3-4).
CTpoeHMe npoeMT&MMHa A (1 ),
BMT&MMHa A (2) M ero npOM3BOAH1>1x (3, 4)
1)
|
|
|
13-KapornH |
19 |
|
20 |
|
9 |
11 |
13 15 CH20H |
|
~10 |
~12 |
~14 |
|
2) |
|
|
|
PernHon |
|
|
|
|
0 |
0 |
|
|
II |
II |
|
|
C-H |
C-OH |
3) |
|
4) |
|
PernHanb |
|
PernHoeeasi KlllCilOTa |
IUMH KapoTMHOH.UhI B aKTMBHYIO <J:>opM)'mrraMMHa A.
Cyro1111aB mrrpe6uocTL BMTaMMHa A B3pOCJioro
'feJIOBeKaCOCTaBJUieT OT 1 ,Il,0 2,5 Mr BMTaMMHa MJIM OT 2 )J;O 5 Mf ~-KapOTMHOB. 06bI'-IHOaK-
Tl1BHOCTh BMTaMMHa A B rrnmeBhIX npo.n;yKTaX Bhipa:lKaeTCH B Me)l(,Il,yHapO.D;HbIX e,IJ,MHMUax;
O,IJ,Ha Me)l()J;yHapO,Il,HaH e;:i:11H11ua (ME) BMTaMMHa A 3KBl1BaJieHTHa 0,6 MKr ~-KapoTMHa M
0,3 MKf BMTaMMHa A.
BHoJiollfllecKHe <l>YHKU,HH eHTaMHHa A. B opraHl13Me peTMHOJI npeBpamaeTCH B peTMHaJib 11 peTMHOeByro KMCJIOTY' y'faCTBYJOIUMe B peryJIHU1111 pf!)J;a <l>YHKU11H (B pocTe M ,Il,tt:<J:><J:>epeHUMPOBKe KJieTOK); OHM TaK:lKe COCTaBJIHIOT <J:>o- TOXMMl1'-1ecey10OCHOBY aKTa 3peHMH.
Ha116onee ;:i:eTaJihHO M3y'feHoyqacT11e BMTaMMHa AB 3pMTeJihHOM aKTe (ptt:c. 3-3). CBeTOqyBCTBMTeJihHhIH annapaT rna3a - ceT'laTKa. Tia;:i:a10IUMH Ha CeT'laTKYCBeT a;:i:cop6tt:pyeTCH M lpaHC- <J:>opMMpyeTCH IlMfMeHTaMM ceT'laTKMB ,Il,pyryro <J:>opM)'3Hepm11. Y qenoBeKa ceT'laTKaco;:i:ep-
)l(MT 2 TMila peuenTOpHbIX KJieTOK: IlaJIO'-IKM11 Kon6o'-IKM. TiepBhie peampyIOT Ha cJia6oe (cy- Mepe'-IHoe)OCBeIUeH11e, a KOJI60'-IKM - Ha xopornee ocBemem1e (;:i:HeBHoe 3peHMe). TiaJio'-1- KM co.n;ep:lKaT 3pMTeJihHhIH IllffMeHT pO,Il,OilCMH, a KOJI60'-IKM - ito;:i:oncMH. 06a ntt:rMeHTa - CJIO:lKHble 6eJIKM, OTJIM'faIOIUMeCHCBoeH 6eJIKO-
Bow 'faCThIO.B Ka'fecrneKo<J:>epMeHTa o6a 6en-
Ka co;:i:ep:lKaT l l-u11c-peTMHaJih, aJih,Il,erH,UHoe npoM3BO,Il,HOe BMTaMHHa A.
PeTHHOeBaH KHCJIOTa, nono6Ho cTepo11.n;HhIM ropMoHaM, B3aHMo;:i:eitcrnyeT c peuenTopaMH B H)J;pe KJieTOK-MMllleHeH. 06pa30BaBIIIMHCSl KOMIUleKc CBSl3hIBaeTCSl c onpe;:i:eneHHhlMH yq.a-
CTKaMH ,IJ,HK H CTHMYJIHpyeT lpaHCKPHilUHIO reHOB (cM. pa3.n;en 4). EenKM, 06pa3YJOIUMeCS1 B pe3yJihTaTe CTHMYJISIUHH reHOB ITO)); BJIHSIHl1eM peTMHOeBOH KMCJIOThl, BJIHSllOT Ha POCT, ,Il,H<J:><J:>epeHUHPOBKY, penpo.n;ym1110 lf 3M6pMOHaJihHOe pa3BHTHe (p11c. .
0CHOBHbie KJIHHH11eCKHe npoJ1BJieHW1 fHDOBHTllMHHOJa A. HaM60Jiee paHHHH 11 xapaKTepHhIH npM3HaK He.n;ocTaTO'-IHOCTHBMTaMMHa A y n10nei1 11 3KCnepMMeHTaJihHhIX )l(lfBOTHbIX - HapyrneHMe CYMepe'-IHoro3peHHS1 (reMepaJion11S1, MJil1 «KYpHHaH» cnenorn). Cneu11<J:>M'IHOMSI aBMTaMHH03a A nopa:lKeH11e rna3Horo S16JI0Ka - KCepo<J:>TaJihMHSl, T.e. pa3BMTHe cyxocTH poroBOH o60JIO'-IKMfJia3a KaK CJie)J;CTBMe 3aKYfiOPKM CJieJHOfO KaHaJia B CBSl3H c oporoBeHHeM 3ITMTeJIMSI. 3To, B CBOIO O'fepe,Il,h,npMBO,Il,MT K pa3BlfTHIO KOHbIOHKTHBHTa, OTeey, H3bSl3BJieHMIO H pa3MHf'feHHIOporoBOH o60JIO'-IKM,T.e. K KepaTOMMHUMM. Kcepo<J:>TaJihMMSI M KepaTOMaJISIUMSI npH OTCYTCTBl1H COOTBeTCTBYIOIUero Jie'feHMSI MOryT npMBeCTH K IIOJIHOH IlOTepe 3peHMSI.
POAOnC~H
(onc111H-11-4111c-peT111HaJlb)
3Heprnsi ceeTa
onc~H
2
1..t111c-peT111HaJlb |
TpaHc-peT111Hanb-onc111H |
|
HepeHblH lllMnynbc
PeT111Hanb- 5
Mo3ra
11130Mepa3a
TpaHc-peT111HaJlb
TpaHc-pernHon
PMC. 3-3. CxeMa 3pMTenbHOrO LIMKll8. 1 - 1..1111c-pernHanb B T0MHOTe C00jll/IHSl0TCSI c 6enKOM OflClllHOM, o6pa3ysi poaonc111H; 2 - nC>,Q jl0MCTBlll0M KBaHTa CeeTa npOlllCXOAMT ctJoTOlll30M0p1113al.llllSI 11-L.llllC-peTlllHanSI B TpaHc-pernHanb; 3 - -rpaHC-peTlllHanb-OflClllH pacnaaaeTCSI Ha TpaHC-peTlltHanb Ill onc111H; 4 - flOCKOflbKY nlllrM0HTbl BCTpoeHbl B M0M6paHbl CB0TO"IYBCTBlllT0flbHblX Kn0TOK C0T"laTKlll, 3TO np111BOAlllT K MecTHOM aenonsip1113al.lllllll MeM6paHbl Ill B03Hl/IKHOB0HllllO HepeHoro lllMnynbCa, pacnpocTpaHS!IOUl0rOCSI no HepBHOMY BOflOKHy; 5 - 3aKnlO- "llllT0nbHblM 3Tan 3TOro npo1..10CCa - pereHepa1..1111si lllCXC>,QHOro n111rMeHTa. 3TO npolllCXOjll/IT np111 yYaCTllllll peT111Hanblll30Mepa3bl Yepes CTajlllllll: TpaHC-peT111Hanb ~ TpaHc-pernHon ~ L.llllC-peTlllHOn ~ 1..1111c-peT111Hanb; nocnejlHlllM BHOBb C00jll/IHSl0TCSI c onClllHOM, o6pasysi poi::ionClllH.
|
|
|
|
|
|
|
|
|
|
|
|
|
PeTMHaJlb __... |
Per1.1Hoeeasi |
|
|
|
|
|
IKapoTMHO-l _.A. |
|
|
|
|
|
t |
KMCJlOTa |
|
|
|
|
~ |
og2 |
GI±IJJ |
|
|
|
|
|
|
|
|
A. AA |
A111cpcpepeH1.1111poeKa |
|
|
8Jll!ISIHl.1e Ha |
Cn1Mynsi41.1si |
111 pereHepa4111si |
|
|
BaKTe |
3nMT9111.181lbHOM TKBHlll |
|
|
penpOAYKLIMIO |
apeHMSI |
pocTa |
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
PMC. 3-4. J:leMCTBMe peTMHOMAOB B opraHM3Me. BeUISCTBa (Ha3BaHllll'IB paMKax) - |
KOMnOH0HTbl nlllUllll. |
y )leTett H MOJIO)lbIX )KHBOTHhIX rrpH aBHTaMHH03e A Ha6JIIO)laJOT oeTaHOBKY poem KOeTett, \Kepam3 3IIHTeJIHaJibHbIX KJieTOK Beex opraHOB H, KaK CJie)leTBMe 3TOfO, M36hJTOqHoe oporoBeHHe KO)KH, rropaxeHHe 3IIMTemrn )l(KT, MoqerronoBott eHeTeMbI H )lbIXaTeJibHOfO arrrrapam. IlpeKpaw:eHHe poem KOeTett qeperra rrpMBO)lMT K IIOBpe)l()J;eHMIO TKaHett llHC, a mIOKe K IIOBbIrneHMIO )laBJiemrn errMHHOM03fOBOH )KH)J;KOeTH.
2. BumaMUHbt zpynnbt D (KMbU,U<jJepOAbt)
KanbIJmpeponhI - rpyrrrra XMMMqeeKH poaCTBeHHbIX COe)lHHeHHW, OTHOCJIW:MXCJI K rrpOM3BO)lHblM cTepttHOB. Hatt6onee 6ttonorttqecKM aK-
THBHbie BMmMHHbI - D2 H Or BttmMMH D2 (3proKaJibIJHcpepon), rrpott3BO)lHOe 3proeTepHHa -
paCTMTeJibHOfO eTepoMD.a, BCTpeqa10w:erocJI B HeKOTOpblX rp116ax, LlPO)JQKaX H paeTHTeJibHblX MaCJiax. IlpH o6nyqeHMM n11w:eBbIX rrpOi:lYKTOB Y<l>O H3 3procTepMHa rronyqaeTCJI BMmMMH D2, 11crroJih3yeMbitt B neqe6HbIX IJeJIJIX. BttmMMH D 3, MMeIO-
W:MttCJI y |
qeJIOBeKa M )!(HBOTHbIX, - |
XOJieKaJihIJM- |
cpepon, |
o6pa3YIOW:Htte5l B KO)((e |
'IeJIOBeKa H3 |
7-)lerlf)lpOXOJieCTepMHa IIO)l )lettCTBHeM y <l>-nyqew (pMC. 3-5).
BmaMHHbI D2M 0 3 - 6enb1e KPMCTaJIJihI, )((HpHhie Ha ow:yrrb, HepaCTBOPMMhie B BO)le, HO xopo-
GbCH,- CH,-CH, - ~~H,
|
|
|
21 |
|
|
26 |
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
CH3 |
|
CH |
|
|
|
|
|
|
I |
|
/ |
3 |
|
|
|
|
|
18 CH - CH2 - |
CH 2-CH 2 |
- CH |
|
|
|
|
|
|
20 22 |
23 24 |
25'cH 3 |
|
|
|
|
|
|
|
27 |
|
c 8 |
|
|
|
16 |
|
|
|
|
|
II |
19 |
D |
|
|
Y<t>O |
|
6c,,.. c 7 |
|
|
|
15 |
|
|
|
|
ll |
CH 2 |
|
|
|
|
|
|
|
|
|
|
2 |
|
|
|
|
01 |
19 |
HO |
|
|
|
|
|
|
7-.QernApOXOJ19CT9pVIH |
|
|
|
|
|
|
|
|
|
|
HO |
|
A
|
|
|
|
|
|
|
CH3 |
CH 3 |
|
c |
|
|
|
|
|
|
|
I |
/ |
|
|
|
|
|
|
|
CH - CH = CH - CH - CH |
|
|
|
|
|
|
|
|
|
|
|
|
|
II 8 |
|
|
|
|
|
|
|
|
|
|
|
19 |
|
c |
D |
|
|
Y<t>O |
|
c / |
c 7 |
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
116 |
CH2 |
|
|
|
|
|
|
|
|
|
2 |
|
|
|
|
|
|
|
|
|
|
19 |
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
HO01 |
|
|
|
|
8procTepVIH |
|
|
|
|
|
|
|
|
HO
BVIT8MVIH D3
Xo11eKaJlb'-IVl¢epo11
CH 3 |
CH3 |
/CH 3 |
I |
I |
|
CH - |
CH = CH - CH - |
CH |
|
|
' CH 3 |
|
|
|
D |
|
|
|
|
|
BVITaMVIH D2
3proKaJlb'-IV1¢epo11
Ei
PMC. 3-5. CxeMa CMHTe38 BMT8MMHOB 0 2 Iii 0 3• npoBf/IT8MfllHbl 0 2Ill 0 3 - CT0pf11Hbl, y KOTOpblX B KOnbl.l0 B AB0 ABOfiiHbl0 CB~3fll. np111 B03Ae~cre111111 ceera e npo1.1ecce ¢oTOXfllMfll4ecKolli peaKL.lfllfll npo111cxoAfllT pacl.llenneH111e Konbl.l& B. A - 7-A0rfllAPOXonecrep111H, npoe111- T&MfllH 0 3 (c111Hrea111pyerc~ 1113 xonecrep111Ha); 6 - aprocrep111H - npoe111raM111H 0 2.
O |
CH3 |
|
|
9H3 |
9H3 |
WCHrCH=C-[CH2-CH2-CHrCHk CH3 |
0 |
B111TaM11H K1 (cp111111ox11HoH) |
W0 CH3
9H3 (CH2-CH=C-CH2)e-H
0 |
B11TaM11H K2 (MeHax111HOH) |
CTBHe Hapyrnemrn BCaChlBaHIUI BHTaMHHa K B KHIIIe'lHHKe,a He B pe3yJihTaTe ero OTCYTCTBHH B IlHlll,e.
Cyro'lllunOTpe6uocTb B BHTaMHHe BJpocJioro
'leJIOBeKaCOCTaBJUieT 1-2 Mr.
BuoJI0111'1ecKaJ1cl>YllKilWI BHTilMHHa K CBH3aHa c
ero yqacTHeM B npouecce c~pThIBaHHH KpoBH
(puc. 3-6). OH yqacrnyeT B aKTuaauHH <f>aKTopoB cBepThIBattHH Kpoau: rrpoTPoM6uHa (<t>aKTOP II), npoKOHBepTHHa (<f>aKTop VII), <f>aKTopa KpHCTMaca ( <f>aKTop IX) H <t>aKTopa CnoapTa (<f>aKTop X). 3-m 6eJIKOBhie <f>aKTOphI CHHTe3uPYJOTCH KaK HeaKTMBHhre npe.uIIIeCTBeHHHKM. 0.UHH H3 3TallOB aKTHBauHH - HX Kap6oKCHJIHPOBaHHe no OCTaTKaM rJIYTaMHHOBOH KMCJIOThl c o6pa30BaHHeM y-Kap60KCHrJIYTaMHHO- BOH KUCJIOThI, Heo6XO)l,HMOH )l,JUI CBH3hIBaHMH HOHOB KaJihUHH (cM. paJ.ueJI 13). BHTaMHH K yqacTByeT B peaKUHHX Kap60KCHJIHPOBaHHH B Kaqecrne Ko<f>epMeHTa.
,I1,.rrn Jie'-leHMHH rrpe.uyrrpe)((,1leHHH fMilOBHTaMHH03a K wcnoJih3YJOT CHHTeTH'lecKHenpoH3BO.U- Hhie Ha<t>TOXHHOHa: MeHa)l,1'\0H,BH.Kacon, CH.H- KaBHT.
Ocuoeuoe npoJ1BJieHHe aemaMHH03a K - CHJih-
Hoe KPOBOTe'!eHHe,qacTO IlPHBOMlll,ee K IIIOKY H rw6eJIH opraHM3Ma.
B Ta6JIHUe 3-3 rrepeqwcJieHhI CYTO'-IHhierroTPe6- HOCTH H 6HOJIOfH'leCKHe <l>YHKUHH )!GlpopaCTBOPHMhIX BHTaMHHOB, a TaJOKe xapaKrepHhie npH3HaKH aBHTaMHH030B.
<l>oc<f>o1111nlll,D,bl MeM6paHbl TpoM60'-IVITOB
|
|
|
|
-,-----r----....-----r- |
|
|
|
|
'' \ |
I / I |
|
\ '\ |
I / / |
|
|
|
|
|
:Ga++ |
|
|
~a++ |
|
|
|
|
I / / |
\ '\ |
I |
/ / I |
\ \ \ |
|
-ooc coo -ooc |
I |
\ |
|
\ |
H N~,...______....._ |
-ooc |
cocr -ooc coo- |
co2 |
H N+_._____, |
H N+_r---------'L.... |
3 |
|
|
|
3 |
|
|
|
|
|
|
|
|
|
|
|
|
|
\ . |
3 |
I |
|
|
|
|
|
|
_ _____.-coo- |
|
______.-coo- |
II, VII, IX,X |
II, VII, IX, X |
KpOBb |
II, VII, IX,X |
|
|
y-Kap60KC11JlrJl)'TaMaT |
|
3peJlble TpOM60'-IVITbl |
npocpepMeHTbl
neYeHb
PMc. 3-6. Ponb BMT&MMHa K a caepTblBaHMM KpoaM.
138
Ta6mn43 3-3. >KMpopacTBOpMMble BMT8MMHbl
|
HueaHue |
CyTO'IHaH |
bHOJJOl'HlfCCKHCcj>yHKU:HH |
XapaKTepHble npu3HaKu |
|
|
noTpe6nocTb, |
|
aeuTaMHH030B |
|
|
Ml' |
|
|
|
A |
1-2,5 |
YqacTByeT B aKTe 3peHH.ll, |
reMepanOTIHR(KYPHHaH |
|
(peTHHOJI) |
|
peryJittpyeT pocT |
CJieTIOTa), KcepoqnanbMHR, |
|
|
|
H .Lltt<P<Pepemi:HpOBKY KJieTOK |
KepaTOManHUHH,KepaT03 |
|
|
|
|
3IlHTeJIHanbHblX KJieTOK |
|
|
|
|
|
|
D |
0,012-0,025 |
PeryJIHUHH o6MeHa <Poc<Popa |
PaxHT |
|
(KaJibUHcliepoJI) |
|
H KanbUHR B opraHH3Me |
|
|
|
|
|
|
|
E |
5 |
AHTHOKCH.L(aHT;peryJinpyeT |
He.L(ocrnrnqHo HJyqeHbI; |
|
(TOKocliepOJI) |
|
HHTeHCHBHOCTb |
H3BeCTHO IlOJIO)l(HTeJibHOe |
|
|
|
CB060.L(HOpa.L(HKanbHbIX |
BJIHHHHe Ha pa3BHTHe |
|
|
|
peaKUHH B KJieTKe |
6epeMeHHOCTH H npn JieqeHHH |
|
|
|
|
6ecnJIO.L(H.ll |
|
|
|
|
|
|
K |
1-2 |
YqacTByeT B aKTHBaunn <PaKTopoe |
HapyrnettHe cBepTbIBaiomeM: |
|
(tta<PTOXHHOH) |
|
cBepTbrnaHnH KpoBn: II, VII, IX, XI |
CHCTeMbl KpOBH |
|
|
|
|
|
61110ClllHTE3 HYKilElllHOBblX KlllCilOT Ill 6EJ1KOB {MATPlll4HblE 61110ClllHTE3bl). OCHOBbl MOJlEKYJlSIPHOlil rEHETlllKlll
HyKJieHHOBhie KHCJIOThI - BhICOKOMOJieKyJI5rpHhie coe,Il,HHeHH51 co CTporo orrpe,11,eJieHHOH JIHHeHHOH IIOCJie,11,0BaTeJihHOCThlO MOHOMepoB. CTpYJCTYPa )],HK H PHK - crroco6 «3armcH HH¢lopMaUHH», 06ecrreqHBa10rutti1 <PopMHPOBaHHe B opraHH3Me ,11,BYX HH<l>OpMaUHOHHhIX rroTOKOB. 0,11,HH H3 IIOTOKOB ocyrueCTBJI5IeT BOCIIPOH3Be,Il,eHHe HH¢lopMaUHH, 3aKJI10qeHHoi1 B MOJieKynax )],HK. Y,11,BOeHHe MoJieKyJI ,UHK Ha3hIBaJOT «penJIHKa~JI». B pe3yJihTaTe 3Toro rrpouecca H rrocne,Il,ylOruero ,11,enemrn ,11,oqepHHe KJieTKH Hacne,Il,ylOT rettoM po,11,HTeJihCKoii KJieTKH, B KOTOpOM CO,Il,ep)KHTC51 IIOJIHhlH Ha6op reHOB, HJIH «HHCTPYKUHH» o CTPOeHHH PHK H Bcex 6eJIKOB opraHH3Ma.
Bropoi1 rroroK HH<l>opMaUHH peanmyeTCH B npouecce )KH3He,Il,e51TeJihHOCTH KJieTKH. B 3TOM cnyqae npowcxo,11,HT «CqHThIBaHHe», HJIH TpaHC-
KpHIIU,IDI, reHoB B ¢opMe nOJIHHYKJieOTH,11,HhIX rrocne,Il,oBareJihHOCTei1 MPHK H HCIIOJih30BaHHe HX B Kaqecrne MaTPHU JJ)I51 CHHTe3a COOTBeTCTBYJOIUHX 6eJIKOB. B rmcne,11,HeM cnyqae ocyruecTBJI5IeTC51 «rrepeB0,11,» (TJ>aHCJHIUIDI) HH<l>opMauHH, 3aKJIJOqeHHoi1 B MPHK, Ha «513hIK» aMHHOKHCJIOT. 3ToT noTOK HH<l>opMauHH OT )],HK qepe3 PHK Ha 6enoK nonyqHJI Ha3BaHIIe «QeHTpa.JlbHasl .uorMa 6uOJioruu». OH xapaKTepeH 1J)I5I Bcex )KHBhIX opraHH3MOB, 3a HCKJI10qem1eM HeKorophIX PHK-co,11,ep)!(aruHX BHpycoB.
MaTpwrna51 rrpwpo,11,a CHHTe3a HYKJieHHOBhIX KHCJIOT H 6eJIKOB o6ecneqHBaeT BhICOKYJO TQqHQCTh BOCIIpOH3Be,Il,eHH51 HH¢lOpMaUHH. TaK, B X0,11,e peIIJlHKaUHH ,11,0qepHHe MOJieKYJihl )],HK CHHTe3HPYJOTC51 Ha HHnIX MaTepttHCKOH )],HK. IlpH o6pa3oBamrn Bcex BH,11,0B PHK, Heo6xo- ,Il,HMhIX JJ)I51 CHHTe3a 6eJIKOB, HH<lJOpMaUH51 06 HX CTpYK'f)'pe«CqHThIBa-
eTC51» c onpe,11,eJieHHhIX reHOB B MOJieKYJiax )],HK. B CHHTe3e HOBhIX MOJieKYJI 6eJIKOB MaTPm.i:eH., C0,11,ep~rueii HH<l>opMaU,HJO 06 HX CTPOe-
HHH, 5IBJI5110TC51 MPHK.
11cripaBJieHHe oum60K, B03HHKaJOIUHX B CTPYKType )],HK IIO)l B03- )letfCTBHeM <PaKTopoB BHenrnei1 H BHYTPeHHei1 cpellhI, ocyruecTBJI5reT erue 0,11,HH MaTPHqHhIH CHHTe3 - penapaQHJI. OH 5rBJI51eTC51 BapHaHTOM orpaHMqeHHOH perrJIHKaUHH H BOCCTaHaBJIMBaeT rrepBOHaqaJihHyIO CTPYKTYPY ,UHK, HCITOJih3Y5r B Kaqecrne MaTPHUhI yqacToK HerroBpe)[(- )leHHOH HMTH )],HK. Ilptt pa3MHO)[(eHHH PHK-co)lep:)l<aIUHX BHpycoB B KJieTKax 3yKapttoTwrecKHx opraHH3MOB HOBhie MOJieKYJihI )l,H K MOryT CHHTe3HPOBaThC51 c rroMOIUhIO rrpouecca, B xo,11,e KOToporo PHK cey- )l{HT MaTpHuetr )lJI5I CHHTe3a KOMIIJleMeHTapHOtt )l,H K, KOTOpa51 MO)l(eT BKJIJOqaThC5l B reHOM BhICllIHX opraHH3MOB (o6paTHaJI TpaHCKPHDQHJI).
I. CTPYKTYPHASI OPrAHlll3AUlllSI
HYKJlElllHOBblX KlllCflOT
B K3)K)J,OM )KllBOM opraHH3Me rrpttcyTCTBYJOT 2
THrra ttyKJieHHOBblX KHCJIOT: pH6ottyKJieHHOBa51 KHCJIOTa (PHK) H ne30KCHpH60HyKJieHHOBa51 KMCJIOTa (LJ:HK). MoneKYJIHpttaH Macca caMo:H «ManetthKOM» H3 H3BeCTHblX HyKJieHHOBblX KMCJIOT -
rpattcrroprno:H PHK (TPHK) cocrnBJUieT rrpHMepHO 25 K,ll,. )],HK - HaH60Jiee KPYIIHhie ITOJIHMepHbie MOJieK)'Jihl; HX MOJieKYJIHpHaH Macca BaphHpyeT oT 1000 no 1000 000 K,ll,. ,ll,HK H PHK cocToHT H3 MOHOMepttbIX enHHHU - HyKJieOTH)J,OB, II03TOMY HyKJieHHOBbie KHCJIOTbl Ha3bIBaJOT IIOJIHHyKJieOTH,JJ,aMH.
A. CTPOEHIIE HYKJIEOTH)].OB
K3)K)J,bIM ttyKJieOTM)l conep)KllT 3 XHMHqecKH pa3JIHqHbIX KOMIIOHeHTa: reTepOUHKJIHqecKoe a30THCTOe OCHOBaHHe, MOHOCaxapH,JJ, (rreHT03y) H OCTaTOK cpoccpoptto:H KHCJIOThI. B 3aBHCHMOCTH OT qHcJia HMeJOIUHXC51 B MOJieKyJie OCTaTKOB cpoccpopHOM KHCJIOTbl pa3JIHqaJOT HyKJie03H,JJ,MOHOcpoccpaTbl (HMCI>), ttyKJie03HMH<PoccpaThl (H,ll,CI>), ttyKJie03H,JJ,TpHcpoccpaThl (HTCI>) (pHC. 4-1 ).
B cocrnB HyKJieHHOBhIX KMCJIOT BXOMT a30THC-
Th1e OCHOBaHHH ,JJ,BYX THITOB: rrypHHOBbie - |
ane- |
HHH (A), ryaHHH (G) H ITHPHMH)J,HHOBhie - |
~o- |
3HH (C), THMHH (T) H ypan;HJI (U). HyMepauHH aTOMOB B OCHOBaHH51X 3aIIHCbIBaeTC51 BHYTPH UHKna (pHc. 4-2). HoMettKJiazypa ttyKJieOTH,JJ,OB rrpHBenetta B rn6n. 4-1.
IleHT03hI B HyKJieOTH,JJ,ax rrpencTaBJietthl JIH6o pH6o3o:H (B cocTaBe PHK), JIH6o ne30KCHpH6o3o:H (B COCTaBe ,ll,HK). qT06bI OTJilfqlfTb HOMepa aTOMOB B rreHT03aX OT ttyMepaUHH aTOMOB B OCHOBaHHHX, 3aIIHCh rrpOH3BO.Il51T c BHenrne:H CTOpOHhI
UHKJia H K UHcppe no6aBJUIIOT IllTPHX (')- l', 2',
3', 4' H 5' (pHC. 4-3).
IlettTmy coe,JJ,HHHeT c octtoBaHHeM N-mHK03H,lJ,- HaB CBH3b, o6pa30BaHHaH C1-aToMoM rrettT03hI (pH6o3hI mm ne3oKcHpH6o3hI) H N 1-aToMoM rrHpHMH,JJ,HHa HJIH N9-aTOMOM rrypHtta (pHc. 4-4).
HyKJieOTH,JJ,hI, B KOTOphlX rreHT03a rrpencTaBJietta pH6030M, Ha3blBalOT pH6ottyKJieOTH,JJ,aMH, a ttyKJieHHOBhie KHCJIOTbl, rrocrpoeHHhie H3 pH60ttyKJieoTH,JJ,OB, - pH60HYKJieHHOBbIMH KHCJIOTaMH, HJIH PHK. HyKJieHHOBbie KHCJIOThI, B MOHOMephI KOTOpbIX BXO.UHT ,JJ,e30KCHpH603a, Ha3hlBaIOT ,JJ,e30KCHpH6ottyKJieHHOBblMH KHCJIOTaMH, HJIH )],HK. HyKJieHHOBbie KHCJIOTbl ITO CBOeMY crpoeHHIO OTHOCHT K
N~NH2
0 0 0 |
.}" |
~N |
<N |
I ) |
-o-PI -0-PI-O -IP-O2 |
-Co |
|
o- o- o- |
0 |
|
OH OH
AAeH03111H
AAeH03111HMOHocpoccpaT (AM<I>)
AAeH03111HA111cpoccpaT (A,Q<I>)
AAeH03111HTp111cpoccpaT (AT<I>)
P1tc. 4-1. HyK11eo3MAMOHO-, AM- M TpMcj>occj>aTbl aAeH03MHa. HyKneOrnAbl - cpoccpopHble :icp11pb1 HyKne0311AOB. OcTaTOK cpoccpopH0'1Kl1CnOTbl np11coeA11HeH K 5'-yrnepOAHOMYBTOMY neHT03bl (5'-cpoccpo:icp11pHa5!CB513b).
4-6),
N~ N~
|
|
;/ |
N |
--...:::::N |
|
|
--...:::::N |
6- |
H |
H |
1';---______N-rnlllK03111,QHafl 6- |
~.....~1'--- N-rn111Koa111,QHafl |
|
H |
|
CBfl3b |
I ~ I |
CBfl3b |
|
OH |
OH |
|
OH |
i:1l...IM<l> |
|
|
AM<l> |
|
|
|
PMC. 4-4. nypMHOBblM It nitpMMMAMHOBblM HyKlleOTMJ:lbl.
|
|
o- |
5'-KoHeu |
|
|
I |
|
|
|
"b- ~ =0 |
|
Bap111a6e11bHb1e |
|
|
o |
|
rpynnb1 |
|
15 |
|
|
|
|
c~H2 OcH0BaH111e |
|
|
3' |
|
|
3',5'-cpoc¢0A111a¢111pHafl |
|
I |
|
|
CBfl3b |
|
"b-P=o |
|
|
|
I |
|
|
|
0 |
|
|
|
5 |
2 |
0CHOB8Hl/le |
|
|
b~H |
3'
0
I
-0-P=O
I
0
~~CHOoaH•e
IH'
0
I
3'-KoHeU
PMC. 4-5. cJ>parMeHT 1.1en1t ,QHK.
IIOCJie)J,OBaTeJihHOCThIO a30THCThIX OCHOBaHHH B
IIOJilfHYKJieOTH)],HOH LJ,eIIH.
Ii. CTPYKTYPA )J;E30KCHPHliOHYK.JIEHHOBOM
KHCJIOThI (,UHK)
IlepBH'llDUIcrpyicrypa ,l(HK - rropH)J,OK qepe-
)J,OBaHJrn )J,e30KCMpM6oHyKJie03H)J,MOHO¢oc¢aTOB
()J,HM<l>) B TIOJIHttyKJieOTH)J,HOH LJ,eTIH.
KIDK.IlM ¢oc¢aTHM rpynna B rroJIHHyKJieOTH)J,- HOH uenH, 3a HCKJIIO'leHHeM¢oc¢opttoro OCTaTKa Ha 5'-KOHLJ,e MOJieKYJihl, yqacrnyeT B o6pa30BaHHH
)J,Byx 3¢HpHbIX CB5!3eH c yqaCTHeM 3'- H 5'-yrne- pO)J,HhIX aTOMOB )J,BYX coce)J,HHX )J,e30KCHpH603, TI03TOMY CB513h Me)l()J,y MOHOMepaMH o603HalJ:aIOT
3', 5'-ct>occf>o)J,H3¢HPHOH.
KoHueBhie HyKJieoTH)J,hI ,lJ,HK paJJIH'laIOT no CTPYKTYPe: Ha 5'-KOHLJ,eHaxO)J,HTC51 ¢ocq>aTHa5! rpynna, a Ha 3'-KoHLJ,eLJ,errH - CBOOO)J,HM OH-rpynna.
3-m KOHLl,hl Ha3bIBaIOT 5'-H 3'-KOHLJ,aMH. JIHHeHHa5!
TIOCJie)J,OBaTeJibHOCTh )J,e30KCHpH60HYKJieOTH)J,OB B
TIOJIHMepHOH uerrH ,lJ,HK 06bl'IHOCOKpameHHO 3a-
IIHChIBaIOT c TIOMOlllblO 0)],H06yKBeHHOfO KO)J,a, HanpHMep -A-G-C-T-T-A-C-A- OT 5'-K 3'-KoHLJ,y.
B Ka)l()J,OM MOHOMepe HYKJieHHOBOH KHCJIOTbl npHCYTCTByeT OCTaTOK <J:>oc<J:>opHOH KHCJIOThl. Ilptt pH 7 ¢oc¢aTHa5! rpynna IIOJIHOCTbIO HOHH3MpOBaHa, TI03TOMY in vivo HYKJieHHOBhle KHCJIOThl CYJLl,e- CTBYJOT B Bl1)],e IIOJIHaHHOHOB (HMeIOT MHOXeCTBeHHhIH OTPHUaTeJihHhIH JapM). OcrnTKH neHTo3 rnxe npo5IBJIHIOT rH)J,pocf>HJihHhie cBolicTBa. A30THCThie OCHOBaHH5I TIO'ITHHepaCTBOPHMhl B BO)J,e, HO HeKOTOpbie aTOMbl nypHHOBOro H I1MpMMl1)],HHOBOfO Ll,HK-
JIOB crroco6Hbl o6pa30BbIBaTb BO)J,Opomn.1e CBJl3H.
BTOpwmaH CT(JYKTYPa ,UHK. B 1953 r. ,l],x. YOT-
cottoM 11 ct>. KpHKOM 6bma npe,ll.JlmKeHa MO)J,eJih npo-
CTpaHcTBeHHOH CTpYKTYPhI )],HK. CornacHo 3TOH MO)J,eJIH, MOJieKYJia )],HK HMeeT <l>OPMY CTIMpaJIM, o6pa30BaHHYJO )J,BYM5J: TIOJIHHYKJieOTl1)],HhlMH ueMMH, 3aKpyqeHHhIMH OTHOCHTeJibHO J],pyr J],pyra H BOKpyr o6metl OCH. )J,BOHHM crrnpaJib npa&03aK.py-
'1eHHaJ1, IIOJIHHYKJieOTH)J,Hhie LJ,eTIH B HeH aHTllila-
PaJIJleJlbllbl (pHC. T.e. eCJIH O)J,Ha H3 HHX opMeHTMpoBaHa B HanpaBJieHHH 3'~5', TO BTOPM - B HanpaBJieHHH 5'~3'. Ilo3TOMY Ha Ka)l()J,OM H3 KOH-
