJie3HH. 0T6op aMHIIOTMtiecKo:H mtt,n;KocTu rrpoBO,n;HT rro.n; KOHTpoJieM yJihTpa3ByKoBoro HCCJie,n;oBaHH.H Ha 14-16-:H He,n;eJie 6epeMeHHOCTH. OnpeaeJteHue 6eJ11w aJtb<jJa<jJemonpomeuHa ( A<PII) B ChrnopoTi~e KPOBH 6epeMeHtto:H HJIH B aMHHOTttt.IecKott mu,n;KOCTH. A<l>II - rJinKorrpoTen.n;, KOTOPhrtt npo,n;yuupyeTc.H rretieHhIO nJio,n;a. HccJie,n;oBaHtte npoBo,n;HT Ha 14-16-:H tte,n;eJie 6epeMeHHOCTH. YpoBeHh 6eJIKa A<l>II nOBhIIIJaeTC.H npn p.H,n;e attoMaJIHtt pa3BHTHH ITJio,n;a, TaKwx KaK aH3ttuecpaJIHH (oTCYTCTBMe M03ra), BHyTpHyTpo6Ha.H rn6eJih rrJio,n;a. CHIDKettwe A<l>II rrpowcxo,n;HT rrptt xpoMOCOMHOtt naToJiornw (cIIH,n;poM .[(ayHa, 3.n;Bap,n;ca II ,n;p.). YJtbmpa3ByKoBoe uccJteaoBaHue nJtoaa ttan6oJiee pacrrpocTpatteHHhltt MeTo.n;. Ott OCHOBaH Ha crroco6HOCTH YJihTpa3BYKOBOM BOJIHhl OTpamaThC.H OT UOBepxHOCTH pa3,n;eJia .D;BYX cpe.n;, OTJIHllaIOil[HXC.H JllllIHOM IIJIOTHOCThlO, tITO noaBoJI.HeT noJiytIMTh H3o6pameHwe Ha 3Kpatte. HccJie,n;oBaHHe rrpoBo,n;HT Ha PaaJIMqHhrx cpoKax 6epeMeHHOCTH. MeTo,n; II03BOJI.HeT .n;warHOCTnposaTh rpy6hre rropoKH M03ra, rropoKH KotteqllOCTeii, rroqeK H .n;p.
06UfaR 6uo;wiuR. I'enemu1w 114 1113MEH'-llllBOCTb. Tlllnbl 1113MEH'-llllBOCTlll. PEnAPAU.lllSI AHK H3Meli'IU6ocmb - 8TO CBOHCTBO oprairn.3MOB ,n;aBaTh OTKJIOHeH11H B npH3Ha- Kax rro.n; .n;eil:CTBl1eM BHelliHl1X 11 BHyTpeHHl1X cpaKTOpOB cpe,n;hI. He1Iae.1eµ;cTsettua.H (MOAlll<j:>lllKal\lllOHHasi 111n111 <j:>eHomnw-iecKasi) Hac.'le;:i;cTBeHHa.H (reHomnw-iecKasi) J{o.1116unamu6HaJ1 I'e111taJ1 nepecmpouJCu I) oeJ1e4u.11; I) ianllOUOU.ll; 2) oyn.w1Ca4u11; 2) no;wn.wuou.11 3) u 1rnepc11J1; 4) mpa11c.w1Ca4u11 ll3MeH•IHBOCTb opraHl13MOB MO:>KHO pa3,n;eJil1Tb Ha .n;Ba B11.n;a: HeHaCJie,n;cTBeHHaH (Mo,n;11cp11Kau;110HHaH 11Jil1 cpeHOTl1IIH'IeCKaH) 11 HaCJie,n;CTBeHHaH (reHOTl1II11qecKaH). Moau<PU1cau,uoHHaJl U3Melil/U6ocmb - :JTO H3MeHeH11H oco611, B03H11Kru11e no.n; B03,n;eil:CTBl1eM cpe,n;hI, He npl1BO.D;HIIJ;l1e K 113MeHeHl1HM B reHOTl1IIe 11 He nepe.n;a10ru;11ecH no HacJ1e,n;cTBY. XapaKTep11cT11K11 Mo,n;11cp11Kau;110HHOil: H3MeHq11BocT11: 1) 2) 3) HeHaCJie,n;yeMOCTb 113MeHeHHOro npl13HaKa; MaCCOBOCTb (npOHBJIHeTCH y MHOr11x oco6eil: B IIOIIYJIHIJ;1111); a,n;eKBaTHOCTb (Bcer.n;a COOTBeTCTByeT 113MeHeHl1HM BHeruHeil: cpe,n;bl: qeM 60Jihllie B03,D;eHCTBl1e, TeM 60Jibllie OTBeTHaH peaKIJ;11H); 4) a,n;aIITl1BHOCTb (nOBhIIllaeT np11cnoco6JieHHOCTb opraH113Ma); 5) .n;11arra30H 113MeHeHl1H .D;JIH oco6eii: ,n;aHHOrO B11.n;a o6yCJIOBJieH reHeT11qe- CKl1 11 xapaKTep113yeTCH HOpMOM peaKIJ;1111. HopMa pea1Cu,uu - 8TO reHeT11qecK11 o6ycJIOBJieHHhIH rrpe,n;eJI pa3BHTHH IIpH3HaKa. HopMa peaKIJ;HH MomeT 6hITh pocT, 11HTemreKT) HJIH rpyrrrra KpOBH). y31Cou wupo1Cou (KOJIHqecTBeHHhie rrpH3HaKH - (KaqecTBeHHhie npH3HaKH - Bee, u;BeT BOJioc, rJia3;
y 3 Men'lu6ocmb. II pwnep1>1 Tun bi u.1.11ien'lu6ocmu. Penapau,,uR If II K 115 Mo.umlrn1rn11;i10H11oii n3MewnrnocnI: 113MeHeH11e III1I'MeHTaU,J1l:I KO./Kl:I IIOCJie 3arapa; IlOBbIWeHI:IC ypOBIUI reMOI'JI06mrn B KpOBl:I B 1\Ba pa3a IIpl1 l(.111'reJJbHOM HaXO./K,'(CHHM B ropax Ha BhICOTe 4000 M. Mow1qn11rn11,Ho1111a.H 113MCH'II:IBOCTh MHorI:Ix npH3HaKOB y pa3Hhrx opra1111 3 MOB IIO,[('IHII.HeTC.H o6II~HM 33KOIIOMepHOCT.HM. ,[(aHHb!M Bl:Ij( II3MeH'll1BOCTI1 l13yqaeTCH B 6oJihIII11X rpyrrrrax opraHI13MOB. CTerreHh rrpOHBJieHHH IIp113HaKa pa3JII1LIHa y 'lJlCHOB 3TOH rpynrrhr. BoJihWHHCTBO IIMeeT cpel(nee 3Ha'Ien11e (poC'r, Bee 11 T. lJ:.). M11HnMaJihHOe II MaKc11MaJibHOe rrpoHBJieHHH npH3HaKa na6J110,z:i;a- IOTCH TO.lbKO y OT1\eJ1bHhIX oco6eiL 311a11eH11e 3HKOHOMepHOCTeii: MO,ll;HQJHKall,HOHHOH II3MeH'Il:IBOCTl1 II03B0Jl.HCT npe,D;cKa3hIB3Tb CTerrenb BbipameHHOCTl:I IIPII3HaKa B 3aBI1ClIMOCTH OT yCJIOBHH cpe,D;hl II HMeeT 60JibWOe 3Ha'ICHIIe B ceJICKll,IIII (coJie-, 3acyxo-, MOp030YCTOH'Il1BOCTb). II HaCJ1eiJcm6eHHaJ1 UJMe1l'tueocm1> rro,D;pa3l(eJ1.HeTCH Ha 1wM6ullamu6Hy10 Mymau,uo11Hy10. IlpH'II1H3Ml1 KOM6HH8TIIBHOH II3MeH'IIIBOCTl1 .HBJI.HIOTCH: KpOCCIIHl'OBep; He- 3aBIICIIMOe pacxom;~eHIIe xpoMOCOM B MeH03e; He3aBIICIIMOe CO'IeTanne raMeT rrpII OIIJIO.[(OTBOpeHIIII. IIpuMepo.'lt KoM6nHaTHBno:ii: H3MeH'IHBOCTn HBJIHeTCH TpeTIIii 3aKoH Mett;i:eJIH (,D;eTII O,ll;HIIX pO,ll;IITeJJe:iI OTJIH'l3IOTC.H ,lJ;pyr OT ,ll;pyra HOT pO,ll;HTCJie:ii:). Mymau,uo1tHaJ1 U3MPH'lU6ocm1> B03H111rneT B pe3yJibTaTe pa3JIII'lHhrx H3MeMymau,uoH1ta.R meopu.R 6bIJia cqiop:viy;rn- HeHIIH HacJJel(cTseHHoro MaTepIIaJia. poBaHa B 1901 r. I'OJIJiaHl(CKHM 6oTaHIIKOM ryro l(e Cl>pII30M. Mymau,uu - aTo pe3Kne cKa11Koo6pa3Hhie H3MeHeHn.H reHOT11na. MyTall,IIH H3MeHHIOT HaCJie,D;CTBeHHhIH MaTepnaJI H MoryT rrpHBO,ll;HTb K pa3.1H'lHbIM rrpo.HBJieHH.HM Ha ypOBHe cpeHOTIUia. CBottcTBa MyTaii:11rr: - B03HHKaIOT BHC38IIHO; - He HarrpaBJieHbI - MOJKeT MYTHPOE3Th JII06aH qacTb reHOTHrra; - 60JihllIHHCTBO MyTau,n:H - peu,eccttBHhre, pe,ll;KO - ,ll;OMHHaHTHhIC; - ,ll;JIH opraHH3Ma 60JihllIHHCTBO MyTall,HH Bpel(Hhie, pe,ll;KO - HeHTpaJihllbie IIJIH IIOJJe3Hbie; - MYTa11,11n nacJre,ll;yIOTc.H; - BhI3bIBHIOT CTOHKne H3MeHeHII.ff HacJie,ll;CTBeHHOrO MaTepIIaJia; - BhI3bIB8IOT Ka'!eCTBeHHbie H3MeHeHH.H rrpn3HaKa, He o6pa3yIOill,He HerrpePhIEHoro p.H,ll;a BOKpyr cpel(HeiI BeJIII'IIIHhr; - MoryT B03HHK8Tb B reHOTHIIe IIOBTOpHO. MYTa11,II11 MOryT B03HHK3Th ITO,ll; ,ll;e:iicTBHeM BHellIHHX n BHyTpeHHHX cpaKTopoB. MymaieH1>1 - 3To cpaKTOpbr BHellltte:H cpe,D;hI, BhI3bIBaIOill,He MyTaU,HII. MyTareHhr rro,D;paa,ll;eJIHIOTCH B 33BHCHMOCTH OT rrpttpO,ll;hI B03l(e:ttcTBI1H tta: XBMnqeCKne (a30THCTa.ff KHCJIOTa, IIIIPIIT, HHTp030MeTIIJIMO'IeBIIH8 II ,lJ;p.), cpnali'lecKne (pa,ll;Hall,11.H, BbICOKa.H TeMrrepaTypa), 6nOJIOI'II'IeCKI1e (a.ueHOBHPYCbI, PeTpoBnpychI).
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#JM€ttl/,U6ocmb. Tunbt U3M€ttl/,U6ocmu. Penapa~uR )~llK 117 Co crJBuzoM pa.MKU c'lumbtBaHUJl (KO.JIIFieCTBO HYKJ'IeOTH.ZWB B reHe MemieTc.fl): ~e.'leUH.fl - Bhma~eHHe; ~yrwmrnun.H - Y~BOCJIHe; Hl!Ccpu;HJI - BCTaBKa. Ilpu.Mepbt: 1. 3aMeHa no THrry TpaHcBepcnH CepnoBH~HOKJieTo<rna.H aHeMH.H 6-ll aMHHOKHCJIOTa HbA - r;1yTaMHHOBaH HbS - HaJIHH --- l.J;AT } KHCJIOTa - IU'T f3-u;euL ,UHK 2. 3aMeHbI mo6oro Tnna MoryT npHBO~HTb K ITO.HBJieHHIO CTOil-KO~OHOB. B pe3YJihTaTe TpaHCJI.Hu;n.H 6eJrna o6pbrnaeTcH. 06pa3yeTCH H3MeHeHHhIH (yKopoqeHHhitt) IlOJ'IHITeITTH~. r~-TaJiacceMHH - HbA HMeeT HOpMaJihHYIO a-cy6'he~i-rnnu;y' HO yKopoqeHHYIO 13-cy6'he~HHHI.I;Y. Xpo.Moco.MHbte Mymau,uu - 3TO H3MeHeHHH CTPYKTYPhI xpoMocoM. 1. ,[(eJieu;nn - Bhma~aeT qacTb xpoMOCOMhI. Ilpwnep. CnH~poM «KOIIIaqhero KpnKa» - ~eJieun.H KopoTKoro rrJieqa Il.HTOH XpOMOCOMbI. 2. ,[(yrrJinKauIIn - y~soeune yqacTKa xpoMocoMhI. IIpuMep. ,[(yrrJinKauna reHa Bar B reHoTnne Myxn ~po3ocpnJihI rrpnBo~HT K yMeHhIIIeHHIO qncJia cpaceTOK r.,1a3. 3. llHBepcnn rrapau;eHTpnqecKIIe ----------- (He 3aTparnBaJOT u;eHTPOMepHbIH paii:oH) -----rrapaueHTpnqecKne (3aTparirnaJOT pattoH r~eHTpoMepbI) c::::JIM::l._ . / c:: ____,. ____ :::> llpu.111ep. Ilpe~ITOJiaraeTC.H, qTo BTOpaH xpoMOCOMa y qeJIOBeKa - 3TO pe3YJibTaT o6'he~nHeHHH, u;eHTpnqecKoro CJIHHHH.H 12-ii: H 13-ii: y rrpe~Ka. 5. TpaHcrro3nu;11n -- rrepeMeru;eHHH Mo6nJihHhIX reHeTnqecKnx 3JJeMeHTOB. llpuMep. TpaHcrro30HhI 6aKTepnn, As n Ds 3JieMeHThI y KYKYPY3hI, M,[(r (Mo6nJibHhie ~ncnepcnpoBaHHhie retthr) ~po3oqrnJihI. I'eno.Mnbte Mymau,uu - 3TO n3MeHeHne qncJia xpoMocoM (B rarrJIOH~HOM Ha6ope - reHoMe).
06UfaH 6uonozun. J'enemu1rn 118 I'eltO.M - ITOJIOBl:IHHhlH Ha6op ,UHK l:I nOJIOBl:IHHhIH Ha6op reHeTI:I'IeCKoro MaTepI:IaJ:ra. 1. 3yrr:IOHl\HH -- yBeJIH'IeHne Ha6opoB xpOMOCOM, Kpanwe n. IIpu.Mep. 3n -- TpI:Irr;10n.n:h1 4n - TeTparnrnM,n:hI II .up. 2. Attayrr.110M,[(MH - He Kpanroe n yBeJIM'ICHI:Ie MJIM yMeHhllleHI:Ie 'IMCJia xpoMOCOM. -2n + 1 - Tpl:ICOMI:IH. IIpu.Mep. TpMCOMHH no 21-n xpoMocoMe - cnH.n:poM ,UayHa (47; + 21-a). XXY - Tpl:ICOMHH y MYiK'IMH no X-xpoMocoMe - CMH,n;poM KJiaiimpeJihTepa. -2n - 1 - MOHOCOMUH. Ilpu.Mep. C1rn.n:poM lllepellleBcKoro-Tepttepa: 45, XO. JKettru;MHa c rroJIOBhrMM xpoMocoMaMM XO. IlpH'IMHa amiynJIOl:I,ll;l:Il:I - Hepacxom,n:eHI:Ie IlOJIOBhIX xpoMOCOM BO BpeMH Menoaa. 1. Hepacxom.n,em1e noJIOBhIX xpoMocoM BO BpeMH Melioaa y MaTepM: P: ~ 46, XX x 46, XY cf' ~- -" r.-~ ,;;;~ G: 24,(_~_:X,,/ 24,1,Q; 23,Q\.) 23,\¥/ F: 47, XXX 47, XXY 45, XO 23, YO TpMCOMMH no Cntt,n:poM CMH,n;poM fM6eJih anrOThI X-xpoMocoMe KJiaiimpeJihTepa llleperneBcKoro-TepHepa 2. Hepacxom.n:em1e noJIOBhIX xpoMocoM so BpeMH Melioaa y 0T1.1,a: P: Y46, XX x 46, XY cf' 24 ,,:::- -"' 2 r.o-;'\ G 2 r;~ : .<~-~ 3,\_)0 F: 47, XXY CI:IH,n:poM KJiai'rn¢leJihTepa 2,\___) 45, XO CnH.n:poM lllepellleBcKoro-TepHepa 3aJWH roMOJIOrH11ec1rnx p.H)J;OB HaCJie)J;CTBeHHOH H3MeH11HBOCTH H. II. BaBHJIOBa I'e1temu'lecKu 6Jtu3Kue euobt u poabt xapaKmepu3y10mc.R «cxoo1tbt.Mu p.Roa.Mu ltllCJleocmee1t1tOU U3.MClt'lU60cmu». 31ta.R U3MCH'lU60Cmb OOltO?O euoa, MOJICHO npeocKa3amb noReJie1-we napaJIJICJlbltbtX <jJopM y (}pyioio pooa uJiu euaa. H. M. BaBMJIOB npocJie,n;MJI M3MeH'IMBOCTh MHomecTBa npI:IaHaKoB y aJiaKOB: pmn, rrrneHHI!hI, oBca, H'IMeHH, KYKYPY3hl. 3aKoH H. M. BaBMJIOBa MMeeT 6oJihrnoe aHa'IeHI:Ie ,n;JIH cem~Kl.l,MH. OH noaBoJIHeT aapaHee npe,n;BI:I,UeTh, KaKI:Ie npH3HaKI:I H3MeHHTCH y Toro HJIH l:IHOro BH.n:a B peayJihTaTe B03,ll;eHCTBHH Ha Hero MyTareHHhIX ¢>aKTOPOB. B HacToaru;ee BpeMH aaKoH roMoJiorM'IecKMX pa,n;oB no,n;TBepm,n:eH He TOJihKO Ha 3Jl8Kax, HO TaKme Ha npnMepe rpM60B, Ml:IKpoopraHH3MOB, iKl:IBOTHhIX. IlpH'IHHhI CXO,ll;CTBa MyTal.l,l:IH 3aKJIIO'IalOTCH B CXO,UCTBe xapI:IOTl:ITIOB y 6JII:I3Kl:IX BH,ll;OB l:I B CXO,ll;CTBe paCTIOJIO;KeHI:IH aJIJieJihHhIX reHOB B xpOMOCOMax.
penapau,un ;n,011,eKyllbi!{HK 119 Penapau;HH MOJieI~YJihI ,r:t;HK B ICICTKe cyw;ecTByeT CHCTC'Ma 3RIIl;llThl HRCJie.n;cTBeHHOro MRTep11a.'Ia ll,HK OT B03HlllO!OBeJHIH MyTau;tt:IT ll IIOBpem.n;eHHM - ClICTeMa penapaILMll. Penapa41ut - 3TO BoccTaHoB.'IeHHe noBpem.n;eHHOM CTPYKTyphr ,UHK (c;romHhIM qiep:vieHTaTHBHbIII npouecc ). BhIJ~eJIHIOT ttecKO.'IhKO pa3HOBM/::(HOCTe:ii: penapaIIMH: I. cf>OTopenapaumi. II. TeMHOBaH penapmi;HH, tt;rn 3KCIIH3HOHHaa (Bh1pe3arow;aH) peuapaIIMH, BIUilO'IalOIIl;aa: .n:opCIIJ1HK3THBHyro penapauttro; IIOCTpeIIJIMKaTHBHYIO penapaumo; penJittKaTHBHYJO penapauttro. III. SOS-penapari;HH. I. «l>0Topenapau;m1 (pHc. 50). IIo.n .n:efrcTBMeM yJLbmparjJuoJZemoeblX JLy'leil ceema B IIen11 ,II.HK Mem.n:y ,nByMH ocHoBaHHHMH T-T o6pa3yroTCH xnMttqecKMe CBH3M (B03HMKaeT T-T ,nttMep), Hapyma10ni:11ii: cqHThIBaHMe MH<l:>opMaIIMll M penJIMK3I::(l1lO .UHK. Ilo.n: ,nettCTBHCM euiJU.MO?O ceema aKTMBMpyeTCH: <I:>epMeHT (<i:>OTOJ1lla3a), KOTOpblll pacw;enJ'IH:eT 3TM ,n:OIIOJIHMTeJibHbie CBH:3H. 8TOT IIPOIIecc H33hIBaeTCH: rpomopenapa4uR. II. TeMHOBaH, HJIH 3Kcu;uauottHaH (Bb1peaarorn;an), penapau;un. 8Ta penapari;tt.fl xapaKTeptt3yeTC.fl ocHOBHhIMH npoueccaMtt: cpepMeHT (:::m.n;ottyKJiea3a) «Y3HaeT» IIOBpem.n;eHHhIM yqacTOK HMTll ,UHK; cpepMeHT (3K30HYH:J'lea3a) «Bh!pe3aCT» IIOBpem.n;eHHhIM yqacTOK; cpepMeHT (,UHK-noJIHMepa3a) CHHTC3HpyeT .UHK no npHHI!HIIY KOMIIJieMeHTapHOCTH; <I:>epMeHT (.'1Hra3a) «ClllIIBaeT» KOHI!hl BHOBb CMHTe3MpOBRHHOrO yqacTKa C OCHOBHOH HHThlO ,[{HK. Ilo BpeMeHM ocyw;eCTBJieHHH TeMHOBa.fl penapau;H.fl MOJKeT npoHCXO.ll:MTh .n:o HJIH IIOCJie peIIJIHK3I!MM ,UHK M BO BpeM.fl penJIMKaIIMM. 1. nopenJtUICllmU61-taJl penapa4UJl - 3TO BOCCTaHOBJieHMe IIOBpem,n:eHHOH HHTM ,UHK .n:o ee y,n:BoeHM.fl B G 1 nep110.n:e KJieToqttoro IIMKJ'la. 2. llocmpenJLu1CamueHa.H penapa4u.H - BoccTaHOBJieHMe ,UHK nocJie penJIMKa!IMM B G 2 nep110.n;e. I I I I I I AfAAl..ll II Ill Ill II T Ll T=T f A I I I \ PMc. 50. BHAHMbU1 C88T t t t + <l> o 10pea n111 ea It l/ljl <l>eJ>MeHT I I I I I I I I AfAAl..ll II Ill II II Ill II ll..lllfA I I I <t>ornpenapa1..wm. TlilM"1HOBbl8 A"1M8Pbl pacw,enmHOTC51, "1 BOCCTaHaBn"1B3IOTC51 BOAOPOAHble CB513"1 A-T
06UfaR 6uo.rwzuR. I'enemuica 120 IlpoHCXO)lHT «Y3HaBaHHe» n curnBaH:ue KOHI~OB .JJ:HR. IrnpymeHtt.H, «Bhipe3aHHe» ,11,ecpeKTa B .JJ:HR l1nor,11,a 6e3 ,11,oCTpa:uBaH:u.H tte.n,ocTa10ruero yqacTKa CTPYKTypa reHa MOJKeT H3MemIThCR. 3. PenJtwcamuenaa penapau,u.R - BoccTanoBJ:re1me .JJ:HR B xo.n;e penJJHKaI.J;Htt. IlpOHCXO,ll,HT y,ll,aJJeHtte H3MeHeHHhlX HYKJJeOTH,ll,OB B XO,ll,e pocTa uerru, HO B HeKOTOph!X CJJyqaax perrJJHKautt.H MO:>KeT H,11,TH no H3MeHeHHOll MaTp:uue. CTpyKTypa re Ha MomeT MeH.HThC.H. CyruecTBYIOT MyTau11tt (y qeJJoBeKa - reHHhte 6oJJe3Htt), BhI3hIBa10ru:ue tta- pyruemrn rJJaBHhIX cpepMeHTOB perrapau:utt. BocCTaHOBJJeH:ue .JJ:HR B Tairnx CJJy'la.HX CTaHOBHTC.H HeB03MO'.IKHhIM. Ilpu.Mepbt 1. Il:urMeHTHa.H Kcepo,ll,epMa. Ilp:u rr:urMeHTHOH KCepo.n,epMe B KJJeTKaX OT- cyTCTByeT cpepMeHT (,ll,e3oKcmrnpHMH,ll,HHcpOTOJJHa3a), Heo6XO,ll,HMhlH ,ll,JJ.H perrapau:u:u ,IJ;HR, noBpem.n;eHHOH yJJhTpacp:uoJJeTOBhlMtt JJyqaM:u. Ilo)l )lettcTBlieM COJJHeqHoro CBeTa IlO.HBJJ.HIOTC.H pacruttpeH:ue Karr:uJJJJ.HpOB, oporoBeHUe 3IlH)lepMHCa, rropameH:ue rJJa3, pa3BHTHe paKOBhIX orryxoJJett KOmH, KOTOphre rrpnBO,ll,.HT K rrpem)leBpeMeHHOH cMepTH. 2. IlpOTHBOilOJJOJKHhIM np:uMepoM qpe3Bhl'l8HHO BhlCOKOH crroco6HOCTH Micrococcus radiodurans. 8TH 6aK- Tep:u:u MoryT BbI)KHBaTh B yCJJOBH.HX YJJhTpacpuoJJeTOBOH pa,ll,:uauHH, B 1000 pa3 KJieToK K perrapautttt .HBJJ.HeTc.H opraHU3M rrpeBbllliaIOruei1: CMepTeJJbHYIO ,ll,03y ,ll,JJ.H ,ll,pyr11x MHKpoopraHH3MOB. m. SOS-perrapaum1 - Bpem)leHHH CTpyKTYPhl 3TO 6bICTpa.H penapau:irn np:u 3Haq:UTeJJhHOM 110- .JJ:HK. Jl:JJ.H Hee xapaKTepHO HeToqHoe co6JJIO)leHHe KOMITJJeMeHTapHOCTH. rEHETM"4ECKME OCHOBbl CEnEKLl,MH >KMBOTHblX, PACTEHMM, MMKPOOPrAH"13MOB. 6"10TEXHononu' CeJte1w u.R 1 HayKa o coa,ll,aHHH HOBhIX :u yJJyqrueHUH cyruecTBYIOJUHX no- po,lJ, ;KHBOTHhIX, copTOB pacTeHHH, IlITaMMOB MHKpoopraHH3MOB. TeopeT:u'leCKOH OCHOBOH ceJJeKu:u:u RBJJ.HeTcf! reHeT:uKa. 3ada'lU ceJle/CU,UU - C03)laH:ue HOBhlX H yJJyqrueH:ue cyrueCTBYIOJUHX no- PO,ll,, copToB, IIITaMMOB pa3JJHqHhIX opraHH3MOB. Jl:JJfl peIIIeHH.H aa.n;aq ceJJeKUHH JJI06hIX opraH:U3MOB Heo6xo.n,HMO: 3HaHHe 3aKOHOMepHOCTeli HaCJJe)lOBaHH.H np:U3HaKOB; H3y'leH:ue HacJJe,ll,CTBeHHOH H3MeHqHBOCTH; H3y'leHHe BJJHf!HH.H cpe.n,bl Ha pa3BHTHe npH3H8KOB (MO,lJ,HcpHKan;:uomrn.fl H3MeH'lHBOCTh); nay'leHne copToBoro, BH.D;OBOro, po.n.oBoro paattoo6paa:ua KYJihTyp; paapa60TKa CTpaTerHH H MeTO,ll,OB HCKYCCTBeHHOro OT6opa. Ilopodbt, copma, mma.M.Mbt - 3TO ncKyccTBeHHO co3.n,aHHh1e qeJJoBeKoM no- nyJif!D;HH opraHH3MOB. Jl:JJR HHX xapaKTepeH Ha6op HHTepecyIOJUHX qeJJoBeKa npHaHaKoB, aaKpenJJeHHhIX ttacJie,n;cTBeHHO.
I'enemu<tecKue ocnoBht ce;ieKlfUU CeJie1n~m1 .[(.'I.fl 121 pacTeHHii perneHH.H 3a_naq CeJ1emJ,Hl1 paCTCHMM o'reHh BaJKHO COpTOBOe l1 BH,l(OBOe pa3H006pa3HC. H. H. BaBHJIOBhIM 6b1Jia co6paHa KOJIJieKIJ,M.H copToB KYJihTYPHhIX pacTe- HHM lI HX ,nHKHX rrpe,nKOB co Bcero 3eMHOI'O rnapa; 6bIJII1 OTKpbl'fbI CCMb IJ,eHTpOB ,npeBHCI'O 3eMJie,neJIH.H, HJIH IJ,eHTPOB MHOroo6pa3I1.H I1 IIPOI1CXOJK,l(eHttH KYJihTypHbIX pacTeHI1M (HMCHHO B 3THX IJ,eHTpax Heo6XO,l(HMO HCKaTb reHOTHIlbl ,nJI.H ceJieKIJ,HH HYJKHhIX rrptt3HaKoB). U:euTpbI npoucxoJK;:i:eu011 RyJILTYPHhIX pacTenuu ,------ --- U:eHTpbl pm-1cxoJK,aeH1u1 - 1. IOarnoA3ttaTCKtttt MecTonoJioJKeHtte --·-·1·--·----------·1 KyJ1hTYPHh1e pacTeHttH -+----- Tporrtt'!ecKaH HH,attH, Hu,aoKttTaii, o-sa IOro- ~~~:~~~· -1 ::~~:~·~ A3ttaTCKtttt BocTO'!Hhlii Kn:Tatt, .HrroHttH, KopeH, TaftsaHh 3. IOro-3arra,11HoA3ttaTCKHii MaJiaH A3HH, Cpe,aH.11.11 A3H.H, HpaH, Acprann:cT~ H, l0ro3arra,11Han f H,!lttH Puc, caxapHhltt TPOCTHHK, 1\ttTpycosh!e, 6aKJia:>KaHbl 11 AP· ( OKOJIO 33 °/.1 KYJihTYPHhIX pacTe11n:H:) CoH, 11poco, rpe'ln:xa, lIJIO,!IOBhle H OBO~Hble KYJihTypw - c.111sa, BHlllH5l 11 AP· (20 'X> KYJihTYPHhIX pacTeHttii) IlIIICHHl\a, pO:>Kb, 6o6osbie KYJihTYPhI, JieH,KOHOlIJIH,perra, '!eCHOK, BHHOrpa,11 H ,lip. (4 % KYJibTYPHhlX pacTeHtttt) !----------+--------------+~-------·--~ 4. CpC,!IH3CMHOMOpCKHH CTpaHhl no 6eperaM Cpe,an:JeMHoro Mopa KarrycTa,caxapHaH CBCKJia, MaCJIHHbl, KJICBep (11 % KYJihTYPHhlX paCTeHHH:) !-----·----+---------------- ---------- 5. 3c}lHOlICKHH 6. U•H,PM>H~· aMepnKaHCKHH 7. AH,11HttCKHii A6HCCHHCKOe narophe AcppHKH IIO*•"' Mm~., I 3arra/(Hoe rro6epemLe IOarnoii AMep1rnn: Tsep,11aH rrllleHttua, H'!MeHh, 6ananb1, RocpeH:Hoe ,n;epeso, co pro KYKYPY" KO<Oo 'w" ea, Ta6aR,'xJiorr'l;THH' H KapToc}leJIL, ananac XHHnoe ,11epeso ~-------~-----------~-------·- I
06~aJt 122 Ben Hoe cKpell.\HBaII11e) 6uo:wzuIL. J'euemu1rn rrepe1\pecTHOonhr.1aeMhIX) AyT6pur11rnr I'H6pHJJ;H3a:u;m1 H H6puiJuni (6JIH3Kopo,n;cTBeHHoe CKperu;ttBam1e) HCIIOJih3YIOT .ll;JJ.H no:ry1.1c- HH.H TaKHX JIUHHH, KOTOphie IIPH IIOCJie,n;yJOru;eM CKpen11rnaHIIH ,n;aIOT a<:\Jc}_)eKT reTepo3Hca. I'emep03UC ( «rtt6ptt.zi;HaH CMJia») - HBJieHMe, nptt KOTOpOM rtt6pH.ll;bl no PH- ,n;y npM3HaKOB M CBOHCTB rrpeBOCXO,ll;JIT po,n;ttTeJihCKMe C\JopMhI. lfaq1rna.H co BTOporo noKOJie1ma reTepo3HC 3aTyxaeT. Aym6puiJuHi - cKperu;ttnaHtte Hepo,n;cTBeHHhIX c}_)opM o,n;noro BIIJJ;a (HeT 06- ru;ttx npe,n;KOB B rrpe,n;eJiax qeThipex - OmiJaJteHHaJl w6puiJu3a4uH - IIIeCTM IIOKOJieHHH). CKperu;1rnamre oco6eiI (paC'reHHH), OTHOCH- IQMXCH K pa3HhIM mr,n;aM. OT,n;aJieHHhie rtt6pn,n;hr o6hPIHO cTepmihHhr, TaK KaK y HMX HapyrnaeTcH MeH03 (,n;Ba raIIJIOll,Zl;HhIX Ha6opa XpOMOCOM pa3HbIX Bll.ll;OB He MOryT IWH'hlOfIIpoBaTb), II raMeTbI He o6pa3yIOTC.fI. B 1924 r. r. ,I(. KaprreqeHKO C03,n;aJI KarrycTHO-pe,n;eqHbIM rII6pn,n;, IIOHBH- JiaCh rrepBaH :\WTO.ll;HKa rrpeo.zi;oneHHH 6ecrrJIO.ll;llH y OT.ll;aJ"IeHHbIX rH6pu,n;oB. YqeHhIH cKpecTHJI pe.n;hKY raMeTbI - n = 9). (2n = 18; n raMeTbI - = 9) n KarrycTy (2n = 18; Ho B rrpou.ecce Meii:oaa xpoMocoMbI He KOH'bIOrnpoBaJIH ~ rH- 6pn.zi;hr CTepMJibHbI. 3aTeM c IIOMOllJ;blO KOJIXIII1HHa (6JIOKHpyeT o6pa30BaHtte MMKPOTpy6oqeK uepeTeua ,n;eJieHHH) y,n;BOHJI xpOMOCOMHblH Ha6op rtt6ptt,n;a BOHJIHCb KaK «pe,n;eqHhie» (9 + 9), TaK n «KarrycTHhie» (4n = 36), IIpll oTOM (9 + 9) xpoMOCOMhI. y;:i;- KoH'hKJ- ral11'IH B03MOJKHa ~ raMeTbl o6pa3yIOTCH, ll ~ IlJIO.ll;OBIITOCTb BOCCTaHOBJ1eHa. CeJieKIJ;HH mHBOTHLIX B ceJieKI111H JKMBOTHbIX MCIIOJib3YlOTC.H Te nw OCHOBHbie MeTO.ll;bI - r116pH- .ll;II3al1II.fI ll OT6op. Oco6ennocmu ceJlelCU,UU JICUBOmllbtX: B OCHOBHOM IIOJIOBOe pa3MHOJKeHne; HeMHOfOqMCJieHHOe IIOTOMCTBO; pe,n;KaH CMeHa IlOKOJieHHH.
Ce.neKijUR. MuKpoopianu3M06. 6uomexno11,ozuJ1. 123 CoepeMeHHbte (iJonoJtuumeJ1bllbte} Jvtemoabt ceJteKu,uu: 11cKyccTBemwe oce- J.14eHeH11e, ropMOHa.'IhHaH cyneposy.'IHIJ,IIH If TpaHcrrnaHTaIJ,IIH 3N16pMOHOB, OT'AaneHHaH rII6pHJJ;II3all,HH. ,Il.oCTHmeHIIH ceneKIJ,HH: rII6pHJJ; il:opKIUMpCKOH 6e.'IOH CBHHbH CBHHhH (?) - (cf') H 6ecnopOJJ;HOH yKpaMHCKOH 6e.irnH CTenHaH CBHHhH; r116pMJJ; K06hinl1IJ;bI c OCJIOM - Myn: BbIHOCJIMB, JJ;OJII'O :>KIIBeT, HO 6ecn.'IOJJ;CH. HHorn;a raMeTorette3 rrpoTeKaeT HOpMaJihHO II nonyqaJOTCH IIJIO,lJ,OBIIThre r 11 6pHJJ;hI. llpuMep. ApxapoMepHHOChr (apxap 11 Mepattoc), 6ecTep (6enyra co cTep.w:- 'AhIO), XOHOP11K (xopeK 11 HOpKa). CeJie1m;m1 MHKpoopraHH3MOB (B OCHOBHOM oaKTepHH H rpHOOB) OcHOBHhrM11 MeTon;aMII ceneKIJ,11H MIIKpoopraHM3MOB HBJIHIOTCH 3Kcnep11l\'leHTaJihHhr:ii: MyTareHe3 II OT6op BhICOKOrrpon;yKT11BHhIX IIJTaMMOB. Oco6eHHOCTH ceneKIJ,1111 M11KpoopraHH3MOB: reHOM 6aKTep11:ii: rarrJI011,1J,HhIH (on;Ha KOJihll,eBaH MOJieKyJia ,UHK), II03TOMY JII06hre MyTaIJ,HH rrpOHBJIHIOTC.H yme B nepBOM IIOKOJieH11II; oqeHh BhICOKa.H HHTeHCHBHOCTh pa3MHO:>KeHHH o6JierqaeT IIOHCK MyTaHTOB. llpuMep. B peayJihTaTe HcKyccTBeHHoro MyTareHeaa 11 oT6opa 6hrJI noJiyqett mTaMM rp116a neHIIIJ,IIJIJia no aHT116IIOTHKY neHHIJ,MJIJIHHY. IIpon;yKTMBHOCTh 6hIJia yBeJI11qeHa B 1000 paa. BnoTexuoJiorHH. reuuan HHmeuepH» BuomexttoJtoZu.fl - ucnOJih30BaH11e :>KMBhIX oprattH3:vIOB u 11x 611onoruqe- CK11x npoIJ,eCCOB B npOM3BOJJ;CTBe Heo6XOJJ;HMhIX qeJIOBeKy Belll,eCTB If npon;yKTOB. 06'beKThI 6HOTeXHOJIOI'HH - 6aKTepuH, rpII6hr, KJieTKH paCTMTeJihHbIX H '1CHBOTHbIX TKaHeil. Hx Bhrpalll,HBaIOT Ha IIHTaTeJihHhIX cperi;ax B cneIJ,MaJihHhIX ycJIOBHHX B 6IIopeaKTopax. 0CHOBHhie MCTOJJ;hl 6HOTCXHOJIOI'MM: KJICTOqHa.H HH:>Kettep11.H (COMaTHqecKaH r116p11,1J,H3all,HH - CJIHHHHe co- MaT11qecKHX KJICTOK, ceJICKll,IIH KJICTOK); XpOMOCOMHaH HH:>KeHepHH; reHHaH HH:>KeHepMH. I'eHHa.R UHJICeHepu.fl - COBOKYIIHOCTh MeTOJJ;HK, II03BOJIHIOII~HX BhIL(eJIHTh HyJK- HLiii reu II3 reHoMa on;ttoro opraHH3Ma 11 BBeCTH ero B reuoM n;pyroro opraHH3Ma. TpattcieHHbte opiaHU3Mbt - :>KMBOTHhie II pacTeHHH co «BCTpoeHHhIM» re- HOM. Tpattc<jJopMupoeaHHbte opiaHU3Mbt - 6aKTep1111 11 rp116hr c BBen;eHHhIM qy- )kepo.n;HhIM reaoM. lipuMep. K11IIJeqttaH rraJioqKa E. coli, m1rnylll,a.H B KttIIJeqtt11Ke qeJioBeKa, ROCJie Tpaacq>0pMaIJ,1111 rettaMH HHCYJIHHa HJIH ropMoHa pocTa rrpIIo6peJia cno006aocTh c1111Te3IIpoBaTh ropMOHhI MHCYJIMH IIJIH coMaTOTporrIIH. HcrroJih3yeT- ea KaK npo.n;yIJ,eHT ropMOHOB.
06UfaJl 6uo;wzuJl. I'enemu1rn 124 UHTOnnA3MATH~ECKOEHACnEAOBAHME HeKoTopaH -.:racT1> aac.1e,11,cTBe1rnoro MaTepnaJrn KJieTKII aaxo,11,IITCH B UIITon.rra3Me. 8To: KOJibUeBaH .IJ:HK MlITOXOH,11,pII:ii:; KOJihIJ;eBaH .IJ:HK IIJiaCTH,11,; BHeH,11,epHbie reHeTnqecmrn sJieMeHThI (nJia3MH,ll,hr). HacJie,11,oBaHHe UIITon.rra3MaTH-.:recKIIX reHOB ae no,11,qHHHeTcH Mett,11,eJieBCKIIM 3aKOHOMepHOCTHM. HacJie,11,oBaHHe UHTOJUia3MaTnqecKHX renoB npoMcxo,11,MT no MaTepHHCKOM JilIHlilI, OHM nepe,11,aIOTCH OT MaTeplI KO BCeM ee IlOTOMKaM. IlplI OilJIO,ll,OTBOpeHHH reHeT11qecKHM COCTaB il;HTOnJia3Mbl B 3IIrOTe onpe,n;eJI.HeT- CH UlITOilJia3MOM .aii:ueKJieTKlI, il;lITOilJia3Ma cnepMaT030H,ll,a B onpe,n;eJieHIIH reHeTMqecKoro COCTaBa npaKTII'leCKlI He yqac·rByeT. llpuMep. HacJie,n;oBaHHe necTpOJIHCTHOCTII y MHorttx pacTeHHtt CB.H3aao c MyTaUH.HMH B .IJ:HK XJ'IOponJJaCTOB II aapyrneHHeM CHHTe3a XJIOpoqmmrn. y qe.JIOBeKa MHTOXOH,ll,pHaJibHbIM reHOM npe,n;cTaBJieH KO.JihUeBOH .IJ:HK, co- CTO.HI.UeM H3 16 569 HyKJieOTlI,ll,OB. qaCTOTa MyTaUlIM B MHTOXOH,11,pliaJibHOM .IJ:HK B ,11,eC.HTb pa3 BbIIIIe, qeM B .a,n;epHOH. MyTaUHlI B MHTOXOH,ll,pHaJihHOH .IJ:HK HBJI.HIOTCH np11q1rnoii: 6oJiee 100 6o.'Ie3ae:H • .IJ:tta6eT, snHJienc1rn, oTcpo- qeHaaa Kap,11,HOMHOilaTIIH, 6oJie3Hb IlapKlIHCOHa rrepe,n;aIOTCH OT MaTepII KO BCeM ee ,n;eTHM. CpaBHlrnaa MlITOXOH,ll,pHaJibHYIO .IJ:HK rrpe,n;KOB lI IIOTOMKOB, MOiKHO ,ll,OKa3aTh po,11,CTBO KaK Mem,11,y OT,ZJ;e.JibHbIMH oco6HMH, TaK lI Mem,11,y IIOIIYJI.H!IlIHMlI. TaKHM o6paaoM no HCKorraeMhIM ocTaHRaM 61>rJia n,11,eHT11qmu11poBaHa ceMh.H pyccKoro uapa HHKOJiaH II.
Bo3HUKIW6enue :JICU3nu na .1eM.rie. Pa.16umue opianu'leCKOU> Mupa 125 3BOJ1IOLl,l1SI Bo3HHKHOBeHHe JICH3HH Ha 3eMJie. Pa3BHTHe opraRH'lec1wro MHpa Bee rn:rroTe3bI o B03HMKHosemn1 iKM3IBI Ha 3el\e1e :vromHo pa3ri;eJIMTh Ha TPM rpynnhl: 1. 2. 3. TeopMR 6ttoreHe3a - .iKMBOe npottcxo,11,MT TOJihIW OT .iKMBOro. TeopMR a6ttoreHeaa Teopmi: naHcnepMMII - .iKMBOe noRBJIHeTc.H OT ttemttsoro. .iKM3Hh Ha 3eMJIIO aatteceHa M3 KOCMoca. B HaCTO.HIUee speM.H Hatt6oJiee npII3HaHHOH .HBJI.HeTCJI rttnoTeaa, BhICKaaattaa.H poccttiicKMM 6noxIIMIIKOM A. CKHM 6ttoJioroM II. OrrapIIHhIM B 1924-1928 rr. H attrJIIIrr- .n;m. XoJI,n;e:HttoM. CyTh rttnoTe3hI B TOM, qTo IIO.HBJieHIIIO .iKII3- HH aa 3eMJie o6.H3aTeJibHO npe,n;mecTBOBaJIO a6Horemrne o6paaosaHtte opraHtt'leCKHX coe,n;IIHeHHH H3 HeopraHHqecKHX. licToqHMKOM SHeprIIH ,D;JI.ll 3Toro cJiyIBHJIO yJihTpaqmoJieTosoe II3JiyqeHHe CoJIHn;a. AMIIHOKHCJIOThI, H ,n;pyrHe coe,n;MHeHM.ll HaKailJIHBaJIIICb CTeHI:J;HIO «rrepBMqHoro ,n;o caxapa Tex nop, IIOKa He rrptto6peJIII KOHCII- 6y JlbOHa ». lIMeHHO B HeM B03HIIK.'Ia .iKH3Hb. 3Ta rHIIO- Te3a .llBHJiaCb OCHOBaHIICM ,D;JI.ll C03,ll;aHHJI TeOpHII 6IIOII033a. TeopuR 6uonoa3a c<t>opMym1posaua B 1947 r. aHrJIIIttCKHM yqeHhIM ,ZJ;m. BepHaJIOM. OH Bh1,n;eJIIIJI TPH CTa,ll;HH 6Horrosaa: 1. 2. 3. A6ttoreHHoe B03HHKHOBetttte 6ttoJiorttqecKIIx MOHoMepos. 06pa30BaHIIe 6ttOJ'IOrHqec1rnx IIOJIHMepoB. ct>opMttposamrn MeM6paHHhIX CTPYKTYP H rrepsttqHhrx opraHH3MOB (rrpo6HOHTOB). IlepBbie ,n;Ba 3Tarra OTHOC.HT K rrepIIo,n;y XHMHqec1w:H 3BOJIIOUHM. c TpeTbero aTana Ha'IHHaeTC.ll 6IIOJIOrII'leCKaJI 3BOJIIOUH.ll. 1. A6uoiei-1Hoe 603HU/CfWBeHue 6uoJtoiu'lecKux .MOHO.Mepo6. 4500 MJIH JICT Ha3a,n; aTMOC<l>epa 3eMJIH Ilo ,l];aHHbIM KOCMOJIOrHH, COCTO.HJia H3 raaoB: OKCII,ll;OB yrJiepo.n;a (CO, C0 2), rrapoB BO.D;hI, MeTaHa (CH 1), aMMIIaKa (NH3) II ,n;p. CBo6o,n;HhIH KIICJiopo,n; (02) OTCYTCTBOBaJI. Ilo,11; ,n;ell:CTBIIeM KOPOTKOBOJIHOBOro YJibTpa<l>IIOJieTOBoro II3JiyqeHII.ll II rp030BblX pa3p.H,D;OB MOrJIII npOHCXO,D;HTh XIIMII'IeCKIIe peaKUIIII, npIIBO,D;.llJUIIe K o6pa30BaHIIIO aMIIHOKHCJIOT, caxapoB H ,n;pyrHX 6IIOJIOrtt'leCKHX MOHOMepoB. B 1953 r. aMepIIKaHcKne IICCJie,n;oBaTeJIII MIIJIJiep II IOpII CKOHCTpyttpo- BaJiu crreu;naJihHhiii arrrrapaT, npoBeJIII sKcrrepIIMeHT II ,n;oKaaaJIH STY B03M:omHocTb. 2. 06paaoBaHue u 360JlfOU,UR 6uonoJLu.Mepo6. ,[t;aJihHell:rua.H XIIMIIqecKa.H 3BOJIIOUII.ll rrpOIICXO,D;IIJia rrpII pe,n;KOM coqeTaHHII 6Jiaronp1rn:THbIX YCJIOBIIH CIIHTeaa 6IIOIIOJIHMepoB. · AMep1maHCKHll: yqeHhIH C. ct>oKc B 1957 r. orrhITHhIM rryTeM ,n;oKaaaJI B03- llomHocTh o6paaoBaHII.ll rrenTII,D;HblX CB.H3eH B pacTBope aMIIHOKIICJIOT. 3KcrrePllllleHTaJibHO 6b1JI0 rroKaaaHo, qTo B BO,D;HOM pacTBope aJiaHIIHa B npHCYTCTBHH OC<>6oro BII,n;a r JIIIH03eMa n A T<I> 06paay10TC.H uenoqKII rroJIIIaJiaHIIHa.
06~aJl 6uoJWZUJl. ;)6(1/1,l()l{UJl 126 Oco6oe 3Ha'!e1rne JJ;.1H 611orroJUIMepoB HMeeT 06paa0Batt11e Mo.1eKy.1, T. qeK B cnoco6tthIX K 1982 caMoBOCIIpo113BOJJ;CTBy. r. oTKph1.1 ptt603HMhI - MoJieKy.1hr PHK PHK, )Ke BpeM.R MOJieKyJia JJ;HK PHK JJ;HK -- o6Jial(aIOil.\He 1rnrn.111- TM'!ecKoii aKTHBHOCTLIO. Ott OIIHcaJI .RBJiettHe caMocnJiaiicnttra -yJJ;a.1eHHH HHTpOHOB H3 MOJ1eKYJ1hI 11 AMeprrn:aHCKHir 6HOXH:vi1n: rrpou;ecca 6e3 yqacTttH 6eJmoB-lpepMeHTOB. B To OFW3aJiaCh J'IY'Illle rrpHCIIOC06.'IeHa AJIH ;:i;o.1rospe- MCHHOI'O xpaHeHH.R HmpopMaD;MH, TaK KaK HBJU:ICTCH 60J1ee YCTOH'!HBOH. 3. <Pop.Mupoea1-tue .ne.M6paHllbtX cmpy1Cmyp u 3eo.1t10u,wi npo6urmmoe. Cw,n:yIOil.\HM rnaroM cTaJio ofrLe,n;MHeHMe no.'IttMepos B .n;ncKpeTHh1e rpyrrIIbI c o6pa3oBaHHeM qacTttu; 6oJiee KpynHoro pa3Mepa - 1wau,epeam1-tbtX 1wne.1b II cpopMnpoBaHHe MeM6pau. B pe3yJihTaTe o6'he.n;ntteHMH Koa:o;epsaTOB c ttyicreHHOBhIMH KHCJIOTaMM o6pa30BaJIMCb crroco6Hhie K caMOBOCIIpOH3Be,n;eHMIO )f~H­ npomo6uo1-tmbt (:u:JIH npo6uo1-tmb1). JKuBbte opiaHU3Mbl - 3TO OTKpbITbIC CMCTeMbl, cnoco6Hbie Bhie opraHH3MhI - K caMOBOCIIpo- H3Be,n:eHHIO, o6MeHHBaIOID;HCC.R co cpe.n;oii BeID;eCTBOM H II0.1y'!a10ID;HC H3 Hee 3HeprHIO. IlepBbie opraHH3MbI 6bIJIH, IIO-BH,!J;HMOMy, reTepoTpocpaMH, IIOJIYtiaBIIIHMH 3HeprHIO 3a C'!CT aHa3p06Horo OKHCJieHH.R opraHHtICCKHX coe,n;HHeHHH. Ilo.RBJICHHe attoKcHrenHoro cpoTOCHHTe3a (rryprryptthie 6aKTepmi, 3e;reHbie H cepuh1e 6aKTepttH) rrpttBeJio K B03HHKHOBeHHIO asToTpocjJnocni. Bo3HHKHOBeHHe OKCHrennoro cpoTOCHHTe3a IIPHBeJIO COBpeMeHHOii aTMOCcpepbl, co.n;epmaID;eii KHCJIOpo,n: c K cpopMHpOBaHHIO (02). MOMeHTa B03HHKHOB€HH.R )KH3HH IJOHBHJ'IaCb CB.R3b Me:m.n;y 6HOJIOI'll'Ie- CKHMH, reOXHMH'!eCKHMH II reoJIOI'HtICCKHMH npou;eccaMH. Pa3BHTHe :tKH3HH Ha 3eMJie B pa3BHTHH :>RH3HH Bbr,n:eJIHIOT TPH 3Tarra: 1. 8BOJ1lOJJ;HH npoKapHOT. 2. 8BOJ'll0ll;HH O)J;HOKJieTOtIHhlX 3yKapHOT. 3. Ilepexo.n; K MHOI'OKJieTOtIHOCTH H 3BOJIIOI.J;HH MHOI'OKJI€TOtIHbIX opraHH3MOB. I. 3eoJlJOU,UR npoicapuom. Bo3HHKHOBeHHe JKH3HH rrponaolllJIO OKOJIO 4 MJip,n; JieT tta3a.n;, H c 3Toro Bpe- Mettn .n;e:HCTBYIOT BaJKHeHlllHe cpaKTOpbI 3BOJIIOI.J;IIH: HacJie,n;CTBeHHaH H3MeH'!HBOCTh H ecTeCTBeHHbIH OT6op. IlocJie IIOHBJieHHH JKH3HH ccpopMnpoBaJ'IHCb TPH rro.n;u;apcTBa: apxe6aKTep1rn, 3y6aKTepHH (BKJIIO'!a.H 11.Hairn6aKTep11n), 3yKapnOTbI (rpn6bI, pacTeHHH, rrpocTeiiurne, :>RHBOTHbie). Apxe6aicmepuu - 3TO IIpoKapHOTbI, KOTopbre HMeIOT oco6b1e csoiicTBa: a) 3K30H-ttHTpOHHYIO CTPYKTYPY rettos; 6) HX KJieTOtIHhie MeM6paHbl COCTOHT H3 H30IIpeHOH,ll;OB (3cjJHpbl yrJieB0,11,0po,n;oB), He BCTpeqaJOID;HXCH y .n;pyrHX opraHH3MOB; B) o6HTaJOT OHH B 3KCTpeMaJibHbIX YCJIOBHHX - B KOHI.J;eHTpnpoBaHHbIX pac- TBopax, ropHtIHX ByJIKaHH'IeCKHX HCTOtIHHKaX, 6eCKHCJIOpO,n;HhIX CJIOHX· IlO-BH,IJ;HMOMy, 3Ta rpyrrna Han6oJiee 6JIH3Ka K npo6HOHTaM. 3y6aicmepuu 6Hoccpepe. Y HHX: 3TO npoKapHOTbl, lllHPOKO rrpe,n:cTaBJieHHhie B COBpeMeHHOli
pa 3 Bumue JICUJnu na 3eM/l,e 127 a) reHOMhI He IIMeIOT :JK30H-HHTpOHHOH CTPYKTyphr; 6) OHII qpe3Bhl'l3HHO pa3H006pa3Hhl no MeTa60JIII3MY. BamHhIH apoMopcpo3 - B03HIIKHOBeHIIe cpoTOCIIHTC3a. 3a C'l8T .n:e.HTeJibHOCTII <PoTOCIIHT83IIpy10ru;nx rrpoKapIIOT' BhI,1.\8.'IRIOil~lIX 1<IICJIOpo.n:, OKOJIO 2 MJIP,ZJ; JI8T Ha3a.n; c<PopMIIpOBaJiaCb aTMoc<Pepa, rro.n:o6Ha.H coBP8M8HHOH. rrpHHIJ;IIrrIIaJihHO HOBhIH 3Tan B 3BOJIIOI~IIII OihWKJtemo'lHbte ayKapuombl - jf{H3HII. CorJiacHo meopuu cuM6uoie1te.Ja, :JyKapIIOThI B03HIIKJIII 6Jiaro.n:ap.H CIIM6II03y Kpyrrttoii aHaapo6ttoii KJieTKII (apxe6aKTepIIII c <PepMeHTaMII rJilIKOJIII3a) c aapo6HhIMII 6aKTepIIRMII. Attaapo6Hhre 6aKTepIIII, crroco6Hbie K OKIICJilITeJrhaoMy qiopMIIpOB3HIIIO, .n:aJIII Ha'laJio MIITOXOH,ZJ;PIIRM. CHM6II03 c IJ;IIaHo6aKTepn.HMH rrpnBeJI K IIORBJieHIIIO xJioporn:racTOB. CnM6Ho3 c 6aKTepmIMH Tuna crrnpoxeT - K o6pa3oBaHuIO JKryTHKOB. B rroJih3Y rHnoTe3hI CIIM6uorette3a roBOPHT CTpOeHIIe opraHeJIJI (.n;ByMeM6paHHOCTb), CTpOeHue IIX ,UHK II 6eJIOKCHHTe3npy10ru;ero arrrrapaTa, a TaK:>Ke crroco6 pa3MHOJKeHIIR II qyBCTBHTeJihHOCTh IC TeM me aHTn6noTnKaM, 'ITO n 6aKTepnu. II. 3e0Jt10u,u.H ayKapuom. 8ToT npou;ecc CBR3aH c p.H.n;oM Ba:arneH:urnx apoMop<Po3oB: noRBJieHneM MnToaa, IlOJIOBoro rrpou;ecca, B03HllKHOB8HHeM MHOrOKJI8TO'lHOCTil (puc. 51). B OCHOBe rrpe.n;cTaBJICHilH 0 <PopMupoBaHHil MHOrOKJieTO'lHOCTH JICJKHT runoTeaa pyccKoro yqettoro 11. 11. Me'lHHKOBa o rrpoMeJKyTO'lHOM npe.n;Ke <Parou;nTeJIJie - wapoo6pa3HOH KOJIOHIIIl JKryTHKOBhIX, cnoco6HhIX K cparor~HT03y. B KOJIOHnn KJieToK B03HnKaJio pa3.n;eJiettue <PYHKIJ;MH (nnru;eBapettne n: nepe,ZJ;BnmeHne) (pnc. 52). -~ E. 1 o, &aKTepH•n ..Ht.1M ¢tOTOCHHTeTHK // MHTOl\\APH• 4 ~T II -.!'&' nepsaA 3 paCTMTenbHaA KneTKa 7 -----b XnoponnacT AHU•po6Hata 6aKrepHA Pile. 5 1. r111noTernyecKasi cxeMa c111Mfa1ornYecKoro 06pa3osaH111si 3yKap111ornYecKOM KJleTKl!I (u,wj:>P&M11t YKa3aH XOA 3BOnlOU,1!11!1)
128 B 70-x rr. HH3M - xx B. 300.JIOrH o6paTH.JIH BHHMairne Ha Me.JIKHW MOpCKOW opra- TpttxorrnaKc. B 1973 r. pyccKHw 300.1or A. B. 11BattoB ycTaHOBH.1, '-no TpHXOII.JiaKC no CBOeMy CTpoeHHIO COOTBeTCTByeT I'HIIOTeTWieCKOW cparou;HTeJ1.rre H HBJJHCTCH rrpoMe)KyToqHblM 3BeHOM MC)K,D;y MHOrOK.JICTOqHblMH H O)J,HOK.JICTOqHbIMH. ,CynMj>onoliyc TepMonna:iwa flllocj>KllW .'i .i' \ ·~ •\.. ~ ~ '"iti Apxe151lcTePa ~. "'•-. Ilporeaon.i PMc. 52. CxeMa B3a1-1MOOTHOweH1<15l ocHOBHblX u,apcTB lK"1Bb1x opraH"13MOB (no 6. M. MeAHl<IKOBy, 198
Ocno6nMe 3manbi 36011-10u,uu opzanu'lec1wzo Mupa 129 OcHOBHbie aTarrhI aBOJIIOD;HH opraHH'lec1wro MHpa HcTop11.H 3eMJIM rro.n;pa3.n;eJI.HeTCH Ha xpyrrHhie rrpoMe)l{yTKM BpeMeHn -reoxpoHoJior11qecxyro IIIKaJiy. Ha1160Jiee xpyrrHhie KaTerop1111 - ::ioHbI, nx BCe·ro ,n;Ba: Kpunm030U (OT rpeq. «CKphITa.H )l{I13Hb») H rpa11epo30U (OT rpetf. «HB· aa.H :IKH3Hb» ). 80Hbl .n;eJIHTCH Ha 3pbl. KpunmOJou: apxeif (.n;peBHeifruaH )l{M3Hh); rrpoTepo3oii (paHHHH )l{H3Hb). <Pa1tepo3ou: rraJieo3oif (.n;peBH.H.H IBM3Hh); Me3o3oif (cpe,n;HHJI )l{H3Hh); Kai-iH030M (HOBaH )l{I13Hb). 8pbI ,n;eJIHTCH Ha rrepuo,n;b1. HaaBaHHH rrep110,n;oB cooTBeTCTBYIOT 11a3BaHHIO MeCT, r.n;e OHH 6bIJIH OIIHCaHbl (rrepMCKMM, KeM6pMHCKMH), JIH6o Bamaeii:llIMM rrpOHBJieHMHM iKM3HH B 3TOT rrepHO,ll; (KaMeHHoyroJibHhIH, MeJIOBOH). HcTopHH 3eMJIM 11ayqaeTcH rro ocTaTKaM .n;peBHHX opraHH3MOB B paaJI11qHbIX reoJior11qecK11x cJIOHX. Coa.n;aeTCH rraJieOHTOJior11qec1rnJI J!eTorr11ch ocHoBHbIX 3BOJIIOil;HOHHhIX co6h1T11ii:. OrpoMHbIH BKJia.n; B pa3BHTHe rrnJieOHTOJIOrH11 c,n;eJiaJI aHrJI11ii:cx11ii: yqeHhIH q. Jia:H:TeJib (1797-1875). OH ccpopMyJmpoBaJI nputtu,un aKmyaJtU3Ma, corJiaCHO KOTOpOMy Ha 3eMJie IIOCTOHHHO .n;ettCTBYIOT cxo,n;Hbie cpaKTOpbI cpe.n;h1. OcHOBHbie co6hlTMH rrMeOHTOJior11qecKoii J!eTorruc11 rrpe,n;cTaBJieHbl B Ta6JI. aa c. 130. Kpy1IHeiinme apoMopcp03bI B aBOJIIOD;HH opraHH'lec1wro MHpa ApxeucKaR 3pa (3500-2500 MJIH JieT Ha3a,n;) 1) IloHBJieHHe m11aH11. iKH3Hh rrpe,n;cTaBJieHa rrpoKap110TaM11. Crroco6 rrHTaHHH reTepoTpOcpHbIH, aHa3po6HbIH. 06MeH Beru;eCTB - rJIMKOJIH3 (Ha1160Jiee .n;peBHHH cpopMa). 2) IloHBJieHHe aBTOTpOcpHoro crroco6a ITHTaHHH - cpOTOCHHTe3a I1 xeMOCHHTe3a, cp11xcan;1111 aTMoccpepHoro aaoTa (oKOJIO 3 MJip.n; JieT Ha3a,n;). 3) IlOHBJieH11e OKCHreHHOro cpOTOCHHTe3a l1 B pe3yJihTaTe HalWrr.1eH11e KHCJIOpo.n;a B aTMoccpepe. 4) BoaHHKHOBeH11e ,D;bIXaHHH - OKMCJIMTeJibHOro cpoccpop11JI11poBaHHH. 5) CcpopMHpoBaJIHCb .u;Ba n;apcTBa 6axTepHif: apxe6aKTep1rn H ::iy6aKTepHH (HCTHHHhie 6aKTepHH). Ilpomepo3oiJcKaR apa (2500-570 MJIH JieT Haaa.u;) l) IlOHBJieHHe ayKapHOT B peayJibTaTe CJIOiKHOro CHM6H03a pa3HblX rpyrrn IIPOKapHOT. 2) IloHBJieHHe MHT03a, Me:H:oaa, rroJioBoro rrpon;ecca. 3) BoaHHKHOBemrn MHOrOKJieToqHoCTH (l,3-1,4 MJip.u; JieT Haaa.u;). cI>opMHpoBaH11e OCHOBHbIX n;apCTB: rrpOTHCTOB, iKHBOTHbIX, pacTeHHH (B0).(0POCJieii), rpH6oB. 4) Bb1xo.u; rp1160B 11 Bo.u;opocJieii: Ha cyruy B BH.u;e JIHlllaHHHKOB. Pa3BHTHe IIPHMHTHBHbIX coo6ru;ecTB Ha cyrue (6aKTepHH H n;11aH06aKTep1111). 5) y iKHBOTHblX IIOJ:!BJIHeTCH BHyTpeHHJ:!J:! IIOJIOCTb l1 CKeJieT (HapyiKHbIH, BHyTpeHHHH, r11.u;pocTaT11qecKHH).
130 06w,aJt 6uO/WZUJt. 360/l/OlfUJt OcuoBHhIC JTanhI naJieOHTOJiorn•rnc1wii JieTonucn -----1 Spa I 0 I Kall:uo3oll:cKaH I 50 Me.no Boil Me303ofrcKaJI IOpcKHil TpHacoBhill: I Co6h1T11e 3BOJUOll,HH qeJIOBeKa Heoreu ( qeTBepTHqHhIH) I <PopMHPOBaHHe COBpeMeHHhlX ceMeHCTB IlTHI.\ H MJieKOIIHTaIOill,HX na.'!eoreu (TpeTHqHhlH) 100 200 IlepHO,[\ >:S: 0 I? 0 A 300 @ IlocJieAHHe AHH03aBphI TiepBhie npHMaThl IlepBhie 11,BeTKOBh!e pacTeHHH )J;HH03aBphI TiepBhie IITHI!hl TiepBbie MJieKOIIHTaIOill,He Be.nHKoe BhIMHpaHHe MopCKHx opraHH3MOB KaMeuuoyroJihHhlll (Kap6ou) TiepBhie perrTHJIHH CeMeHHhle rrarropoTHHKH )J;eBOHCKHH TiepBhle aMqm6HH YBeJin:qeuHe pa3uoo6pa3HH phr6 400 CH.nypHllCKHll TiepBhie Ha3eMHb1e cocy,1v1cThle paCTeHHH 450 OpAOBHKCKHll B3phrn pa3Hoo6pa3HH B ceMellCTBax MHOrOKJieToqHhIX JKHBOTHhlX '9 350 IlaJie030HCKaH 500 550 KeM6pn:iicKn:il 650 >:S: 0 0 ..."' ~ :s:A 700 ~ TiepBhie JJieMeHThI cKeJieTa JJ;oKeM6pHiiCKaH 3AHKapCKHH I I I TiepBhle ph!6hr TiepBhie XOPAOBbre 600 I I II flepMCKHH Cl) ' ,[(HBepreHI_J:HH M.'!eKOIIHTa!Olll,HX 1 fOCIIOACTBO peIITHJIHll 250 I I TiepBhle MHrKOTeJih!e MHOrOKJieToquhre JKHBOTHhie TiepBhle CJieAhI JKHBOTHhIX l I I I 1Ia.1teo3ouc1caR apa (570-230 MJIH JieT Haaa,n;). 1) 2) 3) 4) IlO.HBJieHIIe xop,n;OBhIX (KeM6ptt:H). Bhrxo;:i; pacTeHHM (cocy;:i;ttcThIX) Ha cyruy (cmiyp). 06paaoBaHtte aMqm6tt:H (cTerou;e<t>aJioB) (,n;eBOH). <l>opMHPOBaHHe penTHJIHM (Kap6oH) H IlO.HBJieHHe ceMeHHhIX nanopoTHHKOB.
131 Me303ouc1cafl Jpa (230-67 MJIH JieT Ha3a;:i;) 1) KpynHhIM ap0Mopcj_Jo3 - B03HttKHOBemre TenJioKpOBHOCTH. IlonBJUIJOT- c.H nepBbie MJieKOTIHTalOII~He H HaCTO.Hru;tte KOCTHCTbie pbI6bI. 2) PacnBeT roJioceMeHHhIX, noaBJieHtte nBeTKOBb1x pacTeHttll: • .IJ:aJlhHe:Hmee pa3BHTHe iKH3HeHHbIX cjJOpM H yBeJill'IeHHe TeMTIOB 3BOJIJOntttt. Kai1H030Uc1caJ1 Jpa (65 MJIH JieT Ha3a;:i; - ttacTom1~ee BpeM.H) 1) I'JiaBHhIH apoMopcjJ03 - 2) IlpoHCXO)J;HT cjJopMn:poBaHHe npeCMhIKaroru;n:xc.H, l'OJIOBOHOrn:x MO.JIJIIO- aHTponoreHe3, KOTOpbIM npHBe.1 K IIO.HBJieHHJO COBpeMeHHOI'O qeJIOBeKa. CKOB n: ;:i;pyrux rpyrm ;:i;peBHHX mHBOTHhIX. PacnpocTpaaa10Tc.H MJieKonn:Taroru;ue. 3) 4) IllttpOIW pacceJI.HIOTC.H O,ll;HO,ll;OJibHble TpaB.HHHCTbie pacTeHH.H. 0TCTYTIJieHHe JieCOB, CB.H3aHHOe c IlOXOJIO;:J;aHHeM H cyxoCThlO KJIHMaTa, cTn:Mynn:poBaJio 3Bomonn:ro qeJioBeKoo6pa3HhIX o6e3h.HH (roMn:Hn:;:i;), 'ITO npttBeJio K IlO.HBJieHHIO aBcTpaJIOTIHTeKOB H apxaHTpOilOB. ,IJ;oKa3a TeJibCTBa aBOJIIOD;HH 3BOJIIOnHOHHhIM nponecc n:3yqaeTCSI pa3JIH"<IHhIMH MeTo;:i;aMn:. Kam;:i;hIH H3 MeTO,ll;OB npe,ll;CTaBJI.HeT CBOH ,ll;OKa3aTe.JlhCTBa. 0CHOBHLie /J;Olm3aTeJILCTBa (MeTO/];hl H3yqeHHJI) 3BOJIIOD;HH 1. IlaJieOHTOJIOI'H"<IeCKHe: HCKOnaeMhle nepexo,D;Hhie cjJopMhl; naJieOHTOJiorn:qecKtte p.H,ll;hl. 2. Blrnreorpacpn:qec1me: cpaBHeHn:e qJJiop H cpayH KOHTHHeHTOB; peJUiKTbI. 3. I CTA)Ull 5JJ)~8}5) ~t~~~ Ill CTA.llH.11 MopcpoJiorttlfeCKHe: ~~~ roMoJio- ruSI oprattoB; py,n;ttMeHTh1; aTaBH3Mhl. 4. 3M:6pttOJIOI'H"<IeCKHe (pttc. - 53): «3aKOH 3apo,n;bIIIIeBOI'O CXO,!J;- CTBa~ K. B3pa; - npttttnurr peKamrTy JI.Hnuw 3. I'eKKeJISI IIOBTOpeHue pM6a ca,·u1MaHJ(pa 4epeuaxa KpbICa l.IC.'lOBeK npe,n;KOBbIX cpopM B OHTOreaeae 5. 6. 7. 53. reHeTH'leCKI1e. P111c. BHOXHMHlJeCKHe H MOJieKyJISip- 3M6p1110Hbl scex n03BOHOYHblX Ha paHHlllX 3Ta· 5lsneH111e 3apOAblWesoro CXOACTBa. H0-6nonorH"<IeCKHe. nax pa3Bt.1Tt.1Sl 6onee CXOAHbl APYr c APYrOM, 1Iapa3IiTOJIOI'H"<IeCKHH MeTo,n;. YeM Ha 6011ee no3AH111X (no 3. reKKeJllO, 1866)
132 06UfaJt 6uoJWZUJt. 360./1,IOU,.UJt Ila.JieOHTOJIOrH'leCKHe ,ll;OKa3a TeJihCTBa H cKnnae.MbW nepexoihtbte rpop.'llbt - q>opMbI opraHII3MOB, cotieTaJOn:J,tte rrptt3HaKH 6oJiee µ;peBHIIX tt MOJIO.ll,bIX rpyrrrr. Haxo.ll,KII H orrttcaHtte TaKttx cpopM II03BOJIHIOT BOCCTaHaB.'IHBaTh qmJioreHe3 OT.ll,eJibHbIX rpyrrrr. JipuNepbl. 1. HxTHOCTera - IICKOrraeMaJI cpop:vm, IWTOpaJI II03BOJIJieT CBJI3aTb p1>16 c Ha3eMHhIMII II03BOHO'IHI>IMII. 2. ApxeorrTepttKc (rrepBorrTttI.J.a) - rrepexo.n- HaJI cpopMa OT perrTMJmii: K rrTttu;aM JOpcKoro rrepttoµ;a. IIptt3HaKM perrTuJmi1: ,ll;JllIHHhIH XBOCT c Hecpocrrrn.MIICJI II03BOHKaMII, 6pJOII!Hbie pe6pa, pa3BIITbie 3y6bI. IIptt3HaKH IITHu;: TeJIO IIOKpbITO rrepbRMH, nepe,ll;HIIe KOHe'IHOCTII rrpeBparn;eHhl B THIIH'IHbIC KpbIJlbJI. II a.neonmo.noiu'iecKue pRObL - aTo pR.ll,hI ttcKorraeMhIX cpopM, CBJI3aHHhie µ;pyr c µ;pyroM B rrpou;ecce 3BOJIIOI.J.IIH tt oTpamaJOrn;tte xoµ; qm.noreHe3a. Ilpu.Mep. 8BOJIIOI.J.IIOHHhIH pR.ll, JIOII!a,ll;eH Ha'laT B HCCJie.ll,OBaHHJIX pOCCIIH- CKOrO naJieOHTOJiora B. 0. KoBaJieBcKoro (1842-1883). HaJIII'IHe MHOrttx rro- cJie;:i;oBaTe.'IhHO CMeHRIOll.J.HX µ;pyr µ;pyra cpopM II03BOJIIIJIO IIOCTPOHTh cjJHJioreHeTII'leCKHH PR;:\ OT aorttrrnyca .ll,O COBpeMeHHOH JIOII!a.ll,H. IIaJieoHTOJiorntieCKHe .ll,OKa3aTe.TihCTBa MoryT 6hITh rrpttMeHeHhr K 113yqeHHIO MaKp03BOJIIOI.J.IIH (npoTeKaIOrn;e:H B MaCII!Ta6ax COTeH ThICJI'l JieT) H He HCIIOJ!h3YIOTCJI rrpH H3y'leHHII MIIKp03B0JlI0lJ.HII. Bnoreorpac}>nqecKne .uoKa3a TeJihCTBa BttoreorpacpttR no3BOJIJieT aHaJIH3ttpoBaTh 3BOJIIOI.J.IIOHHhie JIBJieHHJI pa3HhIX MaCII!Ta6oB OT MHKPO- ,Zl;O MaKp03BOJIIOIJ.MM. Cpaaneuue rp.nopbt u rpayHbt. Pa3JHI'IIIJI HJIII cxoµ;cTBa cocTaBa cjJJiophI 11 cpay- HhI MoryT 6hITh CBJI3aHbl co BpeMeHeM reoJIOrII'leCKOro pa3,ll;eJieHHJI MaTepttKOB. IIpuMepbt. 1. ABcTpaJIHJI na rrpOTJIJI<eHHII 6oJiee Jiach c µ;pyrttMH MaTepuKaMtt. B 120 MJIH JieT He coeµ;ttHR- 3TOT rrepttoµ; rrpoucxoµ;HJIO cpopMttpoBaHue oco6oi1: cpayHhI, pa3BHBa.'IttCh cyM'laThie 11 KJIOatIHhie MJieKorr11Ta10rn;tte. 2. CJie- ,ll;hI reoJiorutiecKoro e,n;ttHCTBa IOmHorr AMepttKM, AcppHKII, ocTpoBa Ma,n;aracKap coxpaHJIIOTCJI B coBpeMeHHOH ¢ayHe. HarrpttMep, Rrn;epttIJ.hI-ttryaHhI Ma,n;aracKapa u IOmHott AMepttKII. Pe.nuKmbt. coM PeJIIIKTOBhie cpopMhI - IlpI13HaKOB, aTo HhIHe m11Byrn;11e Btt,n;hr c KOMITJieK- xapaKTepHhIX .ll,.;'IJI ,n;aBHO Bh!Mepll!HX rpyrrrr rrpOIIIJihIX arrox. PeJIHKTOBhie cpopMhI cB11,n;eTeJihCTBy10T o cjJJiope 11 <Payne ,n;aJieKoro rrpOIIIJIOro 3eMJIH. IIpuMepbt. 1. faTTepttR - perrTIIJIIIJI, 0611Ta10rn;aJI B HoBoi1 3eJiaH,n;HH. 8TOT BH,11; JIBJIJieTCJI e,n;HHCTBeHHhIM HhIHe JI<HBYll.J.HM rrpe,n;cTaBIITeJieM IIO,ll;KJiacca IIepBOJirn;epoB B KJiacce PerrTttJIHH. 2. JlaTHMepHR (u;eJioKaHT) - KIICTerrepaJI p1>16a, 06ttTa10rn;aJI B rJiy6oKOBO,n;HhIX ytiaCTKax y 6eperoB BocTOtIHOH A¢pttKH. E,n;ttHCTBeHHhIH rrpeµ;cTaBttTeJih oTpJI.n;a KttcTerrephIX pb16, Hatt6oJiee 6JIH3KHH K Ha3eMHhIM II03BOHO'IHhIM. 3. fttHKro ,ll;BYJIOIIaCTHblll - peJIHKTOBOe pacTe- HHe. B HaCTOJirn;ee BpeMJI pacrrpocTpaHeHo B KttTae 11 .HrroHHH TOJihKO KaK .n;e-
133 yopt/JOlWZU'tec1>ue iloKa3amellbCm6a icopaTMBHOe pacTeHMe. 06JIMK rMHKrO II03BOJUieT rrpe,ncTaBl1Th JT,peBeCHhie <PopJ\{bl, BhIMepume B IOPCKOM rrepMO,D,e. . BMoreorpaqmqecKMe MeTO,ll;hI no3BOJUIIOT rrnyqaTh npol!eCchI KaK MaKpo;rnoJ:IJOI!MH, TaK II MMKpmmOJIIOl!MM. Mop<l>oJiorH11ecKHe ;:i;oKa3a TeJILCTBa Ba3MPYIOTCH Ha OCHOBHOM rrpIIHIJ;IIne: rJiy6oKoe BHyTpeHHee CXO,ll;CTBO opraHH3MOB noKa3hIBaeT po,z:i;CTBO cpaBHMBaeMhIX <j)opM. I'oMOJWZUJl opzaH06. 31'0 opraHhI, IIMe!Oll~IIe CX0,11;- fOMOJIOrIIqHhie opraHhI - Hblff nJiaH CTpOeHIIH, BhITIOJIHHIOIQIIe KaK CXO,ll;Hhie, TaK H pa3JIIIqHhie <PYHKl!HH H pa3BHB3IOIQHeCH II3 CXO,ll;HhIX 3aqaTKOB. IIpuMepbt. 1. Pa3JIJiqHhre no BHeIII- ueMY BH,ll;Y II cpyHKIJ;IIHM KOHeqHOCTII MJieKOIIHT3!0IQIIX IIMeIOT CXO,ll;HhIH IIJ13H CTpOeHIIH H cpopMIIPOBaHIIH: KOCTII IIJieqa, npe,z:i;rrJieqh.H, 33IIHCThH, IIHCTH, cpa- 2. JiaHI' naJihIJ;eB. qepena B pH,z:i;y lfayqeHIIe aHaTOMHII HII3IIIHX II A BhICIIIIIX Il03BOHOqHhIX II03BOJIIIJIO ycTaHOBIITh rOMOJIOrIIIO KOCTeif qepena y pb!6 H CJiyXOBhIX KOCToqeK (pHC. y MJICKOIIHT3!0IQIIX 54). fOMOJIOI'flqHhie opraHhl CJie,z:i;yeT OT- B JIHqaTb OT aHaJIOJ'HqHhIX. AliaJiorHqHhle opraHhl c BHeIIIHHM CXO,ll;CTBOM, aTo opraHhI BhIIIOJIHHIOIQHe CXO,!J;Hhle <l>YHKIJ;HH, HO HMeIOIQIIe pa3HOe npoucxom,z:i;eHIIe. ,[{JIH ycTaHOBJieHHH po,n;- B CTBa H nyTeH 3BOJIIOIJ;HH OHM He HMe!OT 3HatieHHSI. IlpuMepbt. 1. KomoqKH aKau;m1, 6ap6apHca, 60.HphIIIIHHKa H emeBMKH - P1tc. 54. 3TO 3HaJIOrHqHhie opraHhl, rrpOHCXO)J;HT cnyxosbrx KOCTOYeK cpeAHero yxa. CpasH1-1- •a TenbHO-aHaToM1-1YeCK"1H paaJIHqHhIX 3aqaTIWB H TKaHeH, II() BbIIlOJIHSl!OT CXO,ll;HhIC cpyHKIJ;HH 33- JIUITbI pacTeHHH OT noe,z:i;aHHH ;KHBOTlllil:r.t:u. 2. fJia3a qeJioBeKa H KaJihMapa A- AaHCKOro, qTo 6bIJIO IIOK333HO cneu;H- . . . .H HCCJie,n;oBaHHHMH. ~ttPyiJuMenmbt. l'a&LI - 3TO opraHhl, yTpaTHBIIIHe 5 - ll.111X; MOnOTOYeK; CTpeMeYKO; 6- H. H. lllop- KBaApaTHaSl KOCTb - HaKOBanbHSl MneKOn11TalOL1.!HX; Ball KOCTb - npeCMblK8IO- MneKon1-1Tat0ll.lero (113 1979). I - Ha!l KOCTb pe - Py,z:i;HMeHTapHhle op- KOCTHOH pb16b1; 8- llteMOTpSI Ha nopa3HTCJlhHOe CXO)J;CTBO cpaBHHTCJihHO-aHaTOMHqe- nosson11eT HOBn8H"1Sl cnyxOBblX KOCT048K Mn8KOn"1TaIOL1.!"1X: ll.18rGC!l; ·~emm, p11A BOCCTaHOB"1Tb 3BOnlOL\"10HHblH nYTb CTa- bJISIIOTCSI 3H3JIOrHqHhIMH opraHaMH, lllh.abIMH roMononl!Sl opraHOB. r0Monon111 4- 3- 2- C04118HOB- r110MaHAH6y1111- 3y6Ha!l KOCTb; 5- ymo- 6apa6aHHaSl KOCTb MJ18KOnHTalO- rHOHA
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[lalleonmon,oiu..,,ecKue iJoKa3amen,bcm6a npoucxo:HCiJeuun 'len,o6eKa 139 AecmpaJtonumeKu (npe,n;IIIeCTBeHHHKH qe.'IOBeKa); [JumeKaumponbt (,n;peBHeiiIIIne mo,n;H, apxaHTporrbr); HeattiJepma.nbU,bl (,n;peBHHe JIIO,ll;H, rraJieoattTporrhr); qeJtoBeK c0Bpe.Me111wzo muna, ucKonae.Mbtu u coBpe.MeHHbtU (HeoattTpOrrbr). AecmpaJtonumeKu ( «IOmHaH o6e3bHHa») rrpe,n;cTaBJIHIOT co6oii o,n;Hy H3 rrepBhlX CTa,IJ;HM 3BOJII011,HH 'leJIOBeKa. Ilpe,n;IIIeCTBeHHHKH scex HCKOrraeMbIX H coBpeMeHHbIX JIIO,n;e:M:. BBe.'l TepMnH attrJ'IHMCKn:H aHaTOM P. )lapT, Hame,n;mnii B 1924 r. Ha rore AcppnKaHcKoro MaTepnKa OCTaTKH ,n;eTcKoro qeperra. B HacTo.amee BpeMH n3BeCTHO HeCKOJibKO coTeH acppnKaHcKnx aBcTpaJiorrnTeKoB. oTo npoMemyTO'lHbIM THI! Mem,n;y qeJIOBeKoo6pa3HbIMH o6e3bHHaMH H 'leJIOBeKoM. Oco6eHHOCTH aBcTpaJionnTeKoB: o6HTaJIH Ha OTKpbITbIX TeppnTopmi:x, xoponio 6eraJIH, BblCB06o,n;nB nepe,n;HHe KOHe'lHOCTH, BeJin CTa,ll;HblM o6pa3 lliH3HH. lfa-3a )R'.H3HH B caBaHHe H3MeHnJiaCb TepMoperyJIHll,HH: pe,n;y11,npoBaJiaCb niepCTb, yBeJIH'lHJIOCb IIOTOOT,IJ;eJieHne, a TIOTepH c IIOTOM HOHOB HaTpHH KOMneHcnpoBaJiaCb nepexo,n;oM K XHill,HH'leCTBy HJIH IIOTpe6JieHneM TIOBapeHHOH coJIH. He6oJihIIIne KJihIKH n pe311,hI He BhICTynaJm Ha,[\ 06I11,nM ypoBHeM 3y60B. tJeJIIOCTH aBCTpaJIOTIHTeKOB 6bIJIH MaCCHBHee, 'leM y COBpeMeHHOro qeJioBeKa. HB..z:16poBhH BhICTynaJin Bnepe.z:1, o6'beM M03roso:H noJIOCTn He6oJihIIIOH nee 600 CM 3; 3aTblJIO'lHbIH OT,[leJI 3aKpyrJieH. He 60- B CBH3H c rrpHMoxom,n;eHneM aBcTpaJionnTeKn rrpno6peJin « qeJioBe'leCKne » 3a6oJieBaHHH, a 3aTpy,n;HeHHe po,n;oB rrpnBeJ'lO K TOMY' 'ITO rrpenMylll,eCTBO IIOJiyqaJIH ' caMKH, pomaBinHe He,n;oHoIIIeHHhIX ,n;eTe:H. oTo rrpHBeJio K y,n;mrneHnIO rrepHo,n;a ,ll;eTCTBa H crroco6CTBOBaJIO yBeJin'leHHIO cpoKa o6yqeHHH MOJIO,n;oro TIOKOJieHHH. BpeMH CTaHOBJieHHH aBCTpaJIOIIHTeKOB OTHOCHT K nepno,n;y OT 9 ,n;o 5 MJIH. JieT Ha3a,n;. Ilo npO,ll;OJilliHTeJibHOCTH 3TOT nepno,[I 3Ha'lHTeJibHO ,ll;JIHHHee, qeM BCH IIOCJie,n;yroI11,aH ll,HBHJIH3all,nH. Ce:Hqac xoporuo H3yqeHbI ocTaTKn o,n;Horo n3 paHHHX aBcTpaJionnTeKoB act>apcKoro, Ha:H,n;eHHOro B oqmorrHn. 3TO OTHOCHTeJibHO He6oJihIIIaH o6e3bHHa: ,ll;JIHHa TeJia 110-120 CM, Bee 30 Kr, o6'beM M03ra 410 CM 3, npHMOXO,ll;Hlll,aH, ay6bl noxomH Ha qeJioBe'leCKne, o6HTaJia 3,5-4 MJIH JieT Ha3a,n;. BnocJie,n;cT- BHH aBCTpaJIOTIHTeKH pa3,n;eJIHJIHCb Ha ,ll;Be JIHHHH: aecmpaJtonumeK MOW,HblU KopeHacToro TeJiocJ10meHnH, ,n;JinHa TeJia 150-155 CM, Bee 50-70 Kr, qeperr MaCCHBHhIH, o6'beM M03ra 510 CM 3; 3y6hl KPYITHhle, BereTapnaHe11,; '. aecmpaJtonumeK a</JpuKallcKuu nMeJI MeHhIIIne pa3Mephr, ,n;m·rna TeJia ~:20-130 CM, Bee ,ll;O 40 Kr, o6'beM M03ra 450 CM3; BCeH,[IHhIH; TIOJIOmeHne TeJia '4>11 X0,1J;h6e 6hIJIO 6oJiee BbIIlpJIMJieHHbIM. ·lfu ABcTpaJiorrnTeKn 3aHHMaJinch co6npaTeJibCTBOM n oxoTo:H, HCIIOJih30BaJIH ,ll;PY'!Hhie opy,n;HH: naJIKH, ,n;y6nHhl, KaMHH H KOCTn. CTpOHJIH rrpHMHTlIBHhre LITHH - maJiaIIIn H HaBeCbI. OrHH He 3HaJin. Bee OHH IIInpoKo pacnpocTpa- . . JIHch no Acl>pHKaHCKOMY MaTepHKY (B03MOlliHO, rrpOHHKJIH B A3nIO, HO Bbl,.. JIH 1 MJIH JieT Ha3a,n;). :. - · 2-3 MJIH JieT Ha3a,n; mHJin cyI11,eCTBa, eI11,e 6oJiee 6Jin3KHe K qeJioBeKy. Homo
06utan 6uo1wzun. 3601uol{un 140 OcttOBHhie cTa.n:uu :JBOJIIOQUU qeJioBe•rn Dryopithecus Australop1t/1ecus Homo erectus 06e3bRHOnO.O.o0HOe 18-9 MllH ner Yenoee•ooOpaaHao o6e3bRH8; eoapacr OKOJJO 5 MJlH ner 2 MllH-500 TblC. ner J:lpMonITTeK Aecrpa.nomueK PaaMePbl He6011bw11e; 11aaa10U1Me MXOIU'UlMe e nonysblnpRMneHHOM Poer 120-150CM, Macca 20-50 Kr; npsu.AOKOJK,£1ett"'e. pyKa - xearaTenb- >KHBOTMOe; B03pacr 'ienoae• np•MoXOARWMH; eoapacr J:lpeaHei1w11ii 4enoee• (n11reKaHrpon, CHHaHrpon, rellAMb6eprc<1<il '(enoeeK HAP·) Homo sapiens 'ienoee• paayMHblH; aoapacr 500-30 TblC. ner Yenoee• paaYMHb1'1; aoapacr 40 TblC. ner J:lpeeH11o::i 4enoee• HeaH.Qeprane1.1 YenoeeK pa3yMHbl•; lOTblC. neT CoepeMeHHblii ltenoeeK .,;l >S s :i: 3 cc"' :t nOflQ)K6Hl1rt HeeblCOKHl1 pocr. MaCCl'IBHbtl1 KOCTRK HblVI opraH HeeblCOKMH pocr (155-165CM), MaCCHBHbl~ KOCTRK, KOpeHaCTbl'1, nOXOAK8 COrHyraR 'ienoee•oo6pa3Hble liepen H~3KH~. KOCH'! 'iepen 6onee BblCO• 3y6bl noQ>opMe 4epTbl B CTPQeHMM cxo,o,Hbl c 'te11oee4e- Ol{eHb TOflCTble, BblKlo1'1, Jlo6 HH3KMl>1, ~enlOCTll! cicMMH pa><eHHble HaA6POB- CKOWeHHbl", Clll.flbHO Hble AyrM, Macc111e- pa3BMTbl0 HaJl6poaHb1e '"lefllOCT\11. Hble JlyrH. 06beM M03ra 06beMM03ra 06beM MOJra 700-1250 CM 3 430-550CM3 OKOJIO 1400 CM3 Poer AO 180 CM, ¢>\11JH'(ecnu::i nm coepeMeHHOIO yenoeeica '-iepen BblCOKMH, no6 np•MOH, 6ea cnnowHOro HaA6pOBHOro sarwura, pa3svir noA6opoJl04Hbli< Bblcryn. 06beM M03ra 1000-1600 CM 3 06heM MC3f'a 1000-lBOC CM' OpyAM• TPYJl• OTCyrCTBylOT C111creMam-.ecK111 "' ~ "2l. 8 CTaJlHblH o6pa3 >ICM3HM lt13roroemi.io1 np1t1MM- r13rorosm:nor o6pa- lltCOO/lb3Yf()T ecTeCT- TlltBHbre 11:aMetfHble eeHHble npeJlMeTbl <a• opyJ:ll<e TPYJl• (KOCTb. nanKa "T.Jl.) OPYAH• rpyJla (KOnbe, CKOnOTblH KaMeHb, cKpe6oK, py6HJ10) Cr2JlHbli1 o6paa lKH3· 06u;er.reeHHbli< 06HM, COBMeCTH8s:i OXOTa pa3 >KM31iH, noAQep>KaHHe OfHQ lt13roroemnor cno:«V13roroa11nio1 opy- C/10)1(He"1lUHe JPYJl~s:i 11•• rpyw. M3 KOCTM " rpyAa" MexaHMJMbl KaMH• 6ora11Hb'e KaMeHHble Hble COCT86Hbfe OPYA"" TPYA" (OCTpM•. CKpe6KM, HO>KH MAP·) 06w.ec1aeHHbrl>1 o6PoAOBa• o6u;HHa. Ha1.1eblcw1o1e 1locn1pa3 )I013Hlo1; HCnOllbCrpo1o1re11bcreo no- *-0HMR B H8yKe, rex3088HMe OrH•. Crpo- ce11eH1<1'1. no>ieneHHe H\.'!Ke, MCK)'C crne HT8JlbCTBO O"i8rD8 M o6pRJl.OB. B03H'1KHOoeHHe '1CKyCC1Ba, lKHJlb•. nepeble aaxopoHeHMR fOH"i8PHOro Aena, 3eMJ10AellMR
flumeKaHmponbt. JleaniJepma;iblfbl 141 habilis - qe.10BeK yMeJ1hiii HM('Jl l\J1HHY Te.1a llO 120 CM, nee 40---50 Kr, OfrhCM 660 CM3, BCeH,ll;HbIM. 8TH cyru,cCTBa npno6peJrn cuoco6HOCTb 113l'OTaBJII1- JVI03ra BaTb O,ll;HO opy,nue c IIOMOlll,bIO ,L\pyroro, a He c IlOMOU~bIO py1-: 11 3y60B, Kai{ 3TO ):\eJiaIOT o6e3hHHhI. 8To 6bUIH caMhie npocTeiiume opy;111a JCa. Ha3BaHHe opy,n;w:Htto:H KYJihTYPhI - 061w.10Tan ra.'Ib- oJr.D;yBai1cKaH HJIH raJie'maH (TRK Ka1' B peqHhIX ,L\OJIHHax CblpheM p;;rn H3I'OTOB:IeHHH opy.rr,ttii c.1ymtt.ria raJihKa). Opy,!1.HH qeJIOBeKa yMeJIOI'O OKOH'JaTeJihHO BbIBeJIH ero ll3 Mttpa :tKHBOTHhIX. nMTeKaHTpOnbl (Ap0BH0MWM0 fllOAM, apxaHTpOnbl) Homo erectus - qeJioBeK np.RMOXO.L\Hlll,HM. BrrepBbre KOCTHhie cjJparMeHThr nnTeKaHTpona o6Hapy:>KHJI ro.1Jiatt.zi;cKHM Bpaq 8. ,[(J06ya Hao. 5:1.Ba B 1891 r. nume1canmpon (o • .HBa), cu11a11mpon (KHrnli), zeitiJPJ1b6ep2c1Cuu 'LP.wBelC (I'epMamm), amJWHmpon (Am1mp), meJ1a11mpon (IO:arnaH AcjJpHKa), OJ1dyeaitc1wil nume1-caHmpon IfaBeCTHO HeCKOJihKO cjJopM p;peBHeH:umx qeJioBeKoo6pa3Hhix: (1..I;eHTpaJibHaH AcjJpHKa). BpeMH cyru,ecTBoBaHHH y a 1,5-0,5 MJIH JieT Ha3a.D;. paHHHX apxaHTpOIIOB coxpaHHJIHCb IIPHMHTHBHbie o6e3bHHbH rrpH3HaKH, y II03,ll;HHX ITOHBJIHJIOCb 60Jibllie qeJIOBeqecKHX qepT. Oco6eHHOCTH cTpoeHHH: pocT 150-175 70-80 Kr; yse.11wqHJIHCh 750-1400 CM 3 , JieBoe ITOJIY· CM, Bee OT):\eJihI I'OJIOBHOI'O M03I'a, o6'heM M03I'8 ,llOCTHI'aJI mapHe 60JihIIIe rrpaBoro; Y3KHH IIOKaTbIM JI06, MOlll,HO pa3B11TbIH Ha.u,6pOBHblll BaJIHK, MaCCHBHhie qemocTH 6e3 no,n;6opo.z:1oqfforo BbICTyna. IlOHBHJiaCb rrpaBOpyKOCTb, PYKH 6oJiee pa3BHThI; o6pa30B8J'ICJI He60.'IbUIOH CBO)l; CTOIIhI; II03BOHQqHHK rrpuo6peJI Re60JibUIOM H3I'H6; peqh Ha HaqaJibHOH CTa,n;un: cjJopMn:poBaHHH B BH,n;e JieneTa (IIOHBJIHIOTCH OT,D;eJibHhie CJIOBa: «,n;aii:», «Ha» HT. ,n;.); iKHJIH CTa,D;aMn:; 3a60THJIHCb 0 6JIH:>KHHX. IluTeKaHTponbI He 6bIJIH TOJihKO co6l:rpaTeJIHMH. Ha oxoTHHqbnx cTo:H6wru,ax Ha:li:.n;eHhI KOCTH KpynHhIX iKHBOTHhIX. OxoTa Tpe6oBaJia TecHoro cnJioqeHHH ITJieMeHH. T:11rr OXOTbI - o6JiaBhI H 3aca.n;bI. CTOHHKH pa3HhIX TIIIIOB CBn:.n;e- TeJibCTBYlOT o cyru,ecTBOBaHHH pa3.u,eJieHHH Tpy.n;a. Opy,llHH Jia, KOJIYHhI, KOIIbH (3aOCTpeHHbie rraJIKH). KaMeHHbJe py6II- JIIO,n;H 3aHHMaJIHCb pa3HbIMH BH,D;aMH Tpy.n;a. OncpbITbie THIThI CTO.HHO.fC: OXOTHHqbn: Jiarep.H, MacTepCKHe ITO ,n;o6biqe KaMHH, ,n;oJiroBpeMeHHhie CTOHHKH. JKHJIHlll,a CTPOIIJIH ua OTKPhITbIX MecTax H B rreru,epax. OcTaTKH oqaroB roBopHT 06 wcrroJih30BaHHH rrnTeKaHT· POIIaMH OI'H.H. HeaHAepTanb1'bl (APeBHMe nJOAH, naneoaHTponb1) CBoe BH,ZJ;OBOe Ha3BaHirn OHM IIOJiyqn:JIH 6Jiaro.z:1apH Haxo.n;Ke B Jiuae HeaR,L\epTaJib (fepMaHHH). 0611TaJIH 200-35 1856 r. B fJ,O· ThIC. JieT Ra3a,n; B EBpone, .A3uu u AcjJpuKe. BpeM.!I cyru,ecTBOBaHHH II03.L\HHX HeaH,n;epTaJihI1eB cosna,n;aeT c nocJie.n;HHM oJie.n;eHeHHeM. HeaH,n;epTaJihll;hI 6hIJIH Heo,n;Hopo.n;Ho:ii rpyrrno:H. Oco6eHHOCTH CTpoeHHH oTHX .n;peBHHX JIIO,n;e:H: o6'heM M03ra 1200-1700 CMa, 1leperr 6oJiee BbICOKHM H OKpyrJibIH, JI06 HH3KHH, CKOIIIeHHbIH, CIIJihHO pa3BH· Thi Ha,ZJ;6poBHbie .z:IYI'H, HaJIHqHe rro.n;6opo.n;Ka; ,Zl;JIHHa TeJia CllBlloe TeJIOCJIOJKeHue. 155-165 cM; MaC-
142 TexHoJionrn: o6pa6oTKH KaMHH HeaH,n;epTaJihCKHM qeJIOBeKoM OT JIHqaJiaCh OTHOCMTeJibHO BbICOKHM ypoBHeM. 0HH C03,ll;aJIH KYJihTYPY CKpe6eJI H HaKOHeqHHKOB. TexHHKa IIOJiyqeHHH 3aroTOBOK opy,n;HM Tpe6oBaJia OIIhITa, TeXHHqeCKHX 3HaHHH, TOqHOll KOOp,ll;HHau;HH ,ll;BH)KeHHM, 6oJihIIIOrO BHHMaHHH. OrrhIT II03BOJIHJ, ,n;peBHeMy qeJioBeKy COKparu;aTb KOJIHqecTBO BpeMeHH, Heo6xo,n;HMOro ,n;.nH H3rOTOBJWHHH opy,n;Hii. lfa ,n;epeBa H3rOTOB.ITHJIHCh cocy,n;hI ,n;.nH BO,ll;hl H <JJieMeHThl )KH.,llru;. HeaH,n;epTa.IThllhI 6hIJIH HCKYCHhIMH oxoTHHKaMH. Ha HX CTOHHKax o6HapymeHhl CKOIIJ,eHHH KOcTeii: KpyrrHhIX mHBOTHhIX. C.nomHhie oxoTHHqhH ,n;eii:cT- BHH 6br.nH rro,n; CHJIY cor.nacoBaHHoll: rpyrrrre HeaH,n;epTa.nhu;eB. llcrro.nh30BaJIHCh rrpneMhI 06.naB H.ITH roHa mHBOTHhIX, TYIIIH y6HThIX )KHBOTHhIX pa3pe3aJIH n rrpomapHBa.nH Ha orHe. ,Il;e.na.nH MexoByro rrpocTyro o,n;em,n;y. 06HapymeHHhre 3epHOTepKll ll3 KaMHH II03BO.ITHIOT rrpe,n;rroJraraTh, qTO cyru;ecTBOBaJia rrpHMHTHBHaH o6pa60TKa 3epHa. H3MeHHJICH xapaKTep rroceJieHHii. 3aceJIH.'IHCh HaBechr, rpOThI, rreru;ephl. ,Il;JIH 3aiu;MThI KOCTPOB OT BeTpa ycTpaHBaJIHCh BeTpOBhie 3aCJIOHbI. IloHBHJIHCh rrorpe6eHmI - rrepBhre ,u:ocToBepHhre cJie,U:hl 3axopoHeHHH yMepIIIHX. HaqHHaH c py6ema 40 ThIC. JieT HeaH,u:epTaJihIIhI cocyru;ecTBOBaJIH c BIIOJIHe CJIO)KllBIIIHMHCH JIIO,ll;hMH cospeMeHHOro THIIa. 'ienoeeK coepeMeHHoro nma, MCKonaeMblM M coepeMeHHblM (HeoaHTponb1) Homo sapiens - qeJioBeK pa3yMHhIM. MaKCHMaJihHhIM reoJiorHqecKHH B03- pacT IIOJIHOCThIO CQJOPMHpOBaBIIIHXCH B 3BOJIIOllHH HCKOrraeMbIX HeoaHTpOIIOB paHee ou;eHirnaJicH B Ha3BaHHe 40 ThIC. JieT. «Kpo.MaflbOHU,bt» (TaK o6o3HaqaroT B JIHTepaType HCKonaeMhIX HeOaHTpOIIOB) o6H3aHO H3BeCTHOMY Q:>paHIIY3CKOMY rraMHTHHKY BepxHero rraJieOJIHTa - rpoTy Kpo-MaHhOH. ApeaJI pacrrpocTpaHeHHH KpoMaHhOHCKoro qeJIOBeKa orpoMeH. c ero IIOHBJieHHeM BH,ll;OBaH 3BOJIIOIIHH qeJIOBeKa 3aBep- IIIaeTCH. IlpH3HaKH OTJIHqHH qeperroB KpOMaHhOHII,eB OT HeaH,n;epTaJibII,eB: MeHee BhICTyrraroru;HM Brrepe,n; JIHlleBOM OT,n;eJI, BhICOKOe BhIIIYKJIOe TeMH, BbICOKHM rrpHMOH JI06, OKpyrJihIH 3aTbIJIOK, MeHhIIIHe no pa3MepaM qeThipexyroJihHhre rJra3HHIIhI, MeHhIIIe o6ru;.He pa3MephI qeperra. CQ:>opMHpoBaH rro,u:6opo,n;oqHhIH BhICTyrr qeperra: Ha,u:6poBHhIH BaJI.HK OTCYTCTByeT, qeJIIOCTH pa3BHThI CJia6ee, 3y6b1 HMeIOT MaJiyIO IIOJIOCTh. 06'beM M03ra 1000-1600 CM 3 ; pOCT ,n;o 180 CM. CTpoeHHe cKeJieTa Ty.nos11ru;a HCKorraeMhlX HeoaHTporros He OTJIHqaeTcH OT coBpeMeHHoro THIIa, XOTH KOCTH ero 6oJiee MaCC.HBHhI. mHJIH KPOMaHhOHllhI B cpe,n;HeM 30-50 JieT. TaKaH me rrpO,U:OJI)K.HTeJih- ,n;o Cpe,u:H.HX BeKOB. HOCTh m.H3H.H coxpaHHJiach y qeJIOBeKa coBpeMeHHoro THrra KpoMaHhOHII,bI H3roTaBJI.HBaJIH 6oJiee cJiomHhIH Ha6op opy,u:11ll:: HOmH, HaKOHeqH.HK.H Kon11ll:, .HrJihI, raprryHhI HT. ,u:. KocTHHhie opy,u:HH 6hlJIH rrpoqHhIM.H .H 60Jiee ,ll;OJirOBeqHbIMH, qeM KaMeHHhie. I'aprryHhI fl03BOJIHJIH 3QJQJeKTHBHO )l:06hIBaTb phl6y. IlpHeMhI OXOThI c HCIIOJih30BaHHeM 3HaHHH MeCTHOCT.H II03BOJIH-
\ ' 143 CoBpeMennbiii •ie1wBeK u ezo i16011104uR (neoanmponbt) Jiff y6HBaTh 3BepeH TbICffY:aMJL 1Ie.1oneK rro.1ylJMJI B03MO)KHOCTb npo:anrna1rna B TPY.z:IHO.z:IOCTYIIHhIX paiiottax. Jl\HJIH B rreUiepax, KOTOpbie HCIIOJih30BaJIHCb KPYI'JihIH ro.n; I1JII1 ,L\Jla Ce30Haoro npomttBaHM5L TaKme cTpOMJIM mMJIHUia B pe'IHhIX .z:IOJrn1-rnx, r.z:1e CKJiaJ~hr­ BaJIH M3 I\aMHH CTeHbI, a KpbilllY I13I'OTOBJIHJIM M3 IIJKyp. Koqyror.u;ne OXOTHJIKM cTpOMJIM nerKtte lllanaurn:. CypoBhre KJIMMaTH'leCKMe ycJIOBHH KpoMaHhOHI~aivr noMoraJia nepeHOCMTh Ten,1aa o.z:1em,11,a. Bo3JIHKJIO HCKyccTBO - cKy.TJhIITYPhI, ttacTeHHaa JKMBOIIHCh B nelllepax. fpa- BIOPhl JKHBOTHbIX H JII0,11,eH 6hIJU1 c,11,eJiaHhI Ha KaMHHX, KOCTHX, OJieHhMX porax. Ilo.HBHJIHCh o6p.H,11,hI, Bepa B 3arpo6Hy10 iKH3Hh, pttTyaJihHhre rrorpe6em,rn. CoepeMeHHblH ~enoeeK 1r1 ero aeomou,1r1s:i (HeoaHTponb1) CyllleCTBYIOT ,[\Be OCHOBHhre TO'IKH 3peHna nponcxom.z:1etttta qeJioBeKa coBpeMettHoro THrrn: mrroTe3a noJlu~e11mpU3Ma: qeJIOBeK pa3YMHbill B03HHK B HeCKOJibKMX :vrecTax IIJiaHeThI H3 pa3HbIX rrpe,n;KOBbIX cpopM, rrpMHa,n;JielliaUIMX K rraneoattTporraM; rttrroTeaa .11w11o~e11mpu3Ma: cyIIIeCTBOBaJio e.n;ttttoe MecTo B03HHKHOBeHna qenoBe'leCTBa OT KaKoro-To o6r.u;ero rrpe.z:1Ka, nocne qero HalJaJIOCh aKTHBHoe pacceneHHe no rrJiaHeTe. Bee coBpeMeHHoe qenoBelJeCTBO rrpttHa,[IJieJKHT K O,L\HOMY Btt,11,y. 06llltt:H ypoBeHh cptt3HtiecKoro H yMCTBeHttoro paaBMTHH O.z:IHHaKOB y Bcex n10.z:1eii. BttyTpM Homo sapiens BhI,11,en.HIOT 11y10, .Mo1120Jtouo11y10. Kam,n;a.H BH,[la TPH 6oJihlllMe pachi: 11ezpouo11y10, e6poneouo- 113 HHX .z:1enMTCH Ha MaJihie pacbI. PaanH'IHH Me:m.z:1y pacaMM CBO,[IHTC.H K oco6eHHOCTHM ~BeTa KOJKM, BOJIOC, rJia3, cpopMbI Hoca, ry6 M T • .z:I. Bo3HMKnH ::JTM pa3JIH'IMH B rrpo~ecce rrpttcrroco6Jiemrn qenoseqecKHX IIOIIyJIH~HH K MeCTHhIM rrpHpOJJ;HhIM ycJIOBHHM. IlpoMCXOJK,[leHHe pac OCHOBhIBaJIOCb Ha npoaBJieHMH rJiaBHbIX MexaHlI3MOB J13MeHeHlIH reH0$0H.z:1a H IIOIIYJIH~lIH - ecTeCTBeHHOI'O OT6opa lI JJ;pefapa reHOB. 3BOJIIO~lIOHHaa aaBepllleHHOCTb qeJIOBeKa OTHOClITeJihHa JI He 03HaqaeT IIOJIHoro npeKpallleHMH II3MeHeHIIH ero 6nonornqecKnx CBOHCTB. AHTporronorH Ha OCHOBaHIIlI aHaTOMlI'leCKHX Ha6JIIO,[leHlIH rrporH03lIPYIOT IIOHBJieHMe Homo futurus - qeJIOBeKa 6y.z:1yUiero. BttoJiornqecKa.H opraHnaa~n.H coBpe- MeRHoro qeJIOBeKa .z:1aeT B03MOJKHOCTb 6earpaHlilJHOH CO~MaJihHOH ::JBOJIIO~HM. ct>aKTOphl aHTporroretteaa Bo3HUKHOBe11ue npR.MOXOJ1COe11uR rro3BOJIHJIO ocB060.z:1MTb rrepe,ri;Hne KOHe'l- aocTn. TpyiJoBaR iJeRmeJlbHOCmb CmaiJ11btU o6pa3 JICU3HU - cnoco6cTBOBaJia pa3BMTMIO Moara. coBMecTHa.H ,L\e.HTeJihHOCTb rroaBoJiaJia coBeplllaTh CJIOJKHylO T.HJKeJiyIO pa6oTy' KOTOPYIO HeB03MOJKHO 6bIJIO BbIIJOJIHMTb O,[IHOMY. Bo3nUKllOBenue pe'lu u pa36umue R3btKa crroco6cTBOBaJIM rrepe,ri;aqe MHcpop- Man;1u1 n Koop.z:1nHa~nn ,ri;e.HTeJihHOCTH. ' OmKpbtmue u ucnoJLb306anue OZHR rrpnBeJin K aKTMBHOMY ynoTpe6JieHMIO Blactto:H: IllillllI, KOTopaa 6oraTa He3aMeHlIMbIMM aMHHOKMCJIOTaMH, crroco6cT- BYio~nMn pa3BMTMIO M03ra.
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06pa3oBanue Buao6 - pe3y11,bmam Mu«po:J6011,104uu 153 cTpoeHIIeM xpOMOCOM, BbIHBJieHHhIM c IlOMOlll;hlO ,n;m:})cpepeHIJ;Ha.JbHOH OKpacKfl, - KapHOTHilOM. llpwnep. AttaJIH3 xpoMOCOMHoro Ha6opa no3BOJill.'I pa3,n;e.1HTL e,n;HHhitt Btt,n; noJieBKH o6bIKHOBeHHOH ( 46 ( Microtus arvalis) Bn,n; H M icrotus arvalis) Ha ,n;sa no,n;sH,n;a, co,n;epma1QHe 54 ( Microtus Sub-arvalis) xpoMocoMhr. reHeTttt-IeCKH ycTOH'IHBa.H cHcTeMa, coxpatt.H10ru;a.H cB010 IJ;e"1ocT- HOCTh. 06pa30BaHHe BH)l:OB - Buaoo6pa3oeattue - pe3yJihT3T MHKp03BOJIIOIJ;HH aTo pa3,n;eJieHHe (so speMeHH JI npocTpaHcTse) npem,n;e e,n;1rnoro BH,n;a Ha ,n;sa JIJIH HeCKOJihKO. BH,n;oo6pa3oBaHHe npoHcxo.n;HT B pe3y.'IhTaTe nOCTO.HHHO COBeprna10ru;HXC.H BHyTpH Bll,ll;OB npor~eCCOB MHKP03BOJIIOIJ;HH. BbI,n;eJI.HlOT TPJI OCHOBHhIX nyTH JI ,n;sa cnoco6a BH,n;oo6pa3oBaHH.a. BoJihIIIOH BKJia,n; B JI3yqettHe BJI,n;oo6pa3oBaHJIH Bttec 3. Mailp. Ott BhI,n;eJIHJI 56): 1. <PuJtemu'leCICOe euaoo6pa306GHUe. BJI,n; A 11peBpaw;aeTC.H B xo,n;e 3BO.'Il0IJ;JIJI B BJI,n; B. qHCJIO BJI,n;os He H3MeH.aeTc.a. 2. I'u6puaozeHHOe euaoo6pa306GHUe. 3TOT llYTh CB.H3aH CO CJIJI.HHHeM ,ll;BYX cyw;ecTByrow;Hx BJI,n;OB AH B H o6pa3oBaHJieM BH,n;a C. TPJI OCHOBHbIX nyTJI, Be,n;yw;Jie K 110.HBJieHHIO HOBbIX BH,ll;OB (pile. llpuMep. KyJihTypHa.H CJIHBa - ecTecTBeHHhIH TeTpa11JioH,n;, B03HJIKIIIHH JI3 TepHa H aJihl'IJI B pe3yJihTaTe llOCJie,n;y10w;eil llOJIJif!JIOH,n;H3aIJ;JIJI llPH CJIJI.HHJIJI raMeT CJIJIBbl H Teptta. 3. /(uBepzeHmHOC euaoofipa.J06GHUe (paCXOJ/CaeHUe). lfa O,ll;HOrO BJI,n;a B pe- 3YJihTaTe o6pa3yIOTC.H ,n;Ba. 3TOT llYTh, 110-BJI,ll;HMOMy, HaJI60Jiee pac11poCTpaHeHHhIH. 3TOT llYTb 11epBbIM yKa3aJI q, ,n:apBJIH, Bhl,ll;BJIHYB quu: BJI,ll;hl CTpeM.HTCH K npUHL{Un aueepieH- MaKCJIMaJibHOMY 3allOJIHeHJilO MeCT o6HTaHHH H c11oco60B cyw;ecTBOBaHH.H. IlocTO.HHHOe o6pa30BaHJie OTJIJI'IHH H 11pHCllOco6JieHJIH K pa3HhIM MeCTaM o6HTa- HH.H COCTaBJIHeT OCHOBY 11pou;ecca ,D;HBepreHu;HH (pHc. c A B A 57). OcHOBHbte cnoco6bt BH,n;oo6pa3oBa- HHH 6a3HpylOTC.H Ha B03HHKHOBeHHH Pa3JIH'IHo:ll H30JIHIJ;HH BHYTPH BH,n;a. 1. CuMnampu'leCICOe euaoo6pa3o6llHUe (OT JiaT. sym - O.D;HHaKOBhlH, patria - po,n;HHa). BM,n;oo6pa30BaH.He llPOJICXO,ll;JIT B O,n;Horo apeaJia. lfaOJI.HIJ;HH JIOr.HqecKa.a, 3KO- qaw;e reHeT.H'lecKa.a (B03HHKHOBeHHe xpOMOCOMHbIX 11e- B A A P111c. 56. TpH OCHOBHblX nyrn, BeAYlllHe K nm1sneH1<110 HOBblX BHAOB. Cnesa Hanpaso <1mnernyecKOe BHAOo6pa3osaHHe, rn6PHAHOe npOHCXO~AeHHe BHA8, npoHC- pecTpoeK). llpuMep. A 11pe,n;eJiax Ilp.H B03HHKHOBeH.HH TeTpa11JioH,n;Horo pacTeH.HH ( 4n) XO~AeHHe BHAa ( pacxo~eHHfl) nyTeM AHBepreHL\HH
154 06U{UK 6UOJWZUK. J60JUOlfU.ll 7 P1tc. 57. A111sepreH1.1111si nepeAHVIX KOH84HOCTe~ no3BOH04Hb1x .>KVIBOTHblX: 1 - raHTCKoro 6poHeHOCL1a; 3- Pbl.>K8~ B8Y8PHVll.lbl; 4- roplMlllbl; 5- KPOTa; 6- <t>111Hsana; 2 - CVIByYa; 7- rn- llOW(!AVI np.>KeBaJlbCKOrD rH6pH,IJ;bI c ( Nicotiana) (2n) 6y,n;yT cTepm1bHhl. B po,n;e Ta6aKa Solanum) ttcxo,n;Hoe 'IHCJIO xpoMocoM paBHO 12, ,n;HrrJioH,n;HhlM H KapTo<PeJia ( 24, 48, 72 xpOMOCOMaMH. 2. A.nJlonampu'lec1eoe 6uaoo6pa3o6anue (oT JiaT. altos - HO eCTb BH,IJ;bl c pa3Hbiii:, patria - po,n;HHa). 3TOT crroco6 BH,n;oo6pa30BaHHH OCHOBaH Ha rrpocTpaHCTBeHHOH H30JIHD;Hli. HOBbie BH,IJ;hl MOryT B03HHKHYTh B peay.TihTaTe pacrra,n;eHHH IIIHpoKoro HCXO,IJ;Horo apeaJia Ha HeCKOJihKO ,n;pyrHX, MeHbIIIHX. Ma1tpoaBoJ11ou;ua Ma1epoaBOJl/OU,UR - 3TO aBOJIIOD;HOHHhle rrpou;eccb1 Ha Ha,n;BH,IJ;OBOM ypoBHe, IIPHBO,ll;HID;He K o6pa30BaHHIO TaKCOHOB Ha,n;BH,IJ;OBOro paHra. Ilpou;ecc MHKpoaBOJIIOD;HH rrpo,n;oJimaeTca 6ea KaKoro-JI1160 rrepephIBa H Ha MaKp03BOJIIOIJ;HOHHOM ypOBHe. lfaMeHaeTca TOJibKO B03MOiKHOCTh CKpeiu;1rnaHHH Mem,n;y BH,n;aMH. HoBhie BH,ll;hl MoryT BCTyIIaTh B MeiKBH,IJ;OBbie OTHOIIIeHHH, 'ITO BJIHHeT Ha <PaKTOphl 3BOJIIOD;HH H, B CBOIO O'!epe,n;h, ,n;eiiCTByeT Ha MHKp03BOJIIOIJ;HOHHhle rrpou;eccw. Co6wTHH MaKpo3BOJIIOIJ;HH oxBaThlBaIOT MHJIJIHOHhl JieT, H HX H3y'!eHHe OCHOBaHO Ha ,n;pyrHX MeTo,n;ax (rraJieOHTOJIOrHH, Mop<lJo<PH3HOJIOrHH, CTpoeHHH .D:HK H 6eJIKOB pa3JIH'!HbIX rpyrrrr).
155 OcnoBnbte nanpaBRenuJl 360RIOIJUU <PuJIO?ellCJ - 3TO IICTOpIIqecKoe pa3BIITIIe opraHII3MOB, KaK B u;eJIOM TaK- coHe, TaK II B OT,ll;eJibHhIX TaKCOHax. BhI,ll;eJI.lUOT qeTbipe OCHOBHhie cpopMbl <PIIJIOreHe3a: 1. <PuJiemu'leCICa.H 360JI,IOU,U.H - 3TO II3MeHeHII.H, ITpOIICXO,ll;.HIIJ;IIe B O,ll;HOM <PIIJIO- reHeTIIqecKOM CTBOJie (6e3 yqeTa B03MO./KHhlX ,ll;IIBepreHTHbIX OTBeTBJieHIIM). IlpuMep. Pa3BIITIIe ITpe.n;KoB JIOIIrn.n;eii ITO rrp.aMoi1: JIHHIIH: aorIIrrITyc - MHOrHrrITyc - rraparHrrITyc - 2. J1ueepienu,u.H - coBpeMeHHa.H JIOIIIa,ll;h. pacxom.n;eHIIe BeTBeii .n;peBa .JKH3HH OT e.n;irnoro CTBOJia ITpe.n;KOB. IIpuMep. Bo3HHKHOBeHIIe paaHoo6pa3HhIX ITO Mop<PoJiorIIqecKHM oco6eHHO- cT.HM BblOpKOB, HarrpHMep Ha faJiarraroCCKHX OCTpoBax, OT e.n;IIHOro rrpe.n;Ka. 3. KoHeepieHU,U.H - aTo rrpou;ecc qiopMnpoBaHH.H cxo.n;Horo <PeHOTHIIIIqe- cKoro o6JIHKa oco6ei1: .ll:BYX HJIII HeCKOJihKHX rpyrrrr. IIpuMepbl. <l>opMHpOBaHHe CXO,ll;HhIX <PopM TeJia y aKyJIOBbIX, HXTH03aB- poB H KIIT006pa3HhIX. Ilpe.n;cTaBHTeJIH pa3HbIX <PIIJIOreHeTIIqecKHX rpyrrrr HMelOT 60JihIIIOe CXO,ll;CTBO B CTpOeHHH B CB.H3H c IIPIICIIOC06JieHHeM K ./KII3HH B BO,ll;HOM cpe.n;e. Cxo.n;cTBO cTpoeHH.H rJia3 MJieKOIIIITaroiu;nx II roJioBoHornx MOJIJIIOCKOB. 4. IIapaJIJI,eJIUJM - qiopMHpoBaHIIe cxo.n;ttoro o6JIHKa H3HaqaJihHO .n;HBep- reHTHhIMH, HO reHeTIIqecKII 6JIH3KHMH rpyrrrraMH. IlapaJIJieJIH3M - 3TO KOHBepreHIJ;H.H Ha OCHOBe roMOJIOrHqHhIX opraHOB. IlpuMep. Pa3BHTHe ca6Jie3y6oCTH B CTBOJie KOIIIaqhIIX B03HHKaJia Heo.n:- HOKpaTHO. OcHOBHLie HanpaBJieHHH aBOJIIOIJ;HH KpyrrHeMIIIIIe poccniicKIIe SBOJIIOIJ;HOHIICThl A. H. 1. 2. 3. IL IL 58): CeBepu;oB H rayaeH Bbl,ll;eJIHJIH TPH rJiaBHbIX HarrpaBJieHH.H 3BOJIIOIJ;IIH (pHC. IllMaJih- AporeHe3 (apoMop<Po3). AJIJioreHe3 (n.n;noa.n;arrTau;nn). KaTareHe3 (o6iu;a.a .n:ereHepau;n.a). ApoieneJ - KpyrrHhle aBOJIIDU:HOHHhle H3MeHeHH.H, Be.n;yiu;ne K o6iu;eMy yc- JIO.JKHeHHIO CTpoeHH.H H <PYHKIJ;IIH opraHH3MOB, B pe3yJibTaTe qero 3aTpy.n;H.HeTC.H OCBOeHIIe HOBbIX ITpHpO,ll;HhIX 30H, IIO.!IBJieHHe KPYIIHbIX rrpHCIIOC06JieHHH. Apo.;nop<jJ03bl - KpyrrHwe a.n;aITTaU:IIH, rrpHBO,ll;HIIJ;He Ha rryTh aporeHe3a (Ha CXeMe BhIXO,ll; H3 O,ll;HOH IIJIOCKOCTH B .n;pyryro). Ilpu.Mepbl. IIOHBJieHHe JierKHX y Ha3eMHhIX II03BOHoqHbIX II03BOJIHJIO OCBO- RTb HOBYIO cpe.n;y o6IITaHHH, cyIIIy. B SBOJIIDU:HII MJieKOITHTaroiu;nx K apoMopll>o3aM MO./KHO OTHeCTH IIO.HBJieHHe IIIepcTHOro IIOKpoBa, ./KHBOpom.n;eHIIe, BCKapMJIHBaHHe .n;eTeHbIIIIeii MOJIOKOM. AJIJI,o?ene3 - pa3BHTHe rpyrrrrhr BHYTPH o.n;tto:H a.n;arrTHBHOM 30Hhl, a.n;arrTa- n;a:H o.n;Horo MaCIIITa6a. H iJuoaiJanmau,uu - a.n;arrTaU:HH o.n;Horo MaCIIITa6a rrpH ll>opMnpoBaHHH pa3Hhlx rpyrrrr B rrpe.n;eJiax o.n;Ho:H a.n:arrTHBHOH 30HhI (pacxo.n:aIIJ;aeca BeTBH Ha O,ll;HOH IIJIOCKOCTH).
06~aJl 6UO.!WZUJl. ;J60.!llOlfUJl 156 8peMR P111c. 58. CxeMa pa3B111rn5i rpynnb1 no nyrnM annoreHe3a BHyrp111 C1Aam111BHOH 30Hbl (Ha nnocKOcrn) 111 aporeHe3a - IlpuMepbt. c BbiXOAOM B HOBYIO CJAanTlllBHYIO 30HY (B HOBYIO nnOCKOCTb) IloaBJiemrn KphIJia - apoMopcpoa. llaMeHemfe ¢opMhI KphIJia B CBH3H c oco6eHHOCTHMH IIOJieTa IITHll Ka - H.!l;HOa,!l;aIITa1IHH. IlOHBJieHHe llBeT- KPYIIHhIM apoMopcpo3. ll3MeHeHHe B cpopMe u;BeTKa B CBH3H co c11oco60M OilbIJieHHH - H.!l;HOa,!l;aIITau;HH. KamaieHe.3 - aBOJIIOIJ;HOHHhrii: rrpou;ecc, IIPHBO.!l;HillHH K ynpo111eHHIO opra- Hlfaau;HH. 0611.{aJl iJeieHepaU,UJl - 11cqe3HOBeHHe CHCTeM opraHOB H CTpyKTyp, TO eCTb MOpcpocpH3HOJIOrHqecKaH ,!l;erpa,!J;au;HH. IlpuMepbt. Ilapa3HTJfqecKHe cpopMbI COCTaBJIHIOT 4-5 °Ai OT 06111ero q11cJia COBpeMeHHhIX BH.!l;OB :>KHBOTHhIX. Ha Kpa6ax rrapaaHTHpyeT paKoo6paattoe caKKYJIHHa. Bo B3pOCJIOM COCTOHHHH, HaXO.!l;HCb Ha TeJie Kpa6a, OHa HMeeT BH.!I; MeIIIKa, Ha6HTOro IIOJIOBhIMH npO.!J;YKTaMH, c OTXO.!l;HillHMH OT Hero OTpOCTKaMH. ll TOJlbKO H3yqeHHe JIHqHHO'lHblX CTa,!l;HH 1103BOJIHJIO BhlHCHHTb npHHa,!l;- Jie:>KHOCTb aToro BH,!l;a K paKoo6pa3HhIM. "'YTpaTa IIHilleBapHTeJibHOH CHCTeMhl y KJiacca JleHTOqHbie qepBH IlpOH30IIIJia B peayJibTaTe 11apa3HTHpOBaHHH B TOHKOM KHIIIe'lHHKe X03HHHa. BHoJiorH11ecKHH nporpecc H perpecc B aBomou;HH BuoJloiu'tecKuu npoipecc - aTo TaKoii: X0.!1; aBOJIIOI.J;I:IOHHoro npou;ecca, np11 KOTOpOM: IIOCTOHHHO HapacTaeT 'lHCJieHHOCTb oco6eli B IIOIIYJIHIJ;HHX; pacIIIHpHeTCH apeaJI; yBeJIH'lHBaeTCH pa3H006paa11e a,!l;anTaIJ;HH. B110Jior11qecKHH nporpecc opraHH3MOB. aTo ycnex 11 npou;BeTaHHe KaKoli-Jin6o rpynnhr
Ocno6Hbte no.1toJ1Ce1tuR cunmemu'iec1wii meopuu 360.!tlO!!UU 157 Han6o;:iee qacTo 6ttoJior11qecKHH uporpecc o6ecneqHBaeTcH 3a cqeT nOHBJieHMH apoMopcpo3oB. 0.n;HaKo cyr.u;ecTBYIOT rpyrrilhI, npeTepuemrme .n;ereHepaIJ;MIO CMCTeM H opraHOB, HO 3a cqeT 3TOro npttcrroco6HBIIIHCCH K HOBhIM YCJIOBHSiM M urnpOKO pacnpocTpaHMBIIIHeCH. BuoJtoiuttecKuu peipecc - aTo rrpo1~ecc upoTttBono.1omHoro HarrpaBJieHMSi. ,l(JIH Hero xapaKTepHO yMeHhIIIeHHe '-IHCJieHHOCTH oco6eil: B llOIIYJIHI~H­ S{X, COKpar.u;eHHe apeaJia, COKpar.u;eHHe qHCJia BH,ll;OB (BIIJIOTb ;:i:o 3JIHMHHaIJ;MM). BnoJiorttqecKHiI perpecc qacTo CB.H3aH c .n:ereHepautte:iI M Bcer.n;a rrpttBOJJ:HT K yua.n;Ky. B 3TOM 3aKJIIOqaeTcH oTJIHqtte 6ttoJIOI'H'-IecKoro perpecca OT o6u~eil: ,ZJ;ereHepauHH. BttoJiortt'-IeCKHH perpecc MOJKeT 6hITh He CBH3aH c .nereHepautteii:. HaupHMep, BhIMMpaHtte JJ:HH03aBpoB CBH3aHo c Herrpttcrroco6JieHHOCThIO K H3MeHHBIIIeHCH cpe.n;e, 6e3 .nereHepauHH KaKHX-JIH60 CHCTeM. OcnoBHbie rroJio~eHHH CHHTeTHqecKoii TeopHH 3BOJIIOD;HH 1. MaTepttaJI aBOJIIOIJ;HH - 2. 3. 4. 5. 6. 7. 8. HacJie,ncTBeHHhie H3MeHeHMH (MyTauHH). MyTauHOHHaH H3MeHqHBOCTh HOCHT c.'Jyqaii:HhIH H HeHarrpaBJieHHhIH xapaKTep. E,n;HHCTBeHHhIH ,nBHJKyr.u;ttii: cpaKTop - oT6op cJiyqaiIHh1x MyTaunii:, B03HHKaror.u;ttx B reHocpoH.n;e llOIIYJI.HuHH. HattMeHhIIIaH e.n;ttHHua 3BOJII011HH - uorryJIHliH.H. 3BOJll011HH HOCHT JJ:HBepreHTHhlll xapaKTep, TO eCTh OJJ:HH TaKCOH MOJKeT CTaTh upe,nKOM HeCKOJlhKHX ,n;oqepHHX TaKCOHOB. 06pa3oBaHHe HOBoro BH.n;a - aTarr aBOJIIOliHOHHoro upouecca. Btt,noo6pa3oBaHHe upottcxo.n;HT uocTerreHHO H ,nJIHTeJihHO. Btt,noo6pa3oBaHtte IlOCJie,ZJ;OBaTCJihHOCTh CMeHhl uorryJIHliMH. Btt.n; - peaJihHO cyr.u;ecTByIOr.u;aH e.n;HHMua. Btt.n; o6Jia,naeT .n:irnaMHqHocThIO (TOJihKO BHYTPH Bcer.n;a B03MOJKeH o6MeH aJIJieJIHMH). Btt.n; o6Jia,naeT ueJIOCTHOCThIO H 3aMKHYTOCThIO: cyr.u;eCTBYIOT MexaHH3Mhl rro.n;.n;epJKaHHH ueJIOCTHOCTH Ha YPOBHe 11011YJIH11HH (reorpacpH'-IeCKaH, 6HOJIOrWieCKa.H, reHeTttqecKaH H30JIH11HH) H Ha ypoBHe reHocpoH.n;a (rpyrrrrhI cuerrJieHHH, CHCTeMbl peuapauHH). Peupo,n;yKTHBHaH H30JIH11HSi KaK KPHTepttii: BH,n;a HerrpttMeHHMa K rpyurraM opraHH3MOB 6e3 IIOJIOBOI'O upo11ecca (HeKOTOphle rpH6hI), uapTeHOreHeTHqHhIM, arroMHKTHqHhIM cpopMaM. 3ToT KPHTeptti1: HerrpttMeHHM .ll:JIH BhIMepIIIHX rpyuu opraHH3MOB. ,l(JI.H HCKOrraeMhIX cpopM rrpHMeH.HIOT TepMHH «cppaTpHH». MaKpoaBOJIIOIJ;H.H, TO ecTh o6pa3oBaHJrn Ha.n;BH.ll:OBhIX TaKCOHOB, upoHcxo.n;HT Ha OCHOBe Tex JKe 3aKOHOMepHOCTeii:, qTO H MHKp03BOJIIOUl'IH. 9. Jlro6oii: peaJihHhIH TaKCOH HMeeT MOHOcpHJieTifqecKoe npoHCXOJK.n;eHHe. 10. 3BOJIIOuHH Henpe.n;cKa3yeMa, HMeeT HeHarrpaBJieJIHhIH xapaKTep K HeKOH KOHe'-IHOH ueJIH.
158 CaJILTau;HoHHaH TeopHH 3BOJIIOD;HH Ca.llbmau,uoH1taR meopuR B 1899 r. (OT JiaT. pyccKHM 6oTaHHKOM C. H. saltatio - paaoBaHHe u,11,eT aa cqeT pe,11,KHX cKaqKoB ,n;aM rrpHUieJI B HaqaJie cKaqoK) 6hrJia BhI,11,BHHYTa Kopm1rncKHM. Ee cyTh B TOM, qTo BH,11,006MyTa:u;ttii:. K aHaJioruqHhIM Barmi- xx B. roJIJiaH,11,CKHM 6oTaHHK fyro ,11,e <l>pH3, OCHOBOIIO- JIO)RHHK MyTaD;HOHHOM TeOpHH. ,ZJ;eii:cTBHTeJihHO, BH,ll,006pa30BaHHe y pacTeHHM MomeT c 6oJihUIOii: BepOHTHOCThIO rrpottcxo,11,HTh rryTeM IIOJIHIIJIOH,11,HH, 'ITO, 0,11,HaKo, He rrpoTHBopeqHT CHHTeTHqecKoii: TeopHH 3BOJIIOD;HH CT8, paccMaTpHBaroru;eii: BH,ll,006pa30BaHHe KaK CMeHy reHHbIX qacTOT II0,11, ,11,eii:CTBHeM OT6opa. B pH,11,e rpyrrrr opraHH3MOB ,11,0 90 % cJiyqaeB BH,11,006paaoBaHHH HaqttHaeTCH c xpOMOCOMHblX rrepecTpoeK. KoH:u;err:u;ttH CT8 H caJihTaD;HOHHaH TeopHH ,11,orroJIHHIOT ,11,pyr ,11,pyra. CoBpeMeHHaH TeopHH 3BOJIIOD;HH TeopHH 3BOJIIOD;HH Haxo,11,HTCH cero,11,HH B cTa,11,HH ,n:aJihHeii:Uiero pa3BHTHH H CTaHOBJieHHH. 1. CoBpeMeHHaH TeopHH 3BOJIIOD;HH OCHOBhIBaeTcH Ha rrpHHD;Hrrax, aaJio- meHHhIX reHirnJihHbIM 2. B Hell l.J. ,ZJ;apBHHOM. coxpaHHIOTCH OCHOBHhle IIOJIO)ReHHH CHHTeTHqecKOM TeopHH 3BO- JIIOD;HH. B TO me BpeMH COBpeMeHHaH TeOpHH 3BOJIIOD;HH IIpHHHMaeT BO BHHMaHHe Bee HOBbie ,n:aHHhle, OCHOBaHHble Ha COBpeMeHHbIX HCCJie,n:oBaHHHX, paccMaTpHBaeT pOJib rrepeHoca reHOB B rrpe,n:eJiaX O,ll,HOro IIOKOJieHHH (rryTeM TpaHccpopMa:u;1m, TpaHc,n;yKD;HH c yqacTtteM BttpycoB H peTpOBHpycoB), CHM6ttoreHeaa H ,n:pyrttx HBJieHHM.
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177 Ticanu U opzanbl. 6bl.CIUUX pacmenuii PMc. 63. BepxyweYHasi Mep111creMa no6era an0Ae111 : a npOAOnbHbl~ npoAonbH1:>1~ pa3pe3; 6- sHewH111~ BlllA 111 pa3pe3 KOHyca HapacTaHlll5l; B - KneTKlll nepBlllYHO~ Mep111CTeMbl; ,- - napeHXlllMH35l Kner- Ka n111cra, 33KOHYl'1BW35l Alll<l><l>epeHL\1'1POBKy. 1 - KOHYC HapaCT3H1115l ; 2 - 3aYaTOK n111cra IIpocmaR mKaHb COCTOHT H3 KJleTOK O,ll,HHaKOBhIX no cpopMe H cpymrn;mIM (napeHXHMa, CKJiepeHXHMa, KOJIJieHXHMa). CJlOJIC1WJI. mKUHb COCTOHT H3 KJ16TOK pa3JIH'lHhIX no cpopMe I1 cpyHKI.l,H.flM, HO HMeIOJ.I.\HX 061.l\ee npOHCXO>K,ZJ,eHJie (RCHJieMa). Teno BhIClllHX pacTeHHii o6paaoBaHo TKaH.flMH. JIHlIHhie opraHhI pacTeHHH. B Ifa HHX cpopMHPYIOTC.H pa3- aaBHCHMOCTH OT oc- HOBHOH lPYHKI.l,IU'I BhI,ll,eJI.HIOT OCHOBHhie BH,ll,hl TKaHe:ii: o6pa30BaTeJihHhie TKaHH (MepHcTeMhl); OCHOBHhle TKamr; IlOKPOBHhie TKaHH; BhI,ll,eJIHTeJibHbie TKaHH; MeXaHI.f'leC:KHe T:KaHH; npOBO,ll,.Hl.l\He TKaHIL 06pa30BameJtbllble mKanu ( MepucmeMbt) (pHc. 63-64). <l>yH:KI.l,HH: ,n;eJieHHe H ,ll,HlP!PepeHI.T,HpOBKa. XapaKTepHCTHKa: KJieTKH MeJIKirn, He,n;nipipepeHI!HpoBaHHhre, mHBhre, c TOHKOH KJieTOlIHOH 060JIO'IKOH; BaKyoJieH o6bllIHO HeT, '!aCTO ,n;eJI.HTC.11 MHTOTH'leCKH, cpopMHPYIOT Bee OCTaJibHhie 3 TKaHH pacTeHHH. llo BpeM.enu 8031tU1CflOBeHU.R BhI,ll,6JI.flIOT nepBHlI- Bhle n BTOPH'lHhie MepHCTeMhI. IIepBHlIHhre MepncKOTOpbie 3aKJia,ll,blBaIOTC.H PMc. 64. CxeMa pacnonolKe- B TeJie aapo,n;hmra. 8To 3apoJJ.hIIU ceMeHH, 30Hhl poc- Hlll5l pa3nlllYHblX Mep111CT6M B TeMh! - MepHCTeMbl, Ta CTe6JI.H B BblCOTY H 30Ha pocTa KOpH.H. BTopHlIHhle pacTeH111111: 1 - '5'.epHcTeMbl - (am1KanbHa5l) ; 3aKJia,ll,hIBaIOTC.H no3,n;Hee, o6ecneqH- sepxyweYHasi 2 - 111HrepKa- 8aloT POCT CTe6m1 B TOJIJ.I.\HHY (KaM6Hii cocy,n;HCTO- nsipHasi (BCT3BOYH3Sl), 3 - BOJIOKHHCThIX IlY'lKOB H npo6KOBblH KaM6:nii). KOsasi(naTepanbHasi) 60-
BommtuKa 178 II o pacnoJLOJlceHuJO BhI,D;e"i:moT: BepxymeqHyIO (amrnaJibHYIO) MepHCTeMy - B 30He pocTa KOpm'l H KOHyc HapacTamu1 rro6era; 60KOBhie (KaM6Hii:, rrepm.i;HKJI) - o6ecrreqnBaIOT pOCT CTe6Jieii: M KOpHeii: B TOJIIIJ;MHy; BCTaBOqHhie - OCTaTOK BepxyrueqHoii: MepMCTeMhI; pacrrOJIOiKeHbI B OC- HOBaHMH :vrem.n;oy3JIHll CTe6JIH l1 MOJIO,D;hIX JIHCTheB H 06ecrreq11BaIOT pOCT Mem.noy3J1Mll, o6pa30BaHHe rro6eroB, JIMCTbeB, rrpn.n;aTOqHhIX KOpHeii:; paHeBhie - o6pa3yIOT 3aIIJ;HTHblll CJIOii: (rrpo6Ky) Ha.n; MeCTOM IIOBpem.n;e- HMH TKaHeii. 0cH06HaR m1calib ( napeHxu.111a) XapaKTepMCTMKa: KJieTKM KPYIIHhie, HerrpaBMJihHOii: cpopMhr, Mem.n;y HHMM 6oJihlllMe MemKJieTHMKM. Ilo BhlIIOJl:HHeMhIM <}:>yHKIJ;MHM BhI,ZJ;eJIHIOT HeCKOJihKO TMIIOB rrapeHXMMhI: a) aCCMMMJIHIJ;HOHHaH rrapeHXHMa: OCHOBHaH cpyHKIJ;HH - cpOTOCHHTe3; co- .n;epmHT XJIOpOIIJiaCThI; pacrroJiomeHa B JIMCThHX H MOJIO,ll;hIX CTe6JIHX; 6) 3arracaIOrri;aH: HaKorrJieHHe 3anacHhIX Berri;ecTB (6eJIKH, mnphr, KpaxMaJI). IlpHCYTCTByeT BO Bcex opraHax pacTeHMH, B OCHOBHOM B CTe6JIHX, KOpHHX, KOpHeIIJIO,ZJ;ax, KJ1y6HHX, JIYKOBMri;ax, IIJIO.n;ax - B JIMCThHX MeHhllle. KJieTKH KPYIIHhie; crreri;HaJIH3MpoBaHHaH 3anacaIOrri;aa rrapeHXMMa ceMeHM Ha3hrnaeTCH ::rn.n;ocrrepMoM; B) B03,ll;YXOHOCHaH: OCHOBHaH cpyHKIJ;HH - BeHTHJIHIJ;MH; Mem.n;y KJieTKaMM oqeHh 60JihlllMe Me)KKJieTHMKM, CBH3aHHhie c BHelllHeii: cpe.n;oii: qepe3 OTBepCTMH yCThHIJ; H qeqeBMqeK; r) BO,ll;OHOCHaH: crroco6cTBYeT y.n;epmaHMIO BJiarH pacTeHMHMH; KJieTKH co- .n;epmaT B BaKyOJIHX C.HM3HCTbie Berri;ecTBa, y.n;epmHBaIOrri;Me Bo.n;y; xoporuo pa3BMTa y BO,ll;HhIX pacTeHHll My CYKKyJieHTOB (KaKTYChI, aJI03, araBhI); .n;) BCaCbIBaIOrri;aH (3IIH6JieMa, Ha3hIBaeMaH ·raKme PH30,ZJ;epMOll, MomeT 6bITh OTHeCeHa K rrepBHqHoii: IIOKPOBHOll TKaHH KOPHH): KJieTKH 3IIM6JieMhI HecyT KOpHeBbie BOJIOCKM, c IIOMOIIJ;hIO KOTOpbIX npoHCXO,ll;HT rrorJiorri;eHHe MHHepaJibHhIX Berri;ecTB l1 BO,ZJ;hl H3 rroqBhI; o6pa3yeTCH B 30He BCaCbIBaHHH KOpHeii:. II OKPOB!ibte mKallu BwrroJIHHIOT 3a11i;ttTHyIO cpyHKIJ;HIO. 3arri;Hrri;aroT OT BhIChIXaHMH, rreperpeBa, rroe.n;aHMH mHBOTHhIMM, 3apameHMH 6aKTepMHMH H rpH6aMH. Bo3HHKJIH c BhIxo.n;oM pacTeHHii Ha cyllly (HaqMHaH c rrcMJIOcpMTOB B rraJieo3oe). Pa3JIHqaroT: rrepBHqHyro rroKpoBHYIO TKaHh BTOpHqHyIO IIOKPOBHYIO TKaHb TpeTHqHyIO IIOKPOBHYIO TKaHb - :JnuiJepMa ( KOJICUU,a ): :muiJep.My; nepuiJep.My; KOpKy, MJIM pumuiJO.M. KJieTKH mMBhie, IIJIOTHO npMJieraIOT .n;pyr K .n;pyry, JIHUieHbI XJIOpOIIJiaCTOB, IIOKpbIThI KYTHKyJioii:; CJIOmHaH TKaHb COCTOHT M3: 3IlH,ZJ;epMaJibHhIX KJieTOK (y ,ll;BY,ZJ;OJihHhIX Hhie, pa.n;aMH); - eCTb Jie:HKOIIJiaCThI, HeT XJioponJiaCTOB M3BHJIHCTOH cpopMhI, y O,ll;HO,ll;OJihHhIX - rrpHMoyroJih-
TKanu u opzanbL 6bLCuiux pacmenuii 179 3aMb1Ka10IIJ;l1X KJieTOK yCTbHD; - HMelOT 6o60BH,z:t;HYIO cpopMy, Mem.n;y HH- MH yCTbH'IHa.H w;eJih; OKOJIOYCTbH'IHbIX ( rro60'IHbIX) KJieTOK; Tpl1XOM - pa3JIH'IHbIX ITO cpopMe, CTpOeHl1IO l1 cpyHKD;l1.HM BblpOCTOB KJieTOK 3ITl1,z:t;epMbl (BOJIOCKl1, Hble y ceM.HH XJIOII'laTHl1Ka - Ycmbuu,e - u~eTl1HKH, 'leIIIyHKH; Ha1160Jiee ,z:t;JIHH- 5-6 CM). 3TO MemKJieTHHK Mem,n;y ,z:t;BYMH 3aMb1Ka10w;11M11 KJieTKaM11 6o- 6oBl1,z:t;HOM cpopMbl. lIMelOTC.H B OCHOBHOM Ha Jil1CTb.HX, eCTb Ha CTe6Jie. 06bI'IHO paCIIOJI0'1<eHbl Ha HHJKHeH CTopoHe Jll1CTa, y BO,z:t;HbIX pacTeHl1H - IIepuiJepMa - Ha BepxHeii:. cJiomnaH noKpOBHa.H TKaHh CTe6Jieft, KOpHeB11w; l1 KOpHeft NIHoroJieTHl1X pacTeH11ii (peme - o,n;HoJieTH11x). CMeHHeT an11,n;epMy. Ilep11,n;ep- Ma 06pa3yeTcH 113 Mep11cTeMbI - cpeJIJioreHa. <l>eJIJioreH 3aKJia,z:t;hrnaeTc.H B oc- HOBHOH rrapeHx11Me no,n; an11,n;epMoii:. KJieTKl1 cpeJIJioreHa ,n;eJI.HTc.a: Hapymy OTKJia,ZJ;bIBaIOT KJieTKl1 npo6KH (MepTBbie, nponnTaHHhie cy6epl1HOM (T11na BOCKa), a BHYTPh - :>KHBbie napenx11MHbie KJieTKl1 cpeJIJio,n;epMbI c xJioponJiacTaMH. TaKHM o6pa30M, nepH,n;epMa cocTOl1T 113 cpeJIJiorena, npo6KH (cpeJIJieMbI) H <t>eJIJIO,z:t;epMbl. llpo6Ka - COCTOHT H3 MepTBbIX KJieTOK, KJieTO'IHa.H CTeHKa npon11TaHa JKH- pono,n;o6HblM cy6ep11HOM. KJieTKl1 HMeIOT npHMoyroJihHYIO <t>opMy, TIJIOTHO npHJieraIOT .n;pyr K ,n;pyry, pacnoJiomeHhI p.a,n;aM11. Ilpo6Ka coxpan.HeT TKann OT IlOTepH BJiarH, npOHl1KHOBeHl1.H Ml1KpoopraHH3MOB, o6pa3y.H «cpyTJI.Hp». ,lJ;JIH raaoo6Mena no,n; yCTbHD;eM Ha MeCTe B03,z:t;YXOHOCHOH IlOJIOCTH <t>eJIJroreH OTKJia,ZJ;bIBaeT iRl1Bble, pbIXJIO pacnoJiomeHHbie napeHXHMHbie KJieTKH (BbIIlOJIH.HIOIIJ;a.H TKaHb), KOTOpbie pa3pb1BalOT 3IlH,z:t;epMy H C03,z:t;aIOT B03MO:>KHOCTb ra3006MeHa H TpaHCilHpau;n11 c BHeIIIHeM cpe,n;oii:. 8To 06paa0Ban11e Ha3bIBaeTC.H 'te'le6U'lKOU. 0'1eHb KpynHble 'le'leBl1'1Kl1 (.n;o 15 CM) o6paayIOTC.H y 6epe3bl (no- nepe'IHbie IlOJIOCKH l1 qepTO'IKH). KopKa (pumuiJoM) - TpeTH'IHa.H noKpOBHa.H TKaHh. 06paayeTCH y Mnoro- JieTHHX pacTeHHM B KOpHe, CTe6Jie, KOpHeBnw;e. Kam,z:t;hIH ro,n; cpopMHPYIOTC.H 6onee rJiy6oKne CJIOH nepn.n;epM, napymHhie cJIOH H pacnoJio:JKeHHhie Mem,n;y HHMH TKaHH OTMHpaIOT. KopKa o6hI'IHO Tpew;HHOBaTa.H H HepOBHa.H, B OTJIH'IHe OT rJia,n;Koii: nep11,n;epMbI. KopKa aaw;Hw;aeT OT MexaHH'leCKHX noBpem,n;eHHH, pe3KOH CMeHbl TeMnepaTyp, npOHHKHOBeHH.H MHKpoopraHH3MOB. BbtdeJtumeJtbHbte mKanu CJiymaT ,ZJ;JIH naKonJieHHH H BbI,ZJ;eJieHHH Bew;ecTB H3 opraHH3Ma pacTennii:. Ilpe,n;cTaBJieHbI TKaH.HMH napymnoii: H BHyTpenneii: ceKpeu;nn. KJieTKH BhI,z:t;eJIHTeJibHhIX TKaneii: no cpopMe o6bI'IHO napeHXHMHhie, TOHKOCTeHHbie. 0HH .ll;OJiro OCTalOTC.H :>KHBblMH, BbI,z:t;eJI.H.H ceKpeT. TKann llHHMH: Bnympenneu ceKpeu,uu KJieTKH-H,z:t;H06JiaCTbl - npe,n;cTaBJieHhI paaJIH'IHhIMH o6paaoBa- OT,z:t;eJihHbie KJieTKH, co,n;epmaw;ne KpHCTaJIJihl OKCaJiaTa KaJib~H.H, TepneHOH,z:t;bl, CJIH3H; BMecTHJIHw;a BbI,z:t;eJieHHM - 302eHHbte 6.MecmuJtuw,a, IlOJIOCTH B TOJiw;e ,n;pyrnx TKaneii:. lIMeIOTC.H cxu- KOTOPbie B03HHKaIOT B BH,z:t;e MemKJieTHHKOB, 3a cqeT
Eomanu1w 180 pacxom,n;ettiui JKHBbIX 1rneTOK. CeKpeT KJreTOK BhI,D;eJrneTca n IIOJIOCTh MemKJJeTmma (3qmpttbre :-.rncJia, CMOJ!bI, CJIH3b) H JIU3Ucel-tltbte 6MeCmUJIUU{G, B03mmaIOilIHe B pe3yJihTaTe pacrra,lla (JIH3nca) KJieTm< nocJie HaKoIIJiemrn ceKpeTa B MeJKKJ":leTHHKe (HaIIpHMep, :Jq:mpHhie MacJia y !IHTPYCOBblX); CMOJIHHbie XO,llhI - o6pa3yIOTCfl CXH3ore1rno, H3HYTPH BhICTJiaHhI ceKpe- THPYIOI!IHMH KJieTKaMH. MoryT BeTBHThca, co,n;epmaT CMoJiy; MJieqHHKH - c11cTeMa noJiocTeii, npoHH3hIBaIOilIHX see pacTeHne. Co- ,n;epmaT M.rreqtthrii coK - 3MYJihCHIO 6eJioro, peme - opattmesoro !IBeTa (qncTOTeJI), co,n;epmailIYIO CMeCh pa3JIHqHbIX BeilieCTB (HaIIpHMep, KayqyKOHOChI, '!eBeJI, KOK-carbr3). Bhr,n;eJIHTeJihHhie TKaHH e11eurneu ceKpeu,uu (csaaaHhI c noKposaMn pacTemrn): meJie3HCTbie BOJIOCKH (ttanpnMep, repatth :m:eJie3KH - o,n;Ha HJIH HeCKOJihKO ceKpeT11pyIOilIHX KJieTOK sqmpHhre MacJia, coJieBhr,n;eJreHtte); MHoroKJieToqttaa ceKpenrpyrou,a.H CTPYKTypa, Bhr,n;emuo- IIIaa sc}mptthre MacJia; HeKTapHHKH - pa3Hoo6pa3Hhie CTPYKTYPhI, BhI,n;eJIHIOilIHe HeKTap: ca- xapncThIM COK (paCTBOp caxapos, 6eJIKOB, CIIHpTOB, apoMaTHqeCKHX BeilieCTB). PacIIOJIOJKeHbI B OCHOBHOM B !IBeTKax (Ha qarneJIHCTHKax, JieneCTKaX, CTeHKax 3aBH3H, !IBeTOJIOJKe). Mexa11u•tec1Cue mKaHu 06ecne'lHBaIOT onopttyro cpyHK!IHIO, npM,n;a10T npoqttoCTh opraHaM pacTeHHii (Hatt6oJiee pa3BHThI B cTe6Jie). KJieTKH HMeIOT attaqMTeJibHO yToJIilieHHhie KJieToqHbie o6oJioqKH. Pa3JIHqa10T ,n;sa THna Mexatt1111eCKHX TKaHeii: 1. KoJt.1ie11xuMa - TKaHb, KJieTKH KOTopoii HMeIOT ttepasHOMepHo yTo.,1- ilieHHhie KJieToqHhre cTeHKH. KJieTKH BhITHHYThI s,n;oJih OCH opraHa; KJieTKH JKHBbre, co,n;ep:m:aT XJioporrnacThr. B no,n;aeMHhIX oprattax He BCTpeqaroTcH; KJieT011HaH CTCHKa COCTOHT H3 Ile.,1JIIOJI03hl, reMH!IeJIJIIOJI03hl, neKTHHOBbIX seilieCTB. B 3aBHCHMOCTH OT THila YTOJiilieHHH KJieTOqHblX CTeHOK BbI,D;eJIHIOT ,n;sa THna KOJIJieHXHMbl: a) yroJIKOBa.H KOJIJieHXHMa - KJieTKH HMeIOT cpopMy rneCTHyrOJibHOro MHO- rorpaHHHKa. YTOJiilieHMe o6oJIO'lKH rrpottcxo,n;MT c «yroJIKOB» KJieTKH. BcTpeqaeTca B cTe6Jiax ,n;sy,zJ;OJihHhIX pacTeHMii, B qepernKax JIHCTbeB no o6eHM cTopoHaM KPYITHhIX JKHJIOK JIHCTa; 6) IIJiaCTHH'laTaH KOJIJieHXHMa - KJieTKH HMeIOT BH,D; rrapaJIJieJieIIHITe,n;a, YTOJiilieHHyro KJieToqHyro o6oJioqKy ,n;syx rpaHeii. Haxo,n;MTCH B cTe6JIHX ,n;peseCHblX pacTeHHM. 2. CKJtepeHXUMa - BeCHeBillHMH H MexaHHqecKaH TKaHh, COCTOHiliaH H3 KJieTOK c o,n;pe- paBHOMepHo CHJihHO yTOJiilieHHhIMH 060JI011KaMH. KJieTKH MepTBhie, o6oJIO'lKH o,n;pesecHeBaIOT, T. e. npOIIHThIBaIOTCH JIHrHHHOM. BbI,D;eJIHIOT OCHOBHbie BH,D;bl CKJiepeHXHMbI: CKJtepeHXUMHbte 60JlOKHa - KJieTKH BblTHHYTOM ¢opMbI c 3aOCTpeHHbl- MH KOH!IaMH, HMeIOT rropOBbie KaHaJibl B KJieToqHblX CTeHKax. CK.,'IepeH-
TKanu u opzanbt 6bLCmux pacmenuii 181 XlIMHbie BOJIOKHa B ,npeBeCHHe (KCHJieMe) Ha3h!BaIOTCJ! .npeBeCHHHbIMll BOJIOKHaMH (Jrn6pmpopM). CKJiepeHXHMHhie BOJIOKHa B Jiy6e (cpJIOaMe) Ha3b!BaIOTCJ! Jiy6J!HhIMlI BOJIOKHaMH (KaM6llcpOpM). Jiy6J!Hbie BOJIOKHa MOryT 6hITh ll Heo.n;peBecHeBIIIllMll, npo'IHhIMlI ll 3JiaCTlI'tJHhIMlI (HanpllMep, BO.'lOKHa JlhHa wcnOJib3YIOTCJ! B TeKCTlIJlhHOH npOMhIIUJieHHOCTH); CK.'lepellp;bI - B03HHKaIOT lI3 KJieTOK OCHOBHOH napeHXllMbI B pe3yJihTa- Te yToJiu~eHHJ! H JIHrHHcpHKan;HH KJieTO'tJHhIX cTeHoK. KJJeTKH HMeIOT pa3JilI'tJHYIO cpopMy ll BCTpe'laIOTCJ! BO MHOrHX 'laCTJ!X pacTeHHH: CIC.nepeu(Jbt (KaMeHHCThie KJieTKll) - 6paxu- B nJio.n:ax rpyurn:; OCTeOCKJiepe11,nhI HMeIOT pacnmpeHHJ! Ha o6mrx KOHn;ax (BCTpe'laIOTCJ! B JllICThJ!X 'laJI); acTpOCKJiepeH,[l;hI ( cpopMa 3Be3,[l;hl) - B JllICThJ!X KaMeJIHlI ll T • .n. IlpoBOO.RUJ,Ue m/CaHU BhlnOJIHJ!IOT cpyHKIT;HIO npoBe,neHHJ! BO,[l;hI H nnTaTeJihHhIX Bern;ecTB B TeJie pacTeHHH. 06ecne'tJHBaIOT BOCXO,[l;J!rn;Hii ll HHCXO,[l;J!rn;Hii TOK pacTeHHJ!. BocxoiJ.RUJ,UU mo/C - 3TO TOK MHHepaJihHhIX COJieH, paCTBOpeHHhIX B Bo,ne, H,ZJ;yrn;nx OT KOpHeH no CTe6JIIO K .'IlICThJ!M. Bocxo,11;J1rn;Hii TOK ocyrn;ecTBJIJ!eTCJ! no cocyp;aM H Tpaxenp;aM KCHJieMhI (p;peBeCHHhI). HucxoiJRw,uu mo/C - aTo TOK opraHH'leCKHX Bern;ecTB, HanpaBJieHHhIH OT JIHCTheB K KOPIIJ!M no CHTOBH,[l;HhIM aJieMeHTaM cpJIOoMhI (Jiy6a). Kcu.ne.11ta {jJJt03Ma - II <JTO CJIOiKHhie TKaHH, COCTOJ!rn;He H3 Tpex OCHOBHhIX aJieMeHTOB. IlpoBO,ZJ;J!rn;He aJieMeHThI TpaxeH,nhI - /CCUJleMbl (pHC. 65): Han6oJiee p;peBHHe aJieMeHThI. KJieTKH MepTBhie, BhITJIHyToii cpopMbI c 3aOCTpeHHh!Mll KOHn;aMH; HMeIOT op;peBeCHeBllIYIO KJieTO'tJHYIO CTeHKy c pa3Jill'tJHOH CTeneHbIO YTOJirn;eHHJ!; HMeIOT nophI; cocy,ZJ;hl ,ZJ;eJibHhlX nOJihie Tpy6KH, COCTOJ!rn;He H3 OT- 'tJJieHHKOB. .HBJIJ!IOTCJ! OTMeplllllMll KJieTKaMH, pacnOJIOiKeHHhIMH ,npyr Ha,ZJ; p;pyroM; HMeIOT nopbl ll BTOpH'IHhie YTOJirn;eHHJ!. IlpOBO,ZJ;J!IlOie aJieMeHTbl (jJ.n03Mbt: CHTOBH,[l;Uhle Tpy6Kll - KJieTKll iKHBhie, pac- nOJIOiKeHHhJe ,npyr Hap; p;pyroM (OTCYTCTBYIOT J!,[l;pa H BaKyOJih). KottTaKTnpy10T Memp;y co6oii: qepe3 nonepe'tJHhie neperopo,nKH c nopaMH - CHTO- BH,ZJ;Hhie IIJiaCTHHKH; ltJieTKH-CIIYTHHIT;hI - conyTCTBYIOT Kam,noMy lfJieanKy CHTOBH,ntto:H Tpy6KH. KJieTKH iKHBhle, BhIIlOJIHJ!IOT BCilOMoraTeJibHYIO pOJih B TpaHcnopTe opraHH'leCKHX Bern;eCTB. CocyiJucmo-BoJto/CHUCmbte ( npoeoiJ.Rw,ue) nyl{,H:u npoxo,11;J1T qepe3 Bee opraHhI pacTettn:H, o6pa3YI<>T e.n1rny10 npoBO,ll;J1rn;y10 CHCTeMy. CocTOHT H3 npoBO,lJ;arn;e:H ll MeXaHH'leCKOH TKaHH. IlyqKH 6hIBaloT OTKpbITbIMH H 3aKpbIThIMH. PMC. 65. Tpaxell1Abl KCWleMbl (nop111crb1e)
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Bezemamu6nbte opzanbt (1wpenb, cme6e1£b, 1£ucmbK) Ko1w,eumpu'lecKue (u;eHTpOKCHJieMHhre - 183 KCHJieMa B u;eHTpe, cpJioaMa BOKpyr: KOpHeBHill,a nanopOTHHKOB; u;eHTp0cpJI03MHbie - cpJIOaMa B ll,eH- Tpe, KCMJieMa BOKpyr: KOpHemun,a O,ll;HO,D;OJihHhIX). PaiJua.rtbtlbte - KCHJieMa pacxo,n;nTC.H Jiy'!aMH OT u;eHTpa, a cpJIOaMa pac- noJiaraeTC.H Me.m,n;y JiyqaMH. qHCJIO Jiyqeif OT o,n;Horo ,n;o TI.HTM 11 6oJree. XapaKTepHo ,n;Jia nepBwrnoro cTpoeHM.H MOJIO,D;hrx KOpHeM: roJioceMeHHhIX ll TIOKpbITOCeMeHHhIX. TKaHM o6pa3yIOT opraHhI pacTeHH.H. Op2a11 - aTo qacTh pacTeHH.H, cocTomn,a.H H3 TKaHeM: n BhITIOJIH.HIOill,a.H cne- u;naJihHYIO cpyHKll,HIO. OpraHhI pacTeHH.H ,n;eJI.HTC.H Ha BereTaTHBHhre n reHepaTMBHbre. Be2emamu6Hbte op2a11bt - KopeHh, cTe6eJih, JIHCT. I'enepamuettbte opiattbt: u;BeTOK, ceM.H, nJio,n;. B3aHMOCB.H3h Me.m,n;y opraHaMM npoHCXO,D;HT Ha OCHOBaHMH npou;ecCOB IlHTaHM.H 11 ,ll;bIXaHl1.H. BereTaTHBHLie opraHLI (1mpeHL, cTe6eJIL, JIHCTLH) 3TH opraHhI ocyrn,eCTBJI.HIOT npou;eCChI IlHTaHH.H (cpOTOCHHTe3 ll MHHepaJibHOe TIHTaHne), ,ll;hIXaHH.H H ra3006MeHa, a TaK.me TpaucnopT IlHTaTeJibHbIX Bem;ecTB 11 BereTaTHBHoe pa3MHomeune. BnepBhre B npon;ecce aBOJIIOUHH opraHhI IIO.HBJI.HIOTC.H y BhICIIIHX pacTeHl1M. Kopettb - aTo oceBoii oprau, HMeIOrn,Hii ll,HJIHH,n;pnqecKyIO cpopMy H o6Jia,n;aIO- m;nii pa,n;HaJihHOH CHMMeTpneii:. Cnoco6eu K poc- Ty, TIOKa coxpaH.HeTC.H anHKaJibHa.H (BepXYIIIe'IHa.H) MepHCTeMa. MopcpoJiornqecKoe OTJIH'!He OT no6era B TOM, 'ITO Ha KOpHe HHKOr,n;a He B03HHKaIOT JIHCTh.H; annKaJibHa.H MepHCTeMa npHKpbITa KOpHeBbIM qexJIOM. <l>yHKll,HH KOpH.H: MHHepaJihHOe H BO,ll;HOe IlHTaHHe - BCaChIBaHHe BO,ll;bl H MHHepaJib- HbIX Bern,eCTB; 3aI\penJieHHe pacTeHH.H B IIO'!Be - «3a.HKOpHBaHHe »; CHHTe3 opraHH'!eCKHX Bern,eCTB, ropMOHOB; HaKOIIJieHHe 3arraCHbIX Bern,eCTB; BereTaTHBHOe pa3MHOmeHne; CHM6no3 c 6aKTepH.HMH-a30TcpHKCaTopaMH (KJIY6eHhKOBhle 6aKTepnn); CHM6H03 c rpn6aMH ( MHKOpH3a). CnM6Ho3 c 6aKTepH.HMH-a3oT- <l>HKcaTopaMH (pnc. 67). Ha KopH.HX 6o- 6oBhlx H p.H,n;a ,n;pyrnx pacTeHHH o6pa3yIOTC.H KJiy6eHbKH, B KOTOpbIX noceJI.HIOTC.H 6aKTepHH Po,n;a Pn306nyM (Rhizobium), HJIH n;nauo6aKTepHn. 3TH MHKpoopraHH3MbI ycBaHBaIOT PMC. 67. Kny6eHbK~ Ha KOpHS1X
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AnamoMU'tec1we cmpoenue 1wpna 187 l\:CIIJieMa H cpJI03Ma pacnOJIO)IWHbl qepe.n;yIOIU;HMHCH JiylJaMH H3 u;eHTpa. 06pa3yeTCH pa.n;HaJihHhIH npoBo.n;Hru;tt:H nytJoK. B -oT.'rnqHe OT CTe6.11H Kop1rn He nMeIOT cep.n:I1eBHHhI. y O,D;HO,ll;OJlhHhIX pacTeHHH CTpoeHHe KOpmI OCTaeTCH 6e3 3Ha'-IHTeJibHh!X I13- MeHeHHH B TelJeHHe Bce:H )KH3HH pacTeHHH, y .n;By,ll;OJlhHhlX npOHCXO,ll;HT nepexo.n: K BTOPHlJHOMY CTpOeHHIO. Bmopuiuwe cmpoeuue IWpliH y a6yaoJlbliblX pacmeuuu. Pa.n;Ha.'lbHOe pacuo- JIO)KeHHe npOBO,ll;HIU;HX TKaHe:H CMeHHeTCH KOJIJiaTepaJibHbIM. 3To CBH3aHo c ,n;e.aTeJihHOCThlO KaM6H.H, KOTOpbIH B03HHKaeT Mem.n;y JiylJaMH KCHJieMhl n cpJI03MhI. 06pa3yeTCH e.n;HHhr:H CJIOH KaM6HH. R u;ettTpy KaM6HH: OTKJia.n;hrnaeT KJieTKH BTOpHtIHoii KCHJieMhI, a K rrepttcpepHH - BTOpHtIHo:ii cpJI03MhI. Kpo- Me Toro, KJieTKH KaM6HH o6pa3yIOT IIIHpOKHe pa.n;HaJibHhie CBeTJihie JiyqH napeHXHMhI Mem.n;y T.HfKaMH BTOPHlJHOH rrpOBO,D;.HIU;e:H TKaHH - cepou,eBUlillble JlY'lU. 3TO rrepBHlJHbie 0HH o6ecrrelJHBaIOT cpH3HOJIOrHlJeCKyIO CBH3h ueHTpaJih- HOH qacTH KOpH.a c rrepBwrno:H Kopo:H. IIepBHlJHaH Kopa OTMHpaeT n c6pachrnaeTC.H. IloKpOBHOH TKaHhIO cTaHOBHTC.H rreptt.n;epMa. RoMrrJieKc TKatte:ii cHapy- BmopU'lliOU KOpoii. MunepaJLbHoe numauue. Bo.n;y H MHHepaJihHhre jlCH OT KaM6lrn Ha3hIBaIOT Beru;ecTBa KopeHh Bcachrna- eT H3 IIOlJBhI c IIOMOID;hlO KOpHeBhIX BOJIOCKOB. Bo.n;a IIOCTyrraeT B KOpHeBOH BOJIOCOK 3a ClJeT OCMOCa. 3aTeM Bo.n;a npoxO,D;HT no )KHBhlM KJieTKaM rrepBHlJHOH IWphl KOpH.H H rrorra.n;aeT B KCHJieMy u;eHTpaJihHOro oceBoro IIHJIHH,ll;pa KOPHH. MHHepaJihHhre Beru;ecTBa BcachrnaroTc.a B xo.n:e rraccHBHoro HJIH aKTHBHoro TpaHcrropTa qepe3 KJieTOlJHYIO MeM6paHy. B cocy.n;ax pa3BHBaeTc.a rroBhIIIIeHHoe OCMOTHlJeCKOe .n;aBJieHHe. KopHeBoe .n;aBJieHHe H TpaHCilHpau;HH (HcrrapeHHe) o6ecrrelJHBaIOT ,D;BH)KeHHe BO,ll;hl B TeJie pacTeHHH. BamHyIO pOJih HrparoT KaIIHJIJI.HpHhie CHJlhI: MeJK.n;y MOJieKyJiaMH BO,ll;hl H ai)ze3UJl K02e3UH - CHJia CI1errJieHH.H BO,ll;hl K CTeHKaM COCy,ll;OB • .D:JI.a IIHTamrn pacTeHHIO Heo6xo.n;ttMhI: MaKpo;:iJieMeHThI: a30T, cepa, cpoccpop, KaJitt:H, MarHHii, KaJihll;HH, meJie3o; MHKp03JieMeHThI: 6op, Me,ll;h, MapraHeII, K06aJihT, IIHHK H .n;p • .D:JI.a YJIYlJIIIemrn ypoma.a BHOCHT y.n;o6peHH.H. Y,n;o6peHHH pa3.n;cJI.HIOT Ha opraHHlJeCKHe (Topcp, HaB03 H .n;p.) H MHHepaJihHhie (ceJIHTpa, MOlJeBHHa H .n;p.). BH.n;OH3MeHeHH.H KopnenJLOObl - KOpHe:H: BhIIIOJIHHIOT 3arracaroru;yIO cpyHKIIHIO. Bo3HHKaIOT KaK YTOJIIU;eHHe rJiaBHOro KOpH.H (MOPKOBh, CBeKJia, perm H .n;p.). KopneKJLy6nu - 3arracaroru;tte rrptt.n;aTOlJHhie KOpHH (6aTaT, reoprHH). BmHzuewow,ue Kop1w - ocyru;ecTBJI.HIOT norpyJKeHHe B rrotJBY (JiyKoBHqHhle pacTeHH.H). Bo30YUllible KOpllU - BCaChIBaIOT Bo,n;y H MHHepaJihHbie Beru;ecTBa H3 BJiaJKHOro B03,n;yxa (y pacTeHHH-3IIHcpHTOB - opxH.n;en). j(blXameJlbllbte 1wpnu IIO,D;HHMaIOTCH Ha.n; IIOBepxHOCThlO IIOlJBhI (y pacTeHHH 60JIOT). XooyJLbHbte Kopnu - y MaHrpoBhIX pacTeHnii Ha JIHTopaJIH Mop.a. YKpenJI.HIOT pacTeHH.H B rpyHTe.
188 BomanuKa a Pt.tc. 70. reHeparnsHasi nOYKa 6y3111Hbl; B nlllCTbeB; 6 CTpoeH111e pa3nw-1HblX rnnos no4eK: 3- a - 6 serernrnsHasi nOYKa AY6a: reHepaTYIBHaR nOYKa BlllWHlll. 1 - 6 - serernrnsHo- nOY8YHbl8 Yewy111; 2 - 3aYaTKlll 3aYaTOK u,BeTKa CTeoeJIL Cme6eJlb - aTo oceBa.a qacTh rro6era pacTeHH.H, cocTo.arn;a.a H3 Y3JIOB H MeJK,n;oy3JIHH. CTe6eJih qarn;e Bcero HMeeT u;mmtt,n;pttqecKyIO cpopMy H B rrorrepeqHOM ceqeHHH MOJKeT 6bITh OKpyrJihIM, yrJIOBaTbIM, MHOrorpaHHhIM H ,n;p. <I>yHKI.J;HH CTe6JI.a: TpaHCIIOPT BO,Zl;hl H MHHepaJibHhIX Beu~eCTB H3 KOplrn. B JIHCTh.H, opraHHqecKHX Bern;eCTB H3 JIHCTheB B KopeHh; yBeJIHqeHHe IIOBepxHOCTH pacTeHH.H rryTeM BeTBJieHn.a; o6pa30BaHne JIHCTheB ll JIHCTOpacrrOJIO:>KeHHe; o6pa30BaHHe I.J;BeTKOB; 3arracaHHe BO,ll;hl H IIHTaTeJihHbIX Bern;ecTB; BereTaTHBHOe pa3MHO:>KeHHe. Ilo6ez - aTo cTe6eJih c JIHCTh.HMH H rroqKaMH. Ilo6er pa3,n;eJieH Ha y3JihI, B KOTOphIX pacrroJioJKeHhI JIHCTh.H H MeJK,n;oy3JIH.H. MeJK,ZWY3JIH.H 6brnaIOT yKopoqeHHhle H y,n;JIHHeHHbie. IlO'llCa - 3TO 3aqaToqHbIH, He pa3BepHyBilIHHC.H rro6er. I'JiaBHhIH rro6er 06- pa3yeTC.H H3 BepxyllleqHo:H rroqKH, 6oKOBhie rro6ern - H3 rra3yllIHhIX noqeK. IJoqKH 6hIBaIOT eezemamuBHbt.Mu, eezemamueHo-zeHepamumtbt.MU H zeHe- pamuBHbt.MU (pnc. 70). lfa eezemamUBHOU llO'llCU o6paayeTC.H CTe6eJib c JIHCTb.HMH H rroqKaMH. BezemamuBHo-zeHepamueHaR no'l/Ca o6pa3yeT rro6er c u;BeTKOM HJIH cou;BeTHeM. I'eHepamuetta.fl llO'l/Ca o6pa3yeT O,ll;HHQqHbIH I.J;BeTOK HJIH COI.J;BeTHe. BeTBJieHHe no6era BeTBJieHne rro6era Heo6xo,n;HMO ,n;JI.a yaem1qeHH.H rrJiorn;a.n;H TeJia pacTenmr. Bhl,n;eJI.HIOT ,n;Ba THrra BeTBJieHH.H rro6era: 1. Bepxyme'lHoe eemeJteHue. KoHyc HapacTaHH.H ,n;eJIHTc.a Ha ,n;Be qacTH, KaJK,n;a.a H3 KOTOphlx 06pa3yeT rro6er. TaKoe BeTBJieHHe Ha3hIBaeTc.a TaKme xomo.Mu'leCKoe (rrJiayHhI). iJu-
Cmpoeuue cme6nR o(JiwaO/l,bllblX pacmeuuii 189 PMc. 71. Ta.mbl BSTBJ18Hlll51: a -AlllXOTOMlllYeCKOe BeTB11ett111e (BOAOPOCJlb AlllKTlllOTa); 6 - MOHOnOAlllaJlbHOe BeTBJlett111e (no6er COCHbl); Myx111); r - B - ClllMnOA1118JlbHOe BeTBJlett111e no rnny MOHOX8311151 (no6er Yepe- c111MnoA111a11bttoe seTsne>me no T111ny A11txa311151 (no6er c111pett111). L.J,111cppaM111 o6o3ttaYett nop51AOK no6era 2. Bo1C0Boe BemBJteHue. II06er11 06pa3y10TcH H3 rra3yIIIHhIX rroqeK. BoKoBoe BeTBJiem1e 6hrnaeT Tpex THITOB (pHc. 71): 1) MOHOITO,D;HaJibHOe - KOHyc HapacTaHHH coxpaHHeTCH BCIO JKH3Hh, 06ec- lfa rra3yIIIHhlX rroqeK o6pa3ylOTCH 60KOBbie rro6erH (BeT- rreq11BaeT pOCT. BH). 3TOT THIT xapaKTepeH .D;JIH rOJIOCeMeHHhlX (eJih, COCHa H .n;p.), .D;JIH HeKOTOphlX IlOKphlTOCeMeHHhIX (.n;y6, 6yK H .n;p. ); emero.n;Ho BepxyIIIeqHaH rroqKa OTMHpaeT, a poCT 2) CHMITO,D;HaJibHOe - H.D;eT 3a cqeT 60KOBOH rroqKH (TOITOJih, 6epe3a H .n;p.); 3) JIO:>KHO.D;HXOTOMHqecKoe (BapHaHT CHMITO,D;HaJibHOro) - BepxyIIIeqHaH rroqKa OTMHpaeT, a pOCT H,D;eT 3a cqeT .D;BYX 6oKOBhIX rroqeK (cHpeHh). Ilo Timy pocTa rro6ern 6brna10T: rrpaMocToaqne (Bee BpeMH pacTeT BBepx) OOJihIDHHCTBO pacTeHHif; rrp11rro.n;H11Ma10m;Heca (rrJiayH 6yJiaBOBH.D;HhIH); cTeJI10lqHeca (ecJin OH yKopeHaeTCH, TO Ha3hIBaeTCH IlOJI3yqHM, HarrpHMep KJieBep IlOJlQyqH:H, 6y.n;pa IlJIIOW:eBH.D;HaH); BhlOW:HeCH (BblOHOK); u;errmnom;HeCH (xMeJih). CTpoettHe cTe6Jrn: o.n;tto.n;oJILHLIX pacTettu:H B attaTOMHqecKOM cTpoeHHH cTe6JIH TpaBHHHCThIX pacTeHHH Bh1.n;eJIHIOT: no1CpoBHy10 m1CaHb ( anuiJepMa ); nepBu'lny10 1Copy; u,eHmpa.rtbHbtil oceBoil u,u. 4'UniJp (pHc. 72).
Bomanu1w 190 1. J1o1Cp06Ha.H mlCUHb TOHKI:IH c.1oii: TIJIOTHO COMKHYThIX BhlTJIHY- IlepBH"1-IHa.JI Kopa ThIX KJieTOK, l:IMeeT 2. llepBU'lHa.H !Copa - yCThIU~a. o6pa3oBana no.n 3IlI:I,ZJ;epMOH TpeMJI Bl:I,ZJ;a:MI:I TKaneii:: MexaHI:IqecKOH KJieTKI:I; UempanLHh!H: TOJICTOCTeHHhie OCHOBHOH napeHXl:I- MOH (cpoTOCI:IHTe3Mpyro~aJI); 3ana- oceeoH. uw..1HHJlP caro~eii: napeHXl:IMOM ( 0,Zl;l:IH CJIOH KJieTOK). 3. IjeHmpaJlbHblU oceBOU U,UJlUHOp CO- .nepJKI:IT cocy ,Zl;l:ICTO-BOJIOKHl:ICThie nyqKII (scer.na 3aKphIThie). TipocTpattCTBO Mem.ny nyqKa:Ml:I 3am:ITO napeH- B XHMOH. PMC. 72. cocTaB Kam.noro nyqKa BXO,Zl;JIT BOJIOKHa Mexami:qecKOH TKaAHaTOMi1148CK08 CTp08Hi118 CTe6- HH H ml OAHOAOnbHOro paCT8Hi115l Ha nonepe4HOM cpe3e: 1 - 3n111AepMa; 2 - xnopoqmnnoHOCHaH; pi11U,111Kni1148CKafl; Tbl~ ny40K 4 - 3 - napeHx111Ma B CKnepeHX111Ma ne- TKaHh ,ZJ;BYX THTIOB: <jJJl03Ma. cepiJu,eeuna - CKOTIJieHI:Ie KPYTIHhIX KJieTOK c TOHKHMI:I CTeHKa- (4a - cj>no3Ma, 46 - KC111neMa) e 5 - l:I u;eHTpaJihHOH qacTII cTe6JIJI pac- noJiomeHa COCYA111CTO-BOnOKHll1C- CKnepeHXll1MHO~ 06KnaAKe; npOBO,ZJ;JI~aJI ICCUJleMa MH. napeHXll1Ma y 3.JiaKOB Ha MeCTe OTMeprneii: cep.nu;eBI:IHhl ocHOBHaH, 3anaca10w,a51 06pa3yeTCJI B03,ZJ;YIIIHaJI IlOJIOCTh (CTe6eJih - COJIOMHHa). CTpoeHHe cTe6m1 ;:i;peuecH&IX pacTeHHii ,n:epeBhJI H KYCTapHHKl:I HMeIOT MHOroJieTHHe o.npeBecHeBIIIHe CTe6JIH, cnoco6Hhle yTomu;aThCJI. YTomu;eHne, HJIH BTOpnqHhIH pocT CTe6JIJI, nponcxo.n11T 3a cqeT ,ZJ;eJITeJihHOCTH o6pa30BaTeJihHOH TKaHH - B KaM6Irn. CTe6Jie MHOrOJieTHero .npeBeCHOro pacTeHHJI BhI,ZJ;eJIJIIOT OCHOBHhie qacTH, nepuiJepMa, !Copa (nepBnqHaJI n BTopnqHaJI), 1CaM6uu, iJpeeecuHa, cepiJu,eeuna (pnc. 73). llepuiJepMa COCTOHT H3 HeCKOJlhKHX CJIOeB: OTMeprnaJI ann.nepMa (O,ZJ;HH aTo: CJIOH KJieTOK); npo6Ka (nepn.nepMa) - MepTBhie KJieTKH, rrporrnTaHHhie BOCKO- IIO,ZJ;06HhlM Be~eCTBOM cy6epHHOM. I1ep6U'iHa.H !Copa BKJllOqaeT: IIJiaCTHHqaTyIO KOJIJieHXHMY (MexaHnqecKaJI TKaHh); XJIOpocpHJIJIOHOCHYIO rrapeHXHMy; O.Zl;HOPJI,ZJ;HhlH CJIOM KpaxMaJIOHOCHhlX KJieTOK - (< KpaxMaJIOHOCHOe BJiaraJIH~e ». 8TOT KpaxMaJI pacTeHHe He pacxo.nyeT, IIO-BH.Zl;HMOMy, OH nrpaeT pOJih B no.n.nepmaHHH paBHOBeCHJI pacTeHHJI. BmopU'lHa.H !Copa BKJIIOqaeT CJIOH Jiy6a, COCTOJI~HH H3 Jiy6JIHhIX BOJIOKOH (MexaHll'IeCKaJI TKaHh) H CHTOBH.Zl;HhlX Tpy6oK c KJieTKaMH-CilYTHI:ID;aMH, a TaKjKe Jiy6aHyIO rrapeHXHMy. KaM6uu - JICY CJIOH MepncTeMaT11qecKHX KJieTOK, KOTOphIH OTKJia,ZJ;hIBaeT CTe6JIJI 3JieMeHThl emopU'l1WZO Jiy6a, BHympb - Hapy- 3.JieMeHThl BTOpllqHoii .npeBe-
BuiJou3MeJteHUH no6eza 191 cuHbI. KJieToK .n;peBeCHHhI aHa'lttTeJihHO 6oJihllle. fpatttt:u;a KOPhI rrpoxo.n;nT no KaM6HIO. Jl,peeecuna COCTaBJIJieT OCHOBHYIO qaCTb CTe6mI H B OCHOBHOM o6paaoBaHa 3JieMeHTaMH Mbl - rrpOBO,ll;HIIJ;e:i1 cocy,n;aMH. fO,ZJ;H'IHhie CHCTeKOJih:u;a emerO,ZJ;HhIH rrp:ttpOCT .n;peBeCHHhI 3a o,n;uH BereTaIJ;HOHHhIH rrep:tto,n;. Bectto:H:: KaM6HH OTKJia,ZJ;hIBaeT .n;peBeCHHY' KO- 6 Topaa COCTOHT H3 IIIHPOKHX, TOHKOcTeHHbIX KJieTOK (pbIXJiaH, CBeTJiaH). K cepe,n;:tttte JieTa OTKJia,ZJ;hIBaIOTCH MeJIKHe TOJICTOCTeHHhie KJieTKH c 60JihIll IllHM KOJIH'leCTBOM ,n;peBeCHHHhlX BOJIOKOH (MexaHH"'l:eCKaH TKaHh). Ilo ro,!l;HqHbIM KOJihIJ;aM MOiKHO orrpe.n;eJIHTh B03paCT pacTeHH.H. Cepau,eeuna cocTOHT Ha napeHXHM- HbIX KJieTOK. B :u;ettTpe 3TH KJieTKH 60- BH,!J;OH3MeHeHHH rro6era (pHc. y CrpoeHvie APeseCHoro cre6m1 (Ha npviMepe cre6m1 nvinb1): I - 74) KaM6vii;;; MHOrHX pacTeHHH B noqBe o6paay- IOTC.H no.n;aeMHhie rro6erH, B KOTOphIX 1 - orMepwvii;; 3nvi.QepMvic; 2 - nepvi.QepMa; 3- KOnneHXVIMa; 4 - 5- peHXVIMa; Hhle Beru;ecTBa. KpoMe Toro, tta,n;aeMHhie BVIHHbli;; nyy; no6erH MoryT BH,ZJ;OH3MeH.HThCH B opra- .n;. Kapa; II - Ill - .QpesecviHa; IV - cep.Qu.em·1Ha. HaKaITJIHBaIOTC.H aanaCHhie IIHTaTeJih- Hhl IlPHKpenJieHH.H, aaru;HThI HT • 9 IV P111c. 73. Jiee KPYIIHhie H qacTo OTMepIIIHe. aCCVIMVlmlU.VIOHH35l na- yYaCTKVI ¢n03Mbl; 7- HOrD KOnbu.a; 6- C8PAU.8- OC8HH5l5l AP8B8CVIH3 rD,QVIY- 8 - B8C8HH5l5l AP8B8CV1Ha ro9 - nepBVIYHaA KcvineMa ,[\VIYHOrD KOnbu.a; 1. IIoa3e.MHbte no6eiu: KOpHeBHIIJ;e (nanopOTHHK, JiaH,ZJ;bIIII) - MHOrOJieTHHH rro6er, HMeIOIIJ;HM pe- .n;yn;HpoBaHHbie JIHCTh.H B BH,ZJ;e qeIIIyeK, B naayxax KOTOphIX HaXO,ZJ;.HTC.H naayrnHhle noqKH. Ha KopaeBttru;ax o6paayIOTCH np:tt,n;aToqHhle KOPHH. PaattoBH,ZJ;HOCTLIO KOpHeBHIIJ; .HBJI.HIOTC.H CTOJIOHhl KapTocpeJI.H (He,ZJ;OJirOBe'IHhI); KJiy6HH (KapTocpeJih) - qiopMHpyIOTC.H KaK KOHIJ;eBhie HJIH rraayIIIHhie yTOJI- ID;eHH.H no.n;aeMHhIX no6eroB (cTOJIOHOB). Ha KJiy6tt.HX pacno;rometthI rro<JKH (rJiaaKH). K;ry6HH c;rymaT ,ZJ;JI.H BereTaTHBttoro paaMHomeHH.H; KJiy6HeJiyKOBH:a;a (rJia,ZJ;HOJiyc) - 3TO JIHCTBeHHhIH KJiy6eHh, T. e. KJiy6eHh, IIOKphlThlH qeIIIyeBH,ZJ;HhlMH cyxHMH ITJieHqaThIMH JIHCTh.HMH; JIYKOBH:a;a - ca ,!J;OHD;eM. yKopoqeHHhlH rro6er, CTe6JieBa.H qacTb KOTOporo Ha3hIBaeT- K ,n;oH:a;y np:ttKpenJI.HIOTC.H JIHCTh.H H qeIIIyH. Ha ,n;omi;e pacrroJia- raeTc.H BepxyIIIeqHa.H noqKa, H3 KOTOpoii pa3BHBaIOTC.H Ha,n;aeMHhie JIHCTb.H It ll;BeTOHOCHa.H CTpeJIKa. JlyKOBHIJ;hI CJiyiKaT ,ZJ;JI.H BereTaTHBHOrO pa3MH03teHH.H.
Bomanu1w 192 • . . 3 2 1 5 9 7 PMc. 74. COU,B8TMfl) BMHOrpaAa; nb1pes:i; 1- BMAOM3MeHeHMfl (MeTaMopcj:>03bl) no6era: TblH no6er KaKryca c peAyu,11posaHHblML'1 JlMCTbflMM; 7- 5- 3 - o6bl4Hb1Ci YAflMH8HHb1Ci no6er; IlYKOBMU,a nyKa; 4 - 38/leHblH 6e3IlMCTHblH cj:>OTOCMHTe3MPYIOW.MH no6er APOKa: 8- cj:>MJl/lOKilaAMM MrllMU,bl; 2. HaiJ3e.M1tbte no6eiu: KO/ll04Ka meAM4MM; 9- 2- Ms:ic11c- YCbl (BMAOL'13MeHeHHbl8 6- KOpHeBMW,a Kily6eHb Kaprocj:>ens:i KomoqKJf (60.HphIIIIHHK H ,11;p.), ycHKH (BHHOrpa,11; H ,11;p.), JiyKOBHD;hI (qecHOK H ,11;p.), YChI (3eMJI.HHHKa H ,11;p.), CTe6JieBhie KJiy6HH (D;HKJiaMeH H ,11;p.), KoqaH (KarrycTa H ,11;p.) - rHraHTCKa.H noqKa. JIHCT Jlucm - ::!TO H8,11;3eMHhIM, BereTaTHBHhIM opraH pacTeHH.H (pHC. 75). 06Jia,D;aeT ,D;BycTopoHHeii CHMMeTpHeii. PacTeT orpaHHqeHHoe BpeM.H 3a ctieT OCHOB8HH.H JIHCTa. <Pylllw,uu JlUCma: cpoTOCHHTe3, TpaHCIIHpau;H.f! (HcrrapeHHe BO,ll;hI), ra3oo6- MeH, 3anaca10ru;a.H <PYHKD;H.H; BereTaTHBHOe pa3MHO:>KeHHe. OcHOBHhie qacTH JIHcTa: ma, npuJtucmHUICU JtucmoeaR nJtacmu111Ca, 'lepewoK, oc11oea11ue Jtuc- (BhipOCThI OCHOBRHH.H JIHCTa). "IJepeIIIKOB H npHJIHCTHHKOB MO:>KeT H He 6hITh. 0CHOB8HHe JIHCTa MOJKeT pacIIIHp.HThC.H H OXB8ThIB8Th CTe6eJih, IIPH ::!TOM o6paayeTC.f! BJiaraJIHru;e. JlHCThH c qepemKaMH H83hIB8IOTC.f! 'lepeULIC06bl.MU, 6e3 qepeIIIKOB - CUdJl'lU.MU. "IJepem- KH opHeHTHPYIOT JIHCTOBYIO nJiaCTHHKY no OTHOIDeHHIO K HCTOqHHKY CBeTa, co3,D;aBa.H JIHCTOByIO M038HKY - pa3Meru;eHHe JIHCTheB Ha rro6ere, rrpH KOTOpOM OHH He 3aTeHHIOT ,11;pyr ,11;pyra. JlHCTb.H pa3JIHqaroT no p.H,D;Y npH3H8KOB: no pa3MepaM (OT HeCKOJihKHX MHJIJIHMeTpOB ,11;0 20 M - y naJibM), no JKHJIKOBaHHIO, no cpopMe JIHCTOBOM nJiaCTHHKH, no KpaIO JIHCTOBOM nJiaCTHHKH, no pacnoJioJ:KeHHIO Ha CTe6Jie.
193 )ICuJ1,1'06anue Jl,UCma. <PopMa J1-ucma JKHJIKOBaHHe JIHCTa 3TO rrpOBO;:J;HIIVie rryqKH JIHCTa. JKHJIKOBaHHe 6hrnaeT: )/(UJl/CU - ,nyroBoe (JiaH;:J;bIIIJ H ;:i;pyrHe O;:J;HO;:J;OJihHhie); rrapaJIJieJihHOe (JIHCT 3JiaI<a); rraJihqaToe - BCe )KHJIKH CXO,ll;HTCH B O;:J;HOM TOqKe (KJieH TaTapCKHM H .n;pyrHe ;:J;BY;:J;O.lJbHhie); rrepHCTOe - Bhipameua u;euTpaJihHaH )KHJIKa (qepeMyxa H ;:i;pyrHe ,ll;BY;:J;OJihHhie); BHJihqaToe - ;:J;HXOTOMHqecKoe (rHHKro ;:J;BYJIOIIaCTHhIM). <l>opMa JIHCTa Ilpocmou Jtucm - COCTOHT H3 0,ll;HOM JIHCTOBOM IIJiaCTHHKH H o;:i;uoro qepeIIJ- Ka. Ilpu JIHCTorra.n;e oTna;:i;aeT u;eJIHKOM. Jtucm - CJtOJICHbtU COCTOHT H3 HeCKOJihKHX JIHCTOBhIX IlJiaCTHHOK co CBOH- MH qepeIIIKaMH. 06ru;uii qepeIIIOK - paxuc. Ilpu JIMCTona;:i;e Kam;:i;aH JIHCTOBaH UJiaCTMHKa OTrra.n;aeT He3aBMCMMO. IIpocThle JIMCThH .n;eJIHTCH ua (puc. 76): a) JIMCThH c u;eJihHOM JIMCTOBOM nJiaCTMHKOM no cpopMe MOryT 6hITh OKpyrJihle, npo;:i;oJiroBaThle, JIMHeiiHhie u ;:i;p.; a 6 4 4 e PMc. 75. npOCTble nL'1CTbfl: 'iepewKa; I, A - a - B11aranl'1Ll.\Hble; 'tepewKy np1-1111-1CTHl'1KaM1'1; 3 OCHoeaH1-1e YepewKa; 3- YepeWKOBbllii; e - 6 - C1'1Af!Yl'1i1; B - C nOAYW8YKOi1 B OCHOBaHl'11'1 co CB060AHblMl'1 np1'1111'1CTHl'1KaM1'1; )I( - c np1-1pacTa10ll.\1'1Ml'1 K c na3yWHblMl'1 cpaCTalOL1.\1'1Ml'1Cfl npl'1111'1CTHl'1KaMl'1. 1 - a11ara111-1Ll!e; 4- np1-1111-1CTHHKH; 5- yepewoK; 6- n11aCT1'1HKa; 2 - na3ywHafl noYKa
BomanuKa 194 Tpoii'laTo- na11b4aTO- (rpex-) 0: .0 f() :s: c:: Q) Ji f() 0 a c PMC. 76. npoCTble lll1CTbfl C paC4118H8HHOVi lll1CTOBOVi n11aCT11HKOVi 11 CJ10)1(Hbl8 lll1CTbfl 6) JIHCTbH c pactIJieHeHHOM JIHCTOBOM IlJiaCTMHKOM MoryT 6bITh: JIOIIaCTHbie (TpOii'IaTo-, rraJiht:IaTO-, rrepMCTO-); paa,n;eJihHhie (pact:IJietteHHhie, 6oJiee 1/ 2 nmpHHbI); paccet:IeHHh!e (pact:IJieHeHHhie .n;o cpe.n;Heii iKMJIKH). BhI,n;eJIHIOT TPH THrra pacnoJioiKeHHH JIHCTbes: oqepe,n;Hoe, HJIM CIIHpaJibHOe - OT KaiK,n;oro yaJia OTXO,ll;HT TOJihKO O,D;MH JIHCT (.n;y6, 6epeaa, 3JiaKH, 30HTM'IHbie M ,n;p.); cyrrpoTHBHoe - s KaiK,n;oM yaJie rrapa JIHCTbeB (KJieH, cHpeHh, ry6ou;seTHhle H ,n;p.); MyTost:IaToe - B KaiK,n;oM yaJie TPH H 6oJiee JIHCTa (oJieatt.n;p, aJio.n;eH, 1rnatt-t:Iaii H .n;p.). Ilo attaTOMH'IeCKOMY CTpoeHHIO BhI,n;eJIHIOT: 1. Pop3o6enmpaJLbHbte Jtucmb.R - accHMHJIHD;HOHHaH qacTh JIHCTa (cToJI6t:IaTa.H XJIOpocpHJIJIOHOCHa.H rrapeHXMMa) pacrroJIOiKeHa TOJlbKO Ha sepxHeH CTO-
fenepamu6Hble opzaubi (u,.6em01c, nnoiJ, ceMR) poHe JIHCTOBOH IIJiaCTllHKII, ry6'-IaTaH - 195 Ha Hl:DKHeH (caxapnaH CBeKJia). AHaTOMHt.feCKoe cTpoeHne .n;opaoseHTpaJthHoro JIHCTa JI01Cpo61W.H mKaHb - snII.n;epMa IIJIII KOiKIIlla. Rommi;a noKphITa KY- TMKyJioH: II MMeeT yCThMlla, KaK npaBMJIO, Ha HIIiKHeH IlOBepXHOCTH JlllCTa. 3amnmaeT JIMCT CBepxy H CHH3y. Pa6oTa ycThHll 06ecneq1rnaeT nocTyrnreHne yrJieKHCJioro raaa H Bhrne.n;eHHe KHC.'lopo.n;a, a TaKme TpaHcnnpa11mo (HCilapCHMe BO,ll;hl). fa3006MCH B TKaHHX JIHCTa npoTeKaCT IlOCTOHHHO, Tor.n;a KaK CBeTOBaH cpaaa cpoTOCHHTeaa - Me3o</JuJIJI (cpoTOCMHTe3IIPYIOI11aH TKaHb) - TOJihKO ,Zl;HCM. CTOJI6'IaThIM n ry6t.IaThIH. R'.JICTKH CTOJI6t.faToro MeaocpHJIJia 06hI'IHO co.n;epmaT 60JihllIOe KOJIII'ICCTBO XJioponJiaCTOB. fy6t.IaTbIH MC30cpHJIJI HMCCT MHOro MCiKKJICTHMKOB. Cocy8ucmo-60J101CHucmbte ny'lKU (mMJIKH) BKJIIO'IaIOT KCHJieMy (.n;pesec1rny), cpJI03MY (ny6) H MCXaHII'ICCKYIO TKaHb (yroJIKOBYIO KOJIJICHXHMY na.n; ny'IKOM H CKJiepeRXHMY KOJihllOM BOKpyr nyt.IKa). 2. I130JLamepaJibTtbte JlUCmb.H - CTOJI6qaTaH napeHXIIMa c .n;syx CTOPOH JIH- CTa (r.11a.n;nonyc). 3. PaiJuaJibHbte JLucmb.H - xJiopeHXMMa BOKpyr 11eHTpaJihHOH iKMJIKM (XBOH COCHhl). JincTona.n; - ceaOHHaH a.n;anTallMH pacTeHMH K M3MCHCHMHM KJIMMaTa. Oc- HOBHaH npM'IMHa JIMCTOna.n;a oceHblO H 3MMOH. Heo6XO,ll;MMOCTh YMCHbIIIeHIIH ncnapeHMH BO,Zl;hl c IlOHMiKCHMeM TeMnepaTyphl BCaChIBaHMe BO,ll;hl KOPHHMH cHIImaeTcH, pacTeHne MomeT o6easo.n;MThCH n norn6nyTb. C6pachrnaHne JIMCTheB npe.n;oTBpamaeT STOT npo11ecc n yMeHbIIIaeT ncnapenne BO.D;hI. Bn.n;onaMeHeHMH nncTa: KOJIIO'IKM (KaKTyc H .n;p.); ycnKM (ropox n .n;p.); JIMCThH HaCCKOMOH,ll;HhIX paCTCHHH (pOCHHKa H .n;p.). reHepaTHBHbie OpraHbl (I.J;BeTOR, IIJIO)l;, CeMH) IJ;BeTOK z.r6emo/C - 3TO opraH ceMeHHOro pa3MHO.iK8HMH IlOKpbITOCCMeHHhIX pacTe- HMH. IJ;seToK npe.n;cTaBJIHeT co6oft yKopot.IeHHhIH cnopoHoCHhIH no6er, npncnoco6.11eHHhIM ,D;JIH o6paaosaHMH cnop n raMeT. B 11seTKe nponcxo.n;nT onhIJieHMe, OII.110,lJ;OTBOpCHMC, pa3BMTMe aapO,Zl;hIIIIa, o6pa30BaHMe TIJIO.n;a c CCMCHCM. CTpoeHHe u;senta (pl:lc. 77) Il;BeTOK COCTOMT H3 cme6Jle60U 'lacmu (llBCTOHOmKa, llB8TOJIOiKC) H JLucmo60U 'tacmu ('IaUICJIMCTMKM, JierreCTKM, Thl'IMHKH, IICCTHKM). I.{6emoJIOJ/Ce - paCUIMpCHHaH qacTb llB8TOHO.iKKM, K Heii npMKpenJIHIOTCH Bee 'IaCTM llBCTKa. l/ aute1t1Ca (Ca - calyx) - cocTOMT M3 t.IaUie.11ncTMKOB ( o6hI'IHO ae.11eHoro D;BeTa). qaUICJIMCTMKM MoryT 6hITh CB06o.n;HhIMM, TOr.n;a qarnet.IKa Ha3bIBaCTCR paaiJeJLbHOJiucmnou. EcJIM qarneJIMCTMKM cpacTaIOTCH, TO qarueqKa Ha3hIBaeTca cnaunoJLucmnou.
BomanuKa 196 Beu'luK (Co 7 6 8 9 .'3 9 JO --2 -11 15 1.'3 HOCTh corolla) - JieneCTKOB. coBoKyrr- JienecTKM MOryT 6hJTh CB060,ll;Hh1Mll IIJIM CpOClllMMIICR, a seHqIIK, COOTBeTCTBeHIIO, pa38eJlbHOJtenecmHbtU IIJIM cnauttoJtenecmHbtil. BettqIIK o6hlqtto RpKo oKpallleH, IIMeeT pa3ttyro cpopMy, npIIBJieKaeT ttaceiw- MhIX-OIThIJIIITeJiett. 0KOJlOU,6emHUK, o6pa30BaHHhIH qa860UHbt.M. Ilpocmou OKOJlOU,6emHUI( MOJKeT 6b1Th 8eH'lUK06U8Hbl.M (TIOJihIIaH), T. e. llleqKott II BeHqMKOM, Ha3bIBaIOT COCTOHTh TOJihKO II3 JIIICTQqKOB BeHqII- PMc. 77. CxeMa CTpoeH111si useTKa: 1 PblJlbL(e; 2- 3aBSl3b; 3 - CTOJ16111K; 4 - C8MSl3a<iaTOK; 5 - Tbl'lLllHO'lHaSl HLllTb; 6 - CBSl3HLllK; 7- nbtllbHLllK; 8 - nbtJ1bL(8Bbte 3epHa; we11111crnK; Ka; nbtJlbHLllK B pa3pe3e; 9 - 10 - 11enecrnK; 11 - o.ia- 12- L(BeTOJlOlKe; 13- L(B8TOHOlK- 14- npLllL(B8THLllK; 15 - npLllL(B8THLll'l8K Ka, IIJIII 'lGUle'lK06UOHbl.M (KpaIIIIBa). ll,BeTKII 6e3 OKOJIOD;BeTHIIKa Ha3hIBaIOT rOJihlMM (IIBa). I'a.Memo<Pum - rarrJiott,n;ttoe rroJiosoe IIOKOJieHI:Ie, rrpo,n;yu;HpyIOID;ee raMeThI. Cnopo</Jum - ,n;ttrrJioII,n;ttoe rroKoJie- HHe, KOTopoe IIPOH3BO,ll;HT rarrJIOM,ll;Hbie crrophl (MHKpo- H Meracrropb1) y nseTKoBhIX pacTeHHH. Tbt'lUHKa COCTOHT H3 mbl'lUIW't/WU ttumu H nbtJlbHUKa. CosoKyIIHOCTh ThIqHHOK Ha3hIBaIOT aniJpou,ee.M (A). 10 IlbIJlhHHK o6pa30BaH ,D;BYMH IIOJIOBHHKa- 6 MII, coe,n;tttteHHhlMH CBH3HHKOM. 8 ,n;oii: IIOJIOBHHKe IIhIJihHHKa 7 IIhIJihIJeBhlX rtte3,n;a - B Kam- eCTh ,n;Ba .MUKpocnopattzua. llbl.Jf,bU,e- rtte3,n;a IIhIJihHHKOB Ha3bIBaIOT 6bl.MU .Mew1ca.Mu. B HHX o6pa3yroTCH MHKpocrrophl II co3peBaeT IIhIJihIJa. II btJlbU,a - aTo pe,n;yu;ttposaHHhltt MYJKCKOH raMeTOcpIIT' PMc. 78. CTpoeH111e ceMA3a<iaTKa 111 aapo- Abtwesoro MeWKa (lKeHCKOro raMeTOCj>LllTa) noKpbnoceMeHHbtX pacTeH111iii: 1 - no- KPOBbt ceMsiaa<iaTKa, LllJlLll LllHTeryMeHTbl (a Hap~Hbtiii, 6- BHyrpeHHLlliii); 2- ML11Kpon111- 11e; 3 - xanaaa; 4 - Q>yH111Ky11yc; 5 - HyL(e1111yc; 6 - KTleTKa, 3BPOAblW8Bbliii MewoK; 8 - CLllHepfLllAbl, 9 - 7- SliiiL(e- BHTL!lnOAbl; 10- srnp111o.iHoe SlApo; 11 - n11aL(eHTa; 12nposOASll.l.!Llliii ny<ioK COCTOHID;HH H3 ,n;syx KJieTOK. 06e KJieTKH MyiKcKoro raMeTocpttTa ( IIblJlhlJhI) BhIIIOJIHHJOT orrpe,n;eJieHHYIO eezemamuettaa, 06pa3yeT IIhIJihIJeByIO Tpy6Ky. MaJieHhKaa KJieTKa, HJIH zettepamuetta.R, o6pa3yeT ,n;sa crrepMHH - cpyHKIJHIO. BoJihlllaH KJieTKa, HJIH My»CCKHe raMeThI. IIecmwc MOiKeT 6hITh o,n;IIH HJIH aeCKOJihKO. CosoKyrraoCTh rrecTHKOB aa-
197 iuneu,eeM (G ). IleCTHK o6pa30BaH IlJIO,ll;OJIHCTHKaMH. IleCTHK COCTOHT aB.H3U, cmoJL6uKa w pbtJlbU,a. Ha pbIJiblle rrecTttKa rrorra.n;aeT IIbIJihIIa, w rrpo113 3 3bIBaIOT 11cxo,n;MT rrpopacTaHtte. EcJIH IIJIO,n;OJIHCTMK o6paayeT e,n;11HCTBeHHhitt rrecn:IK, MOHOKapnllbtM. ECJIH ,n;Ba H 6oJiee IIJIO,ll;OJIHCTHKa o6pa3yIOT STO anoKapnHbtU iuneu,eu. Ilptt cpacTaHHH HeCKOJlbKHX IIJIO,ll;OJIHCTHKOB B O,IJ;HH rreCTHK 06pa3yeTC.H u,eHOKapnHblU iuneu,eu. Ce.Mano'lKa (pnc. 78) Haxo,n;ttTC.H B 3aB.H3H rrecTHKa. IlpnKperrJI.HeTc.H TO OH Ha3bJBaeTC.H caMOCTO.HTeJibHhJe rreCTHKH - K cTeHKe 3aB.H3M c noMOIIJ;bJO IIJia11eHTbI Ka COCTOHT H3 H B3.ZJ;YTH.H TKaHell: 3aB.H3H. CeM.HIIOq- ce.M.RHOJICKU ( rpynuKyJLyca ), noKp0606 ( 06a UHmeiy.Me1tma) aopa CeM.HIIOqKH, B KOTOpOM <l>OpMHpyeTC.H 3ap0,1J;bIIIIeBbIH MeIIIOK. 3apOObtUle6btU .MeUlOK (COCTOHT H3 BOCbMl:I KJieTOK) - 3TO pe.n;y11npoBaH- Hblll .meHCKHH raMeTO<l>HT, KOTOpbIH pa3BHBaeTC.H H3 Meracrropbl. IloKpOBbl nbtJlbU,eBXOO ( MUKponu.lle). BJin.me K IIbIJiblleBxo.n;y Haxo,n;.HTC.H Hll:11eKJieTKa H .n;Be KJieTKH - cunepiuobt ( n). Ha rrpoTHBorroJio.mHoM KOHI1e - TPH KJieTKH - anmunoiJbt (n}, B 11eHTpe - ou_nJlOUOHOe BmOpU'lHOe .HOpO ( 2n }, KOTOpoe o6pa30BaJIOCb Ha BepxyIIIKe CeM.HIIOqKH He cpacTaJOTC.H H o6pa3yJOT H3 CJIH.HHH.H ,n;Byx IIOJI.HpHbIX .H,n;ep. o6oenOJlbte. Il;BeTKH TOJibKO c TbloononOJlbte (pa3,n;eJibHOIIOJible), HaIIpH- Il;BeTKl:I c TbiqHHKaMH H rreCTHKaMH qHHKaMH HJIH TOJibKO c rrecnrnaMH Mep KyKypy3a, 6epe3a H .n;p. KJiacc11<l>HKa11m1 pacTenHii PacTeHH.H, y KOTOpbIX MymcKne (Tb1qnHOqHb1e) H .meHCKHe (rrecTnqHbie) IJ;BeTKH Haxo,n;.HTC.H Ha o,n;HoM pacTeHHH, Ha3hlBaJOTC.H oonoooMHbt.Mu (KYKYPY- aa). PacTeHH.H, y KOTOpbIX o,n;HOIIOJibJe IIBeTKH Haxo,n;.HTC.H Ha pa3HbIX pacTeHHsx, Ha3bIBaJOTC.H 06YOO.Mllbl.MU (KparrHBa, HBa H .n;p.). Co11seTHJ1 Cou,Bemua - aTo rro6er HJIH cncTeMa rro6eroB, HecyIIJ;HX IIBeTKH. CTBHH c THIIaMH BeTBJieHH.H rro6eroB co11BeTH.H (pHC. 79) B cooTBeT- paa,n;eJI.HJOT Ha: 1) Mononooua.llbHbte ( 6ompuouiJHbte) cou,Bemua 6hlBaJOT rrpocThle H cJiom- IIhle. BoKOBbJe ocn rrpocTbIX co11BeTHtt He BeTB.HTC.H, CJIOJKHbIX - BeTB.HTC.H. IlpOCTLie MOHOIIO,ll;HaJILHLie co11seTHJI. PocrrycK IIBeTOB rrpoHCXO,!J;HT CHH3Y BBepx, ec.rrn co11BeTHe BepTHKaJibHOe (KHCTb, KOJIOC), JIH6o OT rrepn<PepHH K :u;eHTpy, ecJIH IIBeTKH B COIIBeTHH pacrroJio.meHbl ropH30HTaJibHO (30HTHK, KOp3I:IHKa): KHCTb - BCe IIBeTirn Ha y,n;JIHHeHHOH OCH, HMeJOT IIBeTOHOJKKH (qepeMyxa); Ill;HTOK - cxo,n;eH c KHCTbJO, HO IIBeTOHOJKKH pa3HOH ,IJ;JIHHbl H IIBeTKH pacrrOJIOJKeHhl B 0,IJ;HOH IIJIOCKOCTH (rpyIIIa); 30HTHK - IIBeTKH Ha IIBeTOHOJKKaX 0,IJ;HHaKOBOH ,IJ;JIHHhl, BbIXO,!J;.HT H3 o.n;- HOH TQqKH (BHIIIH.H); KOJIOC - IlpOH3BO,ll;HOe KHCTH, HO 6e3 IIBeTOHOJKeK, IIBeTKH CH,ll;.Hqne (no- .n;opomHHK); noqaTOK - KOJIOC c TOJICTOH OCbIO (KyKypy3a);
198 6omanuKa rOJIOBKa - BH,Zl;OH3MeHeHHbIM 30UTMK; OCb pa3pacTaeTCJI, Ha Heii CM,Zl;H'!He IJ,BeTKH (KJICBep); KOP3HHKa - CH,ll;Jl'!He IlBeTKH Ha pacurnpeHHOM KOHUe OCH cou,aenrn ( CJIOJKHOIJ,BeTHbie). CJimKHLie MOHono,11;HaJILHL1e cou.BeTH.JI: MeTeJIKa (CJIOarnaH KHCTb) CJIOJKHhIH KOJIOC - o6pa3oBaHa H3 OT,ri;e.TCbHhlX KHCTeii (cttpeHb); o6pa30B8H H3 OT,ri;eJihHbIX IlpOCTbIX KOJIOCKOB (muemm;a); CJIOJKHhlll 30HTHK - COCTOHT H3 IlpOCThIX 30HTHKOB OT Bepxyunrn rJiaBHOH OCH (MOpKOBh); CJIOJKHhIH ILl,HTOK - COCT8BJieH H3 npOCThIX ILl,HTKOB (mrmMa); 2) CUMnoauaJtbHblf' (U.UMOJHbte} co- U,BemuR. I'JiaBHaJl OCb 38KaHqHaaeTCR: I.J,BeTKOM. Iloc;re,11;y101I.1,He IJ,BeTKH B03HHKaIOT B na3yxax npHll,BeTHbIX JIHCTb- • eB H np0,!1;0JIJK8IOT POCT rJiaBHOH OCH • PocnycK ll,BeTKOB npo11cxo,n;HT caepxy 4 BHH3 mrn H3 I.J,eHTpa K nepmpepHH. Pa3JIH'laIOT TPH THIIa CHMno.n;HaJih- a 8 9 6 \ HhIX COll,BeTHH: ~ a) MOHOX83HH (H3BHJIHHa H 3aBHTOK) - TpeThero rrop.JI.l(Ka H T. ,ZI;., TO c O.!l;HOH, TO c .l(pyroii CTOpOHbI (JIIOTHK e,n;KHH); 38BHTOK ro - T1>1nb1 co4sernj;j: a - HOM ll\lllTOK; 6- 3 - 30HTlllK; noYaTOK; 7- 4 - 6- 5 - 2 - 11 - 3111j;j); 14- IIO,ZI; ll,BeTKOM rJiaBHOH KOJlOC; meHHbie BeTBH (OCH), 3aKaH'lHB8IOILl,HeCJI ll,BeTKOM. Kam,n;aH H3 3THX oceii TaK- 1113BlllJllllH8 {MOHOX83111M}; 13 - pa3BHJIHHa - OCH o6pa3yIOTCH cynpOTHBHO pacrrOJIO- CJlO>t<Hble 6oT- 8 - CJlOJKHblM KOJlOC; 9 - M8T8JlKa; 10- CllO>t<HblM 30HTlllK; B - l_\lllM03Hbl8 (ClllMnO- 3aBl1TOK (MOHOxaa111j;j) ; JIOJKHO,!(HXOTOMH'leCKOe pacrroJiomeuue oce:H: PlllOWJ,Hble: Al18JlbHb18): 38KPY'lHB8HCb 6) ,Zl;HX83HH (pa3BHJIHHa H ,Zl;BOHHOH 38BHTOK) - KlllCTb; rDJlOBKa; KOP3111HKa; HanpaBJieHHH, B 38BHTOK (ue3a6y,Z1;Ka); npocrb1e 6orp111- I - ocb TpeThero nopH,n;Ka H T • .n;., HO IlpH 3TOM OCH OTXO,Zl;HT TOJlhKO B 0,Zl;- (J PMc. 79. OT rJiaBHOH OCH OTXO- ,Zl;HT OCh BTOporo rropH,n;Ka, OT BTopo- ~ Y, 0WJ.Hble (MOHOnOAlllMbHb1e): OT rJiaBHOH OCH OTXO- .l(HT OCh BTOporo IIOPH,ri;Ka, OT BTOporo - ~12 11 O'lepe,ri;Hoe pacnoJiomeHHe oceH:: H3BHJIHHa - 12- pa3BlllJ111Ha (A111Xa- MHoro11Y4eB11K (n11ej;joxaa111M) JKe o6pa3yeT .n;se cynponrnHhIX ocH (rB03.D;HKa); .!l;BOHHOH 3aBHTOK - ITO,ZI; IJ,BeTKOM rJiaBHOH OCH cynpOTHBHO o6pa3yIOTCH ,Zl;Ba rrpoCTbIX 38BHTKa (KapTo¢leJib);
On""nenue. OnnoiJom6openue B) nJieii:oxa3tt:ll - 199 MyToBqaToe pacnoJiometttte oce:ll - no.n: IJ;BeTKOM rJiaB- aoil: OCH pa3BIIBaeTCH MYTOBKa 60KOBbIX oceii: c IJ;BeTKaMH IIJIII COI1BeTHHMH (l\WJIOqaii:). OnbIJieHHe OnbtJlettue - aTo npoIIecc nepettoca TihIJihIJ;hI II3 nhIJihHIIKOB Ha phIJihIIe ne- CTHKa. Pa3JIIIqaIOT caMOOIIbIJieHHe, nepeKpeCTHOe OilhJJieHtte, IICKYCCTBeHHOe OilbIJJeHtte. Ca.MoonbtJleHue MOJKeT npowcxo,n:HTh y pacTeHHH, HMeIOIIIHX ,n:ByTIOJihie I1BeTKH. IlhIJihIJ;a nepeHOCHTCH Ha pbIJihIIe necTHKa BHYTPH o,n:ttoro IJ.BeTKa. CaMOOilhIJIHIDIIIHec.H pacTeHHH - ropox, .fiqMeHb, pOJKh, OBeC II ,n:p. Jlepe1Cpecmttoe onbtJlettue - Ha phIJihIIe nona,n:aeT TihIJihIIa c ,n:pyrwx pacTe- HHH Toro me BII,n:a. IlepeKpeCTHOe OilbIJieHHe OCyIIIeCTBJIHeTCH no.n: B03,n:eil:CTBHeM pa3JIHqHbIX BHeIIIHHX q:>aKTOpOB: aaeMocpttJitt.a - onhrnem1e c noMOIIIhID BeTpa. PacTeHH.H paao 3au;BeTa- IDT, .n:o TIO.HBJieHII.H JIHCTbeB. ll,BeTKH MeJIKHe, HeB3paqab1e. IlbIJibIJ;a MeJIKa.a, cyxaH. Phrnhu;a necTHKOB qacTo nepttCThie. 3To MHOrIIe 3JiaKH, OCOKa, 6epe3a, ,n:y6 H ,n:p.; 3HTOMOcpHJIHH - OTibIJieHHe c TIOMOIIIbID HaCeKOMhIX (nqeJI, IIIMeJieH, oc, 6a6oqeK). l.I;BeTKII KpynHhie, .HpKO OKpaIIIeHhI, eCTb HeKTap H 3anax. PacTeHII.H, OilbIJI.HeMhie HaceKOMhIMH, - IIIIITIOBHHK, H6JIOHH, qepeMy- xa, BHIIIH$I, MaK, poMaUIKa; rw,n:pocpttJIH.H - OTihIJieHtte c noMOIIIhID BO,ll;hI. XapaKTepao ,ll;JI.H MHorwx BO,ll;HbIX UBeTKOBbIX pacTeHHH. HcJCyccmeeHHoe onbtJLettue - aTo npway,n:wTeJibHoe onhrJieawe, npott3Bo,n:w- Moe qeJIOBeKOM c TIOMOIIIbID co6CTBeHHOH IlhIJibUhI u;BeTKa HJIH IlhJJlhIIhl ,n:pyI'HX pacTeHHH. Omrn,n:oTBOpeHHe OnJLoiJomeopettue - aTo npou;ecc CJIHHHHa meacKoil: H MymcKoli noJIOBhIX KJieTOK c o6pa30BaHHeM 3HrOThl (o,n:HOKJieToqHoro 3apO,ll;hIIIIa). y IIBeTKOBbIX pacTeHHH STOT TIPOIIeCC Ha3hIBaeTC.H iJ60UHOe onJLoiJomeope- nue. Oa 6b1JI OTKphIT B 1898 r. PYCCKHM 6oTaHHKOM c. r. HaBaIIIHHbIM. IlhIJihIJ.eBoe 3epao nona,n:aeT aa phIJih1Ie necTHKa. BeiemamueHaR JC.TLemJCa npopacTaeT B BH,D;e Tpy6KH qepe3 TKaHII pbIJihIIa H CTOJI6IIKa no aanpaBJieHHIO K 3aB.H3II. Tpy6Ka npoxo,n:HT qepe3 MIIKponIIJie (rrhIJihIIeBxo.n:) w oKa3hIaaeTc.a BHYTPII 3apo,n:hnueBoro MeIIIKa. I'ettepamueHaR JCJLemJCa 06pa3yeT .n:aa crrepMIIH. 0Htt ,n:BII?KyTc.a B IIhIJihUeBoli Tpy6Ke Ha ee KOHIJ.e. Ilh1Jih1IeBaa Tpy6Ka pa3pbJBaeTCH, crrepMMH OKa3b1BaIOTC.H B 3apO,ll;hJIIIeBOM MeIIIKe. 0,n:HH cnepMHH CJIIIBaeTC.H c .HHIJ.eKJieTKOH, II 06pa3yeTC.H ;wioma - rrepBaH KJieT- Ka 3apo)J;hIIIIa ceMeHH. BTopoil: crrepMIIH CJIHBaeTCH c ,n:HnJIOII.ll:HhIM H.n:poM IJ.eaTpaJihHOH KJieTKH, 06pa3yeTc.a mpunJLouiJHaR JCJLemJCa. TpHIIJIOH,D;Ha.H KJieTKa .n:aeT HaqaJio 3HiJocnep.My (nHTaTeJibHOH TKaHH ceMeHII).
BomanuKa 200 IIocJie onJio.a;oTBope1uu1 H3 ceMJrnaqaTKa 06pa3yeTCH ce,11.fl. CeMH nMeeT ceMeHtty10 1COJ1Cypy, aapoiJbtUJ, :Janae numameJtbllbtX Bew,ecmB. 0.nttoBpeMeHHo pa3pacTaeTC.H 3aBH3b neCTHKa. lfa CTeHOK 3aBH3H 06pa3yeTCH OICOJLOnJIOOllUIC, oKpymaIOID;Hlr ceMH. IlpOHCXO,ll;HT o6pa30BaHHe ITJIO,Lla. flJIO.ZJ; flJtoiJ - opraH pa3MHOJKeHHH IlOKpbITOCeMeHHblX pacTeHHH. IlJIO,ll; co.nep- JKHT ceMeHa 11pa3BHBaeTCH H3 O,LIHoro u;BeTKa (pHc. 80). <l>yHKIJ;HH TIJIO,n;a: cpopMHpOBaHn:e, 3aID;HTa, pacnpocTpaHeHHe ceMHH. CTeHKa nJio.a;a Ha3hIBaeTca nepuKapnue.M ( 0KoJtonJtoiJ1tuKoM) H HMeeT TPH cJioH:3K3oKapnuu,Me301Capnuu,a1tiJ01Capnuu. B COOTBeTCTBHH c THITOM rMHen;eH Bee ITJIO,Llbl .neJIHT Ha MOllOICapnuu, anoKapnuu, u,e1t0Kapnuu H nceBOOKapnuu. flJLOObl MOHOICapnuu pa3BHBaIOTCH H3 u;BeTKOB, MMerouvrx MOHOKapnHhIH rHtteu;eif: a) O,ll;HOJIMCTOBKa - MHOroceMeHHOH ITJIO,D;, BCKphIBaIOID;HHCH TOJibKO no 6pIOllIHOMY llIBy; 6) o,n;HoKocrnHKa - o.nHoceMeHHoif, HeBcKpb1BaIOID;nifca ITJIO,LI. Me3oKap- nnif O,ll;HOKOCTHHKH MOJKeT 6bITb coqHbIM (BHllIHH, CJIMBa H ,n;p.) HJIH cyXHM (M11H,1l;aJih H ,Lip.); B) 606 - O)IHO- HJIH MHOroceMeHHOH rnIO,n;, pacKphrnaeTCH no ,D;BYM JlIBaM (cnnHHOMY n 6pIOllIHOMy). CeMeHa, pacnoJiomeHHbie Ha CTBOpKax (.z:IOHHHK, apaxn:c, aKaIJ;HH H ,n;p.). flJtOObt anoKapnuu pa3BnBaIOTCH n3 anoKapnHoro rnHeu;eR: a) MHOrOJIHCTOBKa - MHoroceMeHHOH c6op- HblH ITJIO,D;, COCTOHT H3 IlpOCTblX JIHCTOBOK (KYilaJihHHIJ;a, KaJiyJKHMD;a H ,n;p.); 6) MHoroopellleK - c6opHbIH MHoroceMeH- HOH ITJIO,D;, COCTOHID;HH 113 OT,D;eJihHblX O,LIHOceMeHHblX HeBCKpbIBaIOID;HXCH IIJIO,ll;HKOB - 2 opellIKOB (JIIOTHKH, JianqaTKa H ,n;p.); B) MHOrOKOCT.HHKa - c6opHblH ITJIO,ll;, COCTO- HID;HH H3 ,n;ByX HJIH MHOrnx ITJIO,ll;HKOB - 3 KOCT.HHOK. Me30Kapn11ii Kam,n;oro nJio.nnKa coqHbIH, 3H,ll;OKap1rn:if - TBep,n;blif (Ma- JIHHa, emeBMKa H ,n;p.). flJLOObl u,eliOKapnuu pa3BlfBaIOTCH H3 u;eHOKapnHoro rnHeu;eH. 3aBH3h o,n;Ha, HO o6paPMc.80.CTpOeHV1ennoAa(oAHOKO- 30BaHa ,D;BYMH H 6oJiee cpOCJlIHMHCH ITJIO,D;O- CHIHKVI) nepcviKa o6blKHOBeHHOro: JIHCTMKaMM: 1-3 - a) Hro,n;a - oKononnoAH~K. vinvi nviiil ( 1 - 3K30Kapn~~. 2 - nepviKapMe3oKap- nvii1, 3 - 3HAOKapn~~); 4 - CeMft; 5 cneA nnoAOHQ)l(KVI; 6 - nnOAOHOJKKa MHOroceMeHHOH HeBcKpbrnaro- ID;Mifca ITJIO,ll; c KOM - coqHbIM OKOJIOITJIO,D;HH- nepMKaprrn:eM (noMn,n;op, KapTo- cpeJih, KPhliKOBHnK 11 ,n;p.);
201 flJ1,oiJ. CeMH 6) TbIKBHHa - IIJIO,ZJ;, 6JIH3KHH K Hro.n;e, HO OTJIHqaroru;HHCH KOIBHCThIM 3K- 30KaprrtteM (ThIKBa, ap6y3, orypeu; H .n;p~); B) Kopo6oqKa - MHOroceMeHHOH IIJIO,ZJ;, HMeeT cyxo:i1: BCKphIBaIOru;HHCH OKOJIOIIJIO,ZJ;HHK (MaK, .n;ypMaH, HarrepcTHHKa H .n;p.); r) cTpyqoK - o6pa3oBaH .ZJ;BYMH cpocIIIHMHCH IIJIO.ZJ;OJIHCTHKaMH, BHYTPH HaXO,ZJ;HTCH rrpo,n;OJihHaH rreperopo,n;Ka, K KOTOpoii rrpHKpeIIJIHIOTCH CeMeHa (KarrycTa, ropqttu;a H ,n;p.). CTpyqKH KJiaccmpttu;ttpy10T Ha BCKph1Ba10ru;ttecH H HeBCKpbIBaIOru;HecH; .n;) CTpyqoqeK - CTpyqOK, y KOTOporo ,!l;JIHHa IIOqTH paBHa IIIHPHHe (rracTy- IIlhH CYMKa H ,n;p.); e) H6JIOKO - MHoroceMeHHOH HeBCKpbIBaIOIIJ;HHCH IIJIO,n;, 3aBH3h KOToporo o6pa3yeTCH cpacTaHHeM OT Tpex ,n;o IIHTH IIJIO,!l;OJIHCTHKOB (H6JIOHH, PH- 6ttHa, 6oHphllllHHK H ,n;p.). ll;wabt nceBa01capnuu - 3aKJia,!l;hIBaeTCH ,n;Ba H 6oJiee IIJIO,ll;OJIHCTHKOB, a pa3BHBaeTCH TOJihKO O,!l;HH, OCTaJihHhJe pe,n;yu;ttpyIOTCH: a) ceMHHKa O,!l;HOCeMeHHOH IIJI0,!1; c KO:IKHCThIM rrepttKaprrHeM, He cpacTaIOIIJ;HMCH c ceMeHeM (rro,n;COJIHeqHHK, BaCHJieK, poMaIIIKa H .n;p.); 6) aepHOBKa - O,ll;HOCeMeHHOH IIJI0,!1; c IIJieHqaThIM rrepHKaprrHeM, cpacTalO- ru;ttMCH c ceMeHeM. XapaKTepeH ,n;JIH 3JiaKOB (rrmeHttu;a, pmKh, THMo<}JeeBKa H ,n;p.); B) opex - O,!l;HOCeMeHHOH, pe,n;Ko ,n;ByCeMeHHOH IIJIO,!I; c ,n;epeBHHHCThIM rre- pttKaprrtteM (Jieru;ttHa, OJihxa, 6epeaa H ,n;p.). Con;waua - aTo coBoKyrrHocTh apeJihIX rrJio,n;oB, B03HHKa10ru;ttx IIYTeM cpacTa- HHH 3aBH3eH u;BeTKOB B rrpe,n;eJiax o,n;Horo cou;BeTHH (aHrurnc, Hmirnp, IIIeJIKOBHu;a). IIJio,n;hl, o6pa3oBaHHble H3 e,n;HHCTBeHHoro necTHKa B u;BeTKe, BaIOT nacmoaw,uMu. EcJIH B <}JopMttpoBaHHH rrJio,n;a Ha3hl- yqacTBYIOT .n;pyrtte 'laCTH u;BeTKa (u;BeTOJIO:iKe, OKOJIOll;BeTHHK, ThJqHHKH), 3TOT IIJIO,ZJ; cqHTaIOT AO:JICltbtM. IIJio,n;, c<}JopMttpoBaBIIIHHCH H3 HeCKOJihKHX rrecTHKOB o,n;Horo u;BeT- Ka, Ha3bIBaeTCH cJtOJICltbt.M. IIJio,n;hI 11 ceMeHa pacrrpocTpaHHIOTCH c rroMOIIJ;hIO: BeTpa - MeJIKHe, JierKHe ceMeHa, HMeIOru;He KphIJiaTKH, XOXOJIKH (ce- Meii:cTBO OpxH,n;Hhle); BO,!l;bl - ceMeHa BO)l;HhIX pacTeHHH (KyBIIIHHKa, Ky6hlmKa H ,n;p.); :lKHBOTHhlX IITHll; - HMeIOru;He KpJOqKH, IIIHIIHKH (qepe.n;a, perreHHHK H .n;p.); HMeIOru;He coqHhie OKOJIOIIJIO)l;HHKH (pH6HHa, KaJIHHa, BHIIIHH H ,n;p.); IIyTeM caMOIIPOH3BOJihHOrO pa36paCh!BaHHH ceMHH (6emeHhlH orypeu;, He,n;oTpora H ,n;p.); qeJIOBe:KOM - B X03HHCTBeHHhIX u;eJIHX HJIH HerrpOH3BOJihHO. CeMa CeMa - opraH rroJioBoro pa3MHO:>KeHHH H pacceJieHHH. Pa3BHBaeTcH H3 orr- Jlo.n;oTBopeHHoro ceMH3aqaTKa. CeMH cocToHT H3 3apoabtut - B aapoabtUta, anaocnepMa, ceMennou 1wJ1Cypbt. OCHOBHaH qacTh ceMeHH (MHHHaTIOpHbIH crropo<}JHT - :e:eM pa3JIHqaroT mpu aapoabtuteBbtX opzana: 2n). 3apo,n;h1II1eBhlH KopernoK;
Bomanu1w 202 3apO.D;bIUieBhIH CTe6eJreK c 38pO.D;bIUieBOH rroqKoii; 3apO.D;bIIIIeBhie JIHCTb.H M.H.D;OJIH (,n;Be - y .D;BY.D;OJlbHbIX, o;:i;na - ce- y O.D;HO.D;OJibHbIX). °YqacTOK CTe6eJibKa B 3apO.D;hIUie BbIUie ceM.H.D;OJie:H H83bIBaeTC.H anu1wmUJlb (Ha,n;ceM.H.D;OJihHOe KOJieHo), Httme ceM.H.D;OJreil: -- zuno1wmu.1lb (rro.n;ceM.H.D;OJihttoe KOJieHO). Ce.Me1t1la.fl KOJICypa MHOrOCJl:OHHa.H, 3aw;11w;aeT 3apO.D;hIIII OT BbICbIX8HI1.H tt npem.D;eBpeMeHHoro rrpopacTaHtt.H. Ce.Mella iJeyiJoJlblibtX 6hIB8IOT (pttc. 81 ): 6e3 3H.D;OcnepMa (CeMeHa ¢aCOJII1, 6o60BbIX, ThIKBeHHbIX, KpecT01IBeTHhIX, CJIOJKHOIIBeTHhIX tt ;:i;p.). BOJihIIIa.H qacTh rrttTaTeJihHhIX Bew;ecTs 38KJIJOqeHa B ceMH.D;OJl.HX; c 3H.D;OCIIepMOM (ceMeHa 30HTI1qHbIX, rracJieHOBhIX, TOMaTa, ¢ttaJIKI1, JIIOTttKa H .n;p.). 8H,n;ocnepM OKpymaeT ceMH.D;OJIH. Mem.n;y ceMa,n;oJI.HMtt KOHyc ttapacTaHtt.H, rroqeqKa ew;e He c¢opMttpoBaHa; c nepHcrrepMoM (ceMeHa rB03.ZJ;J1qHhrx 11 .n;p.). Ilepucnep.M - 3arracaIOIJI8.H TKaHb, o6pa30B8HHa.H TKaHbIO 1tyU,e.1l.1lyca B o6JiaCTI1 IIOKpoBOB ceM.H3aqaTKa. Ce.Mella oiJllOOO.!lbHbtX 6brna10T: 6e3 3H,D;ocnepMa ( ceMeHa qacTyxtt 6oJIOTHOH, p.n;ecTa 11 .n;p. ). CeMa ttMeeT <I:>opMy IlO.D;IWBbI, 3H.D;OCnepM yme «C'be,n;ett»; c 3H.n;ocnepMOM (60JihlllI1HCTBO ceM.HH O.D;HO.D;OJibHbIX, HanpHMep ttpttc H .n;p.). BoJihrnaa qacTh ceMeHH - 3H,n;ocnepM, B Hero norpymeH naJioq- KOBH.D;HhIH 3apo.D;hIIII. CeM.H.D;OJI.H IIHJIMHJJ;pHqecKaa, ee HHiKH.H» qacTh (BJiaraJrnw;e) OXB8ThIBaeT KOHYC HapacTaHH.H; ceMeHa 3JI8KOB. 3apO.D;bIIII B 3epHOBKe conpHKacaeTC.H c 3H.D;OCnepMOM o;:i;noH: cToponoii - ceM.H.D;OJieii HJIH LQHTKOM, npHmaThIM K an;:i;ocnep- MY. Ill,HTOK ( CeM.H.D;OJI.H) BhIIIOJIHHeT cpyHKIIHIO BC8ChIB8HH.H IIHTaTeJihHbIX Beru;eCTB M3 3H,D;OcnepMa. 8H,n;ocnepM co;:i;epiKHT opraHHqecKHe H IlHT8TeJibHbie MHHepaJibHbie Bew;ecTBa. OpranHqecKHe Bew;ecTBa: 6eJioK (KJieiiKOBHHa), yrJieBO.D;hI (KpaxMaJI), iKHphI (paCTHTeJihHbie M8CJia). MHHepaJibHhie Bew;ecTBa: BO,D;a H MHHepaJibHbie COJIH. !Ipopacmallue ce.M.flll - 3TO nepexo.n; H3 COCTO.HHH.H IIOKO.H K pa3BHTHIO H pocTy aapo.D;hIIIIa. ,ll;JI.H npopacTaHHa aeo6xo;:i;ttMhI onpe,n;eJieHHhre ycJIOBH.H: 3 6 7 a Pr.tc. 81. CTpoeH111e ceM5lH ABYAOllbHblX: a - 6 6e3 3HAOcnepMa; B 6- c 3HAOCnepMoM; B - c nepi-1cnep- 1 - KopewoK; 2 - 30111KOTll11lb; 3 - C8M6HHa5l KO)l(ypa; 4 - rnnOKOTll!llb; 5 - C8MSlA0115l; 6 - 3HAOCnepM; 7 - nep111cnepM MOM.
l{naccut/JuKau,uR noKpbtmoceMe1t1tbtX pacmenuii npOHHKHOBeHHe BO)J;hI: IIpHBO)J;HT K 203 Ha6yxaHHIO, aKTHBH3aD;HH <Pep- MeHTOB ll MCTa60JIH3Ma; TeMnepaTypHhlll peJKMM: XOJIO)J;OCTOHKMe paCTeHHJI npopacTaIOT npu TeMnepaType + 1 ... + 5 C (pOJKh, MOpKOBh H ,a;p.); Tern10JII06HBh1e npopacrn+ 15 'C (orypil;hl, TOMaThI, KYKypy3a H .D;p.); IOT IIPH TeMnepaType Bhillle KHCJIOpO)J; B03,D;yxa: .D;hilllaT KaK cyxne, TaK ll npopacTaIOllJ;MC CeMeHa. Ilptt Ha6yxaHHH ceMeHM .D;hrxaHtte ycttJIMBaeTCJI, 06pa3yeTcJI AT<I>, H.D;YT 6ttOCHHTeTuqeCKHe npou;eCChI. IIpopacTaHue 6hrnaeT Hati;3eMHhIM u IIO.D;3eMHhIM. H aiJae.~uwe npopac mm1ue - ceMJI.D;OJIH BhIHOCJITCJI Ha noBepxHoCTh, 3eJie- HeIOT, CTaHOBJITCJI nepBblMll aCCHMHJIMPYIOllJ;MMll JIHCThJIMll (<}laCOJih H .D;p.). IIoiJ3eMIWe npopacmaHue - CeMJI.D;OJIH He BhIXO)J;JIT Ha IIOBepxHOCTh, CMOp- ru;1rnaIOTCJI, OTMttpaIOT. AccttMHJittpyIOllJ;MMH JIMCThHMH cTaHOBJITCJI cJie.D;yIOru;ne 3a ceMH)J;OJIHMll HaCTOJillJ;He JIHCThJI (ropox, .D;y6 H ,D;p.). KJiaccm1>mcau;m1 110Kp1>1ToceMeHHLix pacTeHHii 0T.D;eJI IloKphIToceMeHHhie ,D;eJIHTCJI Ha .D;Ba KJiacca: iJo.TlbHbte. J(ByiJo.Tlbflbte H OiJtto- 8TH KJiaCChI xapaKTepM3YIOTCJI pHti;OM npH3HaKOB. KJiacc 0,1\HO.l\OJibHbie K;rncc .rt:BY.l\OJihHhre 3apo,l\hlIII o6h!'IHO c ABYMH ceMHp;mrnMH, 3apop;bilII o6hI'IHO c 0,11;HOM ceMH,1l;OJieii, KOTOphle IIPH rrpopacTam1H BblHOCHTCH KOTopaSI rrptt rrpopacTaHHH ceMeHtt Hap; aeMJieii OCTaeTCH rrop; 3eMJieii JIHCThH rrpoCThie HJIH CJIO)KHbie, o6b!'IHO JIHCThH BCerp;a rrpOCTbie, o6b!'IHO He '!eTKo paap;eneHhI Ha qepeIIIoK paap;eJieHhl '!eTKO Ha qepernoK H IIJiaCTHHKY ll IIJiaCTHHKY JKHJIKOBaHHe JIHCTbeB 06h!'IHO rrepHCTOe :IB'.ttJIKOBaHHe JlllCTbeB o6hl'!HO HJIH IIaJih'laToe rrapaJIJieJihHOe HJIH p;yroBHp;Hoe XapaKTepHo BToptt'IHOe yTomu;eHtte KaM6ttii B oceBbIX opraHax H BTOpH'IHhlii CTe6JISI (BTOPH'!Hblll pocT) B pe3yJibTaTe HX pocT OTCYTCTBYIOT; rrpOBOASIIIJ;aH p;eHTeJibHOCTH KaM6HSI; rrponop;mu;aH CHCTeMa B BHp;e OTp;eJibHbIX aaKpb!TblX, CHCTeMa CTe6JISI B BHp;e I~HJIHHp;pa; MMe- p;mp<tJyaHo pacnoJiomeHHbIX B crn6Jie eTCH $JI03MHaa rrapeHXHMa; KOpa rryqKOB; $JI03MHaH rrapeHXHMa H cepp;u;eBttHa xoporno OTCYTCTByeT; HCHO BhlpameHHOM KOPhI AH$$epeHu;ttpoBaHhI H cepp;u;eBHHhl HeT IlepBH'!HbIM IWpernoK 06bi'IHO IlepBH'!Hblll KopelIIOK paHO OTMllpaeT' paaBHBaeTCH B rJiaBHblll KOpeHb, OT 3aMeHHHCh rrpHp;aTO'!HhlMH KOPHHMH; KOToporo OTX0,1J;HT 60K0Bble KOpH11; KOpHeBaH CllCTeMa MO'IKOBaTaH KOpHeBaSI CHCTeMa qaru;e CTepmHeBaSI Il;BeTKll B CBoeii OCHOBe 'laru;e Bcero 5- HJill 4-qJieHHhle IJ;seTKll B cnoeii ocHoBe qaru;e ncero 3-qJieHHbie, O'leHb pep;KO 4- llJill 2-'IJieHHbre ~
Bomanu1w 204 OcHOBHbie ceMeHCTBa KJiacca ,ll;By,n;oJibHLie (KpecTou;BeTHbie, Poaou;BeTHbie, BoooBbie, IlacJieHOBbie, CJimKHOIJ;BeTHbie, MaJibBOBbie, MapeBbie, Buaorpa,n;Hbie) CeMeMcTeo KpecToueeTHble (KanycTHb1e) (p1t1c. 82) CeMeiiCTBO HacqMThIBaeT OKOJIO lKU31teltHbte <jJop.Mbl - 3 Thlcaq BM.ll;OB pacTemiiL pacTeHMR o6oerrOJibie, TpaBHHMCThie, 0,IJ;HO- M .n;By- JieTHMKM, MHoroJieTHMKM. BcTpeqaroTCH KycTapHMKM H rroJiyKyCTapHMKH. JlucmbH - rrpOCThie, O'lepe.n;Hhle, 6e3 rrpMJIMCTHHKOB; 6hIBaIOT pacqJie- HeHHble. a eemlCU - o6oeIIOJibie, C ,IJ;BOHHhIM OKOJIOIJ;BeTHHKOM. M3 qeTblpex qaIIJeJIMCTMKOB, 6eH'lUIC - lf aute'llCa rreCTKOB meJITa.H, 6eJiaa, peme <lJMoJieTOBa.H M po3oBa.H. Tbt'lUHOK .n;:Be KOpOTKMe, 'leThlpe ,IJ;JIMHHh!e. COCTOHT M3 qeThlpex JierrecTKOB. 0KpacKa Jie- IIecmuK - IIJeCTh; o.n;HH, M3 .n;Byx cpocIIJMXC.H IIJIO.ll;OJIMCTMKOB. lIJloa - CTpyqoK MJIM CTpyqo'leK. Il;BeTKM co6paHbI B cou;BeTMe KMCTb HJIM MeTeJIKa. XapaKTepHo HaKOIIJieHMe rJIMK03MJJ;OB, KOTOpbie npM pacrn;enJieHMM .n;a10T e,n;KHe 3<lJHpHhle MacJia (ropqJiqHoe MacJio). BamHh!M rrpM3HaKoM .HBJI.HeTc.H orryIIJeHMe BereTaTMBHhIX M reHepaTHBHhIX opraHOB. 3Ha'leHMe: OBOIIl;Hhie M KOpMOBh!e KYJihTYPhI TypHerrc H ,n;p.; MaCJIMqHhre pacTeHMH - KarrycTa, perra, pe,n;MC, rop'lMu;a, pane H .n;p.; ,n;eKopaTMB- 2 4 1 3 PMc. 82. Kanycrn oropOAHas:t ceMeiiicTBa Kpecrnu,aeTHb1x: 1 - cou,serne; 2 nnocKOCTIA; 3-A11arpaMMa U,BeTKa; 4 - cxeMa u,ser;a Ha aHAPOU,ei/i (a) 11 rl1Heu,ei1 (6) U,BeTKa c HeKT8PHl1KOM (a)
OcnoBnbte ce.11ieiicm6a 1aiacca able - ,a ByiJo.nbnMe 205 HO'IHa.H qmamrn H ,n;p.; copH.HKH - cyperrKa, rracTyUih.H CYMKa H ,n;p. <t>opMyJrn u;BeTKa KpecTou;BeTHhix: *Ca,1Co 4A 2 + ,1G( 2). <t>opMyJia IJ;BeTKa (HCIIOJib3yeMbie o603HaqeHH.H): Ca (Calyx) - qaUieqKa; Co (Corolla) - BeHq1rn; p - rrpOCTOM OKOJIOIJ;BeTHHK; A (aH,n;pou;e:ii:) - Thl'HIHKH; G (nrneu;e:ii:) - rrecTHKH; *- t- aKTHHOMOp<)_}HblM u;BeTOK MHOrOIIJIOCKOCTHOM CHMMeTpHH; 3HrOMOp<lJHbIM IJ;BeTOK, o,n;Ha BepTHKaJihHa.H IIJIOCKOCTHa.H CHMMeTpH.H; ( ) - cpocrrrnec.H aJieMeHTbI u;BeTKa; qepTa Ha,n u;w<tJpo:ii: yKa3hIBaeT BepxHee HJIH rro,n; u;w<tJpoii Hm1rnee rroJiomeHHe aJieMeHTOB. CeMeMCTBO Po30L\BeTHbte CeMeiicTBO HacqwThIBaeT 6oJiee /KU3HeHHble </Jop.Mbl - 3 TbIC.Hq BH,D;OB. O,D;HOJieTHHe H MHOroJieTHHe TpaBhI, KycTapHHKH, ITOJIYKYCTapHHKH,,n;epeBb.H. JlucmbR - oqepe,n;Hhie, rrpOCTbie H CJIOJKHbie, c rrpHJIHCTHHKaMH. i.{BemKu - 06oerrOJibre, c ,n;BOMHbIM OKOJIOu;BeTHHKOM. Cou;BeTH.H 6oTpHqeCKHe (KHCTh, 30HTHK, ~HTOK) H U:HM03Hhie (pa3BHJIHHa, H3BHJIHHa). XapaKTepHa.H qepTa - o6pa30B8HHe ?UllaHmUR ( 'lllUlU). fHIIaHTHH B03HHK8IOT H3-38 cpacTaHH.H IJ;BeTOJIOJKa c OCHOBaHH.HMH qarueJIHCTHKOB, JierreCTKOB, ThJqHHOK. fHrraHTHM yqacTByeT B <t>opMHpOBaHHH JIOJKHhIX ITJIO,ll;OB. Il.HTH, HHOr,n;a H3 qeThipex qarueJIHCTHKOB; 6eH'lU1C - £/aute'l1Cll COCTOHT Jl3 H3 Il.HTH (pe,n;KO H3 qeThl- pex) CBo6o,n;HhIX JierreCTKOB. 0KpacKa 6eJia.H, meJITa.H H po30Ba.H. B KOJIHqecTBe KpaTHOM II.HTH. 'IHHKH - cma.Mu1toouu - IIecmuK - Tbt'lUHKU o,n;HH HJIH MHoro. BecrrJio,n;Hhie ThI- 06pa3yroT HeKTapoHOCHhI:ii: BaJIHK. HaceKoMoOrrhI- JI.HeMhie. CeMeHa 6e3 aH,n;ocrrepMa. IIJtoo - cppara (3eMJI.HHHKa H ,n;p.), MHoroKOCT.HHKa (MaJIHHa H ,n;p.), u;wHapo,n;w:ii: (IDHIIOBHHK H ,n;p.), .H6JIOKO (.H6JIOHeBhie H ,n;p.), O,ll;HOKOCT.HHKa (CJIHBOBhie H ,n;p.). CeMe:ii:cTBO Po30IJ;BeTHble ,n;eJIHTC.H Ha 'IeThlpe rro,n;ceMe:ii:cTBa: Po306bte, H6JtoHeBbte H 3Ha'IeHHe: IlJIO,ll;OBhie H .HrO,D;Hhie KYJihTYPhl - .H6JIOH.H, BHIIIH.H, CJIIrna, 3eM- JUIHHKa, MaJIHHa H ,n;p.; JieKapCTBeHHble pacTeHH.H PaTHBHhie - CnupeitHbte, CJtu606bte. IIIHITOBHHK H ,n;p.; ,n;eKO- po3hl H ,n;p. CeMeio1cTeo 606oeb1e (MoTblnbKOBb1e) CeMe:ii:cTBO Bo6oBble HacqHThrnaeT 18 ThIC.Hq BH,D;OB pacTeHHM. 0,n;Ho H3 Ha- H6oJiee MHoroqwcJieHHhrx. }Ku3nennbte </Jop.Mbl - O,ll;HOJieTHHe H MHOrOJieTHHe TpaBbl, KyCTapHHKH, llOJiyKyCTapHHKH,,n;epeBb.H.
Bomauu1w 206 Xa paKTepttCTHKH nonce:ueitcTB --- i-------1---------I MuoromrcTOBKa MuoroopeweK .H6JIOKO 0.n;HOKOCTRHKa MuoroKOCTRHKa I -----IIo.n;qanrn:e lleT HeT H:vieeTCR y Tpas I IIeT IIpe.n;cTaBnTemr Crrnpea Ma,1nua, I .H6JIOHR, CnnBa, MHHae:viJIRHHKa I pa6mra .n;aJI~BHWHR <l>opMyJihI *Ca 5Co 5AJ}= *Ca 5Co 5AxG I *Ca 5Co:,A=Gt 3 _5) *Ca 5Co,,AooG 1 *Ca 5 + 5 Co 5 Ao£~ I I __J_ _ _____.__ ~. ~---·'Tun JIJro.n;a ~- 00 JiucmbJt - CJIOJKHhre (Tpo:i1qaTo- - 1 KJiesep tt .n;p., rraJihqaTo- - momrn H ,Lip., rrepttcTOCJIOJKHhie - ropox H ,Lip.); JIHCTopacnoJiomeHtte oqepe.n;Hoe, peme cyrrpOTHBHOe; c rrpHJIHCTHHIWMH. Il,BemKU - co6paHhI B 6oTpttqeCKHe COllBCTHH (KHCTh, rOJIOBKa, 30HTI1K ll KOJIOC). IJ;seTKH o6oerrOJihie. 0KOJIOllBeTHHK ,ll;BOHHOM. q awe'lKG COCTOHT ll3 Il.HTH cpOClllllXCH qarneJIHCTHKOB, 60l'lUIC - H3 Il.HTH JieneCTKOB, pa3,LleJihHOJierrecTHOH (sepxHHM JieITeCTOK - napyc, ,UBa 60KOBhIX JierrecTKa -- BecJia, ,l(Ba HHJKHHX cpocIIIHXCH - JIO,L101 ma). Tbt'lUIWK - .n;ecaTh; O)J;Ha cso6o.n;HaR, ocTaJihHhle cpocJIHCh B Tpy6oqKy. llecmuK - o.n;ttH. HaceKoMoonhIJrneMhie, caMoorrhIJIHeMhie (ropox H ,Lip.). IJJtoo - MOHOKapnttli:, 606, o.n;Ho-, .n;By- HJIH MHoroceMeHHoli:. CeMeHa 6e3 ::lH,!l;OCnepMa. XapaKTepHo o6pa3oBaHtte KJiy6eHbKOB Ha KOPHRX. KJiy6eHhKH - pe3yJihTaT CHM6H03a c a3oT<f>HKCHpyJOru;HMH 6aKTepHHMH, ycBaHB8JOlll;HMI1 aTMOc<f>epHhIM a30T. B ceMeHax co.n;epJKHTCH MHoro 6eJIKa. 3HaqeHtte: rrww;eBhie pacTeHHH ropox, coa, <PacOJih, qeqeBttU.a 11 .LIP· KopMOBhie pacTeHHH - JIJOI1epHa, JIH.n;BeHeu;, KJieBep H .n;p. Me,LIOHOCHhie - JIJOITHH, ,LIOHHHK. JleKapCTBeHHhie pacTeHHR - KaCCHR, HJIH aJieKCaH.n;pttli:CKHH JIHCT (cJia6ttTeJihHOe), co<f>opa HITOHCKa.H (yKperrneHHe cocy,LIOB) H .LIP· <l>opMyJia u;BeTKa Bo60Bb1x: tca, 5>Co 1, 2, , 2,A, 9>, 1G 1. CeMeHCTBO nacneHOBble CeMeli:cTBO IIacJieHOBhie HacqttThrnaeT 2900 BH,!l;OB. B OCHOBHOM MHOl'OJieTHHe TpaBhl H.'IH IIOJiyKyCTapHHKH, peme KycTapHHKH HJIH ,LlepeBhH (B TpOITHKax). ')/(u3H€HHble </JOpMbl -
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210 BomanuKa Ba1nepHH BaKTepntt 6hJJIH om1caHbI ro"1"1aH,n;CI<HM yqeHhIM AHTOHH BaH JieBeHryKOM B 1676 r. Bee orpoMHoe pa3Hoo6pa3ne 6aKTepttif KJiaccm!rn!IHPYIOT no <t>opMe, OTHOwettmo K Te:vinepaTypHOMY pemnMy, cnoco6y ITHTaHH.H H ITOJIY'IeHH.H 3HeprHH, poJIH B )KH3HH qeJioBeKa (CM. pa3,n;e.1 «U:HTOJIOrmI» ). <l>opMhl 1rneToK 6aKTepuii: naJ10'l!C06UO/-lbte, H,'IH 6au,UJIJlbt: K HMM OTHOCJITC.H KHIIIeqHa.H IIaJIOqKa, ceHHaJI ITa.noqKa ( O,L(HHOqHhie 6aKTepHH), 6aIJHJIJibI CH6npe.H3BeHHhie ( co6paHhI B IJeITO'IKH); !COK/CU (IIIapHKH): Ofrhe,n;HHeHHbie no ,[(Be - ,n;HnJIOKOKKH (B036y,n;HTeJIH nHeBMOHHH); uerroqKH - CTpCITTOKOKKH (B036y,n;HTeJIH aHrHHhI); CKJieeHHbie no qeTbipe (caplI,HHbI); CTaq>HJIOKOKKH o6pa3yIOT «rp03,L(b»; eu6puO/-lbl: HMeIOT H30rHyTyIO ¢opMy (XOJiepHbIM BH6pHOH); cnupu.n.nbt H cnupoxemot: nn:TeBH.L\HhI n: 3aRpyqeHbI ITO crrnpaJin, pa3Jrn:qaIOTC.H ITO IIO,D;BI-IJKHOCTH. BJie,n;naJI cnnpoxeTa - B036y,n;n:TeJib cn:¢HJIHCa. Ilp11 HaCTYIIJieHHH He6JiaronpmITHhIX ycJIOBHM 6aKTep11n: o6pa3yIOT cnopbI c TOJICTOH o6oJioqKoii. CITopbI 6aKTepnM: cJiymaT )J;JI.H rrepeHeceHHJI He6Jiarorrp11aTHhlx yc.noBn:ll cpe,n;br, cnoco6HhI BbI,n;epmnBaTb oqeHh BhIcoKne 11 oqeHh HH3Kne TeMnepaTyphr. Ilo OTHOIIIeHHIO K TeMuepaTypHOMY pe'1CHMY 6aKTepnn ,n;eJIJITC.H Ha HeCKOJibKO rpyurr: mepMorjJUJlbl (o6IITaIOT B KHII.HIIJ;HX KJIIOqax); Me3orjJuJLbt (o6nTaIOT npn TeMrrepaType 36-37 C, 6oJihIIIHHCTBO BhI3hIBaIOT pa3JIHqHhie 3a6oJieBaHH.H); ncuxporjJu.not (o6nTaIOT rrpn 0 ... -1 'C; B ropax «KPOB.HHhre rraTHa» Ha CHery - KOJIOHHH 6aKTepnii-ncnxpo¢HJIOB). Ilo cnoco6y nuTaHHH u noJiyqeunn auepruu 6aKTepnn oqeHh pa3Hoo6pa3HhI. Tttrrhr o6MeHa Be:rn;ecTB y 6aKTepnii onpe,n;eJI.HIOT no rrpnpo.n;e HCToqHHIWB Tpex Heo6xo,n;HMbIX KOMnOHeHTOB JKI13HH: ucmO'l/-lUI( ;mepiuu, ucmff'l/-lUI( yiJLepoaa, ucmo'LHUK eoiJopoiJa (Bo,n;opo,n; HymeH RaK ,n;oHop aJieKTpOHOB: H 2 ~ 2Ht + 2e) H MOJKeT 6hITb 3aMeHeH ,n;pyrHMH 3JieMeHTaMII, OT,n;aIO:rn;IIMl13JieKTPOHhI. Ilo HCToqHIIKY 3Heprnn: rjJomomporjJbl (IICIIOJib3YIOT COJIHeqHbIH CBeT); xe;nompor/Jbt (ncnoJih3YIOT ;rneprmo xnMnqecRnx CB.H3ett B HeopraHnqeCKIIX Be:rn;ecTBax). CJie,n;yeT OTMeTHTh, qTo 6aRTepHII q>oTOTpo¢bl B KaqeCTBe HCTOqHnKa :3JleKTPOHOB mm BO,n;opo,n;a IICIIOJih3YIOT He BO,n;y' a ,n;pyrne coe,n;nHeHH.H. Ilo3TOMY rro6oqHoro npo,n;yKTa KHCJIOpo.n;a, KaK rrpn ¢oTOCHHTe3e pacTeHIIH, He BhI,L(eJIJieTC.H. 3To anoKcuie1-t1-tbtU (6ecKucJLopoiJnotu) rpomocunmea. IJ;naHo6aRTepnn, HanpoTnB, BbI,L(eJI.HIOT KHCJiopo,n; B xo,n;e ¢oTOCHHTe3a (OKCU?e/-l/-lblU rjJomocuume.1). Ilo ncToqHnKy yrJiepo,n;a: aemompor/Jbt (IICIIOJib3YIOT yrJieKHCJibIM ra3 H3 B03,n;yxa);
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