vaxtang qobalia

mcenareTa genetikuri
konstruirebis
meTodebi

quTaisi, 2019
1
 naSromSi ganxilulia genetikuri da ujred-
li inJineriis, rogorc mcenareTa genetikuri apa-
ratis konstruirebis da Sesabamisad axali niSan-
Tvisebebis matarebeli mcenareebis miRebis meTode-
bis arsi, gamoyenebuli teqnologiebis Tavisebure-
bebi, miRweuli Sedegebi, warmoqmnili problemebis
gadaWris gzebi. aq warmodgenilia ujredebis done-
ze in vitro manipulaciebis – izolirebuli protop-
lastebis Serwyma, subprotoplastebis Serwyma, uj-
reduli organelebis Seyvana, mutaciebi da sxva Se-
saZleblobebi; in vitro ujreduli seleqciis Tavi-se-
burebebi da Sedegebi; genebis veqtoruli sisteme-
biT da pirdapiri gadatanis meTodebiT mcenareTa
transformaciis xerxebi, problemebis gadaWris Ta-
viseburebebi da miRweuli Sedegebi; biousafrTxoe-
bis sakiTxebi.
gankuTvnilia agraruli mimarTulebis specia-
lobebis magistraturis da doqtoranturis stu-
dentebisTvis.

red aq tori : akaki wereTlis saxelmwifo universitetis eme-

ritusi juli eta san i ki Ze

recenzentebi: akaki wereTlis saxelmwifo universitetis pro-

fesori, ssmma akademikosi roland kopaliani
akaki wereTlis saxelmwifo universitetis aso-
cirebuli profesori n i n o yi fi an i

© akaki wereTlis saxelmwifo universitetis gamomcemloba

quTaisi, 4600, Tamar mefis 59. tel.: 24 00 21
E-mail: [email protected]

ISBN 9 78-9941-484-18-6 © v. qobalia, 2019

2
 Sesavali

mcenareTa genetikuri konstruirebis qveS igulisxmeba

in vitro pirobebSi xelovnurad Seqmnili an sxva ujredidan
gamoyofili gen(eb)is, dnm-s fragmentis, ujredis calkeuli
organelebis, citoplastebis cocxal ujredSi Seyvanis,
aseve protoplastebis Serwymis, ujredebis mutaciuri cva-
lebadobis da sxva meTodebiT winaswardasaxuli Tvisebebis
matarebeli mcenareTa axali ujredebis, Sesabamisad, axali
mcenareuli organizmebis Seqmna.
dReisaTvis mcenareTa genetikuri konstruirebis ori
bioteqnologiuri meTodi arsebobs – ujreduli inJineria
da genetikuri inJineria. winamdebare naSromSi ganxilulia
am meTodebis arsi, mcenareTa genetikuri gardaqmnis SesaZ-
leblobebi da miRweuli Sedegebi.
bioteqnologiuri gziT sameurneo mniSvnelobiT mcena-
rTa genetikuri konstruirebis era iwyeba 1983 wlis 18 ian-
vridan, roca maiamSi (aSS) konferenciaze (Miami Winter Sym-
posium) mari -del Ci l ton ma (Mary-Dell Chilton) vaSingtonis uni -
versitetidan (sietli, vaSingtonis Stati, aSS), jef Sel ma
(Jeff Schell) da mark van montagium (Marc van Montagu) gentis uni-
versitetidan (belgiა) da robert fralei m (Robert T. Fraley) da
robert horSma (Robert Horsch) kompania “monsantodan” (sent-
luisi, misuris Stati, aSS) warmoadgines moxsenebebi Sesaba-
misad, antibiotik kanamicinisadmi gamZle Tambaqos mcenaris,
kanamicinisa da metatrioqsatisadmi gamZle Tambaqos da ka-
namicinisadmi gamZle petunias mcenaris miRebis Sesaxeb.
ukve 1986 wels aSS-Si Catarda genetikurad konstru-
irebuli - transgenuri mcenaris (virusuli infeqciisadmi
gamZle Tambaqo) pirveli warmatebuli savele gamocda.
1992 wels CineTSi daiwyes mavne mwerebisadmi gamZle
transgenuri Tambaqos samrewvelo moyvana.
3
 1994 wels aSS-Si miRebul iqna pirveli oficialuri
Tanxmoba (registrirebul iqna) genetikurad modificire-
bul sakveb produqtebze – firma “Calgen”-is Seqmnil Sene-
lebuli momwifebis pomodvris jiS “FLAVR SAVR”-ze da
kompania “Monsanto”-s mier miRebul herbicidgamZle soioze,
xolo erTi wlis Semdeg bazarze gamoCnda bioteqnologiu-
ri firmebis mier SemoTavazebuli mTeli rigi genetikurad
Secvlili simindis, kartofilis, Tambaqos, pomidvris, soi-
os, rafsis, bambis da sxva mcenareebi.
amJamad msoflioSi genetikurad modificirebuli mcena-
reebis miRebiTa da gamocdiT dakavebulia aseulobiT ko-
merciuli firma, romelTa jamuri kapitali 100 miliard
dolarze gacilebiT metia. 1996 wels transgenuri mcenaree-
bis qveS dakavebuli iyo 1,7 mln. heqtari, Semdeg wlebSi es
farTobebi arsebiTad gaizarda: 2000 weli – 44,2 mln. ha,
2005 weli – 90,0 ha, 2010 weli – 148,0 mln. ha, 2015 weli –
179,7 mln. ha, 2016 weli – 185,1 mln. ha. 2017 wlisaTvis am
kulturebis warmoeba xdeboda 24 qveyanaSi, sadac maT qveS
dakavebuli iyo 189,8 mln. ha, rac msoflioSi gamoyenebuli
nayofieri miwebis ~12%-s Seadgenda. 2016 wels kultivirebi-
saTvis daSvebuli iyo 29 transgenuri saxeobis 404 jiSi. es
miuTiTebs imaze, rom genetikurad modificirebuli mcena-
reebis moyvana ukve gaxda sasursaTo da sxva sasargeblo
produqtebis warmoebis mniSvnelovani mimarTuleba, rome-
lic izidavs arsebiT adamianur da finansur resursebs.
uaxloes wlebSi mosalodnelia kulturul mcenareTa
transgenuri formebiT dakavebli farTobebis Semdgomi
swrafi gazrda.
Tu Tavdapirvelad praqtikuli mniSvnelobis genetiku-
rad modificirebuli mcenareebis miRebisas adgili hqonda
sxvadasxva araxelsayreli faqtorebisadmi (daavadebebi,
herbicidebi, mavneblebi, gafuWeba Senaxvisas, stresebi) gam-
Zleobis genebis gadatanas, amJamad mcenareTa genetikuri
konstruirebis amocanaa ara imdenad mcenaris ama Tu im ar-
sebuli niSnis gaumjobeseba, rogorc tradiciuli sele-
qciis SemTxvevaSia, aramed medicinaSi, qimiur, farmkologi-
4
ur da sxva dargebSi farTod gamoyenebuli sruliad axali
naerTebis (mag., gansakuTrebuli cximovani mJavebi, Seu-
cvleli aminomJavebis maRali Semcvelobis sasargeblo ci-
lebi, modificirebuli polosaqaridebi, sakvebi vaqcinebi,
antisxeulebi, interferonebi, axali ekologiuri polime-
rebi da mravali sxva) warmoebis mcenarisaTvis swavleba.
praqtikulad mcenareTa genetikuri konstruirebis yve-
la samuSaos saboloo mizania bioteqnologiis amocanebis
gadawyveta. Tanamedrove bioteqnologia ki aris mecniereba
warmoebis intensifikaciisaTvis, an sxvadasxva daniSnule-
bis axali produqtebis misaRebad, genetikur-inJinruli da
ujredul-inJinruli meTodebiTa da teqnologiebiT geneti-
kurad transformirebuli biologiuri obieqtebis Seqmnisa
da gamoyenebis Sesaxeb.

5
 Tavi 1.
Uujreduli inJineria
ujreduli inJineria aris bioteqnologiis meTodi,
romelic laboratoriuli gziT sinjaraSi (in vitro) ujredis
doneze amuSavebs cocxali organizmebis gamravlebisa da
genetikuri aparatis konstruirebis (genomTan manipulire-
bis) xerxebs, am organizmebSi axali sasargeblo niSan-Tvi-
sebebis warmoqmnis mizniT.
ujreduli inJineria dafuZnebulia izolirebuli uj-
redebis, qsovilebisa da protoplastebis kulturis gamoye-
nebaze (sur. 1).
izolirebuli ujredebis, qsovilebisa da protoplas-
tebis kultura ewodeba organizmidan izolirebuli ujre-
debis, qsovilebisa da protoplastebis kultivirebas (ga-
mozrdas) xelovnur sakveb areze sterilur (aseptikur) pi-
robebSi.
ujreduli inJineria, rogorc ujredebis genetikuri
aparatis konstruirebis meTodi, moicavs maT in vitro kulti-
virebas, mutacias, hibridizacias da rekonstruqcias pro-
toplastebis, subujreduli struqturebis Serwymis, agreT-
ve ujredSi ucxo organelebis da sxva Seyvanis gziT.
am SemTxvevaSi ujreduli inJineriis dros adgili
aqvs genetikur-inJinrul manipulaciebs ujredebis doneze.
xelovnur sakveb areze, regulirebad aseptikur piro-
bebSi, izolirebuli ujredebis, qsovilebisa da protop-
lastebis kultivirebis xerxebis SemuSavebam mcenareTa se-
leqciaSi principulad axali teqnologiebis gamoyenebis
winapirobebi Seqmna, romlis safuZvels ujredebis geneti-
kuri konstruirebis meTodebi warmoadgens.
ujredul inJineriaSi gamoyenebuli meTodebi saSuale-
bas iZleva Catarebuli iqnas organizmebis genetikur in-
formaciasTan dakavSirebuli Semdeg manipulaciebi: geneti-
kuri informaciis Senaxva (mikroklonuri gamravleba, Cana-
saxebis, mtvrianebis kultura da sxva); genetikuri infor-

6
 I II
sawyisi
fesvebis mcenare meristemis
kultura kultura
g a n o g e
r n e
o z
i
III
kalusi calkeuli
ujredebis ujredebis
suspenzia kolonia
Canasaxi
Teslkvirti
Teslnergi

eqs pl antati
Rero
foToli fesvi
yvavili
B
dnm protoplastebi Serwyma
mtveri n Teslkvirti n
i
e
m b e z
n
r i o g e

Canasaxi, ylorti

mcenare-regeneranti

s ur. 1 . ujreduli inJineriis o bieqtebi d a p rocesebi

maciis cvlileba (kalusis, suspenziuri kulturebis, pro-

toplastebis in vitro kultivireba, in vitro mutacia da sxva);
genetikuri informaciis gadatana (somaturi hibridizacia,
organelebis Seyvana da sxva).
A yvela es manipulacia axali etapia mcenareTa seleqci-
is ganviTarebaSi. isini, tradiciuli seleqciuri procesiis
gaadvilebiT da axali seleqciuri teqnologiebis SemuSave-
biT mniSvnelovnad Seuwyoben xels sxvadasxva mimarTule-
biT saitereso da efeqtur mcenareTa miRebis process.

7
eqsplant eqsplantati
at i
U1.1. ujredebis genetikuri konstruirebis
ujredul-inJinruli meTodebi

1.1.1. mce naris ujredebis konstruireba

izolirebuli protoplastebis SerwymiT
protoplasti (sur. 2) aris mcenaris ujredi, romel-
sac mocilebuli aqvs celulozis garsi, SemosazRvrulia
citoplazmuri membraniT da inarCunebs ujredisaTvis dam-
axasiaTebel yvela Tvisebas.

s ur. 2 . izolirebuli p rotoplastebi

izolirebuli protoplasti warmoadgens erT-erT yve-
laze Zvirfas obieqts bioteqnologiaSi. misi gamoyenebiT
SesaZlebelia sxvadasxva aqtualuri sakiTxis gadawyveta
ujredis biologiaSi (sur. 3).PTumca misi erT-erTi mTavari
Rirebuleba sxvadasxva genetikuri manipulaciebis, maT So-
ris somaturi hibridizaciis, agreTve, dauzianebeli da
funqcionalurad aqtiuri citoplazmuri struqturebisa da
organelebis gamoyofis Catarebis SesaZleblobaSi, masSi
sxvadasxva umaRlesi mcenaris, prokariotebis, cxovelebis,
maT Soris ZuZumwovrebis genetikuri informaciisa da sxva-
dasxva ujreduli organelebis (sur. 4) SeyvanaSi, Sesabami-
sad axali ujredis genetikur konstruirebaSi mdgomare-
obs.
somaturi ujredebis inducirebuli Serwymis movlenaze
8
 `
somaturi hibridiza- ujreduli
ciis Catareba da saxe-
seleqciisa da mu-
obaTaSorisi da gva-
reobaTaSorisi hibri- mcenareTa tagenezis specifi-
debisa da cibridebis regeneracia kis Seswavla
miReba

ujredis kedlis age- plazmalemis gav-

bulebis, misi Semadge- liT sxvadasxva niv-
neli nivTierebebis
Tierebebisa da io-
biosinTezisa da tran-
sportis Seswavla
nebis transportis
izol ire bu - Seswavla
l i p rot o-
p l a s te bi
organelebis (mi- citoplazmuri
toqondria, qloro- struqturebisa da
plasti) Seyvana organelebis miReba

membranebis eleqtru- ujredebis ge- protoplastebis meta-

li Tvisebebisa da netikuri trans- bolizmis Tavisebure-
maTze fizikuri da bebisa da transkrip-
formacia
qimiuri faqtorebis ciis regulaciis me-
zemoqmedebis Seswavla qanizmebis Seswavla

s ur. 3. p rotoplastebis g amoyenebis ZiriTadi m imarTulebebi

dafuZnebuli hibridizaciis meTodebi ramodenime aTeuli

welia warmatebiT gamoiyeneba cxoveluri ujredebis gene-
tikur konstruirebaSi. analogiuri xerxebis mcenareul
ujredebSi gamoyenebis ZiriTad xelSemSlel faqtors did-
xans warmoadgenda mkvrivi celulozovani garsis arseboba,
romelic mTlianad gamoricxavda Serwymis SesaZleblobas.
pirvelad mcenareTa protoplastebis gamoyofa (izo-
lireba) ganxorcielebul iqna j. klerkeris mier jer kidev
1892 wels meqanikuri meTodiT (sur. 5).
man protoplastebi gamohyo mcenaris foTlis qsovile-
bidan. am dros jer xdeba ujredebis plazmolizi (plaz-
molitikad iyeneben 1 M saqarozisa da 0.02M KCL-is xsnaris
9
 mitoqondria
qromosoma birTvi

qloroplasti

dnm

baqteria

protoplasti

winaswardasaxuli niSnebis
mqone mcenare

s ur. 4. p rotoplastSi s xva ujredis g ene tikur i

s tr uq tur e bis a d a o r g ane l e bis S e y v ana

narevs), Semdeg ki samarTeblis an sxva instrumentis saSua-

lebiT meqanikurad iWreba ujredis garsi da protoplaste-
bi gareT gamodis.
meTods gaaCnia rigi SezRudvebi (SesaZlebelia gamoi-
yos mxolod mcire raodenobis protoplastebi; Znelia
protoplastebis gamoyofa meristemuli da momwifebuli
qsovilebidan; meTodi xangrZlivi da Sromatevadia), ris ga-
moc protoplastebis izolirebis meTodis farTod ganviTa-
reba daiwyo mas Semdeg, rac 1960 wels e. kokingma pirvelad
moaxdina fermentuli meTodiT (sur. 5) protoplastebis mi-
Rebis SesaZleblobis demonstrireba. man protoplastebi
gamohyo pomidoris aRmonacenis fesvis wverodan, ujredebis
ferment celulaziT damuSavebis gziT.
meqanikur meTodTan SedarebiT protoplastebis gamo-
yofis fermentul meTods gaaCnia garkveuli upiratesoba: 1)
10
 p r otoplastebis g amoyofis (izolirebis) m eTodebi

m e q anikur i f e r m e ntul i

K kl erkeri ( 1892 ) K koki n gi ( 1960)

moaxdina mcenare telo- obis sokos kulturaluri
rezas foTlis ujredebis siTxidan miRebuli hidro-
plazmolizi da qsovilebis lizuri fermentebiT daamu-
meqanikuri gaWra Sava pomidoris fesvis wve-
roebi

s ur. 5. p rotoplastebis g amoyofis (izolirebis) m e Todebi

SesaZlebelia erTdroulad gamoiyos didi raodenobis pro-

toplastebi; 2) protoplastebi ar ganicdian Zlier osmo-
sur SekumSvas; 3) ujredebi ufro intaqturi da dauzianebe-
lia; 4) meTodi SedarebiT swrafia. amitom dReisaTvis pro-
toplastebis izolireba ZiriTadad am meTodiT xdeba.
ujredis garsis mosacileblad iyeneben sami tipis
fermentul preparats – celulazebs, hemicelulazebs da
peqtinazebs. maT Rebuloben sokoebis kulturebidan da lo-
kokinas kuWis wvenidan. fermentebis pirveli komerciuli
preparatebi miRebul iqna 1968 wels iaponiaSi.
amJamad damuSavebulia protoplastebis gamoyofis pi-
robebi mcenaris TiTqmis nebismieri organodan: foTlebi-
dan, Reros apeqsidan, fesvebidan, koleoptilidan, nayofebi-
dan, yvavilis furclebidan, mtvrianebidan, tuberidan, en-
dospermidan, aleironis Sridan da sxva. protoplastebi mi-
iReba agreTve kalusidan da suspenziuri kulturebidan
(sur. 6).
yvelaze xSirad protoplastebis misaRebad iyeneben
foTlis mezofils an kalusis qsovils, romelebic iZleva
mraval erTgvarovan protoplasts. foTlebidan protoplas

11
 protoplastebis miRebis wyaro

mcenareTa izolire- suspenziuri kul- kalusuri kulturebi

buli organoebi da turebi
maTi nawilebi

foTlebi

lebnebi

fesvebi

mtvris marcvlebi

hipokotili

s ur. 6. p rotoplastebis m iRebis w y aroebi

tebis gamoyofisas (sur. 7,8) jer acileben epidermiss, Sem-
deg foTols Wrian segmentebad da axdenen enzimologi-

prot opl ast eb i s gamoyofi s t eqn i ka

(mezofiluri protoplastebis magaliTze)

mcenareuli masalis sterilizacia

fermentebis xsnarSi inkubaciisaTvis foTlebis momzadeba (qveda

epidermisis mocileba da plazmatikebis xsnaris foTlis fragmen-
tebis daWra)

G
foTlis fragmentebis inkubacia fermentebisa da plazmoli tikebis
xsnarSi (inkubaciis xangrZlivobis, temperaturisa da pH- sSerCeva
xdeba empiriulad)

izolirebuli protoplastebis garecxva fermentebisa da dauSle-

li qsovilebis narCenebisagan filtraciisa da centrofugirebis
gziT

izolirebuli protoplastebis cxovelmyofelobis dadgena da maTi

gadatana sakveb areze

s ur.7. p rotoplastebis g amoyofis teqnika

12
 f e rmentebis, ujr d i s
a R monaceni kedeli
m a rilebis, c i toplaz
an S aqrebis Semcveli
bi r Tvi ma
l i a re

f o T Toli
F f rag mentebad d aWra

p r otoplastebi c i toplazm ur i
m e m br a n a
g arecx va da centrofugireba

kol on iis g ada-

f i ltris p r otoplastebis
ku ltivireba
T es va 2-3 kviris
qaR aldi
S emd eg

ujr edis k edlis

n a rCenebi y l orttt
w a rmoqmqna
“Z i Za”- ujredebi

yl ortebis g a- sa k vebi a re maRali

c i tokininebiT d a
d arg va d as a-
n i adag S i g ad ar- d a bali a uqsinebiT
f es vianeblad 2-4
g va s ami kvi ri s
S emd eg kvi ris S emdeg

sa k vebi a re citokininebis g areSe

mcen are-regeneranti d a d abali a uqsinebiT

s ur. 8 . p rotoplastebis d a m aTgan m cenare-regenerantebis m iReba

ur damuSavebas. am dros mniSvnelovan rols asruleben os-

mosuri stabilizatorebi. yvelaze xSirad aseT nivTierebe-
bad iyeneben Saqrebs (glukoza, saqaroza, qsiloza, sorbi-
ti, maniti), zogjer CaCL2 , Na2 HPO4 , KCL-is marilebis xsna-
rebs. enzimologiuri damuSavebis Semdeg filtracia-cen-
trofugirebis teqnikis gamoyenebiT protoplastebis suspen-
zia sufTavdeba qsovilebisa da ujredebis narCenebisagan,
scildeba gamoyenebuli fermentebi. miRebuli protoplas-
tebis hibridizaciaSi gamosayeneblad SenarCunebisaTvis au-
cilebelia maTi CaTesva hipertonul (maniti an sorbit da-
matebul) sakveb areSi, rac uzrunvelyofs maT msubuq pla-
zmolizebul mdgomareobaSi yofnas, raTa ar moxdes pro-
toplastebis gaskdoma, aseve am dros muxruWdeba metabo-
lizmi da ujredis kedlis regeneracia.

13
 izolirebul protoplastebs, ujreduli garsis aRdge-
namde mokle droSi, erTmaneTTan Serwymis unari aqvT. es
Serwyma SeiZleba iyos spontanuri da inducirebuli (sur.
9).

protoplastebi s Serw yma

spontanuri inducirebuli

qimiuri meTodi

fizikuri meTodi
– eleqtroSerwyma

s ur. 9 . p rotoplastebis S erwymis m eTodebi

spontanuri Serwyma SeiZleba ganxorcieldes pasiurad

– plazmodesmebis gafarToebis Sedegad, protoplastebis
izolirebisas ujreduli garsis fermentuli daSlis
dros. aseT Serwymas yvelaze xSirad aqvs adgili aqtiuri
dayofis fazaSi myofi ujredebidan da suspenziuri kultu-
rebidan protoplastebis izolirebisas. pirvelad izoli-
rebuli protoplastebis spontanuri Serwyma SemCneul iqna
e. kiusteris mier jer kidev 1909 wels.
inducirebuli Serwyma warmoadgens Serwymis efeqtu-
robis gazrdis mizniT sxvadasxva faqtoris, e.w. induqto-
ris (qimiuri da fizikuri) moqmedebiT protoplastebis Ser-
wymis process.
dReisaTvis protoplastebis inducirebuli Serwyma Se-
iZleba ganxorcieldes ori meTodiT: 1) izolirebuli pro-
toplastebis kultivirebis sakveb areSi Serwymis mastimu-
lirebeli nivTierebebis damateba, anu “fiuzogenis” (Ser-
14
wymis induqtoris) meTodi anu qimiuri meTodi; 2) Serwyma
eleqtrodenis saSualebiT anu fizikuri meTodi.
pirvelad protoplastebis inducirebuli Serwyma
sakveb areSi Serwymis induqtor - natriumis nitratis dama-
tebiT, ganxorcielebul iqna e. kokingis laboratoriaSi
1970 wels. meTodi dabalfeqturi aRmoCnda da amJamad mxo-
lod istoriuli mniSvneloba aqvs.
SemdegSi efeqturi induqtoris e.w. “fiuzogenis” Zie-
bis procesSi mecnierebma daadgines, rom protoplastebis
yvelaze maRali sixSiris da kontrols daqvemdebarebuli
Serwyma xdeba pH maRali (9-11) maCveneblis da Ca++ ionebis
maRali koncentraciis (100-300 mM) mqone xsnarebis gamoye-
nebisa da induqtor polieTilenglikolis (peg) (mol.masa
1500-6000) koncentrirebuli xsnarebiT protoplastebis wi-
naswari aglutinaciis (laT. agglutinatio – Sewebeba) dros.
aRniSnuli meTodis yvelaze farTod gamoyenebul me-
Todikas warmoadgens k. kaos (1977) mier SemuSavebuli meTo-
dika (sur. 10): ori mSoblis protoplastebis narevi calke-
uli wveTebis saxiT gadaaqvT minis zedapirze, toveben da-
leqvisaTvis 5-15 wuTs da umateben peg-is xsnars ujredebis
suspenziis erT wveTze xsnaris sami wveTis angariSiT. peg-
is xsnars amzadeben 1 gr peg-is (mol. masa 1540) damatebiT
2 ml xsnarSi, romelic Seicavs 0.1 mM glukozas, 10.5 mM
CaCL2 da 0.7 mM KH2 PO4 (pH 5.5). 10-15 wuTis Semdeg preparats
umateben pH maRali maCveneblisa (10-10.5) da Ca++ maRali (70
mM) Semcvelobis 0.5 ml moculobis xsnars, Cveulebriv
qloridis marilis (CaCl2 ) saxiT. proceduras imeoreben 15
wuTis Semdeg. kidev 15 wuTis Semdeg protoplastebs recxa-
ven 5 wuTis intervaliT xsnaris frTxili gamowoviTa da
sakvebi aris damatebiT. garecxvisaTvis saWiro sakvebi aris
saerTo moculoba 15-20 ml-ia. kultivirebisaTvis protop-
lastebis suspenzia gadaaqvT petris jamebSi 2-2 ml mocu-
lobiT TiToeulSi, an jer aureven agarizebul sakveb are-
Si (t=400C) da Semdeg asxamen petris jamebSi 5-5 ml mocu-
lobiT TiToeulSi.

15
 protoplastebi protoplastebi

protoplastebis
narevis wveTi

peg-is xsnari

pipetka

protoplastebis
kultivireba
seleqtiur areze

hibriduli ujredebis
gamorCeva

somaturi
hibridi

s ur. 1 0. m cenareTa s omaturi hibridebis m iR e bis s q e m a

protoplastebis Serwymis meqanizmi am dros mdgomare-

obs SemdegSi: protoplastebs Cveulebriv uaryofiTi muxti
aqvT (saxeobebis mixedviT varirebs -10-dan -30-mde mB). uar-
yofiTad damuxtuli protplastebi erTmaneTisagan ganizi-
debian. pH-is maRali maCveneblisa da Ca++ ionebis maRali
koncentraciis dros adgili aqvs uaryofiTi zedapiruli
muxtis neitralizebas, xolo induqtor peg-is monawileoba
iwvevs protoplastebis aglutinacias (sur. 11b), plazmale-
mis gaxleCas (sur. 11g), membranebis ganvladobisa da dena-
dobis gazrdas da Serwymas (sur. 11d).
16
 polieTilenglikoli

Serwymu-
li pro-
toplas-
ti

a b
g

d
s ur. 1 1. p o lieTilenglikolis s aSual e biT p r o to p l as te bis
S e r wy m is s q e m a d a e ta p e bi
(a – protoplastebis populacia; b – protoplastebis aglutinacia;
g – citoplazmuri Wimebis warmoqmna; d – protoplastebis Serwyma)

Serwymis efeqturoba qimiuri meTodis dros Seadgens

10-50%. masze garemo faqtorebis garda, gansakuTrebiT gav-
lenas axdens ujredebis qsoviluri kuTvnileba. advilad
erwymis meristemuli da kalusuri protoplastebi, nakleb
efeqturia Zlier vakuolizebuli ujredebis da qloro-
filganviTarebuli ujredebis (mag. mezofilis) protoplas-
tebis Serwyma. Tambaqos foTlis protoplastebis Serwyma
ufro advilia, vidre kartofilis an stafilos protop-
lastebis. meTodis uaryofiT mxares warmoadgens is, rom
Zalian Znelia didi raodenobiT erTdroulad Serwymuli
protplastebis miReba. garda amisa, induqtoriT damuSave-
bis Semdeg protoplastebis nawili ziandeba da kvdeba, ase-
ve xSirad erwymis ara ori, aramed meti protoplasti, rac
arasicocxlisunariani hibridebis warmoqmnas iwvevs.
amJamad sul ufro farTod iyeneben protplastebis
Serwymis meore, fizikur meTods – Serwyma eleqtrodenis
saSualebiT, romelic damuSavebulia u. cimermanis mier

17
 1974 wels. meTodis principi dafuZnebulia araerTgavrova-
ni eleqtruli velis gamoyenebaze, romelic warmoiqmneba
100 volti Zabvisa da 50 hc sixSiris cvalebadi denis mier
specialur kameraSi, sadac aTavseben izolirebul proto-
plastebs da SehyavT platinis eleqtrodebi.
Serwymis meqanizmi mocemul meTodSi mdgomareobs Sem-
degSi: eleqtruli velis moqmedebisas protoplastebi lag-
debian Zewkviseburad eleqtrodebs Soris, ganicdian aglu-
tinacias (sur. 12b), warmoqmnian citoplazmur Wimebs (sur.
platinis eleqtrodebi

Serwymuli
protoplasti

eleqt- a b g d
ruli
veli, V 1 2 3 4 5

s ur. 1 2. e l eqtrodenis s aSual e biT p r o to p l as te bis S e r wy m is

s q e m a d a e tap e bi
(a – protoplastebi platinis eleqtrodebs Soris; b – dieleqtroforezi
da aglutinacia; g – citoplazmuri Wimebis warmoqmna; d – protoplastbis
Serwyma; 1, 5 – deni ar aris; 2 – cvalebadi deni (100 volti, 50 hc), 3 –
Zlieri impulsi (750 B/sm, 50 mikrowami), 4 – denis dacema)

12g) da damatebiTi impulsis (kvadratuli talRa, Zabva 750

B.sm,-1 xangrZlioba 50 mikrowami) gatarebisas adgili aqvs
Serwymas (sur. 12d). me-13 suraTze warmodgenilia aRniSnuli
meTodebiT sxvadasxva mcenaris protoplastis Serwymis
procesi.
protoplastebis Serwyma SemTxveviTi procesia. orive
meTodis dros SesaZlebelia Serwyma rogorc sxvadasxva sa
18
 1 2

3 4

s ur. 1 3. 1 ,2 – kartofilis d a Tambaqos p rotoplas te bis

S e rwyma q imiuri m e TodiT; 3,4 – p e tunias p rotoplastebis
S e r wy m a f izikur i m e To d iT .

A B

A A A B B B

h omokarioni h eterokarioni h omokarioni

s ur. 1 4. p rotoplastebis S erwymis S esaZlo v ariantebi

19
xeobis (mag., A da B), is erTi da igive saxeobis protoplas-
tebs Soris (A da A; B da B) (sur. 14) . amdenad protoplastebis
Serwymis Sedegad SeiZleba warmoiqmnas rogorc homokario-
ni (AA, BB), anu ujredi, romelSic monawileobs ori an meti
identuri genetikuri bunebis birTvi, ise heterokarioni (AB),
anu ujredi, romelSic monawileobs ori an meti sxvadasxva
genetikuri bunebis birTvi.

1.1.2. mce naris ujredebis konstruireba

subprotoplastebis SerwymiT
ujredebis konstruirebisas mTliani protoplastebis
Serwymis garda, ujreduli inJineriis meTodebiT SesaZle-
belia SerwymaSi monawileobdes erTi an orive mSoblis e.w.
subprotoplastebi anu protoplastebis fragmentebi, ker-
Zod: nukleoplastebi – ujredis birTvi mcire raodenobis
citoplazmiT, citoplastebi – ujredis citoplazmis didi
nawili genetikuri masaliT birTvis gareSe, mikroprotop-
lastebi – ujredis qromosomis nawili mcire rao-denobis
citoplazmiT.
Tumca am mimarTulebiT arsebobs problemebi, rac da-
kavSirebulia intaqturi ujreduli fragmentebis didi ho-
mogenuri populaciebis miRebisa da gamoyofis srulyofi-
li meTodikis uqonlobasTan.
nukleoplastebis gamoyofis erT-erTi meTodika iTva-
liswinebs kalusuri protoplastebis suspenziis damuSave-
bas 370C temperaturis pirobebSi alkaloid citoqalazin B-
Ti (50 mg/ml) da centrofugirebas (40000 g) 50 mg/ml ci-toqa-
lazin B-s Semcvel 1,5 M sorbitolis fenaSi. aseTi gziT
gamoyofili nukleoplastebis Tambaqos protoplastebTan
Serwymis Sedegad miRebul iqna cxovelmyofeli hibriduli
ujredebi.
intaqturi citoplastebis didi raodenobiT gamoyofis
erT-erTi meTodi SemuSavebuli iqna jer kidev 1981 wels.
citoplastebis miReba xdeba kalusuri protoplastebis
centrofugirebiT uwyveti izoosmosuri gradientis (iqmeba
20
СаСl2 Semcveli koloiduri silikagelis gamoyenebiT) sim-
kvrivis, manitolisa da kalciumis qloridis Semcvel xsna-
rebSi.
citoplastebi warmatebiT iqna gamoyofili simindis,
Tambaqos, lencofas da sxva kalusuri protoplastebis
suspenziidan. jer-jerobiT citoplastebis gamoyenebiT uj-
redebis konstruirebis SemTxvevebi arc ise bevria.

1.1.3. mce naris ujredebis konstruireba

ujreduli organelebis SeyvaniT
mcenaris ujredebis konstruireba SesaZlebelia ag-
reTve izolirebul protoplstebis mniSvnelovani Tvisebis
- saTanado pirobebSi calkeuli ujreduli organelebis
(birTvi, qloroplastebi, mitoqondriebi, metafazuri qro-
mosomebi) garemodan STanTqmis unaris gamoyenebiT.
mecnierebis azriT organelebis STanTqma xdeba an en-
docitozis gziT, romlis gaZliereba SeiZleba garegani ze-
moqmedebiT, an protoplastebis izolirebis procesSi
plazmalemaze warmoqmnili bzarebidan pasiuri SeRwevis
gziT.
damuSavebulia protoplastebis mier ujreduli orga-
nelebis STanTqmis induqciis ramodenime xerxi, maT Soris:
- organelebis damuSaveba membranebis modifikatore-
biT (lizocimi, NaNO3 ) da centrofugireba;
- protoplastebisa da izolirebuli organelebis ag-
lutinacia peg-iT da erTdrouli centrofugireba.
STanTqmis efeqturoba am dros damokidebulia ujre-
debisa da organelebis raodenobaze da sakmaod maRal maC-
venebels aRwevs – birTvis STanTqmisas 20%, qloroplaste-
bis STanTqmisas 25%.
e.w. “sendviC-meTodiT” protoplastebSi birTvis1 Seyva-
nas axdenen protoplastis susti deplazmolizis pirobeb-

1.izolirebuli birTvi (petunias) protoplastebSi (Tambaqos) pirvelad

gadatanil iqna jer kidev 1973 wels i. potrikusisa da f. hofmanis mier.
21
Si. Seyvanis win protoplastebs da birTvs amuSaveben li-
zocimiT. Semdeg xdeba birTvis da protoplastis monac-
vleobiT daleqva centrofugirebiT. sinjarebs umateben os-
motik manitols lizocimis gareSe da isev xdeba centro-
fugireba naxevari saaTi 120g-s pirobebSi. toveben 2 saaTi
40C temperaturaze da bolos axdenen naleqis resuspendi-
rebas.
ujreduli organelebis gadatana xdeba aseve liposo-
mebis gamoyenebiT da mikroineqciiT.
ujredis konstruirebis mizniT SesaZlebelia mcenare-
ul proroplastebSi metafazuri qromosomebis gadanergva.
cxovelur ujredebSi am meTods didi warmatebiT iyeneben.
mcenareul ujredebSi muSaobis sirTules warmoadgens is,
rom ujredebis populacia Znelad eqvemdebareba izolire-
buli qromosomebis didi raodenobiT gamoyofis aucile-
bel winapirobas – dayofis fazis sinqronizacias. ujrede-
bis dayofis sinqronizaciis Cveulebriv meTodebs warmoad-
gens auqsiniT, citoqininiT da fosfatiT SimSili an dnm-s
sinTezis droebiTi daTrgunva.
am mimarTulebiT muSaobisaTvis meTodebi aucileb-
lad saWiroebs srulyofas.

1.1.4. mce naris ujredebis genetikuri

cvalebadoba in vitro manipulaciebiT

in vitro pirobebSi kultivirebul mcenareul ujredebs

sakmaod maRali genetikuri heterogenuroba, e.w. somakloni-
ri cvalebadobis2 warmoqmna axasiaTebT. amis mizezi TviT
eqsplantatisaTvis damaxasiaTebel bunebriv genetikur hete-
rogenurobasTan erTad, ujredebis mier araorganizire-bu-
li zrdis stadiis gavlisas in vitro kultivirebis pirobebi,
sakvebi aris komponentebi, sxvadasxva manipulaciebi warmoa

2
ujreduli xazebis mravalferovnebas “somaklonuri variabeloba” ewode-
ba.
22
dgens.
in vitro pirobebSi kultivirebisas ujredebis kultura-
Si Seyvanis, pirveli dayofis induqciis, araorganizebuli
zrdis, xangrZlivi pasirebis procesSi fitohormonebis da
kvebis aris sxva komponentebis, kultivirebis pirobebis, me-
tabolizmis produqtebis, korelaciuri kavSiris gawyvetis
da sxva faqtorebis zemoqmedeba xSirad iwvevs:
a) kariologiur cvlilebebs - erT sinjaraSi gamoz-
rdili erTi da igive qsovilis ujredebi xSirad erTmane-
Tisagan gansxvavdba qromosomuli nakrebiT (diploidebi,
poliploidebi, aneuploidebi);
b) sxvadasxva qromosomul gardaqmnebs, maT Soris krip-
tikul3, citologiuri kvlevisas SeumCnevels – qromosome-
bis mcire nawilis dakargva, duplikacia, translokacia, in-
versia, subqromatiduli gacvla da sxva;
g) multigenuri blokebis eqspresiis cvlilebas – kul-
tivirebis pirobebiT zogjer icvleba genebis blokis eq-
spresiis regulacia iseTnairad, rom geni, romelic adre
eqspresideboda SeiZleba represirdes da geni, romelic ad-
re mdumare iyo eqspresirdes;
d) citoplazmatur cvlilebebs – qloroplastebis da
mitoqondriebis genebis maRali xarisxiT cvlilebas;
e) mobiluri genetikuri elementebis gadaadgilebas, Se-
degad genis struqturis mTlianobis gawyvetas, genis regu-
laciis Secvlas, mdumare genebis gaaqtiurebas, genis uni-ka-
luri Tanmimdevrobebis sxva regulaciis poziciaSi gada-
tanas da sxva;
v) virusebis (mcenaris daavadebis xiluli simptomebis
ar gamomwvevi, magram genotipis cvlilebis gamomwvevi) krip-
tikul eliminacias;
z) genebis amplifikacias an reduqcias;
T) somatur krosingovers da Svileuli qromatidebis

3 berZ. ktypte – saidumlo

23
gacvlas;
i) adre represirebuli genebis aqtivacias da sxva.
rac in vitro kultivirebadi ujredebis morfologiuri,
funqcionaluri da sxva saxis cvlilebebs iwvevs, Sedegad
aseTnairad konstruirebuli ujredebis regenerirebiT Se-
saZlebelia warmoiqmnas mravalferovani genetikurad Sec-
vlili mcenareebi.

1.1.5. mce naris ujredebis mutaciuri

cvalebadoba in vitro kultivirebisas

mcenaris ujredebis genetikuri gardaqmna SesaZlebe-

lia kultivirebis procesSi mutaciuri cvalebadibis ga-
momwvevi fizikuri da qimiuri faqtorebis gamoyenebiT.
M miuxedavad imisa, rom in vitro kultivirebad ujredebs
axasiaTebT genetikuri arastabiluroba, is mainc sakmarisi
ar aris sasurveliDgenetikuri mravalferovnebis misaRebad.
amitom ujredebis kulturaSi miznis miRweva SesaZle-
belia qimiuri da fizikuri mutagenuri faqtorebis gamoye-
nebiT. magaliTad, rauvolfias ujredebis damuSaveba 2.5x103
M koncentraciis azotovani ipritiT iwvevs qromosomuli
aberaciebis zrdas, triploiduri populaciebis ma-tebas.
Cveulebriv mcenareuli ujredebis populaciaSi muta-
ciuri cvalebadoba iSviaTi movlenaa. sxvadasxva kultu-
rebSi, sxvadasxva seleqtiur sistemebze, spontenuri muta-
ciis sixSire 1.10-9 – 1.10-4-is farglebSia. inducirebuli mu-
tagenezis dros milionobiT da miliardobiT mcenareuli
ujredi advilad eqvemdebareba mutagenebis zemoqmedebas da
sakmaod maRali sixSiriT - 5.10-8 - 6.10-2 warmoiqmneba gene-ti-
kurad Secvlili ujreduli xazebi, romlebic Semdeg SeiZ-
leba moTavsdes seleqtiur areSi sasurveli mutantis gamo-
savlenad.
M mutaciis sixSiris amaRleba SeiZleba qimiuri da fizi-
kuri mutagenebiT: nitrozoguanidini, eTilmeTansulfonati,
nitrozomeTilSardovana, N-meTil-N-nitro-N-nitrozoSard-

24
ovana, meTilmeTansulfonati, ultraiisferi gamosxiveba, γ-
gamosxiveba, rentgenis gamosxiveba da sxva.
seleqtiur agentebad gamoiyeneba antibiotikebi, nukle-
inis mJavebis sinTezis inhibitorebi, purinisa da pirimidi-
nis fuZeebis analogebi, fito- da paTotoqsinebi, wylisa da
marilis stresis gamomwvevi agentebi, aminomJavebis analo-
gebi da sxva.
mcenareuli ujredebis in vitro pirobebSi kultivirebisas
inducirebuli mutagenezis gziT SesaZlebelia miRebul iq-
nas pigmenturi mutaciis mqone, antibiotikebisadmi, damla-
Sebisadmi, dabali temperaturisadmi, metalebisadmi, herbi-
cidebisadmi gamZle da sxva ujreduli xazebi.

1.1.6. Secvlili (konstruirebuli) ujredebis

gamorCeva da Senaxva

zemoT ganxiluli gzebiT ujredebis rekonstruqciis

proceduris dasrulebis Semdeg aucilebelia:
1. transformirebuli produqtis gamorCeva, romelic
SeiZleba Catardes ujredebis doneze, an mcenareTa regene-
raciis stadiaze;
2. garkveuli cvalebadobis matarebeli ujredebis (uj-
reduli xazebis) Senaxva (SenarCuneba), romelic tardeba
ori gziT: a) ujredebis (ujreduli xazebis) gamozrda
(kultivireba) Sesabamis sakveb areebze, maTi zrdis perio-
duli SemowmebiT seleqtiuri agentis monawileobiT; b) uj-
redebidan (ujreduli xazebidan) mcenareebis regenerireba
da garkveuli cvalebadobebis SenarCuneba vegetatiurad
mravlebadi ylortebis an Teslebis saxiT.
ufro misaRebs warmoadgens Senaxvis meore xerxi, vina-
idan in vitro pirobebSi ujredebis araorganizebul zrdas au-
cileblad Tan axlavs sxvadasxva memkvidruli cvalebado-
bebis dagroveba, garda amisa xangrZlivi kultivireba iw-
vevs ujredebis morfogenetikuri potencialis dakargvas.
U

25
 1.2. ujreduli asociaciebi
mikroorganizmebTan

1.2.1. ujreduli asociaciebis Seqmnis arsi,

ZiriTadi mizani da tipebi

ujredebis gardaqmnis erT-erT saintereso ujredul-

inJinrul teqnologias warmoadgens ujredebSi an kultivi-
rebadi ujredebis populaciaSi mikroorganizmebis Seyva-
niT, axali xelovnuri ujredebis an ujreduli sistemebis
anu ujreduli asociaciebis Seqmna.
aseTi asociaciebis Seqmnis arsi mdgomareobs imaSi,
rom mikroorganizmebTan gaerTianebiT ujredebma an maTma
populaciebma unda SeiZinon axali Tvisebebi.
asociaciebi SeiZleba iyos Sidaujreduli (endosimbi-
ozuri tipi) da ujredTaSorisi (ekzosimbiozuri tipi).
pirvel SemTxvevaSi mikroorganizmebi SehyavT umaRle-
si mcenareebis izolirebul protoplastebSi. meore SemT-
xvevaSi – adgili aqvs ujredebis an mcenareTa qsovilebis
da mikroorganizmebis erTobliv kultivirebas.
izolirebul protoplastebSi mikroorganizmebis Se-
yvana SeiZleba ganxorcieldes (sur. 15):
a) endocitoziT4 (STanTqma plazmalemis invaginaciis5
gziT). am dros warmoqmnili vezikulebi 6 mikroorganizme-
biT gadadis citoplazmaSi;
b) protoplastisa da mikroorganizmis membranebis
SerwymiT (integraciiT). am dros mikroorganizmis organe-
lebi protoplastis citoplazmaSi ar arian SemosazRvru-
li protoplastis plazmalemiT;
g) mikroorganizmis jer moTavseba xelovnur membrana-
Si – loposomaSi da Semdeg Serwyma protoplastis membra-
nasTan.

4
garedan ujredSi sxvadasxva molekulebis SeRwevis procesi.
5
Sezneqis
6
Mvezikulebi – membranuli buStukebi.
26
 endocitozi

membranebis Serwyma

Pprotoplastis membranis Serwyma liposomasTan

s ur. 1 5. m ikroorganizmebis S eyvana um aRlesi m cenaree bis

izo l ir e bul p r o to p l as te bS i
TiToeul meTods gaaCnia dadebiTi da uaryofiTi mxa-
reebi. pirvel SemTxvevaSi citolazmaSi moxvedrili mem-
braniT SemosazRvruli vezikulebi mikroorganizmebiT SeiZ-
leba Seerwyas protoplastis lizosomebs da Sedegad dai-
Salos mikroorganizmi. meore SemTxvevaSi irRveva mikroor-
ganizmis mTlianoba da citoplazmaSi xvdebian calkeuli
organelebi. mesame SemTxvevaSi citoplazmaSi moxvedri-li
intaqturi mikroorganizmebi protoplastTan qmnian buneb-
riv simbiozur urTierTobas.
izolirebul protoplastebSi Cveulebriv SehyavT mik-
roorganizmebis Semdegi sistematikuri jgufebi: baqteriebi,
safuarebi, cianobaqteriebi, cianelebi.
ujredebis an mcenareTa qsovilebis da mikroorganiz-
mebis erToblivi kultivireba SeiZleba ganxorcieldes Sem-
degi xerxebiT:
a) mikroorganizmebis Tesva uSualod ujredul kul-
turebze;
b) mikroorganizmebis Tesva agaris zedapirze kalusis
gverdiT, Sexebis gareSe;
g) baqteriuli ujredebisa da ujreduli kulturebis
membranebiT gayofa, maT Soris metabolitebis gacvlis uzr
27
unvelyofiT.
ujredebTan an mcenareTa qsovilebTan erToblivi
kultivirebisaTvis ZiriTadad iyeneben: azotfiqsator baq-
teriebs (Rhizobium, Azotobacter, Azospirillum), wyalmcenareebs
(Chlorella) da sxvadasxva saxeobis sokoebs.
ujreduli populaciebis Seqmnis ZiriTadi mizani Teo-
riuli sakiTxebis (eukariotuli ujredebis simbiozuri
warmoSobis hipoTezis dadastureba, mcenareTa da mikroor-
ganizmTa bunebrivi simbiozuri Tanaarsebobis modelireba)
SeswavlasTan erTad, iseTi praqtikuli amocanebis gadaw-
yvetaa, rogoricaa:
1. mcenareTa ujredi-producentebis mier medicinasa
da sxva dargebisaTvis ekonomikurad mniSvnelovani nivTie-
rebebis produqtiulobis amaRleba. am nivTierebebis renta-
beluri warmoebisaTvis aucilebelia mcenareTa ujredebis
kultivireba Sedgenilobis mixedviT martiv sakveb areze.
gamoyenebuli sakvebi areebi ki sakmaod rTulia, Seicaven
vitaminebs, hormonebs, organul nivTierebebs, radganac uj-
redebi kulturaSi warmoadgenen heterotrofebs an gaaC-ni-
aT fotosinTezis SezRuduli unari (kalusuri ujredebi
sibneleSi). amitom aseT kulturebSi ujredebis zrdisaTvis
aucilebeli substratis an sasargeblo nivTierebebis bio-
sinTezis winamorbedebis masinTezirebeli mikroorganizme-
bis Seyvana warmoadgens metad mniSvnelovans.
2. axali Tvisebebis mqone mcenareTa miReba. gansakuT-
rebiT sasoflo-sameurneo mcenareTa gaumjobeseba da mole-
kuluri azotis fiqsaciis unaris mqone (azotfiqsatori mi-
kroorganizmebis SeyvaniT) mcenareTa miReba.

1.2.2. ujredul populaciebSi mikroorganizme-

bis Seyvanis kvle viTi muSaobis Sedegebi

dReisaTvis cxovelmyofeli sistemebi miRebulia kul-

tivirebadi ujreduli populaciebis safuZvelze Seqmnil
ujredul asociaciebSi.

28
 asoci aci ebi koJri s baqteri ebTan . soios kalusuri
qsovilebis Rhizobium-is baqteriebiT inficirebis Sedegad ka-
lusis ujredebSi miRebul iqna infeqciuri Zafebisa da baq-
teroidebis msgavsi warmoiqmnebi. aseT asociaciebSi baqte-
riis ujredebi avlendnen nitrogenazul aqtiurobas, rac
ar iyo wminda kulturaSi. es adasturebda azotfiqsaciis
process. SemdegSi Seqmnil iqna Rhizobium-is baqteriebis
araparkosan mcenarebTan azotmafiqsirebeli asociaciebi. aq
baqteriis ujredebi ZiriTadad lokalizebuli iyo mcena-
ris kalusis ujredebis zedapirze, iSviaTad kalusis siRr-
meSi. ZiriTadad inarCunebdnen Cxirisebur formas baqteroi-
debis gareSe. iyo erTi warmatebuli cda Tambaqos kalusis
Rhizobium-Tan asociaciidan mcenaris regeneraciisa.
asoci aciebi Tavisuflad mcxovreb azotfi qsatoreb-
Tan . mcenaris rizosferoSi mcxovreb azotfiqsator Azotoba
cter-Tan da Azospirillum-Tan efeqturi azotmafiqsirebeli, ua-
zoto sakveb areze zrdis unaris da mcenaris regeneraciis
perspeqtivis mqone asociaciebis Seqmnis eqsperimentebSi ga-
movlinda saxeobaspecifikuroba. Tambaqos da fetvis kalu-
sis ujredebis asociaciebi Azospirillum-Tan funqcionirebda
4 kvira, baqteriebi avlenda nitrogenazul aqtiurobas, mag-
ram Semdeg iRupeboda. Azotobacter-Tan Saqris lerwmis asoci-
acia subkultivirebda 18 Tve, oRond azotis dabali Sem-
cvelobis an saerTod uazoto sakvebi aris pirobebSi. baq-
teriebi aqac avlenda nitrogenazul aqtiurobas.
orive SemTxvevaSi baqteriebi yovelTvis lokalizebu-
li iyo ujredis zedapirze an ujredSorisebSi da ara uj-
redis SigniT.
asoci aciebi mw vane w yal mcen areebTan . qlorelas erT-
erTi StamiT stafilos kalusis inokulirebisas, asociacia
azotis deficitur sakveb areze kultivirebda ramodenime
Tve. sakontrolo mcenareebi iRupeboda. wyalmcenaris uj-
redebi ver aRwevda mcenaris ujredebis SigniT.
asoci aciebi sokoebTan . balaxovani mcenare teganis (Ru-
ta graveolens) ujredebi cal-calke kultivirebda sxva-dasxva
29
sokoebTan. mcenaris ujredebze gavlenas axdenda sokos ga-
monayofebi.
zogjer mcenaris ujredebs umatebdnen sokoebis kul-
turalur siTxes. Botritis allii- sTan kultivirebisas alkaloi-
debis sinTezi sakontolosTan SedarebiT gaizarda 10-jer,
xolo kulturaluri siTxis damatebisas – 50-jer.
A asociaciebi ci anobaqteri ebTan . cianobaqteriebTan mce-
nareTa ujredebis (kalusuri da suspenziuri) asociaciebis
SemTxvevaSi gamovlenilia rogorc erTi partnioris mier
meoris inhibirebis, ise cxovelmyofelobis procesebis ur-
TierT stimulirebis SemTxvevebi.
Tambaqos suspenziuri kulturisa da Anabaena nidu-
lanss asociaciis dros adgili hqonda: zrdis urTierTstimu-
lacias; mikroorganizmebi adsorbirdebian mcenaris ujre-
debis zedapirze, aRweven ujreduli agregatebis SigniT,
magram partniorebis nawili rCeba suspenziaSi Tavisufal
mgomareobaSi; mcenaris ujredebi iyeneben cianobaqteriebis
fotosinTezis produqtebs; mcenarisa da mikroorganizmis
ujredebs Soris myardeba metabolituri urTierTmoqmedeba.
Tambaqos kalusisa da Anabaena nidulanss asociaciis
dros kalusuri ujredebi izrdeboda ara nakleb 2 weli.
antagonizmi ar iyo.
auqsinebisa da citikininebis damatebiT moxerxda kalu-
sidan cianobaqteriasTan asocirebuli Tambaqos mcenaris
regeneracia. regenerantis Reroebi iyo TeTri feris lurj-
mwvane feris baqteriebis Semcveli monakveTebiT. cianobaq-
teriebi iyo agreTve foTlebSi, gamtar sistemaSi, bageebSi.
kvlevebi am mimarTulebiT aucileblad gagrZeldeba.

30
 1.3. ujreduli seleqciiis raoba,
Taviseburebebi da Sedegebi

1.3.1. ujreduli seleqciis zogadi sqema

U ujredebis genetikuri konstruirebis saiTxebTan erTad

mizanewonilia ganxilul iqnas am konstrirebuli ujrede-
bis gamoyenebiT Casatarebeli samuSaoebi, kerZod, ujredu-
li seleqciis sakiTxebi.
ujreduli da genetikuri inJineriis teqnologiebs
udidesi perspeqtivebi gaaCniaT mcenareTa seleqciis pro-
cesSi. am mimarTulebiT erT-erT efeqtur meTods warmo-ad-
gens ujreduli seleqcia7, romlis drosac axali memkvid-
ruli Tvisebebis mqone ujreduli xazebisa da mcenareebis
gamorCeva xdeba in vitro kultivirebadi ujredebis doneze.
aseTi seleqciis dros, tradiciuli seleqciis meTo-
debTan SedarebiT, mcenareTa axali formebis miReba 2-4-jer
ufro swrafad xdeba.
winaswar konstruirebuli, seleqtiur pirobebSi ga-
morCeuli ujredebidan mcenare-regenerantebis miReba SesaZ-
lebelia mcenaris ujredis unikaluri Tvisebis – totopo-
tenturobis8 gamo.
ujredul seleqciaSi mizanmimarTuli seleqciuri
procesis Casatareblad, anu kultivirebadi ujredebis ma-
sidan sasurveli mimarTulebiT Secvlili calkeuli ujre-
debis gamosarCevad, iqmneba specialuri seleqtiuri sis-te-
ma, romelic am ujredebis zrdis saSualebas iZleva.
dReisaTvis ujredul seleqciaSi am mizniT farTod
iyeneben herbicidebis, paTotoqsinebis, aminomJavebisa da
nukleotidebis analogebis, antibiotikebis, marilebisa da

7
ujreduli seleqcia aris axali genotipebis konstruirebisa da gamorCe-
vis procesi kultivirebadi ujredebis doneze seleqtiuri sistemebis ga-
moyenebiT.
8
totipotenturoba (totus [ laT.] – mTliani, mTeli; potenta [laT.] – Zala.)
aris mcenaris ujredis Tviseba, moaxdinos birTvis genetikuri informaci-
is sruli realizacia da warmoqmnas mTliani mcenare.
31
mZime metalebis maRali koncentraciebis, pH-is dabali maC-
veneblebis, eqstremaluri temperaturis da sxva stresuli
faqtorebis mimarT gamZleobis seleqtiur sistemebs. iyene-
ben agreTve fitohormonebis, aminomJavebis, vitaminebis mi-
marT prototrofulobisa9 da auqsotrofulobis10 siste-
mebs da kultivirebis procesSi am nivTierebebis Semcvel
sakveb areze ujredis zrdis unaris an piriqiT, uunarobis
mixed-viT arCeven sasurvel ujredebs.
me-16 suraTze warmodgenilia ujreduli seleqciis
zogadi sqema. Tu gamorCevas atareben romelime nivTierebis
mimarTa gamZleobaze, maSin ujredebs Tesaven am nivTiere-
bis Semcvel sakveb areze. roca tardeba gamorCeva romeli-
me stresuli faqtorisadmi (maRali an dabali temperatura
da sxva) gamZleobaze, kulturaluri WurWeli Tavsdeba Se-
sabamis pirobebSi.
garkveuli drois Semdeg ujredebis nawili iRupeba,
gadarCeba genetikuri an epigenetikuri cvalebadobebis Se-
degad seleqtiuri faqtorisadmi gamZle ujredebi. Semdeg
xdeba maTgan kalusis miReba, organogenezis induqcia da
mcenare-regenerantebis11 miReba, bolos ki – regenerantebis
Semowmeba gamZleobaze.
mravali sasoflo-sameurneo kulturisaTvis dReisa-
Tvis SemuSavebuli mcenareuli ujredebis kultivirebisa
da maTgan mcenareTa regeneraciis xerxebi, saSualebas iZ-
leva efeqturad iqnes realizebuli ujreduli seleqciis
SesaZleblobebi anu gamoyenebul iqnas mcenareTa axali ji-
Sebisa da formebis misaRebad.

9 Oprototrofi – organizmi, romelic Tavisi zrda-ganviTarebisaTvis ar

saWiroebs mza zrdis nivTierebebs.
10 auqsotrofi – organizmi, romelsac ar gaaCnia Tavisi zrda-ganvi-
TarebisaTvis aucilebeli nivTierebebis sinTezirebis unari.
11 ujredul kulturebSi miRebul mcenare-regenerantebs da maT TviT-

damtveril Taobebs aRniSnaven Semdegnairad: R0 - mcenare-regeneranti; R1 R2

da a.S. – R0 -is pirveli, meore da Semdgomi TviTdamtverili Taoba.
32
sxva obi eqtebi mcenare protoplastebi

kalusi

suspenziuri kultura

stresis faqtorebian agarizebul sakveb

areze gadaTesva
(herbicidi, paTotoqsini, antibi-
otiki, marilebi, mZime metalebi,
aminomJavebis analogebi, osmo-
suri stresi, dabali pH, eqstrem-
aluri temperatura da sxva)

seleqti uri faqtori sadmi gamZle ujredebi s

gamorCeva

gamZle kalusi

O organogenezis i nduqcia

M mcenare-regeneranti

Semow meba gamZleobaze

Semow meba gamZleobaze

in vitro stresis kulturaluri ekologiurad stresuli fo-
faqtorisadmi (herbicidi, paTo- nisadmi (cocxali paTogebi, mZi-
toqsini, antibiotiki, marilebi, me metalebi niadagSi, mJave ni-
mZime metalebi, osmosuri stre- adagebi, integraluri tempera-
si, mJave are, eqstremaluri tem- turul-tenianobiTi reJimi)
peratura da sxva)

s ur. 1 6. ujreduli s eleqciis zo gadi s qema

33
 1.3.2. ujreduli seleqciis obieqtebi U

in vitro pirobebSi ujreduli seleqciis samuSaoebis Ca-

tarebisaTvis kvlevis obieqtis saxiT ZiriTadad iyeneben ka-
lusur qsovilebs, suspenziur kulturebs an izolire-bul
protoplastebs. aseve SeiZleba gamoyenebul iqnas somatu-
ri, anu androgenuri embrioidebi, izolirebuli Canasaxebi,
iseTi organogenuri eqsplantatebi, rogoricaa foTlis
segmentebi an mcenaris sxvadasxva meristemuli da Reroseu-
li nawilebi.
kvlevis obieqtis SerCeva damokidebulia ama Tu im sa-
xeobis mcenarisaTvis SemuSavebuli ujreduli teqnologie-
bis arsebobaze da kvlevis saboloo mizanze.
ujredul seleqciaSi xSirad iyeneben kalusur qso-
vils (sur. 17), romelic warmoadgens advilad misawvdom
obieqts.

s ur. 1 7. kalusuri q sovili

kalusuri qsovili anu kalusi – aris dediferencirebu-

li ujredebis masa, romelic warmoiqmneba mcenaris orga-
34
noenoebis ujredebis araorganizebuli proliferaciis12
gziT.
ujredul kulturebSi kalusuri qsovili miiReba
mcenaris eqsplantatis13, yvela sakvebi nivTierebisa da ori
hormonis - citokininebisa da auqsinebis Semcvel, xelovnur
sakveb areze kultivirebiT. kalusis misaRebad eqsplanta-
tis diferencirebulma ujredebma unda ganicados dedife-
rencireba (iwvevs auqsinebi), anu kvlav SeiZinos dayofis
unari – daubrundes meristemul mdgomareobas da Semdgomi
gamravlebiT (proliferaciiT) (iwvevs citokininebi), araor-
ganizebuli anarqiuli zrdiT warmoqmnas kalusuri qsovi-
li.
uSualod eqsplantatidan miRebul kaluss ewodeba
pirveladi kalusi, xolo misgan miRebuls – meoradi (gada-
Tesili)14. kalusis axal sakveb areze gadatanis process pa-
sireba (pasaJi) anu kultivacia hqvia, xolo axal sakveb
areze gadatanili kalusis gamozrdis process - subkulti-
vireba.
feris mixedviT kalusi SeiZleba iyos moTeTro, moyvi-
Talo, moyavisfro, yavisferi, mTlianad an nawilobriv
qlorofiliT pigmentirebuli an antocianuri; konsistenci-
is mixedviT kalusuri qsovili aris fxvieri, saSualo sim-
kvrivis an mkvrivi. formis mixedviT gvxvdeba momrgvalo,
wagrZelebuli. arCeven homogenur (Sedgeba erTi tipis uj-
redebisagan) da heterogenur (Sedgeba ramodenime tipis uj-
redebisagan) kalusebs.
in vitro pirobebSi kalusuri qsovilis gamozrda SeiZ-
leba ganusazRvrelad didi xnis ganmavlobaSi, periodulad
axal sakveb areze gadaTesvis gziT.

12
proliferacia – ujredebisa da qsovilebis axalwarmoqmna ukve arsebu-
lis gamravlebis gziT.
13
eqsplantati (anu eqsplanti) –kalusis misaRebad gamoyenebuli mcenaris
qsovilis an organos fragmenti. amJamad kalusuri qsovilis miReba SeiZ-
leba mcenaris praqtikulad nebismieri qsovilidan an organodan.
14
kalusis nawils, romelic gadaaqvT axal sakveb areze transplantanti
ewodeba.
35
 kalusuri qsovilis Semdgomi ganviTareba SeiZleba wa-
rimarTos sami gziT:
1. meoradi diferencireba, morfogenezi da mTliani
mcenaris regeneracia;
2. kalusuri qsovilis Semdgomi dediferencireba da
mis mier morfogenezisa da regeneraciis unaris daqveiTeba
an mTliani dakargva;
3. kalusuri ujredebis Cveulebrivi onTogenezi,
romelic mTavrdeba maTi daberebiTa da sikvdiliT.
kalusuri qsovilidan regenerirebul mcenareebs kali-
klonebi ewodeba.
U ujredul seleqciaSi, rogorc wesi, iyeneben pirvelad
an pasirebad kaluss, romelsac ara aqvs dakarguli rege-
neraciis unari. Tumca kalusur qsovils, rogorc seleqci-
uri muSaobis obieqts, gaaCnia garkveuli uaryofiTi mxa-re-
ebi: qsovilebis neli zrda, seleqtiur faqtorad gamo-yene-
buli toqsikuri nivTierebis araTanabari moqmedeba ka-lu-
sis yvela ujredze, agreTve genetikuri arastabiluroba da
ujredebis mier regeneraciuli unaris dakargva kulti-vi-
rebis procesSi da a.S.
ra Tqma unda, yvelaze kargia seleqciuri procesis
warmarTva calkeuli ujredebis doneze (suspenziuri kul-
turebi, protoplastebi), magram mravali saxeobisaTvis es
teqnologia jer srulyofilad damuSavebuli ar aris.
amitom naklovanebebis miuxedavad, kalusuri qsovili,
rogorc kvlevis obieqti, zogierTi saxeobisaTvis (sadac
SeuZlebelia kargi suspenziuri kulturebisa da protop-
lastebis miReba) da aseve im SemTxvevaSi, roca seleqtiuri
faqtori cudad ixsneba wyalSi, jer-jerobiT Seucvlelia.
marilebis maRali koncentraciebisa da toqsinebisadmi
gamZleobis, aseve aminomJavebis maRali Semcvelobis mimar-
TulebiT seleqciaSi, kalusuri qsovilis gamoyenebiT miRe-
bulia axali mcenareebi xorbalSi, qerSi, brinjSi, sorgo-
Si, kartofilSi, stafiloSi, pomidorSi da sxva.
ujredul seleqciaSi kvlevis obieqtad yvelaze xSirad

36
iyeneben izolirebul protoplastebs15 (sur. 18). amis Ziri-
Tadi mizezi sxvadasxva genetikuri manipulaciebis, maT So-
ris somaturi hibridizaciis da maTSi sxvadasxva organiz-
mis genetikuri informaciis, sxvadasxva ujreduli organe-
lebis Seyvanis procedurebis Catarebis SesaZleblobaa.

s ur. 1 8. p rotoplastebi

kvlevis obieqtad protoplastebis gamoyenebis sxva upi-

ratesobebidan mniSvnelovania:
1. seleqciuri procesis warmarTva xdeba calkeuli
ujredebis doneze;
2. protoplastebis kultivireba saSualebas iZleva mi-
Rebul iqnas Zalian didi raodenobis cxovelmyofeli uj-
redebi, rac amorCeviTobis saSualebas iZleva;

15
zogadi informacia protoplastebis miRebisa da Serwymis Sesaxeb
motanilia qveTavSi 1.1.1. (gv. 8).

37
 3. kultivirebis specialuri xerxebis saSualebiT Se-
saZlebelia jvaredini kvebis (roca mutanturi ujredebi
inarCuneben aramutantebs, an roca aramutanturi ujredebi
inaqtivireben mTel kolonias, mutantebis CaTvliT) SesaZ-
leblobis minimumamde dayvana, rac praqtikulad gamori-
cxavs qimerebis16 gamoyofas;
4. axalgamoyofili mezofiluri protoplastebi gene-
tikurad da fiziologiurad SedarebiT erTgvarovania (ma-
Ti daaxloebiT 90% imyofeba G0 -periodSi 17);
5. protoplastebi saSualebas iZleva gamoyenebul iq-
nas ujreduli da genetikuri inJineriis sxvadasxva meTo-
debi;
6. protoplastebis miReba SeiZleba ara marti suspen-
ziuri kulturebidan da kalusidan, aramed uSualod in-
taqturi mcenaris qsivilebidanac.
M mcenareebi, saidanac unda miiRon protoplastebi, unda
akmayofilebdnen garkveul moTxovnebs:
Q 1. swrafi zrda in vitro kulturaSi;
2. regeneraciuli unaris SenarCuneba;
3. mokle sasicocxlo cikli;
4. haploidebis arseboba;
5. qromosomebis mcire ricxvi;
6. protoplastebis gamoyofisa da kultivirebis maRa-
lefeqturi meTodikis arseboba;
7. genetikuri ruqis arseboba.
protoplastebis kultivirebisaTvis iyeneben or xerxs:
1) Txevadi wveTebis meTodi da 2) platirebis18 meTodi.
pirvel SemTxvevaSi protoplastebis suspenzia wveTe-
bis saxiT Tavsdeba Txevad sakveb areSi plastikur petris
jamze. meTodi uzrunvelyofs karg gazTacvlas. garda ami-

16 qimera – organizmi, romelic Sedgeba genetikurad sxvadasxvanairi

ujredebisagan.
17
G0 -periodi – aris periodi, roca ujredi wyvets mitozur gayofas
(aRmoCndeba „mitozuri ciklis miRma“) da iwyebs diferencirebas.
18
platireba [ingl. plating] – ujredebis Tesva agarizebul sakvebze.
38
sa, advilia saWiro koncentraciis axali sakvebi xsnaris
damateba. Tumca am xerxiT kultivirebisas xSirad adgili
aqvs yoveli wveTis centrSi protoplastebis agregacias.
didi raodenobiT dagrovebisas isini warmoqmnian mniSvne-
lovani sididis fenolur da sxva toqsikur SenaerTebs,
rac xels uSlis warmatebiT kultivirebas. es meTodi aseve
araxelsayrelia im SemTxvevaSi, roca saWiroa protoplas-
tebis individualuri koloniis ganviTarebaze dakvirveba.
meore SemTxvevaSi protoplastebis suspenziis garkve-
ul moculobas asxamen Txevad sakveb areSi plastikur pet-
ris jamze da umateben imave moculobis 1% agar-agarian
igive sakveb ares. temperatura ar unda aRematebodes 45°C.
Semdeg petris jamebs webaven parafilmiT da aCereben gada-
brunebul mdgomareobaSi 28°C temperaturaze. am dros pro-
toplastebi fiqsirdeba garkveul mdgomareobaSi da fizi-
kurad dacilebulia erTmaneTisagan. am meTods aqvs mniSvne-
lovani upiratesoba. SesaZlebelia davakvirdeT erTi kon-
kretuli protoplastis ganviTarebis yvela procesis Tan-
mimdevrobas. meTodis uaryofiTi mxare mdgomareobs imaSi,
rom SesaZlebelia moxdes protoplastebis garkveuli da-
zianeba Tbil agar-agarTan maTi Serevis dros.
arsebobs am meTodebis ramdenime varianti.
protoplastebSi ujredis garsis warmoqmna iwyeba ma-
Sinve, fermentTaAxsnaris mocilebis Semdeg. zogierT kul-
turebSi garsi Cndeba kultivirebis dawyebidan 8-24 saaTis
Semdeg. zogierTSi ki igi ramdenime kviris an Tvis Semdegac
ar iwyeba. protoplastis garsis sinTezis erT-erT ZiriTad
ganmsazRvrel faqtors sakveb areSi warmoadgens saqaroza.
mis gareSe ujredis garsis sinTezi ar mimdinareobs.
protoplastebidan ujredebisa da mcenareebis regene-
racia warmoadgens gacilebiT rTul process, vidre ujre-
dis garsis formireba. mcenareebis regeneracia xorcielde-
ba an embriogenezis, an kalusis ganviTarebis da Semdeg mis-
gan morfogenezis induqciis gziT. mcenares, romelic rege-
nerirebulia protoplastis kulturidan, ewodeba protok-
loni.
39
 iseve rogorc protoplastebis SemTxvevaSi, calkeuli
ujredebis doneze seleqciuri procesis warmarTva xdeba
suspenziuri kulturebis (sur.19) drosac, roca maTi gada-

s ur. 1 9. s uspenziuri kultura

gadaTesva xdeba agarizebul seleqtiur areze. garda amisa,

ujreduli agregatebis mcire zomebi da kulturis Sedare-
biTi fiziologiuri homogenuroba, didi raodenobis ujre-
debze seleqtiuri faqtorebiT zemoqmedebis saSualebas iZ-
leva. aseve SesaZlebelia ujreduli populaciebis zrdasa
da metabolizmze egzogenuri faqtorebis moqmedebis gamok-
vleva, subkultivirebis procesis simartivis gamo ujredu-
li kulturis xazebis saimedod didxans SenarCuneba. amave
dros SedarebiT advilia aseTi kulturebidan mcenareTa
swrafi regeneracia. advilia aseve standrtuli mikrobio-
logiuri meTodebis gamoyeneba.
am da sxva upiratesobebis gamo, isini farTodaa gamo-
yenebuli sakvlevi obieqtad ujredul seleqciaSi.
amasTan, suspenziur kulturebs aseve gaaCniaT garkveu-
li naklovanebebi, rac dakavSirebulia maT maRal agregire-
basTan, romelic amcirebs calkeuli ujredebis ricxvs da
40
Sesabamisad – miRebuli Sedegebis efeqts.
suspenziuri kulturebis qveS igulisxmeba calkeuli
ujredebis an maTi mcire jgufebis kultivireba Txevad
sakveb areSi Cakidul mdgomareobaSi, maTi aeraciisa da
arevis pirobebis xelSemwyobi aparaturis gamoyenebiT.
suspenziuri kulturebis miRebis Cveulebriv xerxs
warmoadgens kalusuri qsovilis gadatana kolbaSi an spe-
cialur kulturalur WurWelSi (fermentatorSi) moTavse-
bul Txevad sakveb areSi, sadac ujredebis areva warmoebs
sanjRrevelas an sxvadasxva tipis roleris saSualebiT.
warmatebebi suspenziuri kulturebis miRebaSi dida-
daa damokidebuli imaze, Tu ramdenad kargadaa SerCeuli
da momzadebuli kalusuri qsovili: igi unda iyos faSari,
advilad iSlebodes mcire ujredul agregatebad (araume-
tes 10-12 ujredi) da calkeul ujredebad.
suspenziuri kulturebis kultivirebisaTvis iyeneben
Ria an daxurul sistemebs, periodul an uwyvet reJimSi.
rogorc zemoT aRiniSna, ujredul seleqciaSi kvlevis
obieqtad aseve iyeneben somatur, anu androgenur embrioi-
debs, izolirebuli Canasaxebs, foTlis segmentebs, mcena-
ris sxvadasxva meristemul da Reroseul nawilebs. garkve-
uli sirTuleebis gamo, maT farTo gamoyeneba ara aqvT, mag-
ram dReisTvis am mimarTulebiT mopovebulia calkeuli
warmatebebi. magaliTad, in vitro pirobebSi Teslidan izoli-
rebuli Canasaxebis kultivirebiTa da Semdgomi seleqciiT
miRebulia cilebis gaumjobesebuli Semcvelobis da ami-
nomJavebis analogebisadmi gamZle qeris mcenareebi; karto-
filis kulturisaTvis damuSavebulia qlorofildefeqtu-
ri da antibiotikebisadmi gamZle mutantebis misaRebad Re-
roebisa da kalmebis mutagenebiT damuSavebis efeqturi me-
Todi; natriumis qloridis maRali Semcvelobis sakveb are-
ze saSemodgomo xorblis mtvrianebis kultivirebiT war-
moqmnili somaturi embrioidebidan miRebulia damlaSebi-
sadmi gamZle regenerantebi da sxva.

41
1.3.3. in vitro ujreduli kulturebis meTodebi
mcenareTa seleqciaSi

mcenareTa seleqciur procesSi in vitro ujreduli kul-

turebis gamoyenebis ori mimarTulebas gamoyofen:
1) seleqciuri procesis daCqarebisa da gaadvilebis
meTodebi, anu damxmare meTodebi seleqciaSi;
2) axali genotipebis konstruirebis meTodebi, anu sa-
kuTriv ujreduli seleqcia (sur. 20).

seleqciuri procesis daCqa- axali gen otipebis kon strui-

rebi sa da gaadvilebis meTo- rebi s meTodebi , an u sakuT-
debi , an u damxmare meTodebi ri v ujreduli seleqci a
seleqci aSi
1. in vitro ganayofiereba (pro- 1. somaklonuri cvalebadoba;
gamuri SeuTavseblobis daZ- 2. somaturi hibridizacia;
leva); 3. in vitro inducirebuli muta-
2. izolirebuli ganayofie- cia;
rebuli Teslkvirtebis, nas- 4. mcenareTa transformacia:
kvebisa da moumwifebeli hib- a) birTvuli genebis gada-
riduli Canasaxebis kultu- tana,
ra (postgamuri SeuTavseb- b) citoplazmuri genebis
lobis daZleva); gadatana.B
3. letaluri hibridebis qso-
vilebidan mcenareTa regene-
racia (morfogenezi hibridu-
li qsovilebis kulturaSi);
4. haploidebis miReba;
5. axali jiSebis, hibridebis,
xazebis klonuri mikrogamav-
leba (xelovnuri Teslebis
Seqmnis CaTvliT);
6. mcenareuli genofondis
krioSenaxva (ujredebisa da
qsovilebis banki).

s ur. 2 0. ujreduli kulturebis m eTodebi m cenareTa s e l e q ciaS i

pirvel mimarTulebaSi Sedis Semdegi meTodebi: in vitro

42
ganayofiereba; izolirebuli ganayofierebuli Teslkvirte-
bis, naskvebisa da moumwifebeli hibriduli Canasaxebis
kultura; letaluri hibridebis qsovilebidan mcenareTa
regeneracia; haploidebis miReba; axali jiSebis, hibridebis,
xazebis klonuri mikrogamravleba; mcenareuli genofondis
krioSenaxva.
meore mimarTulebaSi Sedis Semdegi meTodebi: somak-
lonuri variantebis gamoyeneba; somaturi hibridizacia; in-
ducirebuli mutantebis miReba; transgenuri mcenareebis mi-
Reba.

1.3.3.1. in vitro ujreduli kulturebis damxmare

meTodebi mcenareTa seleqciaSi

sakuTvriv ujreduli seleqciis meTodebis ganxilvam-

de, mokled damxmare meTodebze. isini ar enacvlebian Cveu-
lebriv, tradiciul seleqciur process, aramed emsaxurebi-
an, exmarebian mas, aCqareben da aadvileben process.

1.3.3.1.1. in vitro ganayofiereba

(progamuri SeuTavseblobi s daZleva)

igi tardeba im SemTxvevaSi, roca SerCeul mSobel

formebs Soris, bunebriv pirobebSi – ZiriTadad Soreuli
hibridizaciis dros, SeuZlebelia ganayofierebis ganxor-
cieleba. es ukanaskneli, romelsac progamuri SeuTavseb-
loba19 ewodeba, SeiZleba gamowveuli iyos ramodenime mize-
ziT: 1) fiziologiuri (mtvris marcvlis momwifebis vadis
acdena, samtvre milis zrdis SeCereba butkos dingze, mam-
robiTi gametis kvercxujredTan Serwymis uunaroba da
sxva); 2) morfologiuri (mokle samtvre mili, misi zrdis
blokireba sxvadasxva etapze, grZeli butkos sveti da
sxva).

19
progamuri SeuTavseblobis ZiriTadi mizezi taqsonomiurad Soreuli
partniorebis genetikuri SeuTavseblobaa.
43
 in vitro ganayofiereba SeiZleba ganxorcieldes ori
gziT: a) aseptikur pirobebSi svet damoklebuli, an mTlia-
nad mocilebuli naskvis xelovnur sakveb areze kultivire-
biT da gadanaWerze momwifebuli mtvris marcvalebis gada-
taniT; b) naskvis gaxsniT da xelovnur sakveb areze placen-
tis20 nawilebis (ganayofierebisaTvis mza TeslkvirtebiT)
kultivirdebiT da maT axlos an uSualod placentas qso-
vilebze momwifebuli mtvris marcvalebis gadataniT21.
vizualurad in vitro ganayofierebis procesis dadgena
martivia, vinaidan ganayofierebuli Teslkvirtebi gaunayo-
fierebeli Teslkvirtebisagan gansxvavebiT, ar gadadian
Sesvenebis mdgomareobaSi da swrafad matuloben zomaSi.
progamuri SeuTavseblobis daZlevis aRniSnuli meTo-
di damuSavebul iqna XX saukunis 50-80-ian wlebSi. kultivi-
rebisaTvis sakveb ared yvelaze metad iyeneben niCis ares
saqarozisa da kazeinis hidrolizatis damatebiT.
in vitro ganayofierebis meTodi ZiriTadad gamoiyeneba
balaxovan mcenareebSi. am gziT miRebul iqna cxovelmyofe-
li Teslebi yayaCosebrTa ojaxis sxvadasxva gvarSi; saxeo-
baTaSorisi hibridebi yayaCosebrTa ojaxSi; saxeobaTaSori-
si da gvareobaTaSorisi hibridebi ZaRlyurZenasebrTa, mixa-
kisebrTa, marcvlovanTa ojaxSi da a.S.

1.3.3.1.2. Teslkvirtebis, naskvebi sa da moumwifebeli

hibriduli Canasaxebis kultura
(postgamuri SeuTavseblobi s daZleva)

Soreuli hibridizaciis dros sirTuleebi Cneba gana-

yofierebis procesis Semdegac (xSirad am dros izRudeba
Teslkvirtebis, naskvebis zrda-ganviTareba, an warmoiqmneba
aRmocenebis uunaro, an dabali gaRivebis unaris mqone Tes-
lebi, an sicocxlisuunaro aRmonacenebi da sxva), mas pos-
tgamuri SeuTavsebloba ewodeba. misi ZiriTadi mizezi igi-

20
placenta – naskvSi Teslkvirtis Casaxvisa da mimagrebis adgili.
21
mas placentarul ganayofierebas uwodeben.
44
vea, rac progamuri SeuTavseblobis SemTxvevaSi (gv. 43), da
vlindeba hibriduli Canasaxebis zrdisa da ganviTarebis
SeCerebis saxiT, Canasaxisa da endospermis ganviTarebis
tempebis gansxvavebulobis an mdedrobiTi mcenaris qsovi-
lebis toqsikurobis gamo; pirveladi fesvis an Canasaxis
kvirtebis zrdis blokirebis saxiT da sxva.
postgamuri SeuTavseblobis dasaZlevad dReisaTvis
yvelaze metad iyeneben izolirebuli ganayofierebuli Tes-
lkvirtebis, naskvebisa da moumwifebeli hibriduli Canasa-
xebis xelovnur sakveb areze kultivirebis meTods.
pirvelad izolirebuli ganayofierebuli Teslkvir-
tebis kultivireba ganxorcielda 1958 wels yayaCos mcena-
reSi.
am mizniT mcenaris damtvervidan 2-6 dRis Semdeg xde-
boda Teslkvirtebis (Seicavdnen zigotas an orujredian
proembrios da endospermis ramdenime birTvs) gamoyofa da
kultivireba sakveb areze, romelic Seicavda mineralur ma-
rilebs niCis mixedviT, vitaminebs uaitis mixedviT, 5% saqa-
rozas (Canasaxebis ganviTarebas aCqarebda sakveb areSi kine-
tinis an kazeinis hidrolizatis damateba).
izolirebuli ganayofierebuli Teslkvirtebis kul-
tura warmatebiT gamoiyeneba bambis, papaias, wiTeli mocxa-
ris, mixakis, petunias da sxva mcenareebis SemTxvevaSi.
ganayofierebuli naskvebis kultivirebis pirveli ga-
mokvlevebi Catarebul iqna 1949-1951 wlebSi pomidoris mce-
nareSi. amJamad ganayofierebuli naskvebis kulturas far-
Tod iyeneben kombostosnairTa (kombosto, mdogvi, rafsi,
Talgamura da sxva) Soreuli hibridizaciis dros.
damtveril naskvebs 3-7 dRis Semdeg gamohyofen deda
mcenaridan, ukeTeben sterilizacias da gadaaqvT kultura-
Si murasige-skugis sakveb areSi kazeinis hidrolizatis da-
matebiT. 3-4 kviris Semdeg naskvidan ganviTarebuli hibri-
duli Canasaxebis gadaitaneba murasige-skugis an hamborgis
sakveb areSi zrdis stimulatorebis gareSe, sadac isini vi-
Tardebian hibridul mcenared.

45
 ganuviTarebeli hibriduli Canasaxebis22 kultivirebas
xelovnur sakveb areze ewodeba embriokultura. pirvelad
moumwifebeli hibriduli Canasaxebis warmatebiT kultivi-
reba Catarebul iqna 1925 wels selis mcenareSi. kultivire-
bisaTvis Canasaxebs iReben axali nayofebis Teslebidan,
Semdeg gadaaqvT murasige-skugis, uaitis, hamborgis da sxva
modificirebul areebze. naxSirwylebidan yvelaze metad
iyeneben saqarozas. Canasaxis zrdasa da ganviTarebaze mas-
timulirebel moqmedebas axdenen aminomJavebi (xSirad iyene-
ben glutamins da kazeinis hidrolizats) da vitaminebi (B,
C, inozitoli, biotini da sxva). rig SemTxvevaSi sakveb
ares umateben qoqosis rZis moumwifebeli endospermis eq-
straqtebs, qeris endosperms, pomidoris, qacvis nayofebis
wvenebs da sxva. kultivirebas axdenen 25. . .30°C temperatu-
ris pirobebSi. zogierT SemTxvevaSi (mag., citrusovnebi) Ca-
nasaxebis normaluri ganviTareba mimdinareobs sibneleSi.
hibriduli Canasaxebis kulturas farTod iyeneben xe-
xilovani mcenareebis, marcvlovnebis, bambis, mzesumziras,
sakvebi balaxebis, pomidoris, xaxvis, Tambaqos da sxva kul-
turebis seleqciis dros.
Canasaxebis kultura, rogorc damxmare meTodi Soreu-
li hibridizaciisas, aseve warmatebiT gamoiyeneba Zvirfasi
hibridebis swrafi mikrogamravlebisaTvis.Aam mizniT mas
iyeneben im SemTxvevaSi, roca hibriduli Canasaxisa da en-
dospermis SeuTavseblobis gamo, bunebriv pirobebSi SeuZ-
lebelia hibriduli Teslidan normaluri mcenaris miReba.
am dros mikrogamravleba mimdinareobs kalusuri qsovili-
dan kalusogenezis, morogenezis induqciisa da mcenare-re-
generantis miRebis gziT. moumwifebeli Canasaxebis kloni-
rebis teqnika mcenareTa Zvirfasi genotipebis sasicocxlo

22
postgamuri SeuTavseblobis dasaZlevad farTod iyeneben agreTve embri-
okulturis meore saxes – izolirebuli momwifebuli Canasaxebis kulti-
virebas. mas iyeneben ganuviTarebeli Teslebidan Canasaxebis gaRivebisaT-
vis, seleqciuri procesis dasaCqareblad Teslebis mosvenebis mdgomareo-
bidan gamoyvanisaTvis, Zvirfasi hibridebis da sxva.
46
ciklis adreul stadiaze gamravlebis saSualebas iZleva.
igi farTod gamoiyeneba merqnian tyis mcenareebSi (muxa,
fiWvi, alvis xe, naZvi da sxva).

1.3.3.1.3. letaluri hibridebis qsovilebidan mcenareTa

regeneracia
(morfogen ezi hi bri duli qsovi lebi s kul turaSi )

zogjer anomalurad ganviTarebuli, letaluri hibri-

debi, saWiroebis SemTxvevaSi, SeiZleba Senaxul iqnes Sem-
dgomi gamoyenebisaTvis. amisaTvis momakvdavi hibridis co-
cxali qsოvilebidan Rebuloben kaluss, Semdeg morfogene-
zis induqciiT – mcenare-regenerants.

1.3.3.1.4. haploidebis mi Reba

Zalian didia haploidebis roli seleqciur muSaobaSi.

haploidebi uzrunvelyofen TviTdamtverili mcenareebis
tolfas, heterozisul seleqciaSi didi mniSvnelobis mqo-
ne, stabiluri homozigoturi mcenareebis daCqarebul miRe-
bas; haploidebi saSualebas iZleva swrafad moiZebnos saWi-
ro kombinacia, amokleben jiSebis gamoyvanis dros da amci-
reben saseleqcio masalis raodenobas; haploidebi moxerxe-
bulia mutagenezisaTvis, radganac haploidur doneze advi-
lia recesiuli mutaciis gamovlena, maTSi dominirebis mov-
lenis uqonlobis gamo, aq yoveli geni warmodgenilia er-
TaderTi aleliT da fenotipi mTlianad asaxavs maT geno-
tips; haploidebi ujredebisa da qsovilebis kulturaSi se-
leqciisaTvis axali unikaluri genotipebis miRebis saSua-
lebas iZleva; haploidebi warmoadgenen Sesaferis sawyis
masalas seleqciaSi farTod gamoyenebuli monosomikebisa
da polihaploidebis miRebaSi da sxva.
Cveulebriv haploidebi steriluria, radganac qromo-
somebis ganaxevrebuli (n) ricxvis gamo maTSi darRveulia
mamrobiTi da mdedrobiTi gametebis formireba, magram uj-
redebis in vitro kultivirebisas spontanurad, an xelovnu-
47
rad, kolxiciniT damuSavebis Sedegad, SeiZleba moxdes
qromosomebis ricxvis gaormageba da fertiluri homozigo-
turi dihaploidebis miReba.
haploidebi SeiZleba warmoiqmnas spontanurad, magram
maTi aseTi warmoSobis sixSire Zalian dabalia da Sead-
gens daaxloebiT 1.10-5 –1.10. -6 haploidebis miRebis sixSiris
gadidebisaTvis iyeneben sxvadasxva meTods. maTgan yvelaze
ufro efeqturs warmoadgens ujreduli kulturebis meTo-
di, romelebic uzrunvelyofen didi raodenobiT, cxovel-
myofeli haploidebis miRebas,
ujreduli kulturebis meTodis gamoyenebiT haploi-
debis miRebis sami xerxi arsebobs:
1. androgenezi – haploidebis miReba izolirebuli
mtvrianebisa da mtvris marcvlebis (mikrosporebis) xelov-
nur sakveb areze kultivirebiT;
2. ginogenezi – haploidebis miReba izolirebuli gau-
nayofierebeli naskvebisa da Teslkvirtebis xelovnur sak-
veb areze kultivirebiT;
3. parTenogenezi – haploidebis miReba mSobelTa qro-
mosomebis SeuTavseblobis gamo mamrobiT qromosoma elimi-
nirebul hibriduli Canasaxis xelovnur sakveb areze kul-
tivirebiT.
ufro xSirad gamoiyeneba pirveli xerxi. pirvelad hap-
loiduri mcenareebi miRebul iqna 1964 wels lemas mtvria-
nebis kultivirebis gziT. mas Semdeg aRniSnuli meTodiT
haploiduri mcenareebi miRebulia 200-ze saxeobis mcenari-
dan. androgenezis Sedegad miRebul mcenareebs ewodeba an-
drogenuli haploidebi.
androgenezi SeiZleba iyos ori tipis:
1) pirdapiri androgenezi – haploiduri mcenareebi
warmoiqmneba mtvris marcvlis embriogenezis procesSi:
mtvrianas SigniT calkeuli mtvris marcvlebi (mikrospore-
bi) asinqronulad iyofa da warmoiqmneba embrioidebi (Cana-
saxismagvari struqturebi), romlebic Semdeg Rivdeba da iZ-
leva haploidur mcenareebs (damaxasiaTebelia ZaRlyurZena-
sebrTa ojaxis zogierTi gvarisaTvis – Tambaqo, lema da
48
sxva).
2) arapirdapiri (anu kalusuri) androgenezi – haplo-
iduri mcenareebi warmoiqmneba mtvrianas an mtvris mar-
cvlis (mikrosporis) kalusuri qsovilis organogenezis an
embriogenezis procesSi: mtvris marcvlis ujredebi dedi-
ferencirdeba, iyofa, warmoiqmneba kalusuri qsovili, ro-
melSic Semdeg an embriodebi viTardeba, an xdeba organoge-
nezis inducireba haploiduri mcenareebis Semdgomi formi-
rebiT (damaxasiaTebelia marcvlovanTaTvis – qeri, brinji,
xorbali, asparagusi da sxva).
haploidebis misaRebad ufro perspeqtiuls warmoad-
gens pirdapiri androgenezi, radganac am dros kalusuri
qsovili SeiZleba warmoiqmnas ara marto mikrosporidan,
aramed mtvrianas somaturi ujredebidanac, ris Sedegad vi-
Tardeba sxvadasxva ploidobis heterozigotuli mcenaree-
bi.
androgenezis dros haploidebis misaRebad yvelaze me-
tad iyeneben mtvrianebs, romlebsac sterilurad gamohyo-
fen kokrebidan ganviTarebis Sesaferis fazaSi da axdenen
kultivirebas xelovnur sakveb areze (mtvrianebis kultu-
ris meTodi). magram mtvrianebis kultivirebas gaaCnia rigi
uaryofiTi mxarec: jer-erTi, am meTodis dros SesaZlebe-
lia mcenareebis warmoSoba ara marto mikrosporis ujrede-
bidan, aramed mtvrianas somaturi ujredebidanac, radganac
mtvrianaSi mikrospora (mtvris marcvali) moTavsebulia mi-
si somaturi ujredebis SigniT, ris gamoc warmoiSoba sxva-
dasxva ploidobis mcenareebi; meore, mtvrianas kedlebs Se-
uZliaT androgenezis mainhibirebeli nivTierebebis produ-
cireba. amitom ufro efeqturs, magram Sromatevads warmo-
adgens mtvris marcvlebis kultivireba, radganac isini Ta-
visufalia mtvrianas somaturi ujredebisagan (mtvris mar-
cvlebis kulturis meTodi).
meore xerxiT haploiduri mcenareebi pirvelad miRe-
bul iqna 1976 wels qerSi. ukanasknel wlebSi mas xSirad
iyeneben mamrobiTi xaziT sterilur da kulturaSi mtvria-
nebis dabali morfo-genetikuri potencialis mqone mcenare-
49
ebSi. ginogenezis Sedegad miRebul mcenareebs ewodeba gi-
nogenuri haploidebi.
ginogenezic SeiZleba iyos ori tipis:
1) pirdapiri ginogenezi – haploiduri mcenareebi war-
moiqmneba Canasaxis parkis ujredebidan (kvercxujredi, an-
tipodebi, sinergidebi) embrioidebis ganviTarebis procesSi
(axasiaTebs xorbals, qers, siminds da sxva mcenares).
2) arapirdapiri (kalusuri) ginogenezi – haploiduri
mcenareebi warmoiSveba Canasaxis parkis ujredebidan (kver-
cxujredi, antipodebi, sinergidebi) kalusuri qsovilis
organogenezis an embriogenezis procesSi (axasiaTebs Saq-
ris Warxals, siminds da sxva mcenares).
dReisaTvis ginogenuri haploidebi miRebulia rig mce-
nareebSi, maT Soris xorbalSi, brinjSi, TambaqoSi, Saqris
WarxalSi, simindSi, mzesumziraSi, SroSanSi, bambaSi, kar-
tofilSi da sxva.
parTenogenezis arsi mdgomareobs imaSi, rom garkveu-
li saxeobebis Sejvarebisas (mag. kulturuli qeri - Horde-
um vulgare) da veluri qeri – Hordeum Bulbosum) hibriduli Ca-
nasaxis ujredebis dayofis dros, genetikuri SeuTavsebu-
lobis gamo, adgili aqvs mamrobiTi qromosomebis elimina-
cias da haploiduri Canasaxis ganviTarebis. magram Semdeg
Canasaxis normaluri ganviTareba deda mcenareze muxruWde-
ba, ris gamoc xdeba misi izolireba da in vitro xelovnur
sakveb areze gamozrda.
am xerxiT haploidebis miRebulia qerSi, xorbalSi,
WvavSi.

1.3.3.1.5. axali ji Sebis, hibridebis, xazebis

klonuri mikrogamavleba
(„ xelovn uri Teslebi s“ Seqmn i s CaT vli T)

ganuviTarebeli hibriduli Canasaxebidan embriokul-

turis meTodiT mcenareebis gamozrdisas sicocxlisunaria-
ni hibridebis gamosavlianoba Zalian mcirea, amasTan isini
xSirad steriluria. amitom mecnierebis winaSe xSirad dgas
50
miRebuli mcenareebis swrafi gamravlebisa da Senaxvis sa-
kiTxi. amave dros xSirad seleqciis meTodebiT miRebuli
sxvadasxva saintereso hibridi, xazi saWiroebs drois mok-
le monakveTSi swraf gamravlebas da sawyisis identurebis
didi raodenobiT miRebas. axali seleqciuri jiSis warmoe-
baSi danergvis gzaze erT_erT arsebiT winaaRmdegobas er-
Ti sezonis ganmavlobaSi didi raodenobiT Teslebis an
sargavi masalis (vegetiurad mravlebadi mcenareebisaTvis)
miRebis SezRuduloba warmoadgens. xSirad Zvirfasi seleq-
ciuri masalis SenarCuneba fitopaTogenuri infeqciebisagan
dazianebebis gamo SeuZlebelia. Semdgomi seleqciuri muSa-
obisaTvis saWiroa jiSebisa da saxeobebis koleqciis Seq-
mna.
aRniSnulebis gadawyvetaSi farTod gamoiyeneba klo-
nuri mikrogamravlebis meTodi. saseleqcio masalis mikro-
gamravleba xdeba gverdiTi kvirtebis meristemis, adventuri
kvirtebis aqtivaciiT, aseve Canasaxebis kalusuri qsovile-
bidan morfogenezis induqciiT mcenare-regenerantebis miRe-
biT.
seleqciur muSaobaSi, gansakuTrebiT Zneladgasamrav-
lebeli (mcire zomis Teslebi, ganuviTarebeli endospermi,
heterozigoturi Tesli, gaRivebisaTvis mikorizuli sokos
saWiroeba, Teslis ganuviTa-
rebloba, Zviri da Sromateva-
di vegetatiuri gamravlebis
teqnologia da sxva) jiSebisa
da hibridebis SemTxvevaSi,
efeqturia agreTve e.w. „xelov-
nuri Teslebis“ (sur.21) ga-
sur. 21. „xelovnuri Teslebi“ moyeneba.
„xelovnuri Tesli“ war-
moadgens damcav polimerul matricaSi moTavsebul mcena-
ris somatur embrioidebs, kenwrul da gverdiT kvirtebs,
Reros da fesvis segmentebs. amJamad aseTi „xelovnuri Tes-
lebis“ miRebisas (ZiriTadad granulebis saxiT) mcenareuli
masalis inkafsulireba xdeba alginatis gelSi, romelic
51
Semdeg ifareba polimeriT. SegniT Tavsdeba Sesabamisi sak-
vebi nivTierebebi hormonebis gareSe, antibiotikebi da sxva.

1.3.3.1.6. mcenareuli genofon dis krioSenaxva

(ujredebi sa da qsovi lebi s ban ki )

seleqciuri muSaobisaTvis mniSvnelovan xelSemwyob

meTods warmoadgens mcenareTa ujredebisa da qsovilebis
in vitro Senaxva Txevadi (-1960С) an gaziseburi (-150°C) azotis
temperaturis pirobebSi, romlis farTo ganviTareba daiwyo
gasuli saukunis 70-iani wlebidan.
igi saSualebas iZleva xangrZlivi droiT in vitro kul-
turaSi Senaxul iqnas mcenareuli genofodi da aseve nebis-
mier dros seleqcioneri uzrunvelyofili iyos saWiro
niSnis matarebeli genotipis mqone masaliT: aucilebeli
mtverianebi da mtvris marcvlebi SejvarebisaTvis; unikalu-
ri da erTeuli Teslebi; cxovelmyofeli hibriduli, mu-
tanturi, transformirebuli ujredebi in vitro morfogenezi-
saTvis; zigoturi da somaturi Canasaxebi, meristemebi, ka-
lusuri qsivilebi da sxva.
dReisaTvis damuSavebulia krioSenaxvis23 pirobebi mra-
vali saxeobis ujredisaTvis, kalusuri qsovilebisaTvis,
izolirebuli protoplastebisaTvis, meristemebisaTvis da
sxva.
krioSenaxvis arsi mdgomareobs imaSi, rom specia-
lur sinjarebSi an ampulebSi moTavsebuli mcenareTa uj-
redebi da qsovilebi garkveuli damuSavebis Semdeg iyineba
da inaxeba Semdgomi galRobiT Zalian dabal temperatura-
ze: gaziseburi azotis (daaxloebiT -150°C) an Txevadi azo-
tis (-196°C) temperaturaze.
am dros ujredebi da qsovilebi gadadis Rrma civi
anabiozis mdgomareobaSi, e.i. Seqcevadad Cerdeba an ukidu-
resad muxruWdeba metabolituri da biologiuri daSlis

23mcenareuli obieqtis gayinul mdgomareobaSi Senaxvas krioSenaxva (berZ.

„Kryos“-sicive, yinva, yinuli) ewodeba.
52
procesebi. garda amisa, am gziT Senaxuli masala rCeba ge-
netikurad stabiluri, adgili aqvs miwis farTobebis, mate-
rialuri da SromiTi danaxarjebis SesamCnev ekonomias.
amasTanave, garantirebulia masalis Senaxva praqtikulad
nebismieri drois ganmavlobaSi.
krioSenaxvis teqnologiuri procesi mravaletapiania.
igi Sedgeba Semdegi mniSvnelovani da urTierTdakavSirebu-
li etapisagan: 1) ujredebisa da qsovilebis momzadeba; 2)
krioproteqtorebis damateba; 3) programuli gayinva; 4) Txe-
vad an gazisebur azotSi Senaxva; 5) galRoba; 6) kriopro-
teqtoris mocileba; 7) ujredebisa da qsovilebis cxovel-
myofelobis gansazRvra; 8) rekultivireba da mcenareTa re-
generacia.
krioSenaxvis warmatebisaTvis didi mniSvneloba aqvs
gayinvis reJims. gamomdinare iqedan, rom swrafi gayinvisas
wyali ver aswrebs ujredidan gamosvlas da mis SigniT
warmoiqmneba letaluri yinuli, xolo neli gayinvisas
vlindeba gauwyloebis (dehidrataciis) mavne efeqti da sak-
veb areSi marilebisa da metabolitebis maRali koncentra-
ciis toqsikuroba, iyeneben programul gayinvas, romelic
uzrunvelyofs ori urTierTsawinaaRmdego moqmedebis da-
mRupveli Zalis gawonasworebas, da maSasadame, ujredebisa
da qsovilebis maqsimalur gadarCenas. programuli gayinva
iTvaliswinebs temperaturis dawevas erT an ramdenime eta-
pad, mkacrad daculi mudmivi siCqariT yoveli etapis far-
glebSi.
dReisaTvis arsebobs gayinvis ramdenime meTodi:
1. neli TandaTanobiTi gayinva – winaswari momzadebisa
da krioproteqtorebiT damuSavebis Semdeg ujredebisa da
qsovilebis nimuSebs anawileben sinjarebSi an ampulebSi
da temperaturis dawevas axdenen mudmivi siCqariT, an 0°C-
ze dabla garkveul temperaturamde, xolo Semdeg sinjare-
bi an ampulebi gadaaqvT Txevad azotSi, an Txevadi azotis
bolo uaryofiT temperaturaze;
2. mSrali gayinva – winaswari momzadebisa da kriopro-
teqtorebiT damuSavebis Semdeg ujredebisa da qsovilebis
53
nimuSebs aTavisufleben ukanasknelisagan, anawileben sinja-
rebSi an ampulebSi da temperaturis dawevas axdenen igive
reJimiT, rogorc winaSi;
3. swrafi gayinva – winaswari momzadebisa da kriopro-
teqtorebiT damuSavebis Semdeg ujredebisa da qsovilebis
nimuSebs anawileben sinjarebSi an ampulebSi da pirdapir
aTavseben Txevad azotSi;
4. zeswrafi gayinva – winaswari momzadebisa da kriop-
roteqtorebiT damuSavebis Semdeg ujredebisa da qsovile-
bis nimuSebs afiqsireben, magaliTad, kanqveSa ineqciis nem-
sebze da aseTi saxiT pirdapir aTavseben Txevad azotSi.
mcenareuli obieqtebis krioSenaxvasTan dakavSirebuli
teqnologiebi Zalian swrafad viTardeba da srulyofili
xdeba.

1.3.3.2. ujredul seleqciaSi genetikuri

mravalferovnebis Seqmnis meTodebi

1.3.3.2.1. somaklonuri cvalebadoba

ujredebisa da qsovilebis kulturaSi ujreduli se-

leqciisaTvis genetikuri mravalferovnebis Seqmnis sainte-
reso meTods warmoadgens somaklonuri cvalebadoba.
somaklonuri cvalebadoba – aris ujredebisa da qso-
vilebis kulturaSi ujredebis mier araorganizebuli
zrdis stadiis gavlis procesSi warmoSobili genetikuri
cvalebadoba.
ujredebisa da qsovilebis kulturidan regenerirebul
mcenareebs ewodeba somaklonebi (somaturi klonebi), xolo
Tu maT axasiaTebT somaklonuri cvalebadoba – somaklonu-
ri variantebi.
didi xnis ganmavlobaSi miaCndaT, rom in vitro kultivi-
rebadi mcenaris ujredebi warmoadgenen genetikurad stabi-
lurs – mkacrad erTgvarovans. magram dadgenil iqna, rom
kalusur, suspenziur da protoplastur ujredebs aqvT ga-
moxatuli genetikuri heterogenuroba (arastabiluroba),
54
rac maTi ujredul seleqciaSi gamoyenebis safuZveli gax-
da.
kultivirebadi ujredebis genetikuri arastabiluro-
bis da maSasadame, somaklonuri cvalebadobis warmoSobis
ZiriTad mizezs warmoadgenen:
1. sawyisi eqsplantatis somaturi ujredebis genetiku-
ri heterogenuroba. bevr mcenareSi diferencirebuli qsovi-
lebi xasiaTdebian sxvadasxva ploidobis, aseve aneuplo-
iduri ujredebis arsebobiT (mxolod mcenaris onTogenez-
Si aqtiurad prolifirebadi ujredebi, iseTebi rogoricaa,
kenwruli meristemebi, kambiumi da sxva, rCebian yovelTvis
diploiduri). qimerizmi 24 da miqsoploidia25 maRali sixSi-
riT vlindeba apomiqtur26 da vegetatiurad mravlebad mce-
nareebSi. xSirad amis mizezi mcenareTa kultivirebis piro-
bebis cvlilebaa, gansakuTrebiT maTi mkveTri gauaresebaa.
aseTi movlenebi SemCneulia dabali da maRali temperatu-
ris, damlaSebis, mineraluri sasuqebis maRali dozebis,
Wrilobebis miyenebis, mcenaris uCveulo saarsebo garemoSi
gadatanis, ucxo mtvriT damtverianebis da sxva pirobebSi .
es da sxva, bunebaSi mudmivad mimdinare faqtorebi, rogorc
Cans, iwveven fiziologiur darRvevebs, romlis erT-erT Se-
degs warmoadgens anomaliuri mitozebi da sawyisisagan ge-
netikurad gansxvavebuli ujredebis miReba.
2. xangrZlivi pasirebis Sedegad ujredul kultureb-
Si genetikuri cvlilebis dagroveba. ujredebis xangrZlivi
pasireba xels uwyobs genetikuri mravalferovnebis warmoq-
mnas. ujreduli kulturebis in vitro xangrZlivi kultivire-
bisas somaklonur cvalebadobas ganapirobebs Semdegi meqa-
nizmebi: uxeSi kariologiuri darRvevebi; qromosomuli gar-
daqmnebi (translokacia, delecia, inversia, duplikacia);

24
sxvadasxva genotipis ujredebis Tanaarseboba erT organizmSi.
25
Uujreduli mozaicizmis forma – erTi saxeobaSi sxvadasxva ploidobis
ujredebSi arseboba.
26
Gganayofierebis gareSe mravlebadi.
55
mobiluri genetikuri elementebis gadaadgileba; genebis am-
plifikacia (anu dozis gadideba) da reduqcia; somaturi
krosingoveri, Svileuli qromatidebis gacvla; virusebis
kriptikuli eliminacia; citoplazmonis (mitoqondrionisa
da plastomis) cvalebadoba da sxva.
3. mcenaris qsovilis izolirebis da sakveb areSi mo-
Tavsebis dros korelaciuri kavSirebis darRveva. in vitro pi-
robebSi mTlian mcenareSi arsebuli mkacri regulaciis
moSla ujredis spontanur cvalebadobas iwvevs.
4. kultivirebis pirobebis gavlena, saxeldobr, fito-
hormonebis mutagenuri moqmedeba. sakvebi aris egzogenuri
fitohormonebi arRveven kultivirebadi ujredebis hormona-
lur balanss. Sedegad icvleba ujredis ciklis, kerZod,
mitozis kinetika da Cndeba qromosomebis sawyisi ricxvisa-
gan gansxvavebuli ricxvis mqone ujredebi. yvelaze ufro
aqtiuri mutagenuri fitohormonebi arian 2.4-d da kinetini.
analogiuri zemoqmedeba kultivirebad ujredebze SeuZli-
aT moaxdinon sakvebi aris sxva komponentebmac: mineraluri
elementebi, saqaroza, vitaminebi, qoqosis rZe da sxva, ag-
reTve kultivirebis reJimmac.
somaklonuri cvalebadoba mniSvnelovnadaa damokide-
buli sawyis genotipze, eqsplantis bunebasa da ganviTare-
bis stadiaze (asakze). zog saxeobas ufro maRali soma-
klonuri cvalebadoba axasiaTebs, vidre sxvas. amasTan es
cvalebadoba damokidebulia ara marto genomze, aramed ci-
toplazmonzec. somaklonuri cvalebadobis gamowvevis ar-
sebiT faqtors warmoadgens eqsplantis ploidurobis done
da heterogenuroba. cvalebadoba maRalia poliploidizire-
buli saxeobebis ujredebSi, vidre arapoliploidizirebu-
lebSi. maRalia cvalebadobis sixSire amfiploidur saxeo-
bebSi. somaklonur cvalebadobaze gavlenas axdens aseve eq-
splantis tipi (foToli, muxlSorisi, ylortis segmenti,
yvavili da sxva); morfogenezis tipi – somaturi embrioge-
nezis dros “ujredi – mcenare” stadiis gavlis dro gaci-
lebiT moklea, vidre organogenezis dros, amitom miRebu-
li mcenarisa da sawyisi mSobeli genotipis msgavsebis xa-
56
risxi SeiZleba gacilebiT maRali iyos; sawyisi masalis
fiziologiuri heterogenuroba, romlis drosac ujredebi
populaciaSi imyofebian sxvadasxva fiziologiur mdgoma-
reobaSi, anu iyofian, izrdebian, berdebian, kvdebian.
K in vitro kultivirebadi ujredebis genetikuri heteroge-
nuroba ganpirobebulia, kultivirebis procesSi maTSi mim-
dinare genetikuri, epigenetikuri da modifikaciuri cva-le-
badobebiT.
genetikuri, anu mutaciuri cvalebadobebi iwvevs geno-
tipis Secvlas da memkvidreobiT gadaecema. dnm-s raodeno-
bisa da struqturis cvlileba xdeba genis, qromosomis da
genomis doneze. genuri mutaciebi iwveven ujredis morfo-
logiuri, bioqimiuri da diziologiuri Tvisebebis Zlier
an sust cvlilebas. qromosomuli mutaciebi (qromosomuli
aberaciebi) warmoiqmneba inversiis, deleciis, duplikaciis
translokaciisa da transpoziciis Sedegad. genomuri muta-
ciebi dakavSirebulia birTvSi qromosomebis ricxvis cvli-
lebasTan, anu kariotipis cvlilebasTan. epigenetikuri 27
cvalebadoba, ar exeba dnm-s srtuqturis cvlilebas, magram
iwvevs (dnm meTilireba, qromatinis remodulireba, histon-
rbis modifikacia, rnm-inteferencia da sxva) genebis aqtiu-
robis memkvidrul cvlilebas. ujredul kulturebSi ara-
memkvidruli cvalebadobas miekuTvneba modifikaciuri cva-
lebadobebi, romlebic umetes SemTxvevaSi adaptur,DSegue-
biT xasiaTs atareben.
namdvili genuri cvalebadobisagan somaklonuri cva-
lebadobebi gansxvavdebian maRali sixSiriTa (sami rigiT
maRali) da cvalebadobis kompleqsurobiT. maTSi Zalian
farToa cvalebadobis speqtri – Cndebian cvalebadobebi
sxvadasxva bioqimiuri, morfologiuri, sameurneo, xaris-
xobrivi, citogenetikuri niSnebiT. udavoa, rom yvela cva-
lebadobebis realizeba ar xdeba – zogierTi maTgani on-
Togenezis procesSi TandaTan qreba. magram somaklonuri

27
epi (berZ.) – damateba.
57
cvalebadobebis umetesoba warmoadgens unikalurs. aseTi
cvalebadobebi dReisaTvis ar SeiZleba miRebul iqnas arc
erTi sxva gziT.
somaklonuri cvalebadobebi sawyisisagan gansxvavdebi-
an ara marto monogenuri da Tvisobrivi niSnebiT, aramed
poligenuri da raodenobrivi niSnebiTac, iseTebiT, rogo-
ricaa, zrdis intensivoba, produqtiuloba, garemos araxel-
sayreli faqtorebisadmi gamZleoba, virusebisadmi gamZleo-
ba da sxva. zogjer maTSi Cndeba niSan-Tvisebebis iseTi Se-
Tavseba, romelTa gaerTianeba erT genotipSi SeuZlebelia
an Zalian Znelia tradiciuli seleqciuri meTodebiT.

1.3.3.2.2. somaturi hibridizacia

somaturi hi bri di zaci i s arsi . ujredul seleqciaSi

didi praqtikuli mniSvnelobis mqone, genetikurad mraval-
ferovani formebis miRebis efeqtur meTods warmoadgens
somaturi hibridizacia.
somaturi hibridizacia aris mSobeli formebis soma-
turi ujredebidan izolirebuli protoplastebis Serwymis
gziT arasqesobrivi hibridebis miRebis meTodi. igi moicavs
oTx etaps: protoplastebis gamoyofa; protoplastebis Se-
rwyma; gamorCeva da mcenareTa regeneracia; mcenare-regene-
rantebis analizi.
mcenareebs, romlebic miRebulia protoplastebis Ser-
wymis gziT ganxorcielebuli somaturi hibridizaciis Se-
degad, ewodeba somaturi hibridebi.
somaturi hibridizaciis arsi mdgomareobs imaSi, rom
hibridizaciaSi iyeneben ara reproduqtiul ujredebs (game-
tebs), aramed mcenaris sxeulis (somatur) ujredebs (pro-
toplastebs). protoplastebis Serwyma uzrunvelyofs uj-
redebis ara marto genomebis, aramed sxvadasxva citoplaz-
mis gaerTianebas.
somaturi hi bri di zaci i s Tavi seburebebi . hibridebis
garda, protoplastebis Serwyma xSirad iwvevs e.w. cibride-

58
bis (citoplazmuri hibridebis) warmoqmnas, sadac organiz-
mi Seicavs erT-erTi mSoblis birTvs da an orive mSoblis,
an alternatiuli mSoblis ci-
birTvi mitoqondriebi toplazmas (sur. 22). cibridebs
qloroplastebi xSirad Rebuloben birTvis da-
Slis mizniT Serwymis win erT-
erTi mSoblis protoplastis
radiaciuli da qimiuri agente-
biT damuSavebis gziT.Ggarda
amisa cibridebi miiReba izoli-
variantebi
rebuli protoplastis cito-
plastTan SerwymiT, mikroineq-
ciiT, liposomebiT gadatanis
gziT.
sur. 22. c ibridis miR eb i s
ZiriTadi gansxvaveba sqesob-
sqema riv da somatur hibridebs Soris
mdgomareobs imaSi, rom sqesobri-
vi hibridizaciis dros adgili aqvs mdedrobiTi da mamro-
biTi birTvuli genomebis Serwymas da mxolod mdedrobiTi
citoplazmonis (qloroplasturi da mitoqondrialuri ge-
nomebis) arsebobas, somaturi hibridizaciis dros – ro-
gorc mdedrobiTi da mamrobiTi birTvuli genomebis, ise
mdedrobiTi da mamrobiTi citoplazmonebis Serwymas. garda
amisa, sasqeso ujredebs rogorc wesi, qromosomebis haplo-
iduri ricxvi aqvT, xolo somatur ujredebs, Tu isini
izolirebuli ar arian haploiduri mcenareebidan – qromo-
somebis diploiduri ricxvi. amitom somaturi hibridebi
warmoadgenen amfidiploidebs da ar saWiroeben hibridebis
fertilurobis aRsadgenad qromosomebis ricxvis inducire-
bul gaormagebas.
protoplastebis Serwyma mimdinareobs rogorc wyvi-
lad, ise sxvadasxva ricxvis monawileobiT. didi mraval-
birTviani Serwymis produqtebi iSleba. izolirebuli pro-
toplastebis Serwyma warmoebs SemTxveviT da amis gamo er-
Tnairi sixSiriT warmoiSoba rogorc homokarionebi, ise he-
terokarionebi. Serwymis Semdeg adgili aqvs ori ujredis
59
citoplazmis Serevas, mere ujreduli garsis regeneraciis
Semdeg ori birTvis sinqronul dayofas maTi Serwymis ga-
reSe, orbirTviani Svileuli ujredebis wamoSobiT, an bir-
Tvebis Serwymas mitozuri dayofis dros da erTbirTviani
Svileuli ujredebis wamoSobas.
somaturi hibridizaciis dros birTvuli genebi SeiZ-
leba memkvidrulad gadaeces rogorc ormSobliani – (hib-
ridSi SenarCunebulia orive mSoblis genomi) – aseT hibri-
debs hqvia “simetriuli,” ise erTmSobliani (hibridSi Senar-
Cunebulia erTi mSoblis genomi) an erTmSobliani meore
partnioris qromosomebis nawiliT – aseT hibridebs hqvia
“asimetriuli.” aseTi hibridebi xSirad warmoiqmneba filo-
genezurad Soreuli organizmebis ujredebis Serwymisas. am
dros, ori sxvadasxvagvari qromosomaTa nakrebis arakoor-
dinirebuli moqmedebebiT gamowveuli, ujredebis araswori
gayofis gamo, rig TaobebSi nawilobriv an mTlianad ikar-
geba (ganicdis eliminacias) erT-erT mSoblis qromosomebi.
erT-erTi mSoblis qromosomebis eliminacia SeiZleba ga-
mowveuli iqnas aseve radiaciuli da qimiuri agentebis ga-
moyenebiT. xSirad asimetriuli hibridebi ufro gamZle, na-
yofieri da cxovelmyofelia, vidre simetriuli.
citoplazmuri genomebis (plastomis, qromosomebSi da
mitoqondriebSi lokalizebulis, mitoqondriebSi lokali-
zebulis) memkvidreobis bedi somaturi hibridizaciis dros
principulad gansxvavdeba aseTisagan sqesobrivi hibridiza-
ciis dros.
rogorc cnobilia, sqesobrivi hibridizaciis dros
citoplazmuri genomebi memkvidreoben mdedrobiTi xaziT,
xolo somaturi hibridizaciis dros isini memkvidreoben
ormSobliani (e.i. hibridSi SenarCunebulia orive mSoblis
citoplazmuri genomi). aseve adgili aqvs maT segregacias
(dacilebas Svileul ujredebs Soris), romelic mimdinare-
obs birTvSi mimdinare procesebisagan damoukideblad. ci-
toplazmaturi genebis mixedviT cvalebadobis damatebiT
wyaros warmoadgens mitoqondriebisa da qloroplastebis
genebis rekombinacia.
60
 ujredis organelebis genomebis eliminaciis an segre-
gaciis (dacilebis) Sedegad
mitoqondriebi
birTvi SeiZleba miRebul iqnas bi-
qloroplastebi
rTvuli da citoplazmuri
genebis sxvadasxvanairi uC-
veulo kombinaciebi (sur.
23), rac genetikuri mra-
valferovnebis Seqmnis mniS-
vnelovani wyaroa somaturi
variantebi hibridizaciis dros.
somaturi hi bri di zaci i s
produqtebi s gamorCeva.
izolirebuli protoplas-
tebis Serwymis procesis
damTavrebis Semdeg mniSvne-
lovan amocanas warmo-ad-
gens Serwymuli protop-
sur. 23. b i r Tvul i da c i t op-
l azmuri gen omeb i s S esaZl o lastebis identifikacia da
komb i n ac i eb i s sqema gamorCeva anu skriningi da-
narCeni protoplastebisa-
gan 28.
amisaTvis yvelaze xSirad iyeneben Semdeg meTodebs:
meqanikuri izolacia. Serwymuli (hibriduli) protop-
lastebis meqanikuri izolacia moicavs ramdenime etaps: 1)
Serwymuli protoplastebis vizualuri identifikacia, 2)
Serwymuli protoplastebis meqanikuri gamorCeva, 3) Serw-
ymuli protoplastebis kultivireba.
es meTodi gamoiyeneba mSobelTa ujredebis morfolo-
giuri kontrastulobis dros, magaliTad, roca erTi mSob-
lis protplastebi mdidaria qlorofiliT (gamoyofilia
mezofilis ujredebidan), xolo meore uqlorofiloa, ufe-
rulia (gamoyofilia kalusis ujredebidan). invertirebul
mikroskopis qveS Serwymuli protoplstebi vizualurad

28 Cveulebriv SerwymaSi monawileobs protoplastebis 3-30%.

61
advili gasarCevia mSobeli ujredebisagan. maTi izolireba
da kultivirebis sakveb areSi gadatana xdeba mikropipetis
saSualebiT. iyeneben agreTve protoplastebis cocxlad Se-
Rebvas fluorecentriuli saRebavebiT.
genetikuri komplmentacia. genetikuri komplementacia
dafuZnebulia ori araaleluri recesiuli letaluri ge-
nis matarebeli mSobelTa formebis gamoyenebaze, romelTa
ujredebi iRupebian kultivirebis garkveul pirobebSi, mag-
ram maTi protoplastebis Serwymis Sedegad miRebuli hib-
riduli ujredebi am pirobebSi gadarCebian genebis komple-
mentaciis29 wyalobiT. magaliTad SeiZleba moyvanil iqnas
ori, sxvadasxva recesiuli homozigoturi genis matarebe-
li, qlorofildefeqturi, intensiur ganaTebaze sicoc-
xlisuunaro, mSoblis protoplastebis Sewyma da genebis
komplementaruli moqmedebis Sedegad sicocxlisunariani,
qlorofildefeqturobis orive alelis mixedviT hetero-
zigoturi, muqimwvane hibridis miReba.
fiziologiuri komplementacia. meTodi dafuZnebulia
hibriduli ujredebis unarze gaizardon da gamravldnen,
an gadavidnen organizmul zrdaze (morfogeneze) kultivi-
rebis iseT pirobebSi, sadac mSobeli ujredebi amas ver
SeZleben. amasTan rig SemTxvevebSi mSoblebis es reaqcia
warmoadgens normalur fiziologiur reaqcias garemo pi-
robebze. erT-erT fiziologiuri komplementaciis meTodis
mixedviT protoplastebis Serwymis Semdeg ujredebis sus-
penzia jer gadaaqvT sakveb areze, sadac erTi mSoblis uj-
redebi ver izrdeba da eliminirdeba30, Semdeg gadaaqvT
sxva alternatiul sakveb areze, sadac ver izrdeba da
eliminirdeba meore mSoblis ujredebi. Sedegad gadarCeba,
gamravlebas da morfogenezze gadadis hibriduli ujredi.
protoplastebis inaqtivacia. meTodi gulisxmobs Serw-

29
genebis komplementacia – araaleluri genebis iseTi urTierTmoqmedeba,
romelic iwvevs sruliad axali niSnis ganviTarebas.
30
aseTi sakvebi areebis dadgena xdeba winaswar.
62
ymis win erTi an orive mSoblis protoplastis inaqtivaci-
as bioqimiuri SxamebiT an X- an γ-gamosxivebiT. magaliTad,
damuSvebulia cibridebis miRebis xerxi, erT-erTi mSoblis
protoplastis γ-sxivebiT winaswari damuSavebis Sedegad ge-
nomis mizanmimarTuli eliminaciis gziT.
fizikuri gamdidreba. meTodi dafuZnebulia masze,
rom centrofugirebis saSualebiT SeiZleba sxvadasxva xve-
driTi simkvrivis mqone protoplastebis fraqciebad dayo-
fa. Serwymul protoplastebs aqvT Sualeduri simkvrive.
somaturi hibridizaciis produqtebis gamosarCevad ga-
moiyeneba agreTve antibiotikebisadmi, herbicidebisadmi, ami-
nomJavebis analogebisadmi gamZleobis mixedviT gamorCevis
da sxva meTodebi.
somaturi hi bridebis an alizi . somaturi hibridi-zaci-
isas Serwymis procesis dros sistema Sedgeba mravalri-
cxovani (103–107) protoplastisagan. maTSi SeiZleba ganxor-
cieldes sxvadasxva gardaqmnebi – poliploidia, genuri,
qromosomuli gardaqmnebi. amitom, arasasurveli Sedegebis
Tavidan asacileblad, aucilebelia miRebuli Serwymis
produqtebis srulyofili analizi, risTvisac SemuSavebu-
lia hibridologiuri analizis, citogenetikuri Seswavlis,
bioqimiuri analizis meTodebi.
H hibridologiuri analizi. am meTodis dros adgili
aqvs winaswar specialurad SerCeuli mSobeli formebis
protoplastebis Serwymis Sedegad miRebuli Taobis feno-
tipur klasebad ganawilebis zust statistikur damuSave-
bas.
igi gamoiyeneba Semdeg dakvirvebebSi mSoblebisa da
maTi Taobebis genetikuri struqturis dadgenisaTvis. maga-
liTad, Tu somaturi hibridebi miRebulia recesiuli mu-
tantebis gamoyenebiT, maSin maTi TviTdamtvervisas mutacie-
bi fenotipurad gamovlindeba, rac hibridebis gamoyofis
saSualebas iZleva.
citogenetikuri Seswavla. meTodi gulisxmobs somatu-
ri hibridebis qromosomebis ricxvisa da morfologiis Ses-

63
wavlas, pirvel rigSi im SemTxvevaSi, roca sawyisi mSobeli
ujredebis qromosomebi erTmeneTisagan gansxvavebulia.
meTodi efeqturia Soreul kombinaciebSi. axlonaTesa-
uri hibridebis qromosomebze informacia birTvis mixedviT
hibridulobaze msjelobis saSualebas iZleva, Tumca nak-
leb sarwmunoa, vinaidan kulturaSi mutaciebis maRali six-
Sirea. citogenetikuri analizi citoplazmatur heterozi-
gotebSi birTvis mixedviT hibridulobis gamoricxvis kargi
meTodia.
bioqimiuri meTodebi. am dros xdeba fermentebis mra-
vali molekuluri formis Seswavla. farTod gamoiyeneba
fermentebis aqtiurobis gansazRvris eleqtroforezi. leq-
troforegramisa da fermentebis aqtiurobis Sedareba mSo-
belTa genebis arsebobis dadgenis saSualebas iZleva.
ase, fotosinTezis ZiriTadi fermentis - ribulozobi-
fosfatkarboqsilazas (rbfk) analizi saSualebas iZleva
erTdroulad miRebul iqnas informacia hibridizaciis
procesSi rogorc birTvuli, ise arabirTvuli genebis bed-
ze, vinaidan fermentis didi subnawilaki kodirdeba qlo-
roplasturi dnm-iT, xolo mcire subnawilaki – birTvuli
dnm-iT. specialuri aparatebis – seqvenatorebis saSuale-
biT dgindeba dnm-s nukleotiduri Tanmimdevroba. Sesaswav-
li dnm-s fragmentSi ama Tu im genis arseboba SeiZleba ga-
irkves hibridizaciis gziT. amisaTvis aucilebelia gasaana-
lizebeli dnm-s hibridizacia Ziebuli dnm-s Semcvel radi-
oaqtiurad niSandebul dnm-fragmentTan – zondTan.
Tu gasaanalizebeli dnm Seicavs Ziebul gens, maSin
masTan moxdeba radioaqtiuri zondis dakavSireba. rac
dgindeba gasaanalizebeli dnm-s radioaqtiuribis gansaz-
RvriT.
somaturi hi bridizaciis mniSvneloba. mcenareTa soma-
turi hibridizacia warmoadgens seleqciisaTvis genetikuri
mravalferovnebis Seqmnis mZlavr xerxs. igi sqesobrivi
procesis SezRudvebis daZlevis da axali mcenareebis kon-
struirebis saSualebas iZleva.

64
 am meTodis upiratesoba mdgomareobs imaSi, rom far-
Tovdeba saWiro genebis Soreul taqsonomiur mcenareebSi ga-
datanis SesaZlebloba. igi saSualebas iZleva gadailaxos
taqsonomiurad Soreuli saxeobebis Seujvareblobis buneb-
rivi barieri. man arsebiTad gaafarTova Soreul hibridi-
zaciaSi tradiciuli kvlevebis SesaZleblobebi.
somaturi hibridizacia uzrunvelyofs:
1. Cveulebrivi sqesobrivi gziT Seujvarebeli, filo-
genezurad Soreuli saxeobebis Sejvarebas;
2. ara marto sqesobrivad Seujvarebeli Soreuli sa-
xeobebis (gvarebis, ojaxebis) genebis SeerTebas birTvSi,
aramed hibridul ujredSi partniorebis citoplazmuri ge-
nebis SeTanawyobas;
3. erT-erTi mSoblis mTliani genomis da meoris ramo-
denime qromosomis (an ramdenime genis, an organelebis da
citoplazmis) matarebeli asimetriuli hibridebis miRebas;
4. erTdroulad sami da meti mSoblis ujredebis Ser-
wymis ganmaxorcielebeli hibridizaciis sistemis Seqmnas;
5. arabirTvuli genebis mixedviT heterozigoturi mce-
nareebis miRebas;
6. erTi mSoblis birTvis da meoris citoplazmis mqo-
ne mcenaris (cibridis) miRebas;
7. generaciuli sistemebis SeuTavseblobiT gamowveu-
li SezRudvis daZlevas;
8. genetikurad mSoblebis jamuri genotipis matarebe-
li hibridebis miRebas;
9. morfogenezSi an gametogenezSi anomaliebis gamo
sqesobrivi gziT Seujvarebeli mSoblebis Sejvarebas;
10. mTlianad steriluri mcenareTa formebis seleq-
ciur procesSi CarTvis SesaZleblobas;
11. mutaciebis heterozigotur mdgomareobaSi gadayva-
nas da amiT cxovelmyofeli formebis miRebas, vinaidan mu-
tagenezi xSirad morfogenezis mixedviT defeqtur mcenare-
ebs iZleva;
12. sxvadasxva epigenetikuri programis matarebeli
ujredebis Sejvarebas da sxva.
65
 somaturi hibridizaciis ZiriTad naklovan mxares
warmoadgens somaturi hibridebis, gansakuTrebiT saxeoba-
TaSorisi, gvareobaTaSorisi, tribaTaSorisi hibridebis, Se-
darebiT dabali regeneraciis unari. ris gamoc mas ufro
farTod iyeneben in vitro pirobebSi maRali regeneraciuli
potencialis mqone saxeobebSi (ZaRlyurZenasebrTa, kombos-
tosnairTa, SroSanisebrTa, qolgosanTa da sxva ojaxi).

1.3.3.2.3. in vitro inducirebuli mutacia

ujredul seleqciaSi genetikuri mravalferovnebis

miRebis farTod gavrcelebul meTods warmoadgens in vitro
inducirebuli 31 mutacia.
inducirebuli mutacia – aris mutagenuri faqtorebiT
anu mutagenebiT32 damuSavebis Sedegad ujredis genetikur
masalaSi uecrad warmoSobili memkvidruli cvalebadoba.
inducirebuli mutaciis Sedegad miRebul, Secvlili
memkvidruli niSnebis mqone organizmebs, inducirebuli mu-
tantebi ewodeba, xolo aseTi mutantebis warmoSobis pro-
cess – inducirebuli mutagenezi.
mutagenezi pirobiTad iyofa bunebriv da xelovnur
(inducirebul) mutagenezad. in vitro pirobebSi bunebrivad
mimdinare mutaciuri cvalebadoba SemTxveviTia da umete-
sad uaryofiTi. amasTan misi sixSire arc ise maRalia. ris
gamoc ver akmayofilebs seleqcionerebis moTxovnebs gene-
tikur mravalferovnebaze.
ujredul seleqciaSi procesis daCqarebisaTvis auci-
lebelia maRali sixSiris da mizanmimarTuli, mravalfero-
vani, dadebiTi mutaciuri cvalebadobebi. amitom mimarTaven
cnobili mutagenuri faqtorebis Segnebul gamoyenebas kon
kretuli amocanidan gamomdinare.

31
sityva “inducirebulis” sinonimebia – “xelovnuri” da “eqsperimentuli.”
32
mutagenuri faqtorebi anu mutagenebi – aris mutaciis gamomwvevi qimiuri
da fizikuri faqtorebi.
66
 inducirebuli mutantebis ujredul seleqciaSi gamo-
yeneba dawyebul iqna arc ise didi xnis winaT. 1959 wels g.
melxersma da l. bergmanma pirvelad gamoaqveynes cnobebi
eqstremaluri temperaturebisadmi gamZle mutanturi ujre-
duli kulturebis seleqciis Sesaxeb.
ujredul seleqciaSi aRniSnuli xerxis efeqturobis
damajerebel dadasturebas warmoadgens 1973-1975 wlebSi p.
maligasa da misi TanamSromlebis klasikuri Sromebi kalu-
suri qsovilis kulturis gziT Tambaqos streptomicingam-
Zle mcenareebis miRebaSi.
ujredebis doneze inducirebuli mutantebis warma-te-
biT miRebisaTvis aucilebelia:
1. inducirebuli mutagenezisaTvis obieqtis SerCeva;
2. obieqtis damuSaveba mutagenuri faqtorebiT, anu mu-
tagenebiT;
3. seleqtiuri sistemebis (seleqtiuri areebis) SemuSa-
veba;
4. damuSavebuli obieqtebis kultivireba seleqtiur
sistemaze (seleqtiur areze) da inducirebuli mutantebis
gamorCeva;
5. cvalebadobis genetikuri bunebis dasabuTeba.
bunebriv pirobebSi mutaciuri cvalebadobis sixSire
mcenaris ujredebSi Zalian dabalia da Seadgens 10-5 -10-8. mi-
si gadidebisaTvis xelovnur pirobebSi, e.i. eqsperimentebSi
iyeneben inducirebul mutagenezs – awarmoeben obieqtis
ujredebis damuSavebas sxvadasxva mutagenuri faqtorebiT
anu mutagenebiT.
farTodaa gavrcelebuli universaluri fizikuri da
qimiuri mutagenuri faqtorebis gamoyeneba, romlebsac gaaC-
niaT ujredis genomze zemoqmedebis farTo speqtri.
universalur fizikur faqtorebs miekuTvneba ultrai-
isferi sxivebi, maionizirebeli gamosxiveba (rentgenis sxi-
vebi, γ-sxivebi), swrafi eleqtronebis, pozitronebis, proto-
nebis, α-nawilakebis, neitronebis tipis korpuskularuli
gamosxiveba, agreTve ultrabgera.

67
 aRniSnuli mutagenuri faqtorebi an uSualod aziane-
ben dnm-s molekulas, an iwveven destruqciul procesebs
dnm-is molekulaSi, romelic warmoiqmneba am dros warmo-
Sobili Tavisufali radikalebis zemoqmedebiT. mutaciebis
yvelaze farTo speqtri SeimCneva mutagenis saxiT maionizi-
rebeli gamosxivebis gamoyenebisas.
qimiur faqtorebs miekuTvneba sxvadasxva qimiuri or-
ganuli da araorganuli SenaerTebi: nukleinis mJavebis Se-
dgenilobaSi Semavali azotovani fuZeebis inhibitorebi
(kofeini, eTiluretani, Teobromini, azaguanini, azaserini
da sxva), nukleinis mJavebSi CarTuli azotovani fuZeebis
analogebi (5-bromuracili, 5-qloruracili, 2-aminopurini
da sxva), alkilirebadi SenaerTebi (dimeTilsulfati-dms,
dieTilsulfati-des, nitrozoeTilSardovana-neS, nitrozo-
meTilSardovana-nmS, ipriti, fenoli da sxva), mJangavebi,
aRmdgenelebi da Tavisufali radikalebi (azotismJava, ze-
Jangebi, aldehidebi, mZime liTonebis marilebi, Jangbadi da
sxva) da akridinuli saRebavebi.
es nivTierebebi Trgunaven calkeuli nukleotidebis
sinTezs, wyveten dnm-s Zafs, iwveven nukleinis mJavebis de-
zaminirebas da a.S., ris Sedegadac ujredSi warmoiSoba mu-
taciuri cvalebadoba.
genetikurad aqtiuri fizikuri faqtorebisagan gansxva-
vebiT, qimiur mutagenebs gaaCniaT ufro maRali toqsikuri
moqmedeba33 . letalobis maRalma procentma, qimiuri mutage-
nebis gamoyenebisas, SeiZleba gamoiwvios ujredebis mTeli
populaciis daRupva, amitom maTTan muSaobisas iyeneben da-
bal dozebs.
maionizirebeli gamosxiveba naklebad toqsikuria. ga-
mosxivebis maRali dozebi iwvevs ujredebis mxolod rep-
roduqtiuli da regeneraciuli unarebis daqveiTebas, maTi
metabolituri aqtiurobis SenarCunebiT.

33
toqsikuri Tvisebebi gaaCnia rogorc qimiur, ise fizikur mutagenebsac,
magram qimiuri mutagenebis toqsikuroba Seudareblad maRalia.
68
 inducirebuli mutaciis maqsimaluri sixSire miiRweva
mutagenuri faqtoris optimaluri dozis dros.
fizikuri mutagenis zemoqmedebis dozebi gamoisaxeba
radiaciis tipis Sesabamisi gamosxivebis erTeulebSi (ren-
tgeni, radi da a.S.), xolo qimiuri mutagenis zemoqmedebis
doza – garkveul temperaturaze mutagenis koncentraciasa
da eqspoziciaSi.
mutagenis dozis SerCevisas orientirebas akeTeben da-
samuSavebeli obieqtis ujredebis gadarCenis unarianobaze,
romelic damokidebulia rogorc mutagenis dozaze34, ise
mocemuli ujredis mutagenis letaluri efeqtisadmi
35
mgrZnobiarobaze . rogorc wesi, seleqciur muSaobaSi iye-
neben iseT dozebs, romlebic uzrunvelyofen ujredebis
gadarCenas 0.1-dan 50-80%-mde diapazonSi.
mutagenur faqtorebTan muSaobisas aucilebelia gar-
kveuli pirobebis dacva: zemoqmedebis qveS myofi biologi-
uri masala unda iyos diskretuli, anu Semdgari calkeuli
nawilebisagan (ujredebisagan); ujreduli suspenzia SeZle-
bisdagvarad unda iyos moklebuli koStebs-konglomera-
tebs, radganac konglomeratis nebismieri ujredis induci-
rebuli mutacia SeiZleba mTlianad daikargos misi sakveb
areSi kultivirebisas; qimiuri mutagenebiT damuSaveba unda
Catardes buferul xsnarebSi, mocemuli mutagenisaTvis pH-
is yvelaze optimalur pirobebSi; qimiuri mutagenebiT da-
muSavebis Semdeg kultivirebamde biologiuri masala unda
gairecxos steriluri wyliT an axali sakvebi ariT.
inducirebuli mutantebis gamoyofisaTvis iyeneben se-
leqtiur sistemebs (seleqtiur areebs). maTi damuSaveba sa-

34
arsebobs ukudamokidebuleba ujredebis gadarCenis unarianobasa da mu-
tagenis dozebs Soris. amitom didi dozebi SeiZleba gamoyenebul iqnas
mxolod iSviaTi mutantebis misaRebad.
35 ujredebis gadarCenis unarianoba damokidebulia ujreduli kulture-
bis ganviTarebis stadiaze (mag. protoplastebis radiomgrZnobeloba
izrdeba kultivirebis dawyebidan da maqsimums aRwevs ujreduli ciklis
G2 periodSi), ujredebis ploidurobaze (haploidebi ufro mgrZnobiarea
mutagenebis mimarT, vidre diplodebi).
69
Sualebas iZleva Seiqmnas iseTi xelovnuri pirobebi, ro-
melSic vlindeba ara saerTod Secvlili mutantebi, aramed
mutantebi, romlebsac gaaCnia garkveuli mimarTulebis cva-
lebadobebi.
dReisaTvis damuSavebulia seleqtiuri sistemebi, rom-
lebic saSualebas iZleva gamovlindes antibiotikebisadmi,
herbicidebisadmi, aminomJavebisa da maTi analogebisadmi,
nukleinis mJavebis sinTezis inhibitorebisadmi, stresuli
faqtorebisadmi da sxva gamZle mutantebi.
obieqtebis mutagenuri faqtorebiT damuSavebis Semdeg,
mimarTaven maT kultivirebas seleqtiur sistemaze (seleq-
tiur areze) da awarmoeben inducirebuli mutantebis ga-
morCevas.
biologiuri obieqtebis kultivirebas seleqtiur sis-
temaze (seleqtiur areze) axdenen Semdegi meTodikiT: 1)
obieqtidan calkeuli ujredebis gamoyofa da maTi gamoz-
rda mikrokameraSi an mikrowveTSi, 2) obieqtis yvela ujre-
dis masobrivi Tesva sakveb areSi.
inducirebuli mutantebis gamorCevisaTvis arsebobs
Semdegi xerxebi:
1. pirdapiri (pozitiuri) gamorCeva, romlis drosac se-
leqtiur sistemaSi gadarCeba ujredebis mxolod garkveu-
li saZebni mutanturi tipi. pirdapir gamorCevas iyeneben ma-
rilebisadmi, herbicidebisadmi, antibiotikebisa da maTi
analogebisadmi, sokoebis toqsinebisadmi da sxva gamZle
mutantebis misaRebad. mutantebis pirdapiri gamorCevis
dros jer xdeba obieqtis miReba da damuSaveba mutageniT,
Semdeg damuSavebuli obieqtis kultivireba seleqtiur are-
ze saZebni mutanturi ujredebis gamosarCevad da bolos,
mutanturi ujredebis kultivireba saregeneracio sakveb
areSi.
2. arapirdapiri (negatiuri) gamorCeva, romlis drosac
iqmneba piroba, sadac SerCeviT iRupeba veluri tipis dayo-
fis fazaSi myofi ujredebi da gadarCeba metabolurad
araaqtiuri mutanturi ujredebi. es xerxi efeqturia piro-

70
biTad letaluri mutantebis – auqsotrofuli 36 da tempera-
turagamZle mutantebis gamorCevisaTvis. veluri tipis uj-
redebis SerCeviTi daRupvisaTvis yvelaze metad iyeneben
nukleinis mJavebis analogebs (5-bromdezoqsiuredani, 5-
ftordezoqsiuredani), aminomJavebis analogebs (ftorfeni-
lalanini) da iseT toqsikur nivTierebebs rogoricaa nat-
riumis arsenati. gadarCenili mutanturi ujredebis kulti-
virebas Semdeg axdenen aminomJavebiT, vitaminebiT, eqstraq-
tebiT da sxv. gamdidrebul sakveb areeebze, sadac isini
mravldebian.
mutanturi ujredebis gamorCeva jer kidev ar warmo-
adgens inducirebuli mutantebis miRebis samuSaos da-
sasruls. aucilebelia kidev somaturi ujredebis doneze
miRebuli cvalebadobis genetikuri bunebis damatebiTi da-
sabuTeba. vinaidan gamorCeul ujredebs Soris SeiZleba
iyos adaptirebuli ujredebi, romelTa zrda seleqtiur
pirobebSi dakavSirebulia ara mutaciasTan, aramed mocemu-
li genotipis reaqciis normis farglebSi populaciaSi uf-
ro gamZle ujredebis gamorCevasTan. aseve SeiZleba iyos
Secvlili ujredebi, romelTa warmoqmna dakavSirebulia
epigenetikur cvalebadobasTan. somaturi ujredebis doneze
cvalebadobis dasadgenad arsebobs rigi kriteriumi: 1. Sec-
vlili ujredebis sixSire unda iyos Zalian dabali (1.10-6-
10-7); 2. is Zalian izrdeba mutagenebis gamoyenebisas (1.10-4-10-
5 ); 3. Secvlil ujredebs aqvT stabiluri xangrZlivi zrdis

unari; 4. Secvlili niSnis stabiluroba narCundeba seleq-

tiuri pirobebis mocilebis Semdegac; 5. vlindeba Secvli-
li genis produqti. Tumca TiToeuli maTgani calke ar
warmoadgens mutaciis damamtkicebels. amitom, radganac
mcenareul ujredebs in vitro pirobebSi gaaCniaT totopoten-
tobis, anu garkveul pirobebSi mcenaris regeneraciis una-
ri, somaturi ujredebis doneze miRebuli cvalebadobis ge-

36
auqsotrofuli mutantebi – mutantebi, romlebic saWiroeben sakvebi aris
damatebiT komponentebs maTi sinTezis unaris dakargvis gamo.
71
netikuri bunebis saboloo dadgena yvelaze efeqturad Se-
saZlebelia mcenare-regenerantebis genetikuri analiziT.
inducirebuli mutagenezis efeqturobis SefasebisaT-
vis xSirad iyeneben mutaciis sasargeblo sixSires, romel-
sac sazRvraven miRebuli mutantebis raodenobis Sefarde-
biT kultivirebuli ujredebis saerTo raodenobasTan.
saerTod inducirebuli mutagenezis upiratesoba se-
leqciis tradiciul meTodebTn SedarebiT, mdgomareobs se-
leqciisaTvis sawyisi masalis ufro swraf gaumjobesebaSi,
rogorc erTi, ise rigi sameurneod Zvirfasi niSnis mixed-
viT.
mcenaris doneze eqsperimentul mutaciasTan Sedare-
biT, in vitro mutacias gaaCnia rigi upiratesobebi: erTi pet-
ris jamSi kultivirebuli ujredebis imdenive raodenobaa,
rac aTas heqtarze darguli mcenareebisa; mutanturi niSne-
bi calkeuli ujredebis doneze ufro swrafad vlindebian;
SesaZlebelia axali tipis mutaciebis miReba, maT Soris bi-
oqimiuri xasiaTis; axali sasurveli niSnis miRebaze gaci-
lebiT naklebi dro da SromiTi resursebi ixarjeba.
U ujreduli mutagenezis warmatebiT gamoyenebis erT-erT
aucilebel pirobas warmoadgens mcenareTa regeneraciis
kargad damuSavebuli sistemis arseboba. aseve Zalian mniS-
vnelovnia kvlevis obieqtad aRebul mcenareSi haploidebis
miRebis SesaZlebloba, vinaidan ujredebis kulturaSi mu-
tagenis efeqturoba izrdeba haploidebis doneze, yvela re-
cesiuli mutaciis gamovlenis gamo. Semdgom seleqciur mu-
SaobaSi gamoiyeneba mxolod is genotipebi, romlebSic mu-
taciebi vlindeba mTliani mcenaris doneze.

1.3.4. gamorCevis meTodebi ujredul seleqciaSi

ujreduli seleqciis CatarebisaTvis iyeneben gamorCe-
vis Semdeg meTodebs:
1. pi rdapiri (pozitiuri) gamorCeva. am dros gadarCeba
mxolod garkveuli tipis Secvlili ujredebi. amisaTvis
sakveb ares umateben calkeul komponentebs, romelTa mi-
72
marT Cveulebriv, ucvlel ujredebs gamZleobis unari ara
aqvT da iRupebian.
2. arapirdapiri (n egatiuri) gamorCeva. igi mdgomareobs
iseTi pirobebis SeqmnaSi (magaliTad, kultivireba dabali
an maRali temperaturis pirobebSi sakveb areze calkeuli
komponentebis gareSe da a.S.), romlis drosac ucvleli (ve-
luri tipis) ujredebis zrda an Cerdeba, an es ujredebi
iRupeba. igi dafuZnebulia dayofis fazaSi myofi veluri
tipis ujredebis SerCeviT daRupvasa da metabolurad ara-
aqtiuri ujredebis gadarCenaze, romlebic saWiroeben maT-
Si cvalebadobis arsebobis damatebiT identifikacias.
3. totaluri gamorCeva. am dros adgili aqvs yvela
ujredis individualur testirebas.
4. vi zualuri da araseleqtiuri gamorCeva. ujredebi s
mTliani populaciidan Secvlili ujredebis identifici-
reba xdeba vizualurad an bioqimiuri meTodebiT (Txel-
Sriani an Txevadi qromatografia, mikrospeqtrofotomet-
ria, radioimunuri analizi da sxva).
yvelaze ufro gavrcelebul meTods warmoadgens pir-
dapiri gamorCeva da ZiriTadad gamoiyeneba sxvadasxva meta-
bolitebis, magaliTad, herbicidebis, antibiotikebis, toqsi-
nebis da sxva mimarT gamZle ujredebis gamosarCevad. aseTi
ujredebis gamosarCevad yovel konkretul SemTxvevaSi, sa-
wyisi masalidan da genotipidan gamomdinare, aucilebelia
sakveb areSi toqsikuri nivTierebebis koncentraciis dadge-
na.
umetes SemTxvevaSi toqsikuri nivTierebebis Semcve-
loba ar unda iyos imaze gacilebiT maRali, romlis dro-
sac mTlianad inhibirdeba qsovilebis zrda. zogierT Sem-
TxvevaSi (magaliTad, citoplazmaturi cvalebadobis, bir-
Tvuli genetikuri cvalebadobis dros) gamarTlebulia sa-
fexuriseburi gamorCeva, roca sakveb areSi toqsikuri niv-
Tierebebis koncentracias zrdian TandaTanobiT. meTodi
warmatebiT iqna gamoyenebuli marilebis, paTotoqsinebis,
aminopternis mimarT gamZle ujreduli xazebis gamorCevi-
sas.
73
 magaliTad, mutantebis pirdapiri gamorCevis zogadi
sqema moicavs Semdeg procedurebs: 1) protoplastebis gamo-
yofa da maTi damuSaveba mutagenebiT; 2) ujredebis kulti-
vireba araseleqtiur pirobebSi 3-5 ujredul gayofamde; 3)
ujredebis kultivireba seleqtiur areebze da ujreduli
variantebis gamoyofa; 4) ujreduli koloniebis kultivire-
ba regeneraciul sakveb areebze (SesaZebelia Catardes uj-
reduli variantebis ganmeorebiTi Semowmeba gamZleobaze);
5) mcenareTa regeneracia da maTi gamravleba (gamZleobaze
testebis, bioqimiuri analizebis Catareba); 6) mcenaris ga-
dargva niadagSi (Teslebis testireba gamZleobaze, gamZle-
obis genetikuri bunebis Seswavla).
arapirdapiri gamorCevis meTodi, romelic efeqturia
pirobiTad letaluri (auqsotrofi 37, temperaturamgrZnobia-
re) mikroorganizmebisa da cxoveluri ujredebisaTvis,
sul ufro farTo gamoyenebas poulobs mcenareul ujre-
debSi.
yvelaze TvalsaCino Sedegebia miRebuli, roca sawyis
masalad warmodgenili iyo protoplastebi.
warmatebiT iyeneben agreTve suspenziur kulturebs.
magaliTad, temperaturamgrZnobiare ujredebis SerCeviTi
daRupvisaTvis (am mizniT ujredebis kultivireba xdeba -320
C temperaturaze) toqsikuri nivTierebebis saxiT sakveb are-
ze ujredebs umateben nukleinis fuZeebis analogebs (5-
bromdezoqsiuridini da 5-ftorfenilalanini) an natriumis
arsenats. Semdeg gadarCenili, potenciurad mutanturi uj-
redebidan, saWiro ujreduli xazebis misaRebad, isini gada-
aqvT kultivirebis sxva pirobebSi (sakvebs amdidreben ami-
nomJavebiT, fuZeebiT, vitaminebiT an kazeinis hidroliza-
tiT, safuarebis eqstraqtiT, kultivirebis temperatura
aris -250C).

37auqsotrofi (berZ. Auxo - vzrdi, trophe - sakvebi) – organizmi, romelsac

ara aqvs zrdisaTvis aucilebeli garkveuli organuli naerTebis sinTezis
unari.
74
 totaluri gamorCevis, anu ujredebis individualuri
testirebis meTodi damuSavebul iqna g. bidlisa da e. ta-
tumis mier 1945 wels Neurospora-Si kvebis moTxovnilebis mi-
xedviT defeqturi mutantebis gamosarCevad. amisaTvis isini
sokos konidiumebs amuSavebdnen mutagenebiT ganayofiere-
bamde, Semdeg warmoqmnili sporebi gadahqondaT gamdidre-
bul sakveb areze. bolos micelarul klonebs individua-
lurad Tesavdnen testareebze defeqturobis gamosavlenad.
meTodis ZiriTad naklovanebas warmoadgens misi maRa-
li Sromatevadoba. swrafi testirebisaTvis (magaliTad, au-
qsotrofebis gamosarCevad) SeiZleba gamoyenebul iqnas rep-
likebis meTodi. amisaTvis ujredebis suspenzia gadaaqvT
agarizebuli sakvebi aris zedapirze da afareben neilonis
bades. zrdis garkveuli periodis Semdeg, neilonis bade
masze miwebebuli nawilakebiT, gadaaqvT sxva petris jamSi
“anabeWdebis” misaRebad. kopireba 80%-mde aRwevs.
vizualuri gamorCeva dafuZnebulia imaze, rom xSirad
in vitro pirobebSi kultivirebadi ujredebi da ujreduli
agregatebi erTmaneTisagan gansxvavdebian garegani niSnebiT.
es meTodi gamoyenebulia iseTi ujreduli xazebis gamosar-
Cevad, romlebic producireben iseT pigmentebs, rogori-caa
qlorofili, karotinoidebi da anTociani. meTodi efeqtu-
ria agreTve qlorofildefeqturi mutantebis identifika-
ciisaTvis.
fotosinTezis mixedviT defeqturi ujredebis gamor-
Cevisas koloniebis kultivireba xdeba regeneraciuli pro-
cesebis da fotosinTezuri aqtiurobis mainducirebel sak-
veb areebze. Secvlili ujredebis gamorCeva SeiZleba Sefe-
rilobis mixedviT. magaliTad, ujredebisa da qsovilebis
Seferilobis mixedviT gamorCeul iqna streptomicinis,
linkomicinis, calkeuli herbicidebis mimarT gamZle mutan-
tebi. amisaTvis gamoyenebul iqna am nivTierebebis iseTi
koncentraciebi, romlebic ar Trgunavda maT zrdas, magram
iwvevda qsovilebis gamwvanebas. gamZle mutantebs hqonda
mwvane feri.

75
 Secvlili ujredebis araseleqtiuri gamorCevisas iye-
neben sxvadasxva bioqimiur meTodebs. individualuri kolo-
niebis bioqimiuri analizi, cilebis an izofermentebis
speqtris mixedviT, Secvlili variantebis gamorCevis saSua-
lebas iZleva. sxva xerxebiT (qromatografia, mikro-peqro-
fotometria, radioimunografia da sxva) SesaZlebelia spe-
cifikuri nivTierebebis, magaliTad, alkalodebis da sxva
meoradi metabolitebis maproducirebeli xazebis gamorCe-
va.

1.3.5. ujreduli seleqciis upiratesobebi,

SemzRudveli pirobebi da faqtorebi

garkveuli siZneleebis miuxedavad, ujredul seleqci-

is meTodebs gaaCnia rigi upratesobebi tradiciuli seleq-
ciis meTodebTan SedarebiT:
1. muSaobaSi sezonurobis ararseboba;
2. gamorCevaSi milionobiT ujredebis gamoyenebis Se-
saZlebloba;
3. mizanmimarTli seleqciis warmarTva seleqtiuri
areebis gamoyenebiT;
4. samuSaoebis Catareba laboratoriul pirobebSi;
5. eqsperimentis procesSi kontrolirebadi pirobebis
Seqmna da saWiroebisas misi kopireba;
6. farTobebis ekonomia – 10 sm diametris petris jamze
SeiZleba 107–108 ujredis kultivireba, igive raodenobis
mcenareebisaTvis ki saWiroa aTas heqtarze meti farTobi;
7. Secvlili niSnebi calkeuli ujredebis doneze
vlindeba SedarebiT swrafad;
8. SesaZlebelia axali tipis cvlilebebis miReba, maT
Soris bioqimiuri xasiaTisac;
9. sasurveli niSnis misaRebad droisa da SromiTi da-
naxarjebis ekonomia.
ujreduli seleqciis warmatebiT gamoyenebisaTvis au-
cilebelia Semdegi pirobebis dakmayofileba:
76
 1. unda arsebobdes mcenareTa regeneraciis kargad da-
muSavebuli sistema;
2. niSani, romlis mixedviTac tardeba gamorCeva, kar-
gad unda vlindebodes ujredul doneze;
3. sakvlevi mcenaridan SesaZlebeli unda iyos haplo-
idebis miReba, vinaidan Semdgom seleqciur muSaobaSi erT-
veba mxolod is genotipebi, romlebSic Secvlili niSnebi
vlindeba mTliani mcenaris doneze.
arsebobs ujreduli seleqciis SemzRudveli rigi
faqtori:
1. mravali sameurneo Zvirfasi niSnis, iseTebis rogo-
ricaa mosavlianoba, marcvlebis raodenoba, pesticidebi-
sadmi gamZleoba da sxva, in vitro kultivirebis pirobebSi mi-
Reba Zneli an praqtikulad SeuZlebelia, radganac isini
ar vlindebian ujredul doneze. arasakmarisadaa agreTve
mTliani mcenaris doneze am niSnebTan korelaciaSi myofi
bioqimiuri da molekuluri markerebi.
2. ujredul doneze seleqtirebadi yvela niSani ar
narCundeba mcenare-regenerantis doneze. mas ramodenime mi-
zezi aqvs: cvalebadobis garkveuli nawili ar exeba ujre-
dis dnm-s (genetikur aparats), amitom ar inaxeba TaobebSi;
genetikurma cvalebadobebma SeiZleba ganicados eliminacia
diferenciaciisa da meiozis procesSi; Secvlili genis
funqcia SeiZleba gamovlindes mxolod kultivirebadi uj-
redebis doneze; erTi genis cvlilebas SeiZleba Tan ax-
ldes izofermentebis makodirebeli sxvadasxva genis aqti-
vacia; genotipis nawils ara gaaCnia normaluri fertiluri
mcenaris regeneraciis unari.

1.3.6.. ujreduli seleqciis Sedegebi da

perspeqtivebi
1.3.6.1. somaklonuri cvalebadoba

arsebobs monacemebi, romlis mixedviTac somaklonur

variantebs SeuZlia 2-jer daaCqaros axali jiSebis miRebis
77
procesi. amJamad somaklonuri variantebi miRebulia brin-
jSi, xorbalSi, simindSi, kartofilSi, TambaqoSi, xanWko-
laSi, selSi da sxva kulturaSi.
pirvelad ujreduli kulturebidan somaklonebi mi-
Rebul iqna Saqris lerwmSi XX saukunis 70-ian wlebSi. mi-
Rebuli mcenareebi gansxvavdebodnen morfologiuri, cito-
genetikuri da izofermentuli niSnebiT. amave kulturaSi
identificirebulia somaklonuri variantebi, romlebic sa-
wyisi formebisagan gamoirCevian daavadebebisadmi maRali
gamZleobiT da mosavlianobis arsebiTi zrdiT.
somaklonuri cvalebadobis meTodi warmatebiT gamo-
iyeneba sakveb, teqnikur, dekoraciul da samkurnalo mcena-
reebSi. aRniSnuli meTodis gamoyenebam dekoraciul mcena-
reebSi SesaZlebeli gaxada miRebul iyo pelargonias axa-
li jiSi.
Kkartofilis erTi-erTi jiSidan miRebulia Zvirfasi
somaklonebi, romlebic gamoirCevian daavadebebisadmi gam-
ZleobiT, tuberebSi proteinisa da saxameblis maRali Sem-
cvelobiT. meore jiSidan gamoyofilia xazebi, romlebic
sawyis jiSs aRematebian mosavlianobis, fotoftorisa da
virusebisadmi gamZleobiT.
1986 wels kartofilis jiS Feltwell-is 235 somakloni ga-
mocdil iqna savele pirobebSi. maTgan prototipze ufro
mosavliani iyo 92, yvela maTgans gaaCnda nematodebis erT-
erTi formisadmi gamZleobis geni, zogierTi maTgani iyo
aseve rezistentuli Y-virusisadmi.
somaklonebis miReba metad mniSvnelovania marcvlovne-
bis seleqciaSi, sadac tradiciuli meTodebiT iqmneba me-
tad viwro memkvidruli cvalebadobebi.
mkvlevarTa sxvadasxva jgufis mier, saSemodgomo da
sagazafxulo xorblis jiSebis kalusidan miRebuli somak-
lonebis analizma aCvena didi cvalebadoba somaklonebs
Soris da aseve maTi gansxvaveba sawyis jiSebTan iseTi mor-
fologiuri niSnebis mixedviT, rogoricaa mcenaris simaR-
le, bartyobis sixSire, ylortebis raodenoba, TavTavis Se-

78
fardeba namjasTan, TavTavis fxianoba da ufxooba, TavTa-
vis forma, Teslebis Seferiloba.
fiziologiuri niSnebis mixedviT mcenareebi gansxvav-
debodnen vegetaciis droiT, dabali temperaturebisa da ga-
dazamTrebis pirobebisadmi gamZleobiT, daavadebebisadmi
gamZleobiT.
somaklonuri cvalebadobis gamoyenebiT miRebulia
pomidori mSrali nivTierebis maRali SemcvelobiT, pomi-
dori da brinji mamrobiTi sterilobiT, maRalmosavliani
brinji, adremwifadi simindi, dabali temperaturisa da daa-
vadebebisadmi gamZle xorbali, Saqris lerwami, kartofili.
miRebulia fuzariozisadmi gamZle kartofilis so-
makloni. gamorCeulia marilebis maRali Semcvelobisadmi
rezistentuli xanWkolas, Tambaqos da kartofilis mcena-
reebi da sxva.
amrigad, somaklonebis praqtikuli mniSvnelobis aR-
niSnuli da sxva mravali dadebiTi magaliTi, seleqciuri
muSaobis praqtikaSi somaklonebis miRebis sxvadasxva xer-
xis ufro efeqturi danergvis aucileblobas adasturebs.
yvelaze realuria somaklonuri cvalebadobis gamoyeneba
ukve arsebuli jiSebisa da xazebis calkeuli niSnebis mi-
xedviT gaumjobesebis saqmeSi.
aRniSnulTan erTad genetikurad Secvlili mcenareTa
formebis miRebis warmodgenil meTods gaaCnia arsebiTi
naklovanebebi. erT-erTi mTavaria – saWiro variantebis mi-
zanmimarTuli seleqciis meTodebis ararseboba. ar aris ga-
moricxuli, rom rig SemTxvevebSi, spontanuri somaklonuri
variantebis miRebis procesi, maTi identifikacia da Sefase-
ba mindvris pirobebSi, sabolood damTavrdes uaryofiTi
SedegiT.
amitom aucilebelia cvalebadobis regulaciis Sem-
dgomi Seswavla, mcenareTa Sesabamisi saxeobebis, genotipe-
bisa da sawyisi eqsplantatis SerCeva, aseve somaklonuri
cvalebadobis speqtris gazrdis pirobebis gansazRvra da
sxva.

79
 1.3.6.2 somaturi hibridizacia

somaturma hibridizaciam arsebiTad gazarda tradici-

uli kvlevebis SesaZleblobebi Soreul hibridizaciaSi.
igi saSualebas iZleva gadawyvetil iqnac mcenareTa seleq-
ciis mravali fundamenturi da praqtikuli amocana.
pirvelad somaturi hibridizaciis gziT somaturi hib-
ridi miRebul iqna 1972 wels karlsonis mier, Tambaqos ori
saxeobis (Nicotiana glauca, 2n =24 + N. langsdorfii, 2n=18) protop-
lastebis Serwymis Sedegad. amis Semdeg dRemde miRebulia
mravalricxovani cxovelmyofeli saxeobisSigniTa, saxeoba-
TaSorisi da gvareobaTaSorisi hibridi.
ganxorcielebul iqna kulturuli kartofilis (Sola-
num tuberosum) erT-erTi jiSis, romelic gamoirCeva msxvili
tuberebiT, magram daavadebebisadmi mgrZnobiarobiT da ve-
luri kartofilis (S. chacoense), romelsac Zalian patara
tuberebi, magram mravali daavadebisadmi gamZlea, protop-
lastebis Serwyma. miRebul saxeobaTaSoris somatur hibri-
debs foTlebisa da buCqis formis, aseve tuberebis zomis
mixedviT hqondaT Sualeduri monacemebi mSoblebTan Seda-
rebiT, xolo gamZleobis mxriv avlendnen maRal imunuro-
bas У virusisadmi, riTac Zlier gansxvavdebodnen am par-
tniorebis sqesobrivi hibridebisagan.
gvareobaTaSorisi somaturi hibridebis Seqmnis pirve-
li mcdeloba ekuTvnis g. melxerss, romelmac 1978 wels mi-
iRo aseTi hibridi kartofilsa da pomidors Soris. Mcena-
re iyo steriluri, morfologiurad anomaluri: msxvili
fesvebi, tipiuri stolonebis uqonloba, defeqturi yvavi-
lebi.
iaponelma mkvlevarebma x. kisaka da sxva, 1997 wels qe-
risa da brinjis protoplastebis eleqtroSerwymiT miiRes
gvareobaTaSorisi somaturi hibridi. brinjis protoplas-
tebi miiRes suspenziuri kulturidan, xolo qeris – axal-
gazrda foTlebidan. hibridi morfologiiT axlos iyo
brinjis mcenaresTan, citologiurad mcenares hqonda brin-

80
jis patara qromosomebi da qeris didi qromosomebi. hib-
rids aseve hqonda mitoqondrialuri da qloroplasturi
dnm-s mSoblebisagan gansxvavebuli axali Tanmimdevroba.
1978 wels i. glebam TanamSromlebTan erTad miiRo
pirveli tribaTaSorisi 38 hibridi – arabidobrasika, arabi-
dopsisis (Arabidopsis thaliana) in vitro kulturis ujredis da
turnefsis (Brassica campestris)FfoTlis mezofiluri ujredis
protoplastebis Serwymis gziT.
es warmoadgenda pirvel kvlevas, romelmac srulad
da damajereblad aCvena somaturi hibridizaciis SesaZleb-
lobebi (da SezRudvebi) filogenezurad Soreuli saxeobe-
bis Sejvarebis dros. mcenare iyo genetikurad da morfo-
logiurad erTgvarovani, ar yvavilobda, maxinji formis,
Zalian bevri yvavilisnairi teratomamsgavsi 39 warmonaqmne-
biT.
1979 wels ganxorcielebul iqna tribaTaSorisi soma-
turi hibridizacia ZaRlyurZenasebrTa ojaxis ori gvaris
saxeobebs lemas da beladonas Soris (Datura innoxia + Atropa
belladonna). yvela regenerirebul mcenareSi gamovlenil iqna
orive mSoblis qromosomebi. amfidiploidebs ar aRmoaC-
ndaT Reroseuli morfogenezis unari. poliploiduri da
aneuploiduri qromosomebis mqone xazebidan Rebulobdnen
anomalur Reroebs. regenerirebuli mcenareebi iyo steri-
luri, lemas msgavsi, magram Seicavdnen beladonas mcire
raodenobis qromosomebs.
beladonas protoplastebis Cinuri Tambaqos kalusis
ujredebis protoplastebTan SerwymiT miRres hibridebi,
romelTa ujredebSi napovni iqna orive mSoblis qromoso-
mebi, Tumca erTi wlis Semdeg ramdenime mcenareSi adgili
hqonda beladonas qromosomebis srul eliminacias.
pirveli tribaTaSorisi fertiluri hibridebi miRe-
bul iqna qlorofildefeqturi Tambaqos protoplastebis

38 triba – taqsonomiuri erTeuli gvarsa da ojaxs Soris.

39 teratoma – simsivne.
81
beladonas protoplastebTan Serwymis Sedegad. sul miRe-
bul iqna 40 fotosinTezisunariani ujreduli kolonia,
maTgan 4 iZleoda beladonas normalur mcenareebs, 4 –
morfologiurad anomalur hibridebs, danarCeni – mwvane,
zogjer WrelfoTola, Tambaqos identur mcenareebs, rom-
lebic yvavilobda da iZleoda Teslebs.
PojaxTaSorisi somaturi hibridebis miRebis pirveli
kvlevebi Catarebul iqna 1976-77 wlebSi k. kaosa da v. vete-
ris mier soiosa da Tambaqos (soio + Tambaqo) Soris. mog-
vianebiT sxva mecnierebis mier Catarebul iqna analogiuri
eqsperimentebi ZaRlyurZenasebrTa, parkosanTa da SroSani-
sebrTa ojaxebis warmomadgenelTa monawileobiT (barda +
Tambaqo da xaxvi + Tambaqo). aseve mecnierebma SeZles Re-
rosmagvari teratomebidan morfogenezis induqcia Nicotinia
tabacum + Vicia faba-s ojaxTaSorisi hibridebis kulturaSi.
P praqtikulad yvela SemTxvevaSi adgili hqonda erT-erTi
mSoblis qromosomebis saxeobaspecifikur eliminacias. ase-
Ti hibridebis kulturaSi iyo bevri mravalbirTviani uj-
redi, ujredebi mini birTviT, giganturi qromosomebiT.
morfogenezi aseT masalaSi SeCneuli ar yofila.
“cxoveli+mcenare” ujredebis samefoTaSorisi hibri-
debis Sewavlam aCvena, rom Serwymis etapze saxeobaspecifi-
kuroba ar vlindeba da SeiZleba cxovelisa da mcenaris
ujredebis Serwyma. magram onTogenezis ufro gvian etapeb-
ze es gansxvaveba Tavs iCens, rac dadgenil iqna arabidop-
sisisa da Tambaqos protoplastebis adamianis limfoci-teb-
Tan Serwymis eqsperimentebis dros. am dros adgili hqonda
citoplazmis Serwymas, magram birTvebis Serwyma ar xdebo-
da.
A amrigad, somaturi hibridizacia Seujvareblobis bari-
erebis gadalaxvis da Sedegad, cocxali organizmebis sxva-
dasxva gvaris, tribis, ojaxis, klasis da TviT samefos sa-
xeobebis genetikuri masalis gerTianebis saSualebas iZle-
va.
Seujvareblobis barieri ar arsebobs protoplastebis
Serwymis etapze, Semdeg etapebze kombinaciebis filogene-
82
zurad dacilebis Sesabamisad Cndeba sxvadasxva citogene-
tikuri barierebi: erT-erTi mSoblis qromosomebis an
mTliani genomis eliminacia, qromosomis fragmentebis war-
moqmna, mravalricxovani translokaciebi da sxva gardaq-
mnebi; klasTaSoriso da ojaxTaSorisi hibridebi arseboben
ujreduli koloniebis doneze, mcenaris regeneraciis una-
ris gareSe; trobaTaSorisi hibridebi nawilobriv an mTli-
anad arafunqcionirebadi, steriluri regeneranti-mcenaree-
bia.
P praqtikuli mniSvnelobis somaturi hibridebi miiReba
mxolod saxeobaTaSorisi da gvareobaTaSorisi somaturi
hibridizaciis dros, romlis zogierTi magaliTi motani-
lia cxrilSi 1.
Y yvelaze mniSvnelovani warmatebebia miRweuli saxeobaTa-
Soris hibridizaciaSi, gansakruTrebiT ZaRlyurZenaseb-rTa
(Solanaceae) da kombostosnairTa (Brassicaceae) ოჯaxis warmomad-
genlebSi.
bolo periodSi kargi Sedegebia aseve mopovebuli par-
kosanTa da marcvlovanTa ojaxis saxeobebSic.
arsebobs agreTve warmatebuli Sedegebi merqniani xexi-
lovani mcenareebis, maT Soris citrusovanTa, saxeobaTa-
Soris somatur hibridizaciaSi.

1.3.6.3. in vitro in ducirebuli mutacia

in vitro mutagenezi warmoadgens genetikurad Secvlili

formebisa da mcenareTa axali jiSebis miRebis efeqtur
xerxs ujredul seleqciaSi. dReisaTvis am gziT miRebulia
mravalricxovani mecnieruli da praqtikuli mniSvnelo-
bis aminomJavebisa da maTi analogebisadmi gamZle simindis,
qeris, brinjis, Tambaqos, petunias da sxva; herbicidebi-
sadmi gamZle forToxalis, ionjas, Tambaqos, pomidoris,
soias, stafilos, da sxva; damlaSebisadmi gamZle Tambaqos,
lemas, brinjis, ionjas da sxva; gvalvagamZle Tambaqos, po-
midoris, soias da sxva; daavadebebisadmi gamZle kartofi-
lis, Tambaqos, brinjis, simindis da sxva inducirebuli mu-
83
somaturi hi bri di zaci i s gzi T mi Rebuli M seleqci urad
Zvi rfasi n i San ebi s matarebeli hi bri debi
cxri li 1
kombi n aci a M seleqci urad Zvi rfasi
memkvi druli n i San i
Brassicaceae:
B. napus (+) Diplotaxis harra, Eruca sativa, -cms, gvalvis, nematodebis mi-
Sinapsis alba, S. Pubescens; marT gamZleoba;
B. oleracea (+) B. napus, B. oleracea (+) -sxvadasxva daavadebebisadmi
Sinapsis alba; gamZleoba;
B. napus (+) B. nigra, B. juncea, B. carinata. -sokovani daavadebebisadmi
gamZleoba.
Solanaceae:
Solanum tuberosum (+) S. bulbocastanum; -nematodebisa da fitoptoro-
zisadmi gamZleoba;
S. tuberosum (+) S. Brevidens; -virusuli da baqteriuli da-
avadebebisadmi gamZleoba;
S. tuberosum (+) S. Nigrum; -fuzariozisadmi gamZleoba;
S. tuberosum (+) S. Commersonii; -sicivisadmi gamZleoba;
S. tuberosum (+) S. etuberosum; -virusuli daavadebebisadmi
gamZleoba;
Lycopersicon esculentum (+) S. Tuberosum; -cms;
Nicotiana tabacum (+) N. repanda, N. rustica. -sxvadasxva daavadebebisadmi
gamZleoba.
Leguminosae:
M . sativa (+) M . coerulea, M . intertexta, -sxvadasxva daavadebebisadmi
M . falcate. gamZleoba.
Gramineae:
Oryza sativa (+) Porteresia coarctata; -damlaSebisadmi gamZleo-
ba;
O. sativa (+) O. perrieri, O. officinalis, -sxvadasxva daavadebebisadmi
O. eichingeri, O. M inuta. gamZleoba.

Rutaceae:
Citrus sinensis (+) Fortunella sp., Severinia -sxvadasxva daavadebebisadmi
disticha, C. Unshiu; gamZleoba; sicivegamZleoba.
Citrus unshiu (+) C. jambhiri, C. junos.
tantebi da a.S.
es realuri gzaa aminomJavebis, gansakuTrebiT Seuc-
vleli aminomJavebis maRali Semcvelobis mcenareebis misa-
Rebad. gamorCeulia sawyis mSoblebTan SedarebiT Seuc-
vleli aminomJava triftofanis 20-30-jer ufri meti sinTe-
zis unaris mqone Tambaqosa da stafilos ujrduli xazebi.
84
am xerxiT miRebulia lizinis, meTioninis, fenilalaninis,
glicinis zesinTezis unaris mqone kartofilis, stafilos,
brinjis rigi ujreduli xazi.
aRniSnuli meTodiT miRebulia helmintosporozisadmi
gamZle simindis da fitoptorisadmi rezistentuli karto-
filis xazebi; Tambaqos mozaikis virusisadmi imunuri Tam-
baqos mcenareebi.
mizanmimarTuli seleqciuri muSaobiT Seqmnilia sxva-
dasxva stresuli faqtorebis – damlaSeba, daWaobeba, daba-
li da maRali temperatura, mimarT gamZle sxvadasxva mce-
naris ujreduli xazebi da mcenareebi.
mutagenezis gziT miiRes kartofilis ramodenime uj-
reduli kloni prolinis40 maRali (2-25-jer meti) produci-
rebis unariT, xolo maTgan regenerirebuli mcenareebidan
erTi – dabali temperaturisadmi gamZleobis maRali una-
riT.
seleqtiur areSi pegi-s monawileobiT Seqmnili wylis
stresis pirobebSi gamorCeulia Tambaqos da pomidoris
gvalvagamZle ujreduli xazebi.
inducirebuli mutageneziT pirdapiri gamorCevis gziT
Seqmnilia vercxliswylisadmi gamZle petunias, aluminisa
da manganumisadmi gamZle stafilos, kadmiumisadmi gamZle
pomodorisa da lemas ujreduli xazebi da sxva.
amrigad, ujreduli seleqcia somaklonuri cvaleba-
dobis, somaturi hibridizaciis da in vitro inducirebuli mu-
taciis meTodebis gamoyenebiT, kvlevis obieqtebze mizan-mi-
marTuli moqmedebiT ujredebis genetikuri aparatis kon-
struirebis Sedegad, memkvidreobisa da cvalebadobis regu-
laciis uzrunvelyofiT, warmoadgens mcenareTa genetikuri
mravalferovnebis Seqmnis, axali jiSebis miRebis an arse-
bulis gaumjobesebis efeqtur saSualebas.

40 warmoadgens araspecifikur proteqtors, romelic damlaSebisa da

eqstremaluri temperaturisadmi gamZle mcenareebis miRebis saSualebas
iZleva.
85
 Tavi 2.
Ugenetikuri inJineria

2.1.GmcenareTa genetikuri inJineriis raoba

da teqnologiis etapebi
genetikuri inJineria aris bioteqnologiis meTodi,
romelic cocxali organizmebis genetikuri aparatis winas-
war dasaxuli gegmiT (e.i. mizanmimarTulad) gardaqmnisa da
maTTvis axali Tvisebebis miniWebis, an principulad axali
organizmebis Seqmnis meTodebsa da xerxebs amuSavebs. igi
laboratoriuli gziT sinjaraSi (in vitro) e.w. rekombinantu-
li (hibriduli) dnm-s molekulebis saxiT uzrunvelyofs
xelovnuri, funqcionalurad aqtiuri genetikuri struqtu-
rebis konstruirebas.
mcenareTa mizanmimarTuli genetikuri konstruirebis
mZlavr bioteqnologiur saSualebas warmoadgens mcenareTa
genetikuri inJineria.
mcenareTa genetikurma inJineriam ganviTareba gacile-
biT gvian daiwyo, vidre cxovelebis da miT umetes, baqte-
riebis.
mcenareSi ucxo genebis gadatanis pirveli eqsperimen-
tebi TariRdeba 1980 wliT. pirveli dadebiTi Sedegebi mi-
Rebul iqna 1983 wels.
cneba “mcenareTa genetikuri inJineria” moicavs samuSa-
oebs ujredul doneze da iTvaliswinebs ujredebisa da
qsovilebis kulturis, molekuluri biologiisa da geneti-
kuri masalis gadatanis (ucxo dnm-s gamoyofa, mcenaris uj-
redSi Seyvana da eqspresia) yvela aspeqts41. aseTi samuSaoe-
bis dasrulebis Semdeg adgili aqvs organizmis memkvidru-

41mniSvnelovan upiratesobas warmoadgens mcenaris ujredis totipoten-

turobis Tviseba, anu calkeuli ujredidan mTliani mcenaris regenerac i-
is unari.
86
li aparatis transformacias42 da miiReba e.w. transformi-
rebuli (transgenuri, genmodificirebuli) organizmebi.
sqesobrivi hibridizaciis dros genetikuri masalis ga-
datanisas Cveulebriv adgili aqvs SeWiduli genebis mTeli
blokebis (klasteris) Secvlas, mcenareTa genetikuri inJi-
neriis meTodebi ki calkeuli genebis gadatanis, Sesabami-
sad, konkretuli niSnis cvlilebis da ukve Camoyalibebu-
li genotipis daSlis riskis Semcirebis saSualebas iZle-
va.
sqesobrivi hibridis memkvidrul aparatSi sasurveli
niSnebi fiqsirdeba SemTxveviTi wesiT, rac seleqcioners
aiZulebs Caataros erTdroulad mravalricxovani Sejvare-
bebi da gamoarCios ufro perspeqtiuli mcenare, rac saWi-
roebs aTwleulebs.
mcenareTa genetikuri inJineriis dros gamoricxulia
aseTi SemTxveviTobebi Sesaqmneli mcenareebis genetikurad
axal kombinaciebSi, radganac am dros uzrunvelyofilia: 1)
sasurveli niSnis miRebaze pasuxismgebeli konkretuli ge-
nis identificireba da gamoyofa, 2) am genis mcenaris ujre-
dis memkvidrul aparatSi CarTva Sesabamisi winaswar damu-
Savebuli meTodebiT, 3) Secvlili genomis mqone calkeuli
ujredebidan srulfasovani mcenaris regeneracia.
amdenad, mcenareTa genetikuri inJineria mimarTulia
mcenareebSi sasurveli niSan-Tvisebebis miRebisaken maTSi
Sesabamisi genebis pirdapiri Seyvanis gziT, nacvlad mra-
valricxovan SejvarebebSi xangrZlivi muSaobisa tradiciu-
li seleqciis dros.
transformirebuli mcenareebis miRebis mcenareTa ge-
netikuri inJineriis teqnologia moicavs Semdeg ZiriTad
etapebs:
1. saWiro genis SerCeva, Zieba, identifikacia da klo-
nireba;

42mcenareTa ujredis genetikuri aparatis mizanmimarTul modifikacias

ucxo dnm-s mcire fragmentis (rekombinantuli dnm-s) mizanmimarTuli ga-
datanis da CaSenebis gziT transformacia ewodeba.
87
 2. mcenare-recipientis genotipis SerCeva;
3. genis Seyvana mcenare-recipientis genomSi;
4. transformirebuli ujredebis regeneracia da tra-
nsformirebuli mcenareebis gamorCeva;
5. transformirebuli mcenaris genomSi genis eqspre-
siis testireba.
Ggenis SerCeva dakavSirebulia mcenarisaTvis garkveuli
sasargeblo niSnis gadacemasTan.
dReisaTvis mcenareTa transformaciisaTvis ZiriTadad
iyeneben genebs, romlebic gansazRvraven iseT monogenur
niSnebs, rogoricaa herbicidebis, pesticidebis, zogierTi
stresuli faqtoris mimarT gamZleoba. am niSnebis ganmsaz-
Rvreli genebis umetesoba gamoyofilia baqteriebis geno-mi-
dan.
bolo dros gamZleobis niSnebis ganmsazRvreli gene-
bis donorebad xSirad iyeneben veluri saxeobebis genomebs.
aseTi genebis mcenare-recipientis genomSi Seyvana sqesob-
rivi hibridizaciis gziT SeuZlebelia sxvadasxva saxeobis,
gvaris da zogjer ojaxis mcenareTa biologiuri SeuTav-
seblobis gamo.
kidev ufro rTulia raodenobrivi niSnebis: produq-
tiulobis, gvalvis, dabali da maRali temperaturisadmi
gamZleobis ganmsazRvreli genebis SerCeva da klonireba.
mcenare-recipientis genotipis SerCevisas idealuri
SemTxvevaa, roca SesaZlebelia iseTi jiSis an xazis mce-na-
ris SerCeva, romelic pasuxobs warmoebis moTxovnebs mo-
savlianobis, nayofebis xarisxis, bioturi da abioturi
stresebisadmi gamZleobis mixedviT, magram aqvs mxolod er-
Ti uaryofiTi Tviseba, magaliTad mavne mwerebisadmi sus-ti
gamZleoba. aseT dros am mcenaris genomSi magaliTad, mwe-
rebze damRupvelad momqmedi prototoqsinis masinTezirebe-
li baqteriuli genis bt2 (cry2A) Seyvana, gamoiwvevs jiSis an
xazis arsebiT gaumjobesebas.
mcenare-recipientis genotipis SerCeva didadaa damoki-
debuli misi ujredebis mier mTliani fertiluri mcenaris
regeneraciis unarze, vinaidan es Tviseba dakavSirebulia
88
genotipTan.
genis Seyvana orlebniani da zogierTi erTlebniani
recipenti-mcenareebis genomSi arsebiTad gaadvilebulia ni-
adagis baqteria Agrobacterium tumefaciens-is Ti-plazmidis aR-
moCenis Semdeg. bolo dros gansakuTrebiT erTlebnian mce-
nareebSi sakmaod xSirad, iyeneben genis Seyvanis biobalis-
tikur meTods.
metad mniSvnelovania Seyvanili genis mcenare-recipi-
entis genomSi efeqturi eqspresiis uzrunvelyofa da axa-
li niSnis TaobebSi stabiluri memkvidreoba. Seyvanili ge-
nis eqspresia damokidebulia mraval faqtorze, maT Soris
genomSi genis integraciis adgilze, promotoruli ubnis
da genis meTilirebaze da sxva.
transformirebuli mcenareebis miRebis efeqturoba
pirdapir kavSirSia mcenare-recipientis ujredebis, anu
transformirebuli ujredebis regeneraciul unarze. Tavis
mxriv regeneracia damokidebulia garkveuli qsovilebis
ujredebis totopotenturobaze, amitom regeneraciis unari
sxvadasxva qsovilebSi sxvadasxvaa. garda amisa, regenera-
ciis unariT erTmaneTisagan gansxvavdebian calkeuli saxe-
obebi da jiSebic ki.
totopotenturoba kargadaa gamoxatuli orlebniani
mcenareebis ujredebSi iseTebSi, rogoricaa kartofili,
Tambaqo, Warxali, soio, rafsi, pomidori, stafilo, kom-
bosto, zogierTi xexilovani, maSin roca gaZnelebulia
transformirebuli mcenareebis regeneracia umetesi parkos-
nis, wiwakis, badrijnis ujredebidan. erTlebnianebSi, gan-
sakuTrebiT marcvlovanebSi, totipotenturobis Tviseba ga-
moxatulia Zalian sustad, amitom am ujredebidan mcenare-
Ta regeneracia Zalian Znelia. Tumca bolo wlebSi aseT
mcenareTa regeneraciis meTodebis srulyofa mudmivad xde-
ba.

89
2.2. mcenareebSi genebis gadatanis meTodebi
mcenareebSi genebis gadatana SeiZleba sxvadasxva me-
TodebiT. isini ZiriTadad iyofa sam jgufad (sur. 24):

m ce nare e bS i g e ne bis g ad atanis

m e To d e bi

g en ebis g ad atan i s G komb i n i reb u l i g en ebis g ad atan i s

meT od eb i veqto- meT od eb i pi rd api ri
reb i s s aS u al eb i T meT od eb i

ag rol i s ti ki s
„ f oT l i s rg ol e- meT od i mcen ari s protop-
b i s “ meT od i l as tebis donorul
d n m-n kokultivacia

protopl as teb S i
d n m- s mi kroi n eq-
ag rob aqteri as T an ci i s meT od i
protopl as teb i s
koku l ti vaci a
protopl as teb i s
el eqtrof oraci i s
meT od i

in vitro eqs pl antate-

b i s i n oku l aci i s d n m-s l iposomeb S i
meT od i C awyob i s meT od i

vaku u m-
in planta tran s f or- i n f i l traci i s
maci i s meT od i meT od i

b i ob al i s ti ku ri
tran s f ormaci i s
meT od i

meqaniku ri i n oku -
l aci i s meT od i

s ur. 2 4. m cenareebSi g e nebis g adatanis m eTodebi

1. genebis gadatanis meTodebi veqtoruli sistemebis

(veqtorebis) saSualebiT;
90
 2. genebis gadatanis pirdapiri meTodebi;
3. genebis gadatanis kombinirebuli meTodebi.
pirvel jgufs miekuTvneba:
- „foTlis rgolebis“ meTodi;
- agrobaqteriasTan mcenaris protoplastebis kokul-
tivaciis meTodi;
- in vitro eqsplantatebis inokulaciis meTodi;
- in planta43 transformaciis meTodi.
meore jgufs miekuTvneba:
- Tavisufali STanTqma anu mcenaris protoplastebis
donorul dnm-Tan kokultivacia;
- protoplastebSi dnm-s mikroineqciis meTodi;
- mcenaris protoplastebis eleqtroforaciis (eleq-
troSokis) meTodi;
- dnm-s liposomebSi Cawyobis meTodi;
- vakuum-infiltraciis meTodi;
B - biobalistikuri transformaciis meTodi;
- meqanikuri inokulaciis meTodi.
Mmesame jgufs miekuTvneba agrolistikis meTodi.

G2.3. genebis gadatana veqtorebis gamoyenebiT

2.3.1. veqtorisadmi wayenebuli moTxovnebi
genetikur inJineriaSi genebis gadatanis ZiriTad ia-
raRs warmoadgens veqtorebi.
veqtori ewodeba dnm-s molekulas, romelsac gaaCnia
recipientis ujredSi ucxo genetikuri informaciis gadata-
nis, gamravlebisa da Senaxvis unari.
veqtori warmoadgens ucxo dnm-s recipientis ujre-
debSi gadatanis Tavisebur satransporto saSualebas (er-
Tgvar molekulur „taqss“)44.
Tavisi rolis efeqturi SesrulebisaTvis veqtori zo-
gadad unda akmayofilebdes garkveul moTxovnebs:

43
ingl. in plant - mcenareSi
44
vehile [ingl] – oTxTvala.
91
 1) iyos mcire zomis, raTa mis dnm-Si SeiZlebodes sa-
surveli genis/genebis CarTva;
2) iyos replikoni, e.i. hqondes replikaciis unari, ra-
Ta stabilurad arsebobdes recipientis (patronis) ujred-
Si;
3) hqondes amplifikaciis e.i. mravaljeradi gaormage-
bisa da didi raodenobiT aslebis warmoqmnis unari, raTa
masSi CarTuli geni ujredSi aseve warmodgenili iyos mra-
vali aslebiT da maSasadame, misi Sesatyvisi cila – didi
raodenobiT;
4) hqondes ama Tu im restriqtazas Secnobis unikalu-
ri saiti, raTa am saitSi CarTos saWiro geni/genebi;
5) unda uzrunvelyos misi molekulis SedgenilobaSi
CarTuli ucxo dnm-s eqspersia (muSaoba);
6) hqondes seleqtiuri markeri, e.i. Seicavdes gens,
romlis funqcia ujreds aniWebs advilad gasarCev niSans
(magaliTad, antibiotikebisadmi gamZleobas), rac xels uw-
yobs ujredebis, romlebmac miiRes ucxo geni, gamorCevis
efeqturobis gadidebas;
7) ucxo dnm-s Seyvanam ar unda daarRvios misi arsebi-
Ti funqciebi;
8) hqondes Seyvanili ucxo dnm-s Sesabamisi organiz-
mis ujredebSi gadacemis SesaZlebloba.
bunebaSi veqtorebis yvelaze xelsayrel kandidatebs
warmoadgenen – plazmidebis, virusebis, maT Soris fagebis,
mitoqondriebis da qloroplastebis dnm-s molekulebi.
uSualod veqtorebad maT praqtikulad ver iyeneben. imisaT-
vis, rom isini pasuxobdnen zemoT aRniSnul moTxovnebs, au-
cilebelia maTi modificireba an kombinireba. amasTan veq-
torebis konstruireba xdeba raRac konkretuli miznisaT-
vis. magaliTad, aris veqtorebi, romlebic emsaxureba mxo-
lod genebis amplifikacias (gamravlebas) an seqvenirebas;
aris veqtorebi dnm-s patronis qromosomaSi gadatanisa da
integraciisaTvis da a.S.
genetikuri inJineriis veqtorisadmi wayenebuli zoga-
di moTxovnebi mTlianad exeba mcenareul veqtorebsac.
92
 idealuri mcenareuli veqtori unda flobdes Semdeg
Tvisebebs:
1) iyos molekulurad klonirebadi;
2) gaaCndes genetikuri informaciis aqceptirebis (miRe-
bis) unari;
3) Seicavdes specifikur saits masSi ucxo genis integ-
raciisaTvis;
4) hqondes patronis ujredebSi gadatanis unari;
5) aucilebelia veqtorisa da patronis ujredSi klo-
nirebisaTvis gankuTvnili dnm-s stabiluroba da SeTavseba-
doba;
6) uzrunvelyos didi raodenobiT aslebis replikacia
da mcenaris ujredebSi ucxo dnm-s eqspersia;
7) Tu saWiroa transformirebuli mcenaris memkvidru-
li cvalebadoba, maSin veqtori stabilurad unda iyos
axal patronSi;
8) veqtoruli molekulis mcenaris ujredSi an mcena-
reSi arseboba advilad unda seleqtirebdes45 , rac veqto-
rul molekulaSi seleqtiuri markeris CarTvis aucileb-
lobaze miuTiTebs;
9) iyos SeTavsebadi normaluri, jansaRi mcenaris fe-
notipTan, e.i. dnm-s es fragmenti ar unda iyos paTogenuri;
10) klonirebadi geni stabilurad unda gadaecemodes
transformirebuli mcenaris Taobas rogorc somaturi, ise
sqesobrivi gziT;
11) hqondes mcenaris SigniT sistemuri gavrcelebis
unari, raTa uzrunvelyos Seyvanili dnm-s gamovlena mcena-
ris sxvadasxva organoebSi;
12) hqondes nebismieri genis (misi warmoSobis miuxeda-
vad) mcenaris ujredebSi Seyvanis, damagrebisa da eqspresi-
rebis unari;
13) veqtorisaTvis metad sasurvel Tvisebas warmoad-
gens patronTa farTo speqtri, raTa is gamoyenebul iqnas

45
seleqturi [laT. selectio] – gamorCeva.
93
mcenareTa farTo wrisaTvis, pirvel rigSi yvelaze Zvirfa-
si sasoflo-sameurneo kulturebisaTvis.

2.3.2. mcenareTa genetikuri inJineriis veq torebi

mcenareebisaTvis potenciuri veqtorebi SeiZleba dai-

yos Semdeg kategoriebad:
1) patron-mcenaris genomSi integrirebis unaris mqone,
bunebrivi mcenareuli veqtorebis safuZvelze miRebuli veq-
torebi – agrobaqteriis plazmidebi (Ti da Ri);
2) mcenareuli paTogenebis safuZvelze miRebuli veq-
torebi, romlebic xasiaTdebian ucxo dnm-s (an rnm-s) mcena-
reSi Seyvanis bunebrivi unariT – fitovirusebi, viroidebi;
3) veqtorebi, romlebsac SeuZliaT arseboba rogorc
damoukidebel replikonebs mcenaris ujredebSi – mitoqon-
drialuri da qloroplasturi dnm;
4) mcenaris genomis arastabiluri struqturebis sa-
fuZvelze miRebuli veqtorebi – transpozonebi.
jer-jerobiT mcenareTa genetikur inJineriaSi yvelaze
farTo gavrceleba moipova agrobaqteriis plazmidurma veq-
torebma.

2.3.2.1. Aagrobaqteriis veq torebi

mcenareTa genetikur inJineriaSi genebis gadasatanad

agrobaqteriuli veqtorebis gamoyenebis teqnologiebis axs-
nisaTvis aucilebelia mcenareebSi agrobaqteriiT gamowve-
uli simsivnewarmoqmnis procesis, anu bunebrivi genetikuri
transformaciis sakiTxis ganxilva.

2.3.2.1.1. mcenareTa bunebrivi transformacia

agrobaqteriebiT

gvi rgvinisebri galebis w armoqmna – gen etikuri i n Ji -

n eri i s bun ebri vi forma. mcenaris ujredebSi genebis gada-
tanis problemis gadawyvetis saqmeSi udidesi garRveva iyo
94
niadagis baqteriis Ti-plazmidebiTNmcenareTa transforma-
ciis bunebrivi sistemis gamoyenebis SesaZleblobis aRmoCe-
na.
cnobili iyo, rom gramuaryofiTi niadagis baqteriis
(Agrobacteria) zogierTi saxeobebi (Agrobacterium tumefaciens) ase-
nianeben orlebnian mcenareebs da iwveven specifikuri sim-
sivneebis, e.w. „gvirgvinisebri galebis“ warmoqmnas (sur. 25).
simsivneebi Sedgeba intensiurad dayofadi, dazianebis adgi-
las mzardi dediferencirebuli ujredebisagan. in vitro pi

s ur . 2 5. g v ir g v inis e br i g a l e bi

robebSi kultivirebisas simsivnuri ujredebi izrdebian

normaluri mcenareuli ujredebis zrdisaTvis aucilebeli
hormonebis gareSe anu axasiaTebT hormondamoukidebeli
zrda, vinaidan axdenen sakuTari fitohormonebis – auqsine-
bisa da citokininebis producirebas. gvirgvinisebri gale-
bis simsivnuri ujredebis hormondamoukidebeli zrda
95
mdgomareobs imaSi, rom normaluri ujredebis zrdasTan
gansxvavebiT isini kulturalur sakveb areze izrdebian ho-
rmonebis – auqsinebisa da citokininebis damatebis gareSe.
gairkva, rom dasenianebis Semdeg agrobaqteriebis mo-
cilebis (inaqtivirebis) SemTxvevaSic gvirgvinisebri gale-
bis ujredebi inarCuneben dasenianebis adgilas araorgani-
zebuli da arakontrolirebadi zrdis unars, anu agrobaq-
teriebis arseboba aucilebelia mxolod simsivnis warmoq-
mnis inducirebisaTvis.
amasTanave dadginda, rom simsivnis induqciis aucile-
bel pirobas warmoadgens potenciuri patron-mcenaris meqa-
nikuri dazianeba (Wrilobis miyeneba). dauzianebel epider-
misze agrobaqteriis gadataniT simsivneebis inducireba
uSedegoa.
Ffitohormonebis gaZlierebuli sinTezis garda, gvir-
gvinisebri galebis simsivnuri ujredebi iwyeben mcenari-
saTvis uCveulo, dabalmolekuluri biologiurad aqtiuri
naerTebis – opinebis (nopalini, oqtopini, agropini da
sxva) sinTezs, romelsac agrobaqteriebi iyeneben azotisa
da naxSirbadis wyarod. TviT agrobaqteriebs opinebis sin-
Tezi ar SeuZliaT, vinaidan sinTezis Sualeduri produqte-
bi iwveven baqteriuli ujredebis zrdis inhibirebas da mce-
nareul ujredebs iyeneben rogorc opinebis warmoebis
„fabrikebs.“ agrobaqteriebiTMmcenareTa dasenianebisas ad-
gili aqvs transformirebuli mcenareuli ujredebis meta-
bolizmis gardaqmnas da isini iwyeben mxolod baqte-rii-
saTvis aucilebeli naerTebis sinTezs. opinebis utili-za-
cia ar SeuZlia mcenareul ujreds da isini gadadian uj-
redSorisebSi, sadac STainTqmebian agrobaqteriebis mi-er. A
eqsperimentuli monacemebis safuZvelze aRmoCnda, rom
agrobaqteriiT gamowveuli gvirgvinisebri galebis warmoq-
mnis procesi aris genetikuri inJineriis bunebrivi forma.
igi Sedgeba rigi etapisagan, moicavs rig faqtorebs da zo-
gadad mdgomareobs SemdegSi (sur. 26): specifikuri plazmi-
dis matarebeli agrobaqteriebi dazianebuli, Wrilobamiye-

96
 a b g d e
qromosomuli dnm

plazmid a qromosoma
Ddnm-
fragmenti
dnm-fragmenti
Wriloba

A.tumefaciens gvirgv.
transformi gali
opinebi
rebuli mcen.
Agrobacterium ujredi
tumefaciens

s ur . 2 6 . g v ir g v inis e br i g al is ind uq ciis e tap e bi

a – A. tumefaciens-is ujredebi arseboben mcenaris rizosferoSi;
b – baqteriis ujredebSi qromosomasTan erTad aris plazmida;
g – agrobaqteriebi dazianebuli, Wrilobamiyenebuli qsovilebidan
SeRwevis gziT asenianebs mcenares;
d – plazmida SeaRwevs mcenaris ujredSi da dnm-s fragmenti CaSende
ba (integrirdeba) mcenaris ujredis qromosomaSi;
e – es iwvevs gvirgvinisebri galebis warmoqmnas da opinebis sinTezs.

nebuli qsovilidan SeRwevis gziT asenianebs mcenares.

plazmiduri dnm-fragmenti Sedis mcenaris ujredSi. igi
amoiWreba plazmididan da integrirdeba (CaSendeba) mcena-
ris ujredis qromosomaSi. Sedegad baqteriis plazmidis
dnm-s fragmenti xdeba mcenaris memkvidruli aparatis nawi-
li (sur. 27) da mcenaris genebTan erTad iwyebs eqspresire-

baqteriis plazmidis
mcenaris dnm mcenaris dnm
dnm-s fragmenti

marcxena marjvena
ganmeoreba ganmeoreba

s ur . 2 7 . baq te r iis p l azm id is d nm - s f r ag m e ntis

CaS e ne ba (inte g r ir e ba) m ce nar is d nm - S i

bas (muSaobas). maTi eqspresia inducirebs simsivnewarmoq-

mnis process. es ki iwvevs mcenaris ujredebSi zemoT aR-
niSnuli ramodenime axali Tvisebis warmoSobas: 1) iwyeba
ujredebis arakontrolirebadi dayofa, ris Sedegadac warmo-
97
iqmneba simsivne; 2) simsivnur ujredebSi sinTezirdeba uCve-
ulo, opinebad wodebuli naerTebi, romelTa mixedviT Se-
saZlebelia simsivnuri ujredebis testireba; 3) iwyeba sim-
sivnuri mcenareuli ujredebis hormondamoukidebuli zrda.
M agrobaqteriis plazmidebis fun qciebi da struqtura.
dadgenil iqna, rom A. tumefaciens-is onkogenuroba ganpiro-
bebulia maTSi didi zomis, sigrZiT 50-80 mkm da molekulu-
ri woniT 112-156 megadaltoni (200-800 aTasi n.w.), kovalentu-
rad Sekruli, avtonomiurad replicirebadi, rgoluri pla-
zmiduri dnm-s arsebobiT. aseTi plazmida Seadgens baqteri-
aSi arsebuli dnm-s daaxloebiT 5-8% da Cveulebriv ujred-
Si arsebobs 1-2 aslis saxiT. swored es didi plazmidebi
monawileoben gvirgvinisebri galebis warmoqmnis procesSi.
plazmidam miiRo saxelwodeba Ti-plazmida46.
sinTezirebadi opinebis tipis mixedviT arCeven 3 tipis
Ti-plazmidebs: 1) „oqtopinuri“ tipis, romlebic akodireben
oqtopins, lizopins, oqtopinis mJavas da sxva; 2) „nopalinu-
ri“ tipis, romlebic akodireben nopalins, nopalinis mJavas
da sxva; 3) „agropinuri“ tipis, romlebic akodireben agro-
pinis, manopinis, manopinis mJavas da sxvas. yvelaze xSirad
gavrcelebulia nopalinis an agropinis makodirebeli pla-
zmidebi. ujredi Seicavs mxolod erTi tipis plazmidas.
M molekulur-genetikurma kvlevebma aCvena, rom yvela Ti-
plazmidas aqvs msgavsi agebuleba (sur. 28) da Seicaven Tan-
mimdevrobebs, romlebic SeiZleba daiyos or jgufad:
1) TviT agrobaqteriis metabolizmisaTvis aucilebeli
(opinebis katabolizmis genebi, plazmidis replikaciis da-
wyebis adgili da sxva);
2) mcenareuli ujredis transformaciisaTvis aucile-
beli (simsivnis induqciaze da opinuri aminomJavebis sin-
Tezze pasuxismgebeli genebi).
pirveli jgufis genebs gaaCniaT prokariotuli tipis

46
Tumor inducing [ingl.] – simsivnis mainducirebeli.

98
 T-dn m
T-dn m

acs

ocs
noc
vir pTiC58 vir pTiAch5

occ
acc
agc

a b

s ur . 2 8 . no p al inur i (a) d a o q to p inur i (b) Ti-p l azm id is

g e ne tiku r i r u q a
(onc – onkogenebi; nos – nopalinsinTetazas genebi; acs – agrocino-
pinsinTetazas genebi; noc, occ, agc, acc – genebi, romlebic axorcie-
leben opinebis – nopalini, oqtopini, agropini da agrocini kata-
bolizms; vir – genebi, romlebic pasuxusmgebelia virulentobaze)

promotori, amitom funqcionireba SeuZliaT mxolod agro-

baqteriis ujredSi, meore jgufis genebs eqspresia SeuZli-
aT eukariotul mcenareul ujredSi.
unda aRiniSnos, rom mcenareTa simsivneebs Agrobacterium
tumefaciens garda aseve iwveven agrobaqteriis sxva virulen-
turi saxeobebi – Agrobacterium rhizogenes da Agrobacterium rubi.
maT mier inducirebul simsivneebs ewodeba Sesabamisad
“wverisebri fesvi” da“Rerosebri gali” (sur. 29).
daavadeba „wverisebri fesvis“ gamomwvevi baqteria A. rhi-
zogenes Seicavs plazmidas, romelsac hqvia Ri-plazmida47. am
plazmidiT gamowveuli daavadeba “wverisebri fesvi” xasi-
aTdeba meoradi fesvebis farTo proliferaciiT48 - warmo-
iqmneba adventiuri fesvebis simsivnuri ujredebi. mas gaaC-

47
Root inducing [ingl.] – fesvmainducirebeli.
48
proliferacia [laT. proles – STamomavloba; fero – damaqvs, davatareb] – or-
ganizmis qsovilis zrda ujredebis gamravlebis gziT.
99
 s ur. 2 9. “wverisebri f esvi“ (zeda) d a „R erosebri g ali“ (qveda)
nia naklebi moculoba, vidre „gvirgvinisebr galebs“. cal-
saxad dadgenilia, rom sindromi „wverisebri fesvi“ war-
moiqmneba „gvirgvinisebri galebis“ sruliad analogiuri
procesis Sedegad. ZiriTadi gansxvaveba maT Soris mdgoma-
reobs imaSi, rom „gvirgvinisebri galis“ daavadebas safuZ-
vlad udevs simsivnuri zrda, xolo „wverisebri fesvis“ –
diferencirebuli qsovilebis proliferacia.
cnobebi A. rhizogenes-is Ri-plazmidebze amJamad iseTi
farTo ar aris rogorc Ti-plazmidis SemTxvevaSi, magram
arsebuli monacemebiT SeiniSneba Zalian bevri msgavsi ka-
nonzomiereba.
klasifikaciis mixedviT Ri-plazmidebi iyofa 2 kla-
sad: 1) „agropinuri“ tipis, romlebic akodireben agropins,
manopins da sxva; 2) „manopinuri“ tipis, romlebic akodire-
ben manopinis, manopinis mJavis da sxva.
Ri-plazmidebs genetikur inJineriaSi Ti-plazmidebTan

100
SedarebiT gaaCniaT garkveuli upiratesoba, radganac mcena-
reebi ukeT regenerireben simsivnuri fesvebidan. Ri-plazmi-
debi ganixileba rogorc plazmidebis bunebrivi uvnebeli
(„ganiaraRebuli“) varianti.
D T-dn m-s struqtura da fun qci ebi . dadgenil iqna, rom
agrobaqteriuli dasenianebiT mcenaris transformaciisas
TviT agrobaqteria ar Sedis mcenaris ujredSi, is Cerdeba
ujredSorisebSi. ar Sedis ujredSi Ti-plazmidac, magram
mcenaris ujredis birTvSi gadaitaneba Ti-plazmidis nawi-
li, romelic Semdeg CaSendeba mcenareul genomSi da iwvevs
transformacias – „gvirgvinisebri galis“ simsivnis warmoq-
mnas. Ti-plazmidis dnm-s am nawilma, romelic kovalenturad
dakavSirebuli aRmoCnda transformirebuli mcenaris ujre-
debis qromosomul dnm-Tan, miiRo T-dn m-s 49 saxelwodeba. am-
Jamad T-dnm kartirebulia, masSi identificirebulia sxva-
dasxva tipis Ti-plazmidisaTvis saerTo maRalkonservatiu-
li ubani (sur. 30).

s ur. 30. T-p lazmida m niSvnelovani g enebiTa d a ubnebiT

49
T ransforming DNA (ingl.) – matransformirebeli dnm. Tu T i-plazmidis es
nawili mdebareobs baqteriul ujredSi, maSin hqvia – T -ubani (raioni an
monakveTi), xolo Tu transformirebuli mcenaris ujredSi maSin – T -dnm.
101
 T-dnm-s zoma SedarebiT mcirea, varirebs plazmidis
tipis mixedviT da aRwevs 10 – 30 aTas n.w. (4.5-15 megadal-
toni), rac plazmidis mTeli genomis 5-8%-s Seadgens.
T-dnm-s (akodirebs 10-15 cilas) ZiriTad funqcias war-
moadgens mcenaris genomSi gadasvla, CaSeneba (lokalizeba
birTvSi) da ujredis transformireba (misTvis ucxo pro-
duqtebis sinTezis unaris miniWeba)50.
T-dnm-s (T-DNA region) (sur. 30) boloebi SemosazRvru-
lia 25 wyvili nukleotidisagan Semdgari pirdapiri maRal
konservatiuli kidura ganmeorebebiT (sur. 31), romlebic
Seadgenen e.w. marcxena (Left border repeat) da marjvena (Right
border repeat) mosazRvre raionebs (sur. 30). isini aucilebe-
lia plazmididan T-dnm-s amoWrasa da mcenaris genomSi in-
tegraciisaTvis. gansakuTrebiT mniSvnelovania am saqmeSi
marjvena sazRvris roli. transformaciisaTvis mniSvnelo-
vania aseve marjvena sazRvris gareT mdebare mokle (24 n.w.)
Tanmimdevroba, romelsac overdraivi (overdraive) ewodeba.

GCTGGTGGCAGGATATATTGTGGTGTAAACAAATT 1
GTGTTTGACAGGATATATTGGCGGGTAAACCTAAG 2
AGCGGCGGCAGGATATATTCAATTGTAAATGGCTT 3
CTGAC TGGCAGGATATATACCGTTGTATTTGAGCT 4

s ur . 31 . T-d nm - s kid ur a g anm e o r e be bi

(1 – nopalinuri T -dnm-s marcxena sazRvari; 2 – nopalonuri
T -dnm-s marjvena sazRvari; 3 – oqtopinuri T-dnm-s marcxena
sazRvari; 4 – oqtopinuri T -dnm-s marjvena sazRvari)

T-dnm-s SigniT lokalizebulia 7 geni (sur. 30): uCveu-

lo aminomJavebis – opinebis (Opine syntesis) [oqtopinsin-Tata-
zas (ocs), an nopalinsinTetazas (nos)] masinTezirebeli geni,
agreTve simsivnewarmoqmnis niSnebis makodirebeli eqvsi ge-
ni, romelTagan ori (iaaM da iaaH) akodirebs fito-hormon
auqsinis sinTezSi CarTul fermentebs, xolo erTi (iptZ) –

50
simsivnis ujredebSi imyofeba T -dnm-s erTidan xuTamde asli.
102
fitohormon citokininis sinTezs. am genebis eqspresiis Se-
degad transformirebul ujredebSi icvleba hormonaluri
statusi, rac iwvevs maT dediferencirebas da simsivnis
warmoqmnas.
unda aRiniSnos, rom T-dnm-Si lokalizebuli arc er-
Ti geni ar transkribirdeba agrobaqteriis ujredSi, maTi
aqtiuroba vlindeba mxolod mcenaris ujredSi. aseve mniS-
vnelovania is, rom T-dnm-s gadatanasa da integraciaze pa-
suxismgebeli yvela geni ganlagebulia am ubnis gareT.
plazmidebis vir-ubnis funqcia da struqtura. agrobaq-
teriuli transformaciisaTvis T-dnm-s arseboba aucilebe-
lia, magram sakmarisi ar aris. Ti-plazmididan T-dnm-s amoW-
rasa da mis mcenaris genomSi gadatanaze pasuxismgebelia
Ti-plazmidis sxva, T-dnm-s gareT ganlagebuli ubani, ro-
melsac vir-ubani (virulentobis ubani) ewodeba. igi gansaz-
Rvravs simsivnuri transformaciisaTvis aucilebel proce-
sebs: baqteriis urTierTqmedebas mcenaris ujredebTan, Ti-
plazmidis T-dnm-s SeRwevas mcenaris ujredSi, T-dnm-s in-
tegracias mcenaris genomSi, e.i. virulenturobis (vir) fun-
qciebs.
vir-ubani warmoadgens did segments zomiT daxloebiT
35 aTas nukleotidis wyvili da Ti-plazmidis saerTod mi-
Rebul ruqaze lokalizebulia marcxniv T-dnm-s ubnidan
(sur. 28).
vir-ubani Sedgeba (sur. 30) xuTi vir-operonisa – virB, virC,
virD, virE, virH da sami calkeuli vir-genisagan – virA, virG da virF.
operonebi Tavis mxriv moicavs ramodenime gens, amitom
Tvlian, rom vir-ubani Sedgeba 40-mde genisagan.
Ffunqciis mixedviT vir-genebi SeiZleba daiyos or kla-
sad:
1. virA da virG genebi – regulatoruli genebi, romelTa
eqspresia agrobaqteriul ujredSi mcire raodenobiT mainc
mudmivad xdeba;
2. virB, virC, virD, virE, virF genebi – akodireben cilebs,
romlebic uSualodaa CarTuli T-dnm-s mcenareSi gadatanis
103
procesSi da eqspresirdebian mxolod maTi promotorebis
uSualo aqtivaciis Semdeg.
cxrilSi 2 motanilia agrobaqteriuli simsivnuri
transformaciis procesSi monawile zogierTi vir-genis pro-
duqtebis (Vir-cilebis) ZiriTadi funqciebi.

A. tumefaciens-i s Ti-plazmi di s vir-gen ebi T kodi rebuli

zogi erTi Vir-ci li s Zi ri Tadi fun qci ebi
cxri li 2
aminomJ.
cil a aveb i s f unqcia
rao -b a
VirA 829 orkomponentiani (VirА/ VirG) regulatoruli sistemis
sensoruli cila (membranuli receptori)
VirB1 239 IV tipis sekreciis sistemis aparatis formirebis pir-
vel etapze peptidglikanazas gaxleCa

VirB2 121 Т -pilusis struqturuli cila

gare membranaze asocirebuli cila, romelic xels
VirB3 108
uwyobs Т -pilusis awyobas

VirB4 789 IV tipis sekreciis sistemis aparatis awyobaSi monawi-

leoba, atf-is gaaqtiureba

VirB5 220 gansazRvravs Т -pilusis mimagrebas gare membranasTan

da awyobas

VirB6 295 monawileobs IV tipis sekreciis sistemis aparatis

awyobaSi da substratebis transportirebaSi
lipoproteini, ukavSideba gare membranas, gansaz-
VirB7 55
Rvravs Т -pilusis lokalizavias
gansazRvravs gare membranaze arxSi Sesasvlelis
VirB8 237
lokalizavias
sekretoruli arxis formireba periplazmatur ubanze
VirB9 293

VirB10 377 VirВ8 cilasTan erTad axdens sekretoruli arxis pe-

riplazmatur nawilTan VirВ11 cilis mierTebis stabi-
lizebas
sekretoruli arxis Sesasvlelis formireba Sida mem-
VirB11 343
branaze, atf-is gaaqtiureba

VirС1 231 xels uwyobs cila VirD2-is mimagrebas Т -dnm-Tan

VirС2 202 xels uwyobs Т -kompleqsis gadaadgilebas mcenaris

ujredSi

104
 cx r ili 2 -is d asasruli
VirD1 147 moqmedebs cila VirD2–Tan erTad Т -kompleqsis for-
mirebis dros
endonukleaza, T -dnm-s erTi jaWvis 5′-bolosTan da
VirD2 424 kavSireba da nikireba (Т -kompleqsis formireba);
Т -kompleqsis transportireba mcenaris ujredis bir-
TvSi

VirD4 656 Т -kompleqsis orientacia IV tipis sekreciis sistemis

arxSi Sesasvlelad
dakavSireba T -kompleqsis erTjaWvian dnm-Tan mcena-
VirE2 533 ris ujredis citoplazmaSi da misi transportireba
mcenaris ujredis birTvSi
Saperoni (damxmare cila) VirЕ2 cilis mcenaris uj-
VirЕ3 redSi transportirebisaTvis, dakavSireba mcenaris
ujredebis saerTo transkripciul faqtorTan (pBrp)

VirF 202 mcenaris ujredis birTvSi importirebuli T -kompleq-

sis momzadeba qromosomul dnm-Si integraciisaTvis

VirG 241 vir-operonebis transkripciis regulatori orkompo-

nentiani (VirА/ VirG) sensoruli sistemidan

agrobaqteriuli transformaci i s meqan i zmi . agrobaq-

teriis gamoyenebiT transformirebuli mcenareebis miRebis
teqnologiuri procesi zogadad moicavs Semdeg samuSaoebs
(sur. 32): plazmidis gamoyofa agrobaqteriidan da T-dnm-s
gaWra restriqciis fermentebiT (restriqtazebiT), ucxo
dnm-s amoWra igive fermentebiT (restriqtazebiT), ucxo
dnm-s CaSeneba plazmidis T-dnm-Si gaWris adgilas da re-
kombinantuli plazmidis miReba, plazmidis gadatana isev
agrobaqteriaSi, baqteriis saSualebiT ucxo genis gadatana
mcenaris ujredis qromosomaSi, ujreduli kulturis meTo-
diT transformirebuli ujredidan mcenaris regeneracia,
ucxo genis eqspresia transformirebul mcenareSi.
agrobaqteriis saSualebiT mcenareTa transformacii-
saTvis aucilebelia sami pirobis Sesruleba:
1. agrobaqteriis oTxi qromosomuli genis (chvA, chvB,
chvE, pscA) aqtivacia.
isini baqteriuli ujredis zedapirze axdenen, mcena-
ris ujredTan baqteriis dakavSirebisaTvis aucilebeli,
105
 CaSenebuli T-dnm
ucxo geniT
plazmidis
gadatana isev
baqteriaSi
baqteriebi gamoiyene-ba
restriqciis T-dnm-s ucxo ge-niT
saiti CasaSeneblad ujredis
genomSi
pTi mcenareTa
T-dnm ujredebis
rekombinantuli kultura
plazmidis gamoyofa Ti-plazmida
baqteriidan da T-dnm-s
gaWra restriqtazebiT
ucxo dnm-s CaSeneba
plazmidis T-dnm-Si
erTi ujredidan miRebuli mcenare. misi
ucxo dnm-s amoWra yvela ujredi Seicavs ucxi dnm-s,
imave restriqtazebiT Sesabamisad gaaCnia axali Tvisebebi

s ur. 32 . ag robaq te r iis g am o y e ne biT tr ans f o r m ir e bul i

m ce nar e e bis m iR e bis zo g ad i s q e m a.

specialuri polisaqaridebis sinTezs. simsivnuri trans-

formaciis procesSi monawile A. tumefaciens-is qromosomuli
genebis produqtebis (cilebis) ZiriTadi funqciebi motani-
lia cxrilSi 3;

si msi vn uri tran sformaci i s procesSi mon aw i le

A. tumefaciens-i s qromosomuli gen ebi s produqtebi s
Zi ri Tadi fun qci ebi
cxri li 3

cil a f unq cia

MiaA tRNA izopenteniltransferaza, gavlenas axdens vir-ope-

ronebis induqciaze
Ros virC- da virD-operonebis negatiuri regulatori
ChvA β1,2-glukanis komponentebis transporti ujredis ze-
dapirze
ChvB monawileoba β1,2-glukanis komponentebis sinTezSi
ChvE vir-operonebis qemotaqsisisa da induqciis dros mono-
naxSirwyalbadebis damakavSirebeli cila

106
 ChvG fosfatur SimSilze mareagirebeli orkomponentiani
sensoruli sistemis sensoruli cila

ChvJ fosfatur SimSilze mareagirebeli orkomponentiani

sensoruli sistemis transkripciis aqtivatori
ExoC glukozofosfatizomeraza, gavlenas axdens ekzopo-
(PscA) lisaqaridebis sinTezze

Att gareTa membranis sami cila, monawileoben A. tumefaci-

ens-is mcenaris ujredis zedapirze mimagrebaSi
AchB mcenaris ujredSi Т-kompleqsis gadaadgilebaSi monawi-
leoba
Cel celulozis sinTezis induqcia mimagrebis saitebSi

2. Ti-plazmidis vir-ubnis aqtivacia. igi aucilebelia

mcenaris ujredis birTvSi T-dnm-s gadatanis iniciacisaT-
vis;
3. T-dnm-s arseboba.
agrobaqteriuli simsivnuri transformaciis procesi
(sur. 33) iwyeba adreul stadiaSi CarTuli agrobaqteriis
qromosomuli genebis (chvA, chvB, chvE, pscA) da Ti-plazmidis
virA da virG genebis eqspresiiT.
agrobaqteria mcenares emagreba Wrilobis adgilas.
Wirlobis miyenebisas mcenaris ujredi gamoyofs fenolis
warmoebulebisa da Saqrebis mcire molekulebs, maT Soris
specifikur fenolur SenaerTs – acetosiringons.
rogorc ukve aRiniSna, virA genis eqspresia agrobaqte-
riis ujredSi xdeba mudmivad. cila VirA warmoadgens ace-
tosiringonis (sasignali molekulis) membranul recep-
tors, cila ChvE-Tan qmnis kompleqss, romelic iWreba mem-
branaSi.
mcenaris mier gamoyofili acetosiringons cnobs ci-
la VirA, ukavSirdeba mas, rac iwvevs cila VirA-s aqtivacias
TviTfosfolirirebis (fosfaturi naSTis translokacia
VirA cilis erT ubnidan meoreSi) gziT. es Tavis mxriv iw-
vevs agrobaqteriis ujredSi mudmivad eqspresirebadi meore
cilis – VirG-s aqtivacias. cila VirG regulatoruli

107
 T-dn m
Ti-plazmida

agrobaqteriis
ci toplazma

T- kom-
pl eqsi

vir-uban i

ag robaqteriis ujredis g arsi

mcenareuli ujredis
sasignalo molekulebi

patron-mcenaris uj-
redis kedeli

m ce naris
ujr edi T-d nm

b i rTvi

s ur. 33. ag robaqteriul i tr ans f o r m aciis Zir iTad i e tap e bi

( I – VirA sensoruli cilis (membranuli receptoris) aqtivacia mcenaris ujredis
kedlis hidrolizis produqtebiT (sasignalo molekulebiT); II – vir-genebis tran-
skripciis aqtivator VirG-s fosfolirireba da vir-genebis eqspresia; III – VirD1 da
VirC1-is saSualebiT VirD2-is mierTeba T-dnm-Tan da erTjaWviani wyveta marjvena
sazRvris (RB) raionSi, erTjaWviani Т-dnm-s amoWra marjvena da marcxena (LB)
sazRvrebs Soris; IV – Т-jaWvTan VirD2 cilis dakavSireba da mTeli kompleqsis (T-
jaWvi+VirD2) transporti arxisken; V – virB operonis produqtebiT formirebuli arxi
agrobaqteriis ujredis kedelSi da cilebis kompleqsTan (VirE2 da VieD2) erTad Т-
jaWvis gadasvla mcenaris ujredSi ; VI – Т-dnm-s integracia mcenaris qromosomaSi)

108
cilaa da warmoadgens danarCeni vir-genebis – virB, virC, virD,
virE, virH, virF transkripciis aqtivators. gaaqtiurebuli ci-
la VirG ukavSirdeba am genebis promotorul ubnebs da iw-
vevs maT eqspresias.
vir-genebis – virB, virC, virD, virE, virF cilebi ukavSirdebi-
an T-dnm-s, iwveven mis amoWras Ti–plazmididan da CaSenebas-
mcenaris genomSi.
es procesi iwyeba VirС1, VirD1 da VirD2 cilebiT. Ziri-
Tads warmoadgens endonukleaza VirD2, romelsac SeuZlia
dnm-s orjaWviani molekulis erT jaWvis gawyveta. is cnobs
kidura Tanmimdevrobebs, magram efeqturi mimagrebisaTvis
sWirdeba cilebi VirD1 da VirС1. cila VirD1 uSualod urTi-
erTmoqmedebs VirD2-Tan da gansazRvravs mis swor orienta-
cias T-dnm-s orjaWvian molekulaze.
cila VirС1 emagreba marjvena kidura Tanmimdevrobas-
Tan axlos – overdraivis ubanze da VirD2-Tan erTad iwvevs
T-dnm-s erTjaWviani fragmentis amoWris inicirebas. endo-
nuleaza VirD2 marjvena kidura Tanmimdevrobis me-3 da me-4
nukleotidebs Soris akeTebs erTijaWvian specifikur Wras
da iwyeba am jaWvis mocileba misi komplementaruli jaWi-
dan marcxena kidura Tanmimdevrobis mimarTulebiT. amasTan
TviT endonuklaza mimagrebuli rCeba mosacilebuli jaW-
vis 5′-boloze.
imavdroulad adgili aqvs T-dnm-ze warmoqmnili rRveu-
lis Sevsebas axali nukleotidebiT. ase, rom Zveli jaWvis
sruli mocilebis Semdeg (amisaTvis marcxena kidura Tan-
mimdevrobasTan mimagrebuli cila VirD2-is sxva molekula
akeTebs masze erTjaWvian specifikur Wras) T-dnm rCeba
iseTive rogoric iyo Tavidan.
amoWrili erTjaWviani mobiluri T-dnm (T-strand) 5′-bo-
loze mimagrebuli VirD2 ciliT qmnis moumwifebel T-kom-
pleqs (Immature T-complex), romelic miemarTeba mcenaris uj-
redSi gasasvleli arxisken.
VirB cilebidan (sul 11) minimum sami – VirB2, VirB5 da

109
VirB6 warmoqmnian specifikur transujredul gasasvlel
arxs – T-piluss (T-pilus) (sur. 34), romliTac moumwifebeli
T-kompleqsi gadaitaneba agrobaqteriidan mcenaris ujred-
Si.
aRniSnul kompleqsTan erTad am arxis saSualebiT
mcenaris ujredSi gadadis cilebi VirЕ2, VirЕ3 da VirF.

s ur . 34. s p e cif ikur i tr ans ujr e d ul i g as a s v l e l i

ar xi – T-p ilusi ( T-pilus) ag r o baq te r iis ( Agrobacterium) d a
m ce nar is (Plant cell) ujr e d e bs S o r is

cila VirЕ3 warmoadgens Saperons (damxmare cilas) VirЕ2

cilis mcenaris ujredSi transportirebisaTvis. aseve igi
gadadis birTvSi, ukavSirdeba mcenareuli ujredis tran-
skripciul faqtors pBrp da gavlenas axdens mcenareuli ge-
nebis eqspresiaze (sur. 35).

110
 yvelaze mniSvnelovan funqcias mcenaris ujredis Sig-
niT asrulebs cila VirЕ2. misi molekulebi mTel sigrZeze
ukavSirdeba moumwifebel T-kompleqs, riTac icavs erTja-
Wvian dnm-s mcenareuli endonuklezebiT gaxleCisagan, mii-
Reba momwifebeli T-kompleqsi (Mature T-complex) da VirD2 ci-
lasTan erTad gansazRvravs T-kompleqsis transports uj-
redis citoplazmidan birTvSi.

3. s i msivnis genebis transkripcia

2. 2. g axleCva 4. 3 . p atron-mc. Ggenebis

5. E eqspresiis Secvla

1. 1 .importi birTvSi

6. h i stonebis g enebis transkripcia

7. 4. T-d nm-s
8. i ntegracia

s ur. 35. agrobaqteriuli vir- cilebis aq tiuroba m cenaris ujredSi

VirD2 cila ukavSirdeba or mcenareul cilas, rom-

lebmac miiRo Semoklebuli 51 saxelwodeba, Sesabamisad –
VIP1 da VIP2 (sur. 35). es cilebi mcenareul cila importin-
Tan (importin) urTierTmoqmedebiT uzrunvelyofen T-kompleq-
sis kon-taqts birTvis forebis aparatTan, rac gansaz-
Rvravs T-kompleqsis gadaadgilebas citoplazmidan nukleo-
plazmaSi.

51
sityvidan VirE2-interacting proteins.
111
 VirD2 da VirЕ2 cilebi aseve monawileoben T-dnm-s qromo-
somul dnm-Si integraciis procesSi, oRond mas Semdeg,
rac agrobaqteriidan gadmosuli VirF cila birTvis prote-
osomaSi daSlis (gaxleCs) mcenareul VIP1 cilas da misgan
gaaTavisuflebs T-kompleqss.
birTvSi erTjaWviani dnm, rogorc Cans, aRidgens or-
jaWvian struqturas da homologiuri rekombinaciis meqa-
nizmiT CaSendeba mcenaris genomSi. amisaTvis aucilebelia
2-5 nukleotidis sigrZis mikrohomologia T-dnm-s kidura
Tanmimdebrobasa da mcenaris genomis ubans Soris.
amrigad, mcenareuli ujredis agrobaqteriuli geneti-
kuri transformaciis Zalian rTuli, jer kidev bolomde
daudgeneli procesis ZiriTadi molekuluri meqanizmi ase-
Tia (sur. 36):
1. procesi iwyeba agrobaqteriis mier patron-mcenaris

T-pilus

s ur. 36 . agrobaqteriuli g e netikuri tr ansf ir m aciis m o d e l i

112
(mcenare-recipientis) ujredebis specifikuri receptorebis
(Bacterial and plant receptors) gamocnobiT da agrobaqteriis qro-
mosomis genebis mier kodirebuli cilebis saSualebiT baq-
teriis mimagrebiT mcenaris ujredze;
2. mcenaris ujredis specifikuri sasignalo moleku-
lebis (Plant signals) gamocnoba agrobaqteriis signalis gada-
cemis orkomponentiani (VirА/VirG) regulatoruli sistemis
mier;
3. regulatoruli cila VirG-s mier agrobaqteriis vir-
ubnis (vir-region) virulentobis (vir) genebis aqtivacia;
4. erTjaWviani mobiluri T-dnm-s (T-strand) warmoqmna;
5. moumwifebel T-kompleqsis (Immature T-complex), sxva
ramodenime Vir cilasTan erTad, gadatana patron-mcenaris
ujredis citoplazmaSi;
6. momwifebeli T-kompleqsi (Mature T-complex) warmoqmna
da misi gadaadgileba ujredis birTvisaken;
7. T-kompleqsis aqtiuri importi ujredis birTvSi;
8. T-dnm-s gadatana genomur dnm-Tan integraciis ad-
gilamde;
9. T-dnm-s gaTavisufleba Tanmxlebi cilebisagan;
10. T-dnm-s CaSeneba (integrated T-DNA) mcenaris genomSi
homologiuri rekombinaciis meqanizmiT.

2.3.2.1.2. veqtorebi agrobaqteriis plazmidebis sa-

fuZvelze

veqtorebi Ti-plazmidis safuZvel ze. mcenareTa geneti-

kuri transformaciis unikaluri biologiuri Tviseba Ti-
plazmidas xdis mcenareebSi genebis gadatanis idealur bu-
nebriv veqtorad. mas gaaCnia patronTa farTo wre, SeuZlia
T-dnm-s CaSeneba mcenaris ujredis qromosomaSi, romelic
Semdeg am ujredSi da dayofiT warmoqmnil ujredul Tao-
bebSi stabilurad replicirebs da misi genebi translire-
ben Sesabamisi cilebis warmoqmniT, anu SeuZlia Seqmnas uj-
redSi Seyvanili genetikuri informaciis realizaciis da
rig TaobebSi am informaciis stabilurad SenarCunebis pi-
113
robebi da rac mTavaria T-dnm-s boloebi SemosazRvrulia
marjvena da marcxena 25 n.w. sigrZis kidura ganmeorebebiT,
romelTa SigniT moTavsebuli nebismieri dnm-s Tanmimdevro-
ba, veraviTar gavlenas ver axdens agrobaqteriidan mcenaris
ujredSi T-dnm-s efeqtur gadatanaze, anu am ganmeorebebs
Soris moTavsebuli ucxo dnm-s nebismieri fragmenti warma-
tebiT iqneba gadatanili da integrirebuli mcenaris ujre-
dis dnm-Si.
magram arsebobs rigi seriozuli teqnikuri SezRudve-
bi, ris gamoc Cveulebrivi bunebrivi Ti-plazmidebis veqto-
rad gamoyeneba SeuZlebelia: Ti-plazmidas gaaCnia Zalian
didi zoma (200-800 aTasi n.w.), rac tradiciuli meTodebiT
masSi saWiro genis CaSenebas warmoudgenels52 xdis; T-dnm-s
simsivnewarmomqmneli genebis (eqvsi geni) aqtiuroba Trgu-
navs transformirebuli ujredebis regeneraciis unars,
amitom SeuZlebelia aseTi ujredebidan mcenareTa sruli
regeneracia; T-dnm-s opinebis masinTezirebeli genis arsebo-
ba aqveiTebs saboloo biomasis warmoqmnis unars, vinaidan
mcenaris resursebis nawili daixarjeba maT sinTezze; T-
dnm-Si ar aris seleqtiuri da reportiuli 53 markerebi –
genebi, romelTa aqtiurobiT SesaZlebeli iqneba transfor-
mirebuli ujredebis advili gamorCeva da transformirebu-
li ujredis genomSi Seyvanili dnm-s fragmentis arsebobis
dasabuTeba da sxva.
amitom mimarTaven maT modificirebas genetikur-in-
Jinruli saSualebebiT da transformaciisaTvis sasurveli
veqtorebis xelovnurad Seqmnas (konstruirebas). bunebriv
Ti-plazmidas acileben (Wrian) simsivnewarmoqmnisa da opine-
bis sinTezis makodirebel genebs, sxva veqtoris klonirebi-

52
didi zomebis gamo plazmidas gaaCnia restriqtiuli endonuklea-
zebiT gaxleCis Zalian bevri saiti. rac praqtikulad gamoricxavs mis
srulyofil awyobasa da uSualod masSi ucxo genebis klonirebis SesaZ-
leblobas – restriqtazebiT gaWrisa da Sesabamisi ucxo genis Seyvanis
Semdeg funqcionirebadi srulyofili plazmidis awyobis Sansi Zalian um-
niSvneloa.
53
ix. gv. 142.…
114
saTvis araarsebiT fragmentebs, umateben (miakereben) seleq-
tiur markerebs, replikaciis iniciaciis saitebs, polilin-
kerebs da a.S.
T-dnm-s gadatanaze pasuxismgebeli yvela cila kodi-
rebulia vir-genebiT. xelovnurad konstruirebul veqtoreb-
Si vir-ubani da T-dnm SeiZleba iyos erT da igive, an sxva-
dasxva plazmidaSi.
amrigad, mcenareTa transformaciisaTvis Ti-plazmi-
dis safuZvelze Seqmnili yvela veqtori unda Seicavdes
Semdeg struqturul elements:
- T-dnm-s marjvena da marcxena kidura ganmeorebebs;
- T-dnm-s mcenaris ujredis genomSi gadatanisa da Ca-
SenebisaTvis aucilebel Tanmimdevrobebs (vir-ubani);
- transformirebuli ujredebis gamorCevisaTvis auci-
lebeli seleqtiuri markeris genis Tanmimdevrobebs (anti-
biotikebisadmi, herbicidebisadmi gamZleobis genebi da
sxva);
- mis SemadgenlobaSi, T-dnm-s kidura ganmeorebebs So-
ris ucxo (saWiro) genis CaSenebis gaadvilebisaTvis auci-
lebel restriqciis erT an ramodenime saits (polilinke-
ruli 54 Tanmimdevrobebi);
- CaSenebuli genis eqspresiisaTvis aucilebel promo-
torul Tanmimdevrobebs (nopalin (nos)- da oqtopin (ocs)
sinTetazuri genis promotoruli Tanmimdevrobebi);
- baqteriul ujredebSi (Е. coli da sxva) replikaciisaT-
vis aucilebel replikaciis iniciaciis saits.
dReisaTvis agrobaqteriis plazmidebis safuZvelze,
zemoT aRniSnuli principebis dacviT, konstruirebuli veq-
torebis ori tipi arsebobs – kointegraciuli (Sualedi)
da binaluri.

54
linkerebi - sinTetikuri oligonukleotidebi, romlebic warmoadgenen
restriqciis saitebs garkveuli restriqtazebisaTvis (mag. EcoRI, HihdIII,
BamHI).

115
 koi n tegraciuli (Sualedi) veqtori s gamoyenebis arsi
mdgomareobs SemdegSi (sur. 37). ucxo (saWiro) genis mcena-
Ti-p l azmida

Sualedi
veqtori

T-d nm

ucxo geni

ucxo genis CaSeneba T-dnm-s ubanSi

dasenianeba,
koniugacia

hibriduli
plazmida

homologiuri
rekombinacia

ucxo
geni

T-dnm ucxo geniT

Ti-plazmida CaSenebuli integracia mcenaris
ucxo geniT genomSi

s ur. 37 . kointegraciul i v e q to r is g am o y e ne ba
m ce nar is ujr e d S i g e nis g ad as atanad

ris genomSi Seyvanis mTeli procesi iyofa or etapad: 1)

mcenaris genomSi CasaSenebeli genis klonireba55 ; 2) sakuT-
vriv mcenaris ujredis transformacia. es ori procesi
tardeba sxvadasxva veqtorebis gamoyenebiT. damxmare (Sua-
led) veqtorad, romelSic xdeba ucxo genis klonireba,

55
genis gamoyofis da in vitro mravalricxovani aslebis miRebis procesi.
116
iyeneben E. coli-is safuZvelze Seqmnil veqtorebs (mag.
pBR322). 56

Cla I
EcoR Hind
I III Bam
0
HI
Tcr Sal I
Hin 4
dI
Pvu Apr pBR 322 1
I 4.3 aT. n.w.
Pst I
3
Ava I
ori 2
Bal I

Pvu
II
s ur. 38 . v e q to r pBR 322-is f iziku r i r u q a

jer restriqtazebis saSualebiT Ti-plazmididan amoW-

rian T-dnm-s kidura TanmimdevrobebiT da integrireben 57
damxmare (Sualed) veqtorSi. Semdeg veqtors T-dnm-iT amrav-
leben (E. coli-s gamravlebis gziT) da tradiciuli meTode-
bis (restriqtazebis) saSualebiT masSi SehyavT ucxo geni
da seleqtiuri markeris geni da isev amravleben (E. coli-Si)
ukve Canergili geniT. Semdeg rekombinantuli (hib-riduli)

56
veqtorი pBR 322 (sur. 38) konstruirebulia sami plazmidis (CoI E1, pSC 101
da RI drd) safuZvelze. იგი Seicavs antibiotikebis – tetraciklinisa da am-
picilinisadmi gamZleobis or gens, agreTve ramdenime restriqtazas – Hind
III, Sal I, BamH I, EcoRI, Pvu I, Pst I da sxva Secnobis saits. pirveli sami res-
triqtazas Secnobis saiti mdebareobs tetraciklinis genis (T cr) dnm-Si,
xolo restriqtazebis Pst I, Pvu I – ampicilinis genis (Ap r) dnm-Si. es saSua-
lebas iZleva Catardes transformantebis gamorCeva erTi antibiotiki-
sadmi gamZleobis mixedviT, xolo klonebis rekombinantuli dnm-iT – me-
ore antibiotikisadmi gamZleobis mixedviT. veqtoris sigrZe Seadgens 4,3
aTas n.w.
57 damxmare veqtorSi aseve SehyavT seleqtiuri geni, transformirebuli

mcenareebis gamorCevisaTvis.
117
veqtoruli plazmida koniugaciis gziT (E. coli-iT daseniane-
bis Sedegad) SehyavT A. tumefacienns-is ujredSi, romelsac
Tavisi vir-ubnis, T-dnm-s kidura Tanmimdevrobebis da Suale-
di veqtoris analogiuri Tanmimdevrobebis mqone Ti-plazmi-
da gaaCnia. baqteriis ujredSi or plazmidas T-dnm-s homo-
logiur Tanmimdevrobebs Soris ormagi krosingoveris (ho-
mologiuri rekombinaciis) Sedegad rekombinantuli (hibri-
duli) plazmidas – Sualedi veqtoris nawili ucxo geniT
da seleqtiuri markeris geniT CaerTveba baqteriis Ti-plaz-
midas T-dnm-s adgilas, anu miiReba kointegraciuli veqto-
ri - Ti-plazmida CaSenebuli ucxo geniT (sur. 37). bolos,
agrobaqteriebs aseTi kointegraciuli veqtoriT gamoarCe-
ven seleqtiur areze da maTTvis damaxasiaTebeli Cveuleb-
rivi dasenianebis meTodiT iyeneben mcenareTa tra-nsforma-
ciisaTvis.
bi naluri veqtori ucxo dnm-s mcenaris genomSi Seyva-
nis ufro martivi da amitom ufro xSirad gamoyenebuli me-
Todia. rogorc aRniSnuli iyo, agrobaqteriebiT mcenaris
ujredis dasenianebisa da transformaciisaTvis aucilebe-
lia T-dnm-s gadatanaze pasuxismgebeli vir-ubani da T-dnm-s
kidura Tanmimdevrobebi. dadginda, rom procesis warmate-
biT ganxorcielebisaTvis sruliad araa aucilebeli es mo-
nakveTebi imyofebodnen erT da igive plazmidaSi.
binaluri veqtorebis sistemis (sur. 39) Seqmna dafuZne-
bulia imaze, rom mcenaris transformaciisaTvis gamoyene-
bul agrobaqteriul ujredSi erTdroulad imyofeba ori
plazmida. erTi Seicavs T-dnm-s kidura Tanmimdevrobebs, xo-
lo meore – vir-ubans. orive plazmidas SeuZlia damoukide-
beli replikacia58 agrobaqteriis ujredSi, magram cal-
calke ver iwveven mcenaris transformacias. kointegraci-
uli veqtoris msgavsad ucxo geni da seleqtiuri markeris

58
plazmidas T -dnm-s kidura TanmimdevrobebiT aseve gaaCnia E. coli-Si rep-
likaciis unari, rac ucxo genis am baqteriis ujredSi klonirebis saSu-
alebas iZleva.
118
 T-DNA
vector

s ur. 39 . agrobaqteria o r i p lazmidiT

geni SehyavT T-dnm-Si da Semdeg aseTi rekombinantuli pla-
zmida E. coli-is ujredSi koniugaciis gziT SehyavT agrobaq-
teriis ujredSi, romelsac ukve aqvs meore plazmida vir-ub-
niT. kointegraciuli veqtorisagan gansxvavebiT am or pla-
zmidas Soris homologiuri rekombinacia da erT veq-to-
rul molekulad gaerTianeba ar xdeba. aseTi binaluri veq-
toriT mcenaris ujredis dasenianebisas erTi plazmidis
vir-genebis mier eqspresirebuli cilebi amoWrian da mcena-
ris ujredis genomSi CaaSeneben meore plazmidis T-dnm-s
ucxo geniT.
veqtorebi Ri-plazmi di s safuZvel ze. kidev erT mcena-
ris transformirebis unaris mqone plazmidas warmoadgens
„wverisebri fesvis“ mainducirebeli Agrobacterium rhizogenes-is
Ri-plazmida (sur. 40). transformirebuli fesvebis induqci-
is meqanizmi gvirgvinisebri galebis warmoqmnis meqanizmis
analogiuria: virulentur plazmidas gaaCnia Ti-plazmidas-
Tan homologiis didi ubnebi, aqvs Tavisi T-dnm, romelic ga-
daitaneba da integrirdeba mcenare-patronis genomSi. igi
iwvevs gaZlierebul fesvwarmoqmnas da Ti-plazmidis msgav-
sad, opinebis sinTezs. Ri-plazmida warmoadgens bunebriv
uvnebel araonkogenur veqtors da mcenaris ujredebis tra-
nsformaciis Semdeg, jansaRi cxovelmyofeli mcenareebis
regeneraciis unari aqvs. amJamad Ri-plazmidis safuZvelze

119
 s ur. 40. Ri-p lazmida m niSvnelovani g e nebiTa d a ubnebiT

aseve miRebulia mravali veqtori mcenareTa genetikuri in-

JineriisaTvis.

2.3.2.2. veq torebi fitovirusebis safuZvelze

mcenareebis mainficirebeli fitovirusebidan mxolod

1-2% miekuTvneba dnm-Semcvel virusebs. mxolod isini war-
moadgens rekombinantuli dnm-s teqnologiebSi gamoyenebis
yvelaze xelsayrel obieqts da amitom ganixilebian mcena-
reebSi genebis gadatanis veqtorebis Seqmnis ZiriTad kandi-
datebad.
am mxriv yvelaze perspeqtiuls warmoadgens jvaro-
sanTa ojaxis mcenareTa daavadebis gamomwvevi kombostos
mozaikis virusi CaMV59 (sur. 41). am virusis nawilakebis60
diametri daaxloebiT 50 nm-ia da Seicaven 8 aTasi n.w. sig-
rZis rgolur dnm-s. mis SemadgenlobaSi gamoyofen genebis
7 jgufs: I-akodirebs cilas, romelic uzrunvelyofs viru-
sis nawilakebis gadatanas erTi ujredidan meoreSi

59
inglisur enaze misi saxelwodebidan – cauliflower mosaic virus.
60
sanam virusi imyofeba ujredis gareT, an ujredis dasenianebis proces-
Si, is arsebobs damoukidebeli nawilakis saxiT.
120
 s ur. 41. ko mbostos m ozaikis v ir us is g e no m is s q e m a

plazmodesmebis gziT; II- da III-akodireben virionebis erTi

mcenaridan meoreSi gadatanaze pasuxismgebel P2 da P3 ci-
lebs; IV-akodirebs kapsiduri cilebis winamorbedebs; V-
akodirebs proteinazebis, ukutranskriptazasa da rnmaza H-
is winamorbedebs; VI-akodirebs CarTvis cilas – aucile-
bels inficirebuli mcenaris ujredSi genomuri dnm-s sin-
Tezisa da virusis nawilakebis awyobisaTvis; VII- funqciebi
daudgenelia. genetikuri inJineriisaTvis gansakuTrebiT sa-
interesoa ori genTaSorisi ubani (sqemaze Sav ferSi), rom-
lebSic imyofeba 19S RNA (mcire ubani ~700 n.w. sigrZis) da
35S-RNA (didi ubani ~1500 n.w. sigrZis) promotorebi.
CaMV-s genomis mcire zoma, virusul dnm-Tan, rogorc
baqteriul plazmidasTan in vitro manipulirebis da Semdeg
misi foTolSi Sezelvis gziT mcenareSi Seyvanis saSuale-
bas iZleva. mcire raodenobis ujredebis inficireba mTeli
mcenaris dasenianebas iwvevs, radganac virusi swrafad
vrceldeba ujredidan ujredSi gadacemis gziT. erTi mce-
naris inficirebisaTvis 1-5 mkg dnm-s gamoyeneba, infeqciis
TiTqmis 100%-ian efeqturobas iwvevs, rac gacilebiT maRa-
lia, vidre agrobaqteriuli dasenianebis SemTxvevaSi.
CaMV-s dnm-s E. coli-Si klonirebisaTvis E. coli-is plazmi-
dur veqtorebs, mag., pBR 322-s. CaMV-s da pBR 322-s dnm-s Se-
121
sabamisi restriqtazebiT damuSavebisa da ligirebis Semdeg
warmoiqmneba E. coli-Si amplifikaciis unaris mqone hibridu-
li plazmida.
infeqciis Semdeg patronis genomSi CaMV-s dnm-s pir-
dapiri integracia ar dafiqsirebula. magram SemuSavebulia
agroinfeqciis meTodi, romelic inokulaciis Semdeg viru-
suli dnm-s pirdapiri CaSenebis saSualebas iZleva. amisaT-
vis CaMV-s genomi CaSendeba T-dnm-Si da misi saSualebiT
integrirdeba mcenaris birTvul genomSi. amasTan CaMV-s
Semadgenlobidan winaswar amoiWreba misi virulentobis
ganmsazRvreli Tanmimdevrobebi.
ZaRlyurZenasebrTa ojaxis mcenareebis genetikur inJi-
neriaSi iyeneben Tambaqos mozaikis virusis safuZvelze Seq-
mnil veqtorebs, parkosanTa ojaxis mcenareebSi – bardis
mozaikis virusis safuZvelze Seqmnil veqtorebs. ukanas-
knel aTwleulSi veqtorebad gamocdil iqna kartofilis
X-virusi, bambukis mozaikis virusi, papaias mozaikis virusi,
kitris mozaikis virusi da sxva.
virusebis safuZvelze Seqmnili veqtorebis upirateso-
bas warmoadgens: genomis mcire zoma, rac virusuli dnm-Tan
advili manipulirebis saSualebas iZleva; dasenianebuli
mcenaris ujredSi virusuli dnm-s maRali kopireba (50 aTa-
samde ujredSi); Zlieri promotoris arseboba, rac ucxo
dnm-s efeqtur eqspresias uzrunvelyofs.
virusuli veqtorebis uaryofiT mxareebidan yvelaze
mniSvnelovania veqtoris mcire moculoba (800 n.w.-ze nakle-
bi) da patronTa SezRuduli wre – jvarosanTa ojaxi. veq-
toris moculoba SeiZleba gaizados 800-1000 n.w.-iT Tu vi-
rusiT davainficirebT mcenaris protoplastebs. am dros
infeqcia ar gadaecema ujredidan ujredSi da araa aucile-
beli dnm-s SefuTva nawilakebSi. amitom virusul nawila-
kebSi SefuTvaze pasuximgebeli virusuli genomis nawili
SeiZleba mocilebul da Canacvlebuli iqnas ucxo geniT.
virusebis safuZvelze Seqmnili veqtorebi dReisaTvis
naklebad gamoiyeneba mcenareTa transformaciisaTvis, Tum-
ca virusuli promotorebi, da pirvel rigSi promotori
122
35S-RNA CaMV, farTod gamoiyeneba ucxo genis eqspresiisaT-
vis sxva veqtorul sistemebSi. igi Zlieri promotoria,
garda amisa aqtiuria ara marto jvarosanTa, aramed sxva
ojaxis mcenareTa ujredebSi, ar avlens qsovilspecifiku-
robas da eqspresirdeba transformirebuli mcenaris yvela
ujredSi.
veqtorebi SeiZleba Seiqmnas viroidebis safuZvelzec.
isini iwveven mcenareTa zogierTi saxeobis daavadebebs da
warmoadgenen 246 dan 399 nukleotidis sigrZis rnm-s mcire
zomis, kovalenturad Sekrul wriul molekulas. sakmaod
farToa maTi patronebis wre – farTod gavrcelebuli sa-
soflo-sameurnei kulturebi (kartofili, pomidori, badri-
jani, vaSli, msxali, vazi da sxva), tropikuli da subtro-
pikuli kulturebi – avokado, citrusebi, qoqosis palma,
atami, dekoraciuli mcenareebi.

2.3.2.3. veqtorebi mcenaris qloroplasturi da

mიtoqondriuli dnm-s gamoyenebiT

qloroplastis genomi Sedgeba saSualod 120-220 aTasi

n.w. sigrZis dnm-s rgoluri, kovalenturad Sekruli mole-
kulisagan. rac Seesabameba daaxloebiT 150 cilas. 61 erTi
saxeobis qloroplastis SigniT ramodenime aTeuli aseTi
wriuli molekulaa. ujredi Seicavs ramodenime aTeul
qloroplasts. amdenad, ujredSi rgoluri molekulebis
saerTo ricxvi ramodenime aseuls Seadgens. rgoluri mo-
lekulis SemadgenlobaSi Sedis ribosomuli da satran-
sporti rnm-s genebi, agreTve genebi, romelTa produqtebi
aucilebelia qloroplastebis normaluri funqcionirebi-
saTvis.
qloroplasturi genomis replikacia da transkripcia
mimdinareobs avtonomiurad, rac qloroplastis dnm-s veq-

61.qloroplastebSi gacilebiT meti cilis arseboba aixsn eba imiT, rom

maTi umetesi nawili kodirdeba birTvuli da ara plastiduri genomiT.
123
torad gamoyenebis saSualebas iZleva. Tu baqteriul plaz-
midas „mivakerebT“ 62 replikaciis dawyebis ubnis da tran-
skripciis iniciaciisa da terminaciis Semcvel qloroplas-
tis dnm-s fragmentTan, maSin SeiZleba miviRoT binaluri
veqtori. aseT veqtors prokariotul ujredSi replikaciis,
xolo eukariotul ujredSi ara marto replikaciis, ara-
med ucxo genetikuri informaciis eqspresiis unaric gaaC-
nia. aseTi veqtorebi funqcionireben rogorc episomebi, anu
qromosomisagan avtonomiurad.
transplastomuri mcenareebi, anu plastidur genomSi
ucxo genCaSenebuli mcenareebi, am gens inarCuneben rigi
Taobebis ganmavlobaSi da eqspresirebuli niSani TesliT
gamravlebisas gadaecema dedis xaziT, anu ar gadaecema
mtvrianebiT.
binaluri veqtorebis konstruirebisaTvis SeiZleba
gamoyenebul iqnas agreTve mitoqondriuli dnm, romelTa
sigrZe varirebs did farglebSi – 200-dan 2400 aTas n.w.-mde.

2.3.2.4. veqtorebi mo biluri elementebis

(transpozonebis) safuZvelze

ukanasknel wlebSi organizmebSi aRmoCenil iqna geneti-

kuri elementebi, romlebic iwveven genomis arastabiluro-
bas, monawileoben genebis aqtiurobis regulaciaSi, ujre-
debs aniWeben sxva damatebiT Tvisebebs. am elementebis mTa-
var ganmasxvavebel Taviseburebas warmoadgens maTi genomis
SigniT, an sxvadasxva genomebs Soris damoukidebeli gada-
adgilebis unari. maT ar SeuZliaT damoukidebeli replika-
cia – amas axdenen mxolod im replikonis farglebSi, ro-
melSic isini CaerTvebian. aseT genetikur elementebs tran-
spozonebi ewodeba.

62plastidis genomSi ucxo genis Seyvanis strategia mdgomareobs Sesayvan

dnm-sa da plastidis rgoluri molekulis romelime ubans Soris h o-
mologiur rekombinaciaSi.
124
 transpozonebi – aris damoukidebeli Sigaqromosomu-
li da qromosomaTaSorisi gadaadgilebis63 unaris mqone
dnm-s segmentebi. amitom maT kidev uwodeben moZrav, mo-
bilur anu transpozirebad genetikur elementebs.
dnm-s erTi saitidan meoreSi transpozicias anu gada-
adgilebas isini TviTon akontroleben sawyisi saitidan
amoWrisa da axal saitSi CaSenebis gziT.
dnm-ze gadaadgilebiT transpozonebs gadaaqvT genebi
(moTavsebuli or transpozons Soris) axal saitebSi. tran-
spozonebis gadaadgilebis specifikuri rekombinantuli me-
qanizmebi ganpirobebulia specialuri genebisa da transpo-
zonebis boloebze pirdapiri da Sebrunebuli, anu inverti-
rebuli ganmeorebebis arsebobiT.
transpozonebis amoWrisa da CaSenebis molekuluri me-
qanizmi bolomde araa Seswavlili. dnm-s gadatana xdeba
ferment transpozazas saSualebiT, romelic kodirdeba
transpozonis SuaSi ganlacebuli ~20 nukleotidiT. igi
specifikurad urTierTmoqmedebs transpozonis bolo inver-
tirebul ganmeorebebTan da SeuZlia misi amoWra. erT-erTi
modelis mixedviT (sur. 42) transpozonebi transpoziciis
(CaSenebis) procesSi replicirdebian da recipientul qro-
mosomas gadascemen replikas (replikaciis Sedegad warmo-
qmnil asls). es xdeba xdeba Semdegnairad: Sesabamis saitSi
specifikuri endonukleaza Wris dnm-s komplementur jaWvs,
warmoqmnis „mwebvare“ boloebs da transpozoni CaSendeba
„mwebvare“ boloebs Soris, xolo masTan mierTebuli dnm-s
erTjaWviani molekulebi dnm-polimerazas saSualebiT da-
Sendebian orjaWvianamde. transpoziciis meqanizms safuZ-
vlad udevs sait-specifikuri rekombinacia.
mcenareTa ujredebis transformaciisaTvis transpozo-

63
“transpozonebi gadaadgildebian” mTlad zusti gamoTqma ar aris, rad-
ganac transpozonis axal lokalizaciaSi gamovlenisas, pirvelsawyis
mdgomareobaSi is ar qreba. mSobliuri transpozoni sawyiss aZlevs Tavis
axal asls, romelic organizmis qromosomis axal adgilSi vlindeba.

125
 recipientuli
qromosoma
safexurisebri “gaWra”
qromosomaSi

transpozoni

s ur . 42 . tr ans p o zo nis Ca S e ne bis m o d e l i

nebis safuZvelze Seqmnili veqtorebis gamoyeneba daiwyes

gasuli saukunis 90-ian wlebSi. yvelaze farTod gamoiyene-
ba Ac da Ds veqtorebis sistema.
Ac (Activator) – mcenareTa mobiluri elementia, romelic
simindis genomis specifikur ubnebSi iwvevs qromosomul
wyvetas. igi miekuTvneba genomSi aqtiuri transpoziciis
unaris mqone mobilur elementebs. simindis garda aqtiuria
sxva mcenareebis – kartofili, pomidori, stafilo, arabi-
dopsisi, brinji da sxva genomSic.
genomSi aqtiuri transpoziciis transpozonebis ojaxs
miekuTvneba agreTve Ds-elementi, romelic Tavisi pirveladi
struqturiT Ac-elementis msgavsia, magram Sida raionebis
deleciis gamo genoSi damoukidebeli gadaadgilebis unari
ara aqvs. transpozicia SeuZlia mxolod specifikuri ci-
lis - Ac-elementis transpozazas monawileobiT.

2.3.3. veqtorebis saSualebiT genebis gadatanis

meTodebi

„ foTli s rgolebi s“ meTod i . SedarebiT martivi, mo-

xerxebuli da sakmaod xSirad gamoyenebuli meTodia. misi
arsi mdgomareobs imaSi, rom steriluri foTlebidan Wrian
rgolebs, aTavseben myar an Txevad sakveb areze da aseniane-
126
ben agrobaqteriiT (e.w. Ramis kulturiT) da erTad kulti-
vireben.
agrobaqteria xvdeba Wrilobis adgilas da uerTdeba
ujreds. Wrilobamiyenebul mcenaris fragmentTan kontaqti
acetosiringonis induqciiT “CarTavs” agrobaqteriis vir- da
Т-ubnebs. ~ 4-5 saaTis Semdeg daiwyeba Ti-plazmidis Т-dnm-s
gadatana foTlis rgolebis ujredebSi da damTavrdeba baq-
teriis mcenaris ujredze moxvedridan ~ 8-9 saaTis Semdeg64.
inkubaciis Semdeg rgolebidan baqteriebi Camoirecxeba mi-
krobebis elimenaciis mizniT antibiotik damatebuli ste-
riluri Txevadi sakvebi ariT an wyliT da filtris qa-
Raldze odnav SeSrobis Semdeg Tavseba seleqtiuri agen-
tis Semcvel regeneraciis sakveb areze. daaxloebiT erTi
Tvis Semdeg warmoqmnili regenerantebi gadaaqvT uhormono
sakveb areze. bolos dafesvianebul regenerantebs utarde-
ba Sesabamisi testirebebi. meTodi sxvadasxva modifikaciiT
warmatebiT gamoiyeneba sxvadasxva mcenareebis genetikuri
transformaciisaTvis.
agrobaqteriasTan mcen ari s protoplastebi s k o-
k ultivaciis meTod i . simsivnis warmoqmnis procesSi agro-
baqterias gaaCnia WrilobiT miyenebuli dazianebis sapasu-
xo aqtiuri dayofis fazaSi myofi ujredebis transforma-
ciis unari. intaqturi mcenaris ujredebi, rogorc wesi, ag-
robaqteriis infeqciaze maqsimalur onkogenur unars dazi-
anebidan pirvel – mesame dRis periodSi avlenen. sxvadasxva
saxeobebs es “kompetenturi” faza emTxveva Wrilobis sapa-
suxo pirvel ujredul dayofas. Wrilobis sapasuxo reaq-
cia SeiZleba imitirebul iqnas ujredebis kulturaSi, TuF-
foTlis mezofilis mosvenebis fazaSi an suspenziuri kul-
turis aqtiur dayofis fazaSi myofi ujredebidan miviRebT
protoplastebs da Sesabamis sakveb areSi movaxdenT ujre-
dis kedlis regeneraciis da ujredis dayofis inducirebas.

64 kultivirebis vadebi da pirobebi kulturebis mixedviT sxvadasxvaa.

127
protoplastebi kultivirebis garkveul fazaSi kompetentu-
ria agrobaqteriiT dasenianebisaTvis.
meTodis mixedviT or-sam dRiani ujredis garsis rege-
nerireba da dayofa dawyebuli protoplastebi gadaaqvT
kultivirebis sakveb areSi da umateben agrobaqteriis axa-
li Ramis kulturas. erToblivi kokultivireba grZeldeba
36-48 saaTis ganmavlobaSi. am periodSi baqteriis ujredebi
uerTdeba mcenaris protoplastebs da adgili aqvs T-dnm-s
mcenaris genomSi gadatanisa da integraciis process. ko-
kultivirebis dasrulebis Semdeg protoplastebis kultu-
ras umateben fermentul xsnars protoplastebis gamosayo-
fad. Semdeg protoplastebs orjer recxaven Sesabamisi
xsnariT da Semdgomi kultivirebis mizniT gadaaqvT axal
sakveb arSi. protoplastebis kultivireba xdeba standar-
tuli meTodkiT sami-oTxi kviris ganmavlobaSi. Semdeg mi-
krokoloniebi gadaaqvT agarizebul sakveb areze, bolos ki
- regeneraciul sakveb areze. sul bolos miRebul regene-
rantebs utardeba Sesabamisi testirebebi. meTods gaaCnia
garkveuli SezRudvebi gamoyenebisas, vinaidan jer erTi,
yvela mcenareSi ver xerxdeba protoplastebis gamoyofa da
meore, mravali mcenaris protoplastebs Znelad gadaaqvT
erToblivi kokultivacia baqteriasTan da iRupebian.
in vitro eq splantatebis i n okulaciis meTod i . nakle-
bad gamoyenebuli meTodia. am dros adgili aqvs mcenaris
qsoviluri eqsplantatebis (mag. stafilos rgoli, Tambaqos
Reros nawili, arabidopsisis axalgazrda fesvebi, karto-
filis muxlSorisi, pomidvris hipokotili) erTobliv kul-
tivirebas sterilur pirobebSi agrobaqteriebis suspenzieb-
Tan. transplantaciis Semdeg eqsplantatebs acileben baq-
teriebs da kultivireben kalusis warmoqmnis xelSemwyob
sakveb areze. bolos kalusidan xdeba mcenareebis regeneri-
reba da miRebuli regenerantebis testireba.
in planta tran sformaci i s meTod i . veqtorebis saSua-
lebiT genebis gadatanis meTodebi pirobiTad SeiZleba dai-
yos in vitro meTodebad, romlebic iTvaliswinebs mcenaris

128
ujredebisa da qsovilebis kultivirebas da Semdgom mcena-
ris regeneracias da in planta meTodebad, sadac es stadiebi
ar aris. am dros transformirebuli mcenareebi miiReba in-
taqturi mcenaris ujredebis transformaciis gziT in planta
pirobebSi, kultivirebis areSi mcenaris regeneraciis sta-
diis gareSe.
mcenaris ujredebis in planta pirobebSi transformacia
gulisxmobs intaqturi mcenris qsovilebze (Cveulebriv fo-
Tlis an Reros) winaswar Wrilobis miyenebas an Reros de-
kapitacias (wveros moWras) da Semdeg mis dasenianebas ag-
robaqteriis Sesabamisi kulturiT, sayvavile kvirtebis CaS-
vebas agrobaqteriis kulturis suspenziaSi, naskvze agro-
baqteriis kulturos gadatanas, Teslebis inkubacias ag-
robaqteriis suspenziaSi da Semdgom aRmocenebis xelSewyo-
bas, yvavilebis vacuum-infiltracias suspenziiT, generati-
uli organoebis damuSaveba agrobaqteriis suspenziiT uSu-
alod damtverva-ganayofierebis procesis dros, mtvrianas
damuSavebas da sxva.
Yyvela zemoT ganxiluli meTodis (xSirad maT agro-
infeqciis meTods uwodeben) dros gamoiyeneba agrobaqteri-
is Stami, romelsac gaaCnia konkretuli transformacii-
saTvis misaRebi veqtoruli konstruqcia (binaluri, koin-
tegraculi an sxva). veqtori unda Secavdes funqcionalur
ucxo gens, romlis SeyvanisTvisac tardeba samuSao. genis
warmoSobas (prokariotuli Tu eukarioruli) transforma-
ciisaTvis arsebiTi mniSvneloba ara aqvs, magram is auci-
leblad unda iyos mcenaris ujredSi eqspresirebis unaris
mqone promotoris kontrolis qveS. funqcionaluri genis
garda veqtori unda Seicavdes markerul (seleqtiuri an
reportiori) gens, romlis saxiT Cveulebriv iyeneben anti-
biotikebis – kanamicinis, higromicinis da sxva, herbicide-
bis – qlorsulforonis, fosfinotricinis da sxva mimarT
gamZleobis, luciferazas da sxva genebs.
amJamad, meTodebis srulyofis Sedegad, moxerxda ag-
robaqteriis bunebrivi StamebisaTvis damaxasiaTebeli erT-
lebniani mcenareebiis inficirebis uunarobis daZleva da
129
SesaZlebelia mcenareTa TiTqmis yvela saxeobis warmatebu-
li transformireba. miuxedavad amisa, axali veqtoruli
sistemebis gamoyenebis SesaZleblobebis Ziebis procesi ma-
inc grZeldeba.
43-e suraTze warmodgenilia agrobaqteriuli transfor-
maciis gziT transgenuri mcenareebis miRebis zogadi sqema.

2.4. genebis pirdapiri gadatanis meTodebi

mcenaris ujredebSi genebis pirdapiri gadatanis meTo-

debis arsi ucxo genis matarebeli izolirebuli „SiSveli“
dnm-s sxvadasxva xerxebiT mcenaris protoplastebSi 65 Sey-
vanaSi mdgomareobs. am dros SeiZleba gamoyenebul iqnas ne-
bismieri ucxo dnm matarebeli veqtori, radganac ar aris
aucilebeli mas hqondes transformaciis specialuri bio-
logiuri signalebi da funqciebi (T-dnm-s kidura Tanmimdev-
robebi, vir-ubani). pirdapiri meTodebiT gadatanili ucxo
geniM(mas aucileblad unda hqondes eukariotuli promoto-
ri) ujreduli rekombinaciis meqanizmiT integrirdeba mce-
naris birTvis dnm-Si da iwyebs eqspresias. maT ZiriTad ua-
ryofiT mxares warmoadgens transformaciis dabali sixSi-
re.
Tavisufali STanTqmis meTodi an u mce n ari s pro -
toplastebis d onorul d n m-Tan k ok ulti vaci a msgavsia
adre cxovelebisaTvis damuSavebuli analogiuri meTodis
da mdgomareobs mcenaris protoplastebze garkveuli kon-
centraciis (Cveulebriv 2-106 uj/ml) izolirebuli tran-
sformirebadi dnm-s (maT Soris agrobaqteriis, sxva veqto-
ruli plazmidebi), polieTilenglikolis (40%-iani xsnari;
mol. masa. 1500) da matareblis saxiT Timusis dnm-s damate-
baSi. polieTilenglikoli xels uwyobs dnm-s protoplaste

65 garda vakuum-infiltraciis da meqanikuri inokulaciis meTodebisa.

130
 Agrobacterium
tumef aciens T-dnm
m c enare
Ti-p l azmida

q r omosoma

k o k ultivacia
mc enaris
u jredi

vir-u b ani

t r ansformacia
t r ansformirebuli
u jredebi

f o T l eb i s mo T avs eb a s el eqt i u r ar ez e.
.ar at r an s f o r mi r eb u l i u jr ed eb i s d aR u p va

t ransgenuri mcenaris r egeneracia

mc enareTa gamravleba

molekulur-biologiuri analizi

savele gamocda

s ur. 43. tr ansgenuri m cenareebis m iRebis s qema

131
bTan dakavSirebas, romlebic Semdeg pinocitoziT66 (sur.
44) STanTqaven dnm-s ujredis SigniT. aseT areSi protopla-

p l az mat u r i memb r an a

ci toplazma

s ur . 44. p ino cito zi m ce nar is ujr e d S i

stebs inkubireben 30 wT-is ganmavlobaSi 26°C temperatura-

ze, Semdeg TandaTanobiT acileben polieTilenglikols da
kultivireben sakveb areze jer sami dRis ganmavlobaSi sib-
neleSi 25°C temperaturaze, xolo Semdeg sinaTleze. erTi
kviris Semdeg ujredebis mikrokoloniebs seleqtiur areze
gadargaven da iwyeben transformirebuli kalusebis, xolo
Semdeg mcenareebis gamorCevas.
protoplastebSi d n m-s mikroineqcii s meTod i . mik-
roineqciis teqnika pirvelad damuSavebuli iqna cxovelur
ujredebSi dnm-s SeyvanisaTvis. amJamad mas warmatebiT iye-
neben ucxo dnm-s mcenaris protoplastebSi SeyvanisaTvis.
meTodi dafuZnebulia mikronemsebis gamoyenebaze, romelTa
saSualebiT dnm-s molekula pirdapir gadaitaneba protop-
lastis citoplazmaSi an birTvSi. igi potenciurad warmo-
adgens umaRles mcenareebSi ucxo genebis Seyvanis yvela-
ze universalur saSualebas, radganac recipientad SeiZle-
ba gamoyenebuli iqnas obieqtebis farTo wre – rogorc
calkeuli kulturaluri ujredebi, ise mtvris, embrionebis
da sxvadasxva qsovilebis somaturi ujredebic. mcenaris

66
pinocitozi (berZn. pino - vsvam , Seviwov da cito – ujredi) – aris ujredis
mier siTxis mitaceba da STanTqma.

132
protoplastebis mikroineqciis ganxorcielebisaTvis Cveu-
lebriv iyeneben „eleqtroduli milis“ mikronemsebs gare
diametriT 2 mkm da Siga diametriT 1-1,25 mkm. mikroineqciis
proceduris (sur. 45) mixedviT izolirebuli proplaste-

protoplastebis
suspenziis wveTi
Ddnm-s wveTi

protoplaste-
bis suspenzia

mikronemsi

dafiqsirebuli
mowyobiloba
ineqcia

protoplasti dnm

kalusis gadatana
myar seleqtiur
areze

ineqtirebuli protoplastis
kultivireba dakidul wveTSi

s ur. 45. m cenaris p r o to p l as te bis m ik r o ine q ciis

p r o ce d ur a
bis kultivirebis me-2-5 dRes minis mikronemsiT hidravli-
kur sistemiani mikromanipulatoris saSualebiT protopo-
lastis citoplazmaSi an ukeTes SemTxvevaSi uSualod bir-
TvSi (dafiqsirebuli specialuri mowyobilobiT) SehyavT
0,1-1,0 mkl izolirebuli dnm-s (mag. izolirebuli veqtoru-
li plazmidebi) Semcveli buferuli xsnari (ineqciis mocu-
133
loba yovel protoplastSi 108-109 ml). protoplastebi ineq-
ciamde da ineqciis Semdeg mikrokoloniebis miRebamde kul-
tivirdeba sinaTleze 3-4 kviris ganmavlobSi saerTod miRe-
buli meTodikis Tanaxmad. Semdeg mikrokoloniebi sididiT
0,5-1,0 mm gadaaqvT seleqtiur areebze da axdenen transfor-
mireuli mcenareebis gamorCevas. transformaciis efeqturo-
ba Seadgens 10-15 %-s. meTodi misaRebia gamoyenebisaTvis ro-
gorc orlebnian, ise erTlebnian mcenareebSi. mis naklovan
mxares miekuTvneba procesis maRali Sromatevadoba da sak-
maod rTuli, ZviradRirebuli mowyobilobebis aucileb-
loba.
mcen ari s protoplastebi s eleq troforaci i s
(eleq troSok i s) meTod i . es meTodi dafuZnebulia imaze,
rom maRali Zabvis eleqtruli impulsebi Seqcevadad zrdi-
an biomembranebis SeRwevadobas, romlis drosac SesaZle-
belia ucxo genis moxvedra mcenaris ujredSi (sur.46). igi

d en is wyaro

ujr edi

protoplasti

d n m ucxo geniT d n m ujredis ujr edi ucxo

S i g niT g e niT

s ur. 46. p rotoplaste bis e l e q tr o f o r aciis zo g ad i

sqema
mikroineqciis meTodis msgavsad aseve pirvelad damuSave-
bul iqna cxovelTa ujredebisaTvis. aRmoCnda, rom eleq-
troporaciis procedura Zalian efeqturia mcenaris pro-
toplastebisTvisac. protoplstebisa da izolirebuli uc-

134
xo dnm-is (mag. veqtoruli plazmidebis) narevi gadaaqvT
eleqtroforaciis kameraSi da atareben maRali Zabvis den-
is mokle impulsebs (Cveulebriv sami impulsi 10 wm-is in-
tervaliT, Zabva 200-350 volti, xangrZlivoba 1-30 wami). de-
nis gavlis periodSi protoplastebis membranaSi warmoiq-
mneba xvrelebi (forebi), romelTa gziT eleqtruli Sokis
procesSi an mis Semdeg dnm-s molekulebi SeaRweven ujre-
dis SigniT. Semdeg protoplastebi gadaaqvT Sesabamis sak-
veb areSi kultivirebisaTvis. eleqtroforaciis efeqturo-
ba dgindeba eleqtroSokidan 24-48 saaTis Semdeg. gadarCeni-
li ujredebi qmnian mikrokoloniebs, romlebic 10-14 dRis
Semdeg gadaaqvT seleqtiur areebze da arCeven transformi-
rebul mcenareebs.
d nm-s li posomebSi C awyobis meTod i . am meTodis arsi
mdgomareobs mcenaris ujredSi Sesayvani ucxo dnm-s inkaf-
sulirebaSi loposomaSi – sferosebur warmonaqmnSi, rom-
lis garsi Sedgeba fosofolipidebisagan (sur. 47). misi mi-

f o sfolipidis
l i piduri mo l ekula
b i Sre

s ur. 47. liposoma

Reba SeiZleba fosfolipidebis wyliani emulsiis mkveTri

SenjRrevis an ultrabgeriT damuSavebis Sedegad. aseTi
liposomebi SehyavT protoplastebSi liposomebisa da pro-
toplastebis narevis polieTilenglikoliT damuSavebis
gziT. es ukanaskneli xels uwyobs liposomebis protop-
lastebTan dakavSirebas, romlebic Semdeg pinocitoziT
STanTqaven liposomebs da maT SigTavss.
mcenareebSi genebis Seyvanis axali sistemis upirateso-
bas warmoadgens: liposomebis dabali toqsikuroba mcena-
135
ris ujredebisaTvis; sxvadasxva saxeobis mcenareebSi misi
gamoyenebis SesaZlebloba; liposomebSi Cawyobili nuklei-
nis mJavebis dacva sxvadasxva fermentebis (nukleazebis)
damSleli moqmedebisagan.
amJamad transformaciis aRniSnuli meTodi naklebad
gamoiyeneba teqnikuri sirTuleebisa da dabali transfor-
maciuli aqtiurobis (0.5–1%) gamo.
vak uum-infiltraciis meTod i . aRniSnuli meTodi da-
muSavebul iqna arc Tu ise didi xnis winaT da transfor-
maciisaTvis gamoiyeneba rogorc Reroebi, foTlebi, ise
Teslebi. meTodi dafuZnebulia imaze, rom axalgazrda mce-
naris kenwruli nawilebi an winaswar damuSavebuli (nawi-
lobriv daSlili perikarpiumiT) Teslebi Tavsdeba ucxo
genis matarebeli agrobaqteriis ujredebis xsnarSi (sur.
48). WurWels xsnariTa da eqsplantatiT aTavseben kameraSi

s ur. 48. f o Tlebis d a Te slebis v ak uum - inf il tr acia

ag r o baq te r iiT
(mag. vakuum-eqsikatorSi), sadac vakuum-tumbos saSualebiT
iqmneba uaryofiTi wneva. Sedegad agrobaqteria aRwevs mce-
naris eqsplantatis ujredSorisebSi, rac SemdegSi ujre-
debis transformacias iwvevs. vakuum-infiltraciis meTodi

136
sakmaod iafi da efeqturia, radganac am dros ar gamoiyene-
ba protoplastebi da damatebiTi reaqtivebi.
B bi obalistikuri transformaciis meTodi. biobalis-
tikis67 meTodi dReisaTvis warmoadgens erT-erT yvelaze
efeqtur meTods erTlebnianebis transformaciaSi. igi Sei-
Zleba aseve warmatebiT iqnas gamoyenebuli orlebnianebSic.
transformaciisaTvis sawyis masalad iReben suspenziur
kulturebs, kalusur qsovils an erTlebnianebis moumwife-
bel Canasaxebs.
meTodis arsi mdgomareobs imaSi, rom volframis,
platinis an oqros umciresi nawilakebi diametriT 0.6-1.2
mkm imtvereba (ifareba) mcenaris ujredSi Sesayvani ucxo
genis matarebeli veqtoris dnm-iT. dnm-s matarebeli vol-
framis, platinis an oqros nawilakebi plastmasis saTav-
sis saSualebiT Tavsdeba biobalistikuri “zarbaznis” 68
SigniT (sur. 49). kalusi an ujredebis suspenzia gadaaqvT
petris jamze agarizebuli sakvebiT da Tavsdeba bioplas-
tikuri “zarbaznis” qveS 10-15 sm manZilze.

d aWirxnu l i
ai r i
f i ltri
amC qareb el i
mi l i

heli
u mi

kar triji dnm-

mi krosferoebiT

a b
g amos u l i
h aer i

s ur. 49. “genuri p istoleti” (a) d a m isi m uSaobis s qema (b)

“zarbazanSi” vakuum-tumbos saSualebiT mcirdeba wneva

0.01 atm-mde. wnevis gaSvebisas volframis, platinis an oq-
ros nawilakebi didi siCqariT gamoityorcnebian “zarbaz-

67 SemoklebiT biolistikis meTodi. aseve uwodeben mikronawilakebiT

dabombvis meTods.
68 mas asevev “genur pistolets” uwodeben.

137
nidan” da ujredis kedlis gaxvretis Sedegad Sedian ci-
toplazmaSi da birTvSi. Cveulebriv centrSi mdebare ujre-
debi maRali wnevis gamo iRupeba, xolo misgan 0.6-1.0 sm
gverdiT mdebare ujredebi transformirdeba. vinaidan mem-
branaSi da ujredis kedelSi gakeTebuli mcire zomis xvre-
lebi swrafad ivseba, isini ver arRveven ujredis mTliano-
bas. nawilakebi ki imdenad mcirea, rom protoplazmaSi dar-
Cenilebi, xels ver uSlian ujredSi mimdinare bunebriv
procesebs. transformirebuli ujredebi Semdeg frTxilad
gadaaqvT Semdgomi kultivirebisa da regeneraciis sakveb
areebSi.
meq anikuri i nokulaciis meTod i . mcenareTa viruse-
bis safuZvelze Seqmnili veqtoruli molekulebis gamoye-
nebisas tardeba meqanikuri inokulacia – Sesabamisi nukle-
inis mJavebis molekulebis Sezelva foTlis firfitaSi.
amasTan gasaTvaliswinebelia, rom mcenaris ujredebis e.w.
pirdapiri transformacia (nukleinis mJavebis SeRweva uj-
redis garsis gavliT baqteriebis an virusis nawilakebis
monawileobis gareSe) arsebiTad gaZnelebulia. dnm-s an
rnm-s molekulebisaTvis ZiriTad winaaRmdegobas ujredSi
SeRwevis gzaze warmoadgens garsi. misi qimiuri Sedgenilo-
ba da Semadgeneli komponentebis ganlagebis Tavisebureba
iwvevs nukleinis mJavebis SeboWvas, riTac zRudaven ujre-
dis membranasTan maT misawvdomobas. Tumca meqanikuri ino-
kulaciisas sakmarisia virusis veqtoris nukleinis mJavis
Tundac Zalian mcire raodenobis molekulis SeRweva cal-
keul ujredebSi, radganac Semdeg iwyeba veqtoris moleku-
lis swrafi gavrceleba.

2.5. genebis gadatanis kombinirebuli meTodebi

agrolistikis meTod i . es meTodi warmoadgens mcena-

reSi genebis gadatanis pirdapiri da agrobaqteriuli veq-
toruli meTodebis kombinacias, romelic bolo wlebSi
warmatebiT gamoiyeneba. igi dafuZnebulia eukariotuli
promotoris regulirebis qveS myofi ucxo genis, veqtoris
138
T-dnm-s, markeruli genis da virulentobis agrobaqteriuli
genebis Semcveli dnm-s molekulis mcenaris ujredSi pir-
dapiri meTodiT, mag. biobalistikuri meTodiT gadatanaze.
amMdros virulentobis genebis droebiTi eqspresia mcena-
ris ujredSi iwvevs cilebis sinTezs, romlebic sworad
amoWrian plazmididan T-dnm-s da CaaSeneben mas patronis
genomSi, rogorc Cveulebrivi agrobaqteriuli transforma-
ciis SemTxvevaSi.
transformaciis Semdeg ujredebi gadaaqvT in vitro piro-
bebSi fotohormonebis da seleqtiuri agentis Semcvel sak-
veb areSi. warmoqmnili kalusidan regenerirebul trans-
formirebul mcenareebs utardeba Sesabamisi testireba.

2.6. mcenaris ujredis transformaciis

calkeuli sakiTxebi

2.6.1. mcenareSi ucxo genis eq spresiis

regulreba

genis erT-erTi mniSvnelovani Tvisebaa eqspresiis anu

muSaobis unari. am Tvisebaze pasuxismgebelia sxvadasxva ge-
netikuri elementi, romlebic unda CaSendes ucxo genis ma-
tarebel veqtoris molekulaSi.
mcenaris genomSi gadatanili ucxo genis efeqturi eq-
spresiisaTvis aucilebelia, rom am gens hqondes Sesabamisi
regulatoruli Tanmimdevrobebi, romelsac miekuTvneba:
a) promotori – Tanmimdevroba, romelsac ukavSirdeba
fermenti rnm-polimeraza. ujredSi aris 3 saxis rnm-polime-
raza, Sesabamisad 3 saxis promotori. rnm-polimeraza I da
rnm-polimeraza II-is promotorebi ganlagebulia transkrip-
ciis iniciaciis ubnis win, xolo rnm-polimeraza III-is
promotori – struqturuli genis CarCos farglebSi;
b) enhanserebi (gamaZliereblebi) da modulatorebi –
transkripciis donis gamaZlierebeli dnm-s Tanmimdevrobe-
bi;

139
 g) terminatorebi – transkripciisa da translaciis
Sewyvetis specifikuri Tanmimdevrobebi.
P promotori SeiZleba iyos Zlieri da susti. Zlieri
promotori i-rnm-s sinTezis inicirebas axdens xSirad. pro-
motori SeiZleba iyos regulirebadi da araregulirebadi.
ufro moxerxebulia regulirebadi Zlieri promotorebis
(inducibeluri) gamoyeneba.
E eukariotuli organizmebis promotoris agebuleba ma-
Ralkonservatiuli da sakmaod universaluria, amitom mce-
naris ujredSi warmatebiT eqspresirebs ara marto nebismie-
ri saxeobis mcenaris ucxo geni, aramed ZuZumwovrebis, sa-
fuarebis da sxva eukariotebis organizmidan aRebuli
ucxo geni. magram prokariotuli organizmebis (baqteriebi)
promotoris agebulebis gansxvavebulobis gamo, aqedan aRe-
buli genis warmatebuli eqspresiisaTvis aucilebelia pro-
motoris Secvla Sesabamisi eukariotuli promotoriT. ase-
ve aucilebelia sxva gardaqmnebi (“iSviaTi” kodonebis Sec-
vla, “kriptikuli” signalebis mocileba da sxva).
mcenareTa transformaciaSi gamoyenebuli promotore-
bis mravalferovneba didia.Okonstitutivuri promotorebi-
dan 69 orlebnian mcenareebSi efeqtur promotorebs warmo-
adgenen yvavilovani kombostos virusis (CaMV) 35S-promo-
tori, agrobaqteriis nopalin-sinTetazas genis nos-promo-
tori, erTlebnian mcenareebSi – simindis alkoholdehidro-
genazas genis (Adh) promotori, brinjis 1 aqtinis genis (Act)
promotori.
K konstitutivuris garda cnobilia mravali specifiku-
ri promotori, romlebic aqtiuria mxolod mcenaris cal-
keul organoebSi, qsovilebSi an ujredebSi, an mcenaris
onTogenezis calkeul stadiaze. magaliTad, kartofilis
patatinis genis promotori praqtikulad muSaobs mxolod

69 constitutive promoter (frang. Constitutive-ganmsazRvreli) araregulirebadi

promotori, romelic uzrunvelyofs mis mier kotrolirebadi genis mud-
miv transkripcias.
140
tuberebSi. iyeneben inducibelur promotorebs, romlebic
aqtivirdebian temperaturis, ganaTebis, fitohormonebis kon
centraciis da sxva mxolod garkveul pirobebSi.
G genis eqspresiis done damokidebulia mraval faqtorze.
pirvel rigSi, imaze, Tu birTvis qromatinis romel ubanSi
moxvdeba Seyvanili geni. rogorc wesi, genis eqspresia maRa-
lia Tu igi moxvdeba aqtiuri qromatinis ubanSi. eqspresia
ecema, roca birTvis genomSi CaSenebisas geni ganicdis ar-
sebiT cvlilebebs – gardaqmnebs, duplikaciebs, inversiebs
da a.S.
transformaciis procesSi gamoyenebuli calkeuli
procedurebi garkveul uaryofiT zemoqmedebas axdenen pat-
ronis genomze.
ucxo genis CaSenebam SeiZleba daarRvios patronis ro-
melime genis pirveladi struqtura da amiT gamoiwvios misi
inaqtivacia (es movlena arc ise iSviaTia). ucxo genis agro-
baqteriuli an pirdapiri gadatanis dros mcenaris genomSi
xSirad fiqsirdeba sxvadasxva saxis gardaqmnebi qrosomebis
fragmentebis translokaciis CaTvliT. aRniSnulma SeiZle-
ba Secvalos mcenaris genomis normaluri funqcionireba.
K kidev erT problemas warmoadgens genebis “gaCumeba” (ge-
ne silencing)70 . dadgenil iqna, rom sakmaod bevr transformi-
rebul mcenareSi Seyvanili geni garkveuli drois Semdeg
kargavs Tavis aqtiurobas, Tumca geni fizikurad arsebobs
genomSi, anu mcenares gaaCnia ucxo genis eqspresiisadmi aq-
tiuri dapirispirebis unari.
genebis “gaCumebis” problemas gaaCnia didi praqtikuli
mniSvneloba, vinaidan genetikurad transformirebul sa-
soflo-sameurneo kulturebSi ucxo genebi stabilurad un-
da funqcionirebdnen.
ukanasknel wlebSi dadgenil iqna genebis “gaCumebis”
meqanizmi (rnm-interferenciis procesi) da misi xelSemwyobi

70genis eqspresiis gamorTva an darRveva transkripciis an translaciis

doneze.
141
faqtorebi. erT-erT ZiriTad faqtors warmoadgens genomSi
CaSenebuli genis identuri aslebis raodenoba. genis asle-
bi rac meti da grZelia, miT maRalia “gaCumebis” albaToba.
amitom transformirebul ujredSi haploidur do-neze ar
unda iyos erT CaSenebul genze meti, aseve genis regulato-
rul nawils ar unda hqondes grZeli homologiebi patro-
nis genomTan da genidan SesaZleblobis farglebSi mTlia-
nad unda iyos mocilebuli veqtoruli plazmidis an viru-
sis dnm-s fragmentebi.
mcenareebSi Seyvanili genis eqspresiis done da stabi-
luroba SeiZleba amaRldes genze specialuri Tanmimdev-
robebis – terminatorebis mierTebiT. ase, transkripciis
terminatorebis damateba xels uSlis ferment rnm-polime-
razas arasasurvel gadaadgilebas gadatanil genze, roca
patronis genis promotori SemTxveviT axlosaa CaSenebis
adgilTan. SedarebiT mokle (>300 n.w.) bolo monakveTis da-
matebisas, CaSenebul gens qromatinis71 damoukidebeli mar-
yuJis warmoqmnis an transkribirebadi euqromatinis72 Se-
madgenlobaSi CarTvis meti SesaZlebloba aqvs. orive Sem-
TxvevaSi es iwvevs CaSenebuli genis eqspresiis donis amaR-
lebasa da stabilizacias.

2.6.2. markeruli (reportiori, seleqtiuri) genebis

gamoyeneba

mcenareTa transformciis warmatebiT ganxorcielebi-

saTvis aucilebelia mcenaris genomur dnm-Si ucxo dnm-s
integrirebis (eqspresiis) dadasturebis da transformire-
buli ujredebis gamorCevis efeqturi saSualebebis arse-
boba. amisaTvis iyeneben e.w. markerul73 genebs, romlebsac
miakereben mcenaris matransformirebeli genebis regulato-

71 birTvis nivTiereba, warmoadgens dnm-s, rnm-s da cilebis kompleqss.

72 aqtiuri qromatini.
73
marquer (frang.) – aRniSvna.
142
rul ubnebTan.
M transformirebuli ujredebis maidentificirebeli mar-
keruli genebis or jgufs gamoyofen: 1) seleqtiuri genebi
da 2) reportiori genebi.
seleqtiuri genebi 74 warmodgens antibiotikebis (kanami-
cini, tetraciklini, neomicini da sxva), toqsikuri nivTie-
rebebis (herbicidebi, mZime metalebi) da sxva agentebis mi-
marT gamZleobis makodirebel genebs, romelTa eqspresia
advili SesamCnevia organizmSi.
isini transformirebul ujredebs aniWeben aratran-
sformirebuli ujredebis zrdisa da dayofis mainhibirebe-
li garkveuli nivTierebebis Semcvel seleqtiur sakveb are-
ze zrdisa da dayofis unars.
am genebis produqti – niSani, maTi matarebeli ujrede-
bis gamorCevis saSualebas iZleva.
bolo wlebSi transformirebuli sasoflo-sameurneo
mcenareebis misaRebad seleqtiur genebad ZiriTadad iyene-
ben herbicidisadmi – fosfinotricini, qlorsulforoni da
sxva gamZleobis genebs.
reportiori geni 75 warmoadgens ucxo genTan dakavSi-
rebul gens, romelic akodirebs advilad gamosavlen pro-
duqts, Cveulebriv ferments.
igi advilad vlindeba (testirdeba) da amitom misi sa-
SualebiT SesaZlebelia mcenareSi gadatanili matransfor-
mirebeli genis aRmoCena, anu imis garkveva, rom ucxo geni
gadatanilia Tu ara ujredSi. reportiori geni transfor-
maciis Sedegebis vizualizaciis saukeTeso saSualebaa.
reportiori genebis saxiT yvelaze xSirad iyeneben β-
glukuronidazas (GUS), mwvane fluotescenciis cilas
(GFP), luciferazas (LUC), qloramfenikolacetiltransfe-
razas (CAT) genebs.
β-glukuronidazas (GUS) geni (sur. 50) E.coli-dan modi

74
laT. selectio-SerCeva, gamorCeva.
75
ingl. reporter - reportiori laTinuridan report – gadacema.
143
 a b g

d e v

s ur . 50. r e p o r tio r i g e n β-g l uku r o nid azas e q s p r e s ia

m ze s um zir aS i
( a- moumwifebeli yvaviledis W rili; b-yvavili; g-g aRivebuli T eslebi; d-mcenaris
s x vad as x va qs ovi l i ; e- T es l i s l eb an i ; v- mou mwi f eb el i C an as ax i )

ficirebul gens. mas SeuZlia bunebrivi da xelovnuri glu-

kuronidebis farTo speqtris hidrolizi, rac aadvilebs
misi aqtiurobis speqtrofotometriul an flurometriul
gamsazRvras. fermenti sakmaod stabiluria – uZlebs gacxe-
lebas da detergentebis moqmedebas. ar kargavs aqtiurobas
gayinva-galRobis procesSi.
mwvane fluotescenciis cilas (GFP) geni (sur. 51) aRmo-
Cenil iqna meduzaSi Aequorea victoria. pirvelad kloni-rebul
iqna 1992 wels. amJamad farTod gamoiyeneba rogorc proka-
riotebSi, ise eukariotebSi. es gamowveulia ultraiisferi
grZeltalRiani dasxivebisas speqtris xi-lul (mwvane) uban-
Si misi fluorescentrirebis unariT. igi inarCunebs stabi-
lurobas genur-inJinrul samuSaoebSi, aseve stabiluria
igi cocxal ujredebSi.
bolo wlebSi miRebulia am genis mravali warmoebu-
li. magaliTad, zRvis anemona Discosoma sp.-dan gamoyofil
iqna cila DsRed, romelic fluorescentrirebs wiTel
ferSi.

144
 a b

g d

s ur. 51 . r eportiori g e n mwvane f l uo te s ce nciis cil as

e q s p r e s ia x o r bal S i
( a- moumwifebeli C ansaxi; b ,g-kalus uri qsovilebi; d-aratrans f ormi reb u l i
kal u s i ( s akon trol o)

luciferazas (LUC) geni (sur. 52) klonirebulia baq-

teriidan da cicinaTeladan. cila transformirebul uj-
redebSi iwvevs naTebas.
misi aqtiurobis gansazRvrisaTvis aucilebelia spe-
cialuri mowyobilobebi – fluoremetri da sinaTlis
signalis amplifikatoriani cifruli videokamera. fermen-
ti detergentebisa da maRali temperaturis moqmedebisas
kargavs aqtiurobas.
qloramfenikolacetiltransferazas (CAT) geni (sur.
53) gamoyofilia baqteria Escherihia coli-dan. is iwvevs anti-
biotik qloramfenikolis inaqtivacias misi acetilirebis
gziT.
fermentis gansazRvra xdeba histoqimiurad, Sesabami-
si substratis damatebisas qsovilis Seferilobis cvli-
lebis mixedviT. xSirad iyeneben veqtoris replikaciis Se-
safaseblad.
145
 s ur . 52 r e p o r tio r i g e n l ucif e r azas e q s p r e s ia
Tam baq o S i

s ur. 53. r eportiori g en q loramfenikolacetiltr ans f e r azas

e q s p r e s ia d r o zo f il as e m br io nS i

reportior genebs sxva mimarTulebiTac iyeneben. 54-e

suraTze naCvenebia kalanxoe Starlight Avatar - pirveli komer-
ciuli luminescentriuli mcenare.Mmis genomSi zRvis mo-
cimcime baqteria Photobacterium leiognathi-dan gadatanilia
fluorescentrirebadi cilis geni.
markeruli genebis SerCeva damokidebulia konkretuli
146
 s ur . 54. kal anx o e S tarlight Avatar - p ir v e l i ko m e r ciul i
l um ine s ce ntr iul i m ce nar e

eqsperimentis bunebaze. Tu aseTi geni xels uSlis ujredis

normalur zrdas an safrTxes uqmnis garemos dabinZurebas
an adamianis janmrTelobas, is unda Seicvalos sxva geniT.
bolo wlebSi xSirad seleqtiur genebs cvlian reportio-
ri genebiT, vinaidan garemosa da adamianis janmrTelobi-
saTvis es ukanasknelni potenciur riskebs praqtikulad ga-
moricxaven.

2.6.3. transformirebuli mcenareebi mar keruli

seleqtiuri genebis gareSe

rogorc ukve aRiniSna, ucxo genTan erTad mcenaris

genomSi SehyavT seleqtiuri markeruli geni. miuxedavad
imisa, rom dRemde ar dafiqsirebula am genebis raime arasa-
surveli zemoqmedeba adamianze, cxovelebsa da garemoze,
sazogadoebaSi mainc gaCnda SeSfoTeba im mosalodnel ua-
ryofiT Sedegebze, romlebic potenciurad SeiZleba gamo-
iwvios mcenareSi CarTulma seleqtiurma markerulma geneb-
ma. magaliTad, zogierTi seleqtiuri genis produqti SeiZ-
leba iyos toqsikuri an alergiuli, xolo antibiotikebi-

147
admi gamZleobis genebi SeiZleba moxvdnen paTogenur niada-
gis mikroorganizmebSi. garda amisa, seleqtiuri genebis ar-
seboba teqnikurad arTulebs transgenuri mcenareebis
transformacias axali ucxo geniT, vinaidan erTi seleqti-
uri markeri ar SeiZleba gamoyenebul iqnas orjer. amitom
SemuSavebul iqna markeruli genis garSe transformirebu-
li mce-nareebis miRebis meTodebi.
umarkero transgenuri mcenareebis miRebis erT-erTi
midgoma moicavs ori sxvadasxva genis mqone mcenaris er-
Tobliv transformacias, sadac erTi atarebs markerul
gens, xolo meore – ucxo gens. am dros 30%-dan 80%-mde
SemTxvevaSi mcenareebi Seicaven orive gens, magram isini in-
tegrirebuli arian qromosomuli dnm-s sxvadasxva saitebSi.
transformantebis gamorCevis Semdeg markeruli seleqtiu-
ri geni SeiZleba mocilebul iqnas trangenuri mcenaridan
Cveulebrivi Sejvarebis gziT.
sxva midgomiT, markeruli seleqtiuri geni CaSendeba
mcenareul mobilur elementebs – transpozonebs (Tn-ele-
mentebi) Soris da aseTi konstruqcia ferment transpoza-
zas genTan erTad SehyavT T-dnm-Si. fermenti amoWris dnm-s
Tn-elementebs Soris da gadaitans sxva qromosomul saitSi.
T-dnm-is patron-mcenaris dnm-Si CaSenebis procesSi, Sem-
Txvevebis 90%-Si, or Tn-elementebs Soris moTavsebuli mar-
keruli seleqtiuri geni qromosomuli dnm-s sxva saitze
aRmoCndeba, amasTan, 50% albaTobiT es saiti sawyisidan
Sors iqneba. amrigad, markeruli seleqtiuri geni SeiZleba
gamoyenebul iqnas transformirebuli mcenareebis identi-
fikaciisaTvis, xolo Semdeg mocilebuli – Cveulebrivi Se-
jvarebis gziT.

2.6.4. Tambaqos transformacia „foTlis

rgolebis“ meTodiT

transformirebuli mcenareebis miRebis teqnologiu-

ri procesis ukeT gacnobierebis mizniT saWirod CavTvaleT

148
warmovadginoT Tambaqos „foTlis rgolebis“ meTodiT
transformaciis calkeuli samuSaoebi erT-erTi meTodikis
mixedviT.
transformirebuli mcenareebis misaRebad foTlis
rgolebs Rebuloben in vitro pirobebSi gamravlebul an saT-
buris pirobebSi gamozrdil mcenareebs. Sesabamis sakveb
areze kultivirebisas foTlis rgolebi swrafad gadadian
kalusis formirebis fazaSi Wrilobis adgilas da Tu maT
regeneraciis areSi gadavitanT, Zalian mokle droSi – 10-15
dReSi Wrilobis adgilebSi daiwyeba ylortebis formireba.
sakvebi areebi, romlebic astimulireben foTlis rgolebi-
dan mcenaris regeneracias, Cveulebriv gamoirCevian citoki-
ninebis maRali SefardebiT auqsinebTan.
transformaciisaTvis gamoyenebuli agrobaqteriis veq-
torSi markerul seleqtiur genad CaSenebulia neomicin-
fosfotransferazas (NPT) geni. igi fosfolirebs da inaqti-
virebs antibiotik neomicins/kanamicins, romelic Cveuleb-
riv toqsikuria umaRlesi mcenareebis ujredebisaTvis. ami-
tom Wrilobis adgilas NPT gen maeqspresirebeli tran-
sformirebuli ujredebis gamorCeva SeiZleba aRniSnuli
antibiotik Semcvel sakveb areze.
agrobaqteriis mocileba xdeba mcenarisaTvis aratoq-
sikuri antibiotikebiT, magaliTad cefotaqsimiT. Semdeg
inokulirebuli eqsplantati Tavsdeba regeneraciis sakveb
areze, sadac ylortebi viTar-deba uSualod transformire-
buli ujredebidan.
meTodikis mixedviT transformaciis procedura moi-
cavs Semdegi etapebs:
1. Tambaqos axalgazrda, srulad gaSlili foTlebi
Tavsdeba sterilur WurWelSi da esxmeba 10%-iani sarecxi
saSualeba Domestos. foTlebis dauzianeblad steriluri
SpateliT frTxilad aureven da toveben 15 wuTi;
2. guldasmiT recxaven 400 ml wyalSi 4-jer;
3. foTols acileben Sua ZarRvs da sterilizaciisas
dazianebul nawilebs da steriluri lancetiT Wrian 0,5-1
sm2 zomis kvadratebad. regularulad asterileben spirtiT
149
da spirtquris aliT. Rebuloben foTlis 200-mde frag-
ments. Semdeg 10-12 cals aTavseben 20-mde petris jamSi mu-
rasige-skugis (MS) sakvebi ariT da mwebvare lentiT akeTe-
ben hermetizacias. jamebis inkubireba xdeba 2 dRe-Ramis
ganmavlobaSi 24-260C temperaturis da dabali ganaTebis
(2000 luqsi, 16 sT fotoperiodi) pirobebSi.
4. procedura inficirebis procesis mimdinareobis Se-
samowmeblad moicavs sakontrolo variantis arsebobas. ami-
saTvis iReben arainokulirebul 8 petris jams da or-or
maTgans aTavseben 4 sxvadasxva sakveb areze: MS; MS+cefo-
taqsimi (Cx); MS+kanamicini (Km); MS+ Cx+ Km.
inokulacia xdeba 12 petris jamSi. amisaTvis foTlis
fragmentebs aTavseben 1:50 proporciiT ganzavebul agrobaq-
teriis Sesabamisi Stamis Ramis kulturis76 suspenziaSi da
asxamen murasige-skugis (MS) sakveb ares. mwebvare lentiT
petris jamis gverdebis hermetizaciis Semdeg xdeba inkuba-
cia oTxi dRe 24-260C temperaturis da susti ganaTebis (2-4
aTasi luqsi) pirobebSi;
5. agrobaqteriasTan inkubaciis Semdeg 2 dRe-Ramis
ganmavlobaSi an xiluli baqteriuli koloniebis gamoCe-
nisTanave inokulirebuli foTlis fragmentebi gadaaqvT
Semdeg sakveb areebze: 6 petris jami MS+ Cx+ Km; 6 petris
jami MS+ Cx;
6. inkubacia xdeba dabali ganaTebisa (2-4 aTasi luqsi)
da 24-260C temperaturis pirobebSi, ylortebisa da kalusis
ganviTarebaze perioduli dakvirvebiT.
Pproceduris Semdeg etapze adgili aqvs ganviTarebuli
ylortebis gadatanas seleqtiur sakveb areze dafesvianebi-
sa da transformirebuli mcenareebis gamosavlenad. kanami-

76 Ramis kulturis miiReba Semdegnairad: baqteriis kolonias Tesaven

Sesabamisi seleqtiuri antibiotik damatebul sakveb areSi. inkubacia
xdeba Ramis ganmavlobaSi 370C temperaturaze, intensiuri njRrevis piro-
bebSi.

150
cinis Semcvel seleqtiur areze fesvebs inviTareben da
mTlian mcenared regenerirdebian mxolod transformire-
buli mcenareebi.

2.6.5. mcenaris plastomuri genomis tran sformacia

plastidebi warmoadgens citoplazmatur organelebs
da monawileoben yvela mcenareul ujredSi. evoluciurad
plastidebi warmoSobilia cianobaqteriebidan eukariotu-
li ujredebis mier maTi STanTqmis Sedegad da dRemde in-
arCuneben prokariotuli ujredebisaTvis damaxasiaTebel
mraval niSans, iseTebs rogoricaa promotoris prokario-
tuli tipi, genebis gaerTianeba operonebSi, specifikuri ge-
netikuri kodi, transkripciisa da translaciis meqanizme-
bis prokariotuli tipi.
plastidebis genomi warmodgenilia 120-180 aTasi n.w.
sigrZis dnm-s rgoluri molekuliT. plastiduri genebi
dnm-s molekulaSi ganlagebulia Zalian kompaqturad da
funqcionalurad iyofa or ZiriTad klasad: fotosinTeti-
kur aparatTan dakavSirebuli genebi da genetikuri sisteme-
bis genebi (r-rnm-is, cilebis, t-rnm-is, rnm-polimerazas da
sxva genebi).
plastiduri genebis umetesoba organizebulia opero-
nebSi 77 da eqspesirdebian erTad giganturi policistronu-
li 78 m-rnm-s warmoqmniT.
foTlis mezofilis yoveli ujredi Seicavs 100-mde
plastids. yoveli plastidi poliploidia da Seicavs rgo-
luri dnm-s 10-100 asls. amdenad yoveli ujredi SeiZleba
Seicavdes plastiduri genomis yoveli genis 10 aTasamde
asls.
B bolo dros genetikur-inJinruli kvlevebis gazrdili

77 operoni – erTmaneTis gverdiT ganlagebuli regulatoruli ubnebisa

da struqturuli genebis erToblioba, romelTa transkribirebisas warmo-
iqmneba m-rnm-s erTiani molekula.
78 cistroni – termin “genis” sinonimi.

151
interesi plastidebisadmi dakavSirebulia cilovani sinTe-
zis maRal donesTan plastomSi, memkvidreobis dedis xaziT
gadacemasTan da birTvul genomSi transgenebis eqspresiis
arsebiTad Semaferxebeli epigenetikuri procesebis (genebis
gaCumeba da sxva) ar arsebobasTan.
Tumca plastidebis genetikur inJineriaSi aqtiur ga-
moyenebas zRudavda rigi faqtori, pirvel rigSi, plasti-
dis ormag membranaSi ucxo dnm-s Seyvanis sirTule, aseve
plastiduri genomis poliploiduroba, vinaidan genetiku-
rad stabiluri transformantebis misaRebad aucilebelia,
transgeni CarTuli iyos plastiduri genomis yvela aslSi.
mcenaris qloroplastebis genomSi ucxo genis stabi-
luri integracia pirvelad ganxorcielda 1990 wels, roca
biobalistikis meTodiT Tambaqos qloroplastebis genomSi
Seyvanil iqna antibiotik streptomicinisadmi gamZleobis
geni aadA da miRebul iqna pirveli transplastomuri mce-na-
re.
amJamad qloroplasturi transformacia warmoadgens
mcenaris birTvuli genomis transformaciis seriozul al-
ternatvas. amis mizezi cilebis producirebis ufro maRa-
li efeqturoba da ufro meti garemos biousafrTxoebaa.
qloroplasturi transformaciis ZiriTadi upirateso-
bebi Semdegia:
1. CaSenebuli plastomuri transgenis eqspresiis donis
10-50-jer gazrda. genis maRali eqspresiis mizezs warmoad-
gens yoveli plastidis maRali poliplioduroba da uj-
redSi maTi didi raodenoba – qloroplastur genomSi
transgenis Seyvanisas misi aslebis raodenobam erTi ujre-
dis cotoplazmaSi SeiZleba Seadginos 5000-10000. maSin ro-
ca ujredis birTvSi igi mxolod erTia.
2. transgenis eqspresia qloroplastebSi ufro stabi-
luria, vidre birTvuli transformaciis dros. birTvul
genomSi transgenebis integracia xdeba SemTxveviTi wesiT
da qromosomis araprognozirebad adgilebze. es xSirad iw-
vevs eqspresiis doneebis arsebiT sxvadasxvaobas, transge-
nuri cilis sinTezis donis mniSvnelovan cvlilebas, mis
152
mTlian Sewyvetasac ki. qloroplasturi transformaciis
SemTxvevaSi transgenis CaSeneba xdeba qloroplasuri geno-
mis eqsperimentatoris mier SerCeul adgilebSi, rac gamo-
ricxavs transgenis eqspresiasa da transgenuri cilis sin-
Tezis Semcirebaze qloroplsturi dnm-s mosazRvre frag-
mentebis gavlenis SesaZleblobas.
3. erTi veqtoruli konstruqciiT miRebuli yvela
transplastomuri mcenare yovelTvis iqneba identuri gene-
tikurad da fenotipurad. Eesec dakavSirebulia imasTan,
rom transgenis integracia qloroplastis dnm-Si mimdina-
reobs mxolod veqtorul konstruqciasa da qloroplastis
genomSi arsebul garkveul Tanmimdevrobebs Soris homolo-
giuri rekombinaciis gziT. birTvuli transformaciis dros
transgeni birTvis qromosomaSi Tavsdeba SemTxveviTi wesiT
da ori damoukidebeli transformaciis miReba birTvuli
dnm-s erT da imave ubanze praqtikulad SeuZlebelia integ-
raciis potenciuri adgilebis didi raodenobis (ase-ul
aTasze meti) gamo.
4. transgenebis multigenuri eqspresiis SesaZlebloba
qloroplastebSi erTi policistronuli m-rnm-dan. baqte-
riuli genebis msgavsad qloroplasturi genebisaTvis dama-
xasiaTebelia ramodenime genis erTi promotoris qveS er-
Tad transkrobireba erTi m-rnm-is saxiT. es SesaZlebels
xdis transformaciis erTjeradi aqtiT ganxorcieldes ra-
modenime genis eqspresia. garda amisa, am dros arsebiTad
izrdeba transgenuri mcenareebis biousafrTxoeba, vinaidan
aseT multigenur veqtorul konstruqciebs ar SeuZliaT
muSaoba eukariotul genomebSi, mag. mwerebSi, romlebic po-
tenciurad SeiZleba CarTuli iyvnen transgenebis horizon-
talur gadatanaSi.
5. transplastomuri mcenareebis SemTxvevaSi transgene-
bis gavrcelebis riski minimaluria. es dakavSirebulia i-
masTan, rom momwifebuli mtvris marcvali ar Seicavs
plastidebs, romlebic gadaecemian mxolod kvercxujredi-
dan Canasaxs. amdenad, damtvervis Sedegad transgenebis sxva
mcenareebSi gadatanis riskis ar arsebobis gamo, arebiTad
153
izrdeba aseTi mcenareebis biousafrTxoeba.
qloroplasturi transformaciisaTvis veqtoruli kon-
struqciebi iqmneba E. coli-s plazmidis safuZvelze da Seica-
ven seleqtiuri markeris gensa da ucxo gens/genebs. garda
amisa, vinaidan plastomis transformaciis dros qlorop-
lastur genomSi integracia xdeba homologiuri rekombina-
ciis meqanizmis mixedviT, gadasatani ucxo genis/genebis
Semcveli veqtoruli plazmidis monakveTi flankirebulia
(SemosazRvrulia) qloroplasturi dnm-s ori fragmentis
homologiuri TanmimdevrobebiT.
seleqtiur markerad Cveulebriv iyeneben antibiotik
streptomicinisadmi gamZleobis baqteriul gens aadA. am
dros transformantebis gamorCevisas sakveb areSi strepto-
micinis damatebisas aRniSnuli genis Semcveli ujredebi
rCebian mwvane feris, xolo aratansformirebuli ujredebi
– TeTri.
bolo wlebSi seleqtiur markerad efeqturad iyeneben
ispanaxidan gamoyofil, mcenarisaTvis toqsikuri glici-
naldehidis aratoqsikur glicinbetainSi gadamyvani fermen-
tis betain aldehiddehidrogenazas gens BADH.
P dReisaTvis qloroplastebis transformaciis mizniT
iyeneben sam meTods: biobalistikurs, transformacias po-
lieTilenglikolis saSualebiT da transformacias mikro-
ineqciis saSualebiT. yvelaze farTod gamoyenebulia bioba-
listikis meTodi, romelic birTvuli genomis transforma-
ciis analogiurad tardeba.
transformirebuli mcenareebis gamorCevis Semdgomi
etapebic birTvuli transformantebis gamorCevis etapebis
analogiuria. arsebiT sxvaobas warmoadgens is, rom plas-
tiduri transformaciis dros gamorCevisa da regeneraciis
procesebi tardeba ramodenimejer. es dakavSirebulia geno-
mis ploidurobasTan da ujredSi plastidebis mraval as-
lianobasTan. vinaidan veqtoruli konstruqciis integracia
Tavidan xdeba mxolod ramodenime qloroplastSi, ramode-
nimejer (Cveulebriv 3-4 jer) regeneraciis Catareba mkacri
gamorCevis pirobebSi iwvevs imas, rom gamravldebian mxo-
154
lod transformirebuli qloroplastebi da Sedegad uj-
redSi yvela qloroplasti da Sesabamisad mcenare iqneba
veqtoruli konstruqciis fragmentis matarebeli.
dReisaTvis plastomuri genomis transformaciis gziT
miRebulia Tambaqos, kartofilis, pomidoris da sxva kul-
turebis sxvadasxva bioturi da abiotiri faqtorebisadmi
gamZle transplastomuri mcenareebi (cxrili 4).

Q mcen areTa plastomuri gen omis transformaciiT miRebuli

bi oturi da abi oturi faqtorebi sadmi gamZleoba
cx r il i 4

gen i ci la n i San i
cryAc toqsini Bacillus thuringien- mwerebisadmi gamZleoba
sis
cry2Aa2 toqsini Bacillus thuringien- mwerebisadmi gamZleoba
sis
aroA EPSPS herbicidebisadmi gamZle-
oba
tps1 treqalozofosfatsinta- gvalvagamZleoba
za
msi-99 M SI-99 sokoebisa da baqteriebis
mimarT gamZleoba
bar fosfinotricinacetil herbicidebisadmi gamZle-
transferaza oba
BADH betainaldehid dehidro- gvalvagamZleoba
genaza
transplastomuri mcenareebis gamoyenebis calke mimar-
Tulebas warmoadgens sakurnalo preparatebis da saWmeli
vaqcinebis warmoeba (cxrili 5).
ase, Tambaqos mcenaris qloroplastur genomSi Seyva-
nili geni GH1 iwvevs cila somatotropinis (zrdis hormo-
nis) sinTezs, romlis raodenoba foTlebSi Seadgens mTeli
xsnadi cilebis 7%-s. amasTan igi warmoadgens maRali bio-
logiuri aqtiurobis cilas. Tambaqos qloroplastebSi
sinTezirebuli qoleris toqsinis beta-suberTeuli gamoi-
yeneba vaqcinis saxiT. igi warmoadgens bunebrivi toqsinis

155
identurs.

mcen areTa plastomuri genomis transformacii T mi Rebuli

vaqci n ebi da samkurn alo preparatebi
cx r il i 5

gen i ci la n i San i
ctxB qoleris toqsinis beta-su- vaqcina
berTeuli
pa cimbiris wylulis sawina- vaqcina
aRmdego vaqcina
gvgvp-120 bioelstiuri cilovani medicinaSi gamoyenebu-
polimeri li
GH1 adamianis somatotropini samkurnalo preparati
(zrdis hormoni)
proinsulin proinsulini, alfa- da samkurnalo preparati
beta jaWvi
INSa5 a5 interferoni samkurnalo preparati

2.6.6. polimerazuli jaWvuri reaqcia

polimerazuli jaWvuri reaqcia (pjr) da masze dafuZ-
nebuli meTodebi dReisaTvis warmoadgenen genomis anali-
zis erT-erT yvelaze farTod gamoyenebul dnm-teqnolo-gi-
as. igi gamoiyeneba transformaciis procesSi genebis klo-
nirebisaTvis, dnm-s nukleotiduri Tanmimdevrobis gan-saz-
RvrisaTvis (seqvenirebisaTvis), genomSi Ziebuli genis arse-
bobis dadasturebisaTvis, dnm-s mutaciuri cvalebadobis
gamartivebisa da daCqarebisaTvis, genomuri biblioTekidan
genebis an genebis fragmentebis gamoyofisaTvis da sxva.
Mmolekuluri biologiis es meTodi 79 SemuSavebul iqna
keri mulisis80 TanamS.Mmier 1985 wels. meTodis arsi mdgo-

79 ingl. – Polimerase Chain Resction ( PCR)

80 keri mulisma maikl smitTan erTad 1993 wels nobelis premia miiRo
qimiis dargSi .
156
mareobs dnm-s fragmentis mravali aslis swraf miRebaSi
anu amplifikaciaSi.Amisi saSualebiT SesaZlebelia dnm-s
fragmentis aslis raodenobis milionjer gazrda. pjr-s sa-
fuZvlad udevs dnm-s replikaciis meqanizmi da gamoiyeneba
replikaciis analogiuri fermentebi. Tumca aq gamoyenebu-
lia specifikuri dnm-polimeraza – geizeris Termofiluri
baqteriidan Thermus aquaticus (sur. 55a) gamoyofili fermenti
Taq-polimeraza (sur. 55b).

a b

s ur. 55. Term o f il ur i baq te r ia Thermus aquaticus (a)

da Taq-p o l im e r azas m e s am e ul i s tr uq tur a (b)

sxva dnm-polimerazebisagan ferment Taq-polimerazas

ZiriTadi gansxvaveba mdgomareobs imaSi, rom misi inaqtivi-
reba ar xdeba 950C temperaturamde gacxelebisas. am fermen-
tis mier dnm-matricaze komplementuri jaWvis sinTezis
temperaturuli optimumi Seadgens 68-720C. ise rogorc yve-
la dnm-polomerazas, ferment Taq-polimerazas sinTezuri
aqtiurobisaTvis aucileblad sWirdeba erTjaWviani dnm-
matrica da praimeri – mcire sigrZis (15-25 n.w.), dnm-fragmen-
tis (matricis) erT-erTi bolos komplementuri erTjaWvia-
ni oligonukleotiduri Tanmimdevroba.
amitom, pjr-is Casatareblad aucilebelia amplifici-
rebadi dnm-fragmentis boloebis nukleotiduri Tanmimdev-
robis codna. praimerebi pouloben erTjaWian dnm-matrica-
ze komplementur ubnebs da reaqciuli narevis garkveul
157
temperaturaze, romelic winaswar dgindeba yoveli praime-
ruli wyvilisaTvis, ukavSirdebian mas orjaWviani monakve-
Tis warmoqmniT.
Semdeg matricaze komplementuri jaWvis daSenebis
process agrZelebs da amTavrebs Taq-polimeraza.AaRniSnu-
li fermentis kidev erT Taviseburebas warmoadgens is, rom
misi gamoyenebiT Sesalebelia mxolod 3000 n.w.-mde sigrZis
dnm-fragmentis amplificireba.
polimerazuli jaWvuri reaqcia tardeba Semdegnairad:
mcire zomis polipropilenis sinjaraSi, garkveuli marile-
bis, Mg2+ ionebis, buferuli xsnaris Semcveli reaqciuli
xsnariT, umateben pjr-is sawyis komponentebs (sur. 56) –
dnm-matricas, oTxive dezoqsinukleozidtrifosfats

f ermenti
dNTP dn m matrica Taq-pol i meraza

praimerebi

s ur .B56. p o l im e r azul i ja W v ur i r e aq ciis s awy is i

ko m p o ne nte bi
(dNTP) – dezoqsiadenozintrofosfats (dATP), dezo-qsigua-
nozintrofosfats (dGTP), dezoqsicitozintrofosfa-ts
(dCTP), dezoqsiTimidintrofosfats (dTTP), or praimers da
ferment Taq-polimerazas.
polimerazuli jaWvuri reaqciisaTvis iyeneben specia-
lur aparatebs – Termociklerebs (sur. 57), romlebic war-
moadgenen Tavisebur gamacxelebel danadgarebs, maT SeuZ-
liaT mokle droSi swrfad (10C/wT-ze meti siCqariT) avto-
maturad Secvalon temperatura: gaacxelon an gaacivon sin-
jarebi reaqciuli nareviT an moaxdinon maTi inkubacia
garkveul temperaturaze garkveuli drois ganmavlobaSi.
A polimerazuli jaWvuri reaqciis yoveli cikli Sedgeba
sami etapisagan (sur. 58):
158
 s ur . 57 . pjr-i s Casatarebli aparatebi -
Te r m o cikl e r e bi

I etapi – dnm-matricas denaturacia 92-950C-ze. am dros

adgili aqvs orjaWviani dnm-matricas denaturacias or er-
TjaWvian molekulad. misi xangrZlioba, garda pirveli ci-

I etapi
denaturacia
93 -950C

g as amravlebeli dnm-s f ragmenti

II etapi
an elingi
praimerebi
50-650C

III etapi
si nTezi
( elongacia)
70-720C

s ur . 58 . am p l if ikaciis cikl is s q e m atur i

g am o s ax ul e ba
159
klisa, Cveulebriv Seadgens 0.5-1 wuTs. pirveli ciklis xan-
grZliobas ki zrdian 3-4 wuTamde – dnm-matricas (Cveuleb-
riv genomuri dnm-s) garantirebuli denaturaciis Catarebis
mizniT;
II etapi – praimerebis anelingi. am etapze, e.w. anelingis
temperaturaze, adgili aqvs praimerebis dakavSirebas kom-
plementarobis principiT pirvel etapze miRebul erTja-
Wvian dnm-matricasTan da warmoiqmneba e.w. „falia“ Taq-po-
limerazasaTvis. anelingis temperatura dgindeba winaswar
yoveli praimerisaTvis individualurad nukleotidebis Se-
madgenlobisa da sigrZis gaTvaliswinebiT. Cveulebriv igi
meryebs 50-650C temperaturis farglebSi. etapis xangrZli-
oba Seadgens 15-60 wams;
III etapi – sinTezi (elongacia). 70-720C temperaturaze81
fermenti Taq-polimeraza ukavSideba „falias“ da dnm-mat-
ricaze 5' bolodan 3' bolos mimarTulebiT iwyebs replika-
cias – komplementarobis principiT meore jaWvis sinTezs
(jaWvis elongacias). Sedegad yoveli sawyisi jaWvidan sin-
Tezirdeba axali orjaWviani dnm. etapis xangrZlioba damo-
kidebulia dnm-fragmentis sigrZeze da Cveulebriv Seadgens
0.5-2 wuTs.
polimerazuli jaWvuri reaqciis Catarebisas didi rao-
denobiT dnm-fragmentebis misaRebad sakmarisia 25-30 cikli.
ufro meti TvalsaCinoebisaTvis ganvixiloT 59-e su-
raTze warmodgenili sqemiT Catarebuli Tambaqos genomis
markeruli gen nptII-s amflipikaciis procesi.
pjr-is Casatareblad aucilebelia praimerebis Seqmna.
gen nptII-is nukleotiduri Tanmimdevroba cnobilia. amaze
dafuZnebiT SemuSavda da xelovnurad sinTezirebul iqna 21
n.w. da 23 n.w. sigrZis A da B praimerebi, romlebic komple-
mentarulia aRniSnuli genis 5' da 3' boloebis. praimerebis

81es temperatura warmoadgens ferment T aq-polimerazas sinTezuri aqti-

urobis yvelaze optimalur temperaturas.
160
 g as amravlebeli dnm-s f ragmenti

ampl ifikaciis
pi rveli cikli D d enaturaciis etapi

D prai mereb i s an el i n g i s etapi

D s i n T ezi s etapi
d n m-s 2
mol ekulis
mi Reba

ampl ifikaci- D d enaturaciis etapi

i s meore
ci kl i

D prai mereb i s an el i n g i s etapi

ssinTezis
i n Tezis etapi
etapi

d n m-s 4
mol ekulis
mi Reba
ampl ifikaci-
i s me-3 – 3 0
ci kl i

sa Wi ro r aodenobis
d n m -fragmenebis
a m plifikacia

ampl ificirebadi f ragmentis dnm -s raodenoba 2 n, s adac n –ciklebis raodenoba

gen nptII-s fragmenti

praimeri A
praimeri B

s ur. 59 . Pp olimerazuli jaWvuri r e aqciis s q ema

161
sinTezirebuli TanmimdevrobebisaTvis anelingis temperatu-
ra erTnairia da Seadgens 540C.
sinjaraSi Tavsdeba yvela komponenti buferuli xsna-
ri, dezoqsiribonukleotidebi, Taq-polimeraza, A da B prai-
merebi da Tambaqos genomuri dnm, romelic iqneba matrica
pjr-Tvis. sinjaras aTavseben daprogramebul Termocikle-
rSi.
amplifikaciis pirveli cikli. denaturaciis etapi.
940C temperaturaze xdeba Tambaqos genomuri dnm-s – orjaW-
viani dnm-matricas denaturacia. imisaTvis, rom denatura-
cia bolomde dasruldes saWiroa procesi mimdinareobdes
3-4 wT.
A anelingis etapi. TermociklSi temperatura icvleba igi
xdeba winaswar gaangariSebuli 540 C. am dros erTjaWiani
praimerebi erTjaWvian dnm-matricaze pouloben homologiis
ubnebs da komplementarobis principiT ukavSirdebian maT
orjaWviani fragmentis warmoqmniT.
sinTezis etapi. 720 C temperaturaze Taq-polimeraza, pra-
imerze inicirebiT, iwyebs replikacias. Sedegad yovel saw-
yis jaWvze sinTezirdeba dnm-fragmenti.
amplifikaciis meore cikli. 940C-mde temperaturis
cvlileba isev iwvevs rogorc sawyisi, ise axalsinTezire-
buli dnm-s jaWvis denaturacias. Semdeg 540C-ze A da B prai-
merebi ukavSirdeba yvela jaWvis boloebs. 720 C-ze Taq-poli-
meraza dezoqsinukleozidtrifosfatebis gamoyenebiT isev
replicirebs rnm-fragments.
amplifikaciis me-3 – 30-e cikli. 940C-ze denaturirdeba
yvela matrica. anelingis temperaturaze A da B praimerebi
ukavSirdeba yvela matricis boloebs. sinTezis temperatu-
raze Taq-polimeraza isev replicirebs dnm-fragments.
P polimerazuli jaWvuri reaqciisYyoveli ciklis Sem-
deg dnm-fragmentis – gen nptII-s raodenoba matulobs 2n six-
SiriT, sadac n aris pjr-is ciklebis raodenoba.

162
2.7. mcenaris transformaciis dadastureba
mcenareSi ucxo genis gadamtani veqtoris Semadgenlo-
baSi garda ucxo genisa Sedis seleqtiuri markeris geni.
Cveulebriv es antibiotikebis (mag.Kkanamicini, higromicini)
an herbicidebis gamZleobis genia, amitom trans-genuri mce-
nareebis pirvelad gamorCevas atareben Sesabamisi antibio-
tikis an herbicidis Semcvel sakveb areze. aseT sakveb are-
ze regenerireben mxolod is mcenareebi, romelTa genomSi
aris seleqtiuri markeri. magram seleqtiur areze gadarCe-
na ar SeiZleba iyos mcenaris transgenuri bunebis absolu-
turi dadastureba. mcenaris genomSi ucxo genis Semcveli
T-dnm-s arsebobis sruli dadasturebisaTvis tardeba tran-
sformantebis testireba (analizi) polimerazuli jaWvuri
reaqciis da transgenuri mcenaris qromosomuli dnm-s
blot-hibridizaciaze dafuZnebuli molekuluri analizis
saSualebiT. aseTi analizis Catareba Sromatevadi da Zvi-
ria, magram miRebuli Sedegebi transformaciis ganxorcie-
lebaSi darwmunebis da CaSenebuli T-dnm-konstruqciis ro-
denobis gansazRvris saSualebas iZleva. garda amisa, Sesa-
bamisi m-rnm-s an cilis gamovlenis meTodiT tardeba gada-
tanili ucxo genis eqspresiis damatebiTi analizi.

2.8. mcenaris genomSi ucxo genis eqspresia

(funqcionireba)
ucxo genis eqspresiis dros mcenaris genomSi warmoiq-
mneba sxvadasxva problema. mcenareTa transformaciisaTvis
gamoyenebuli pirveli genebi gamoyofili iyo baqteriebidan
da ar SeiZleboda maTi pirdapir gamoyeneba mcenaris ujre-
debis transformaciisaTvis. imisaTvis, rom baqteriuli ge-
nebi transkribirdes eukariotul ujredSi, aucilebelia
maTi sawyisi baqteriuli promotoruli Tanmimdevroba Se-
icvalos mcenaris genebis promotoriT, an sxva, mcenaris
ujredSi transkripciis mainicirebeli promotoriT. garda

163
amisa, baqteriuli genis З'-Tanmimdevrobas aucilebelad un-
da miuerTdes poliadenilirebis fragmenti. aseTi modifi-
kacia aucilebelia imisaTvis, rom eukariotulma fermentma
rnm-polimerazam SeZlos baqteriuli Tanmimdevrobebis
transkripcia da Semdeg m-rnm-ma – baqteriuli cilis tran-
slacia mcenaris ujredSi.
P baqteriuli genebis eqspresiisaTvis promotoris saxiT
yvelaze xSirad iyeneben yvavilovani kombostos mozaikis
virusis (CaMV) 35S promotors. aseTi arCevani wyvets or
problemas: 1) orlenianebis promotori rig SemTxvevaSi ar
muSaobs erTlebnianebis genomSi da piriqiT, xolo 35S pro-
motori uzrunvelyofs transkripcias mcenareTa nebismier
genomSi; 2) umetesi eukariotuli promotori warmoadgens
sakmaod susts, xolo CaMV 35S promotori ganapirobebs mi-
si kontrolis qveS myofi genis Zalian maRali donis eq-
spresias.
CaMV 35S promotori aris konstitutiuri, rac uzrun-
velyofs ucxo genis mudmiv da Zlier eqspresias transfor-
mirebuli mcenaris yvela ujredSi. amitom igi dReisaTvis
warmoadgens yvelaze gamoyenebads mcenareTa transformaci-
is samuSaoebSi.
bolo periodSi sul ufro met mniSvnelobas iZens
mcenareuli genebis qsovilspecifikuri promotorebi. maTi
upiratesoba 35S promotorTan SedarebiT mdgomareobs ima-
Si, rom maTi kontrolis qveS myofi genebi eqspresirdebian
mxolod garkveul qsovilebSi. ase, patatinis promotoris
qveS kontrolirebadi nebismieri geni eqspresirebs mxolod
tuberebSi, vinaidan cila patatini aris kartofilis tube-
ris samaragi cila da misi geni eqspresirdeba mxolod tu-
berebSi.
qsovilspecifikur promotorebs iyeneben maSin, roca
ucxo genis cila saWiroa mcenaris garkveul organoSi. ma-
galiTad, romelime niadagis paTogenisagan dacvis SemTxve-
vaSi moxerxebulia fesvspecifikuri promotorebis gamoyene-
ba, rac gamoiwvevs damcavi cilebis sinTezs mcenaris mxo-
lod miwisqveSa nawilebSi – paTogenis uSualo kontaqtis
164
adgilebSi. aseTi promotorebis gamoyeneba mcenarisaTvis
ufro misaRebs warmoadgens.
aseve iyeneben e.w. inducibelur promotorebs. konsti-
tutiuri CaMV 35S promotorisagan gansxvavebiT, aseTi pro-
motoris kontrolis qveS genebi eqspresirdebian ara mudmi-
vad, aramed mxolod garkveul pirobebSi. magaliTad, pro-
motorebi, romlebic inducirdebian Wrilobis miyenebiT, me-
tal-inducibeluri promotorebi – inducireben metalebis
ionebis (spilenZi, rkina da sxva) monawileobiT. mravali
qsovilspecifikuri da inducibeluri promotorebis uaryo-
fiT mxares warmoadgens maTi susti aqtiuroba.
USeyvanili ucxo genis eqspresiaze promotoris garda
gavlenas axdens mcenaris genomSi misi integraciis adgili.
miuxedavad imisa, rom iTvleba T-dnm-s Tanmimdevrobas SeuZ-
lia CaSendes mcenaris genomis nebismier adgilas, arse-
bobs informacia, rom es ase ar aris. Zalian xSirad T-dnm-s
CaSeneba xdeba heteriqromatinul82 ubnebSi, sadac genis eq-
spresia ar xdeba da ucxo geni nebismieri promotoris qveS
ar transkribirdeba. aucilebeli xdeba ganmeorebiTi tran-
sformaciis Catareba.

2.9. genetikuri inJineriis miRwevebi

genetikur-inJinruli meTodebiT mcenareTa genetikuri

konstruirebis ZiriTadi mizani sasoflo-sameurneo mcena-
reTa iseTi genomebis konstruirebaa, romlebic uzrunvel-
yofen ufro maRal produqtiulobas, nayofebis xarisxs da
araxelsayreli faqtorebisadmi gamZleobas.

82 heteroqromatini – qromatinis ubnebi, romlebic ujreduli ciklis

procesSi imyofebian kondensirebul (kompaqtur) mdgomateobaSi. hete-
roqromatinuli dnm-s Taviseburebas warmoadgens Zalze dabali tran-
skripciis unari.
165
2.9.1. herbicidebisadmi gamZle transgenuri
mcenareebis miReba
herbicidebi warmoadgens fitotoqsikur nivTierebebs,
romelTa gamoyenebis gareSe Tanamedrove soflis meurneo-
bas ganviTareba ar SeuZlia da maTi warmoeba mudmivad
farTovdeba. samwuxarod, praqtikaSi gamoyenebuli herbici-
debis umetesoba, maTi ricxvi amJamad 100-ze metia, grovdeba
niadagSi, xvdeba wyalsatevebSi, xolo zogierTi maTgani na-
wilobrivi degradaciis Sedegad gardaiqmneba ufro toqsi-
kur nivTierebad. praqtikaSi gamoyenebuli herbicidebidan
bevri warmoadgens specifikurs konkretuli saxeobis an sa-
xeobaTa jgufis sarevela mcenareebis mimarT, rac maTi
kulturuli mcenaris aqtiuri vegetaciis dros gamoyenebis
saSualebas iZleva. magram am dros aucilebeli xdeba mce-
nareebis ramodenimejer damuSaveba sxvadasxva herbicide-
biT, rac aZvirebs produqciis warmoebas da xels uwyobs
garemos met dabinZurebas.
U ufro ekonomiur da ekologiurs warmoadgens moqmede-
bis farTo speqtris herbicidebis gamoyeneba, romlebic er-
Tnairi warmatebiT Trgunaven rogorc erTlebniani, ise or-
lebnTa klasis mcenareebs. maTi gamoyeneba erTjeradi damu-
SavebiT garkveul agrocenozSi arsebuli sarevela mcena-
reTa mTeli kompleqsis mospobis saSualebas iZleva. magram
maRali efeqturobis misaRwevad aseTi herbicidebiT damu-
Saveba unda ganxorcieldes kulturul mcenareTa aRmoce-
nebamde, vinaidan aqtiuri vegetaciis periodSi aucilebelia
an isini gamoviyenoT wertilovani wesiT, rac aZvirebs war-
moebas, an damatebiT gamoviyenoT sxvadasxva moqmedebis viw-
ro speqtis herbicidebi, rac zemoT aRniSnul problemebs
iwvevs.
moqmedebis farTo speqtris herbicidebis gamoyenebis
am problemis mogvarebis SesaZlebloba gaCnda mcenareTa
genetikuri inJineriis ganviTarebis Sedegad. kerZod, am her-
bicidebis mimarT gamZle (tolerantuli) kulturul mcena-
reTa Seqmnis Sedegad.
166
 gentikur-inJinruli meTodebiT herbicidgamZle mcenare-
Ta miReba pirvel rigSi dafuZnebulia herbicidebisadmi
tolerantobis warmoqmnis molekuluri meqanizmis codnaze
da am niSnis ganmsazRvreli genebis arsebobaze da moicavs
Semdeg etapebs: mcenareul ujredSi herbicidis moqmedebis
bioqimiuri samiznis gamovlena, e.i. herbicidis moqmedebis
meqanizmis codna; gamZleobis genebis wyarod konkretuli
herbicidisadmi gamZle organizmebis gamoyofa; herbicidgam-
Zleobis niSnis ganmsazRvreli genebis gamoyofa da kloni-
reba; mcenaris ujredebSi ucxo warmoSobis herbicidgam-
Zleobis genebis eqspresiis (muSaobis) Seswavla transgenur
konstruqciebSi gamosayeneblad.
dadginda, rom herbicidebis moqmedebis pirvelad bio-
qimiur samiznes warmoadgens fotosinTezis procesTan, an
aminomJavebis biosinTezTan dakavSirebuli mcenaris fermen-
tebi, Sedegad adgili aqvs fotosinTezis procesis bloki-
rebas (atrazini, simazini, diuroni) an aromatuli aminomJa-
vebis (glifosati) an datotvili jaWvis aminomJavebis (qlo-
rsulfuroni, sulfometuronmeTilis biosinTezis daTr-
gunvas (inhibirebas)83.
H herbicidebisadmi gamZleobis amocanis gadasawyvetad sa-
mi strategiuli mimarTuleba arsebobs: 1) fermenti-samiz-
nis, romelzec moqmedebs herbicidi, mgrZnobiarobis Sec-
vla, e.i. fermentis genSi mutaciis gamowveva, Sedegad Sxams
ar SeuZlia Secvalos fermentis aqtiuroba da maSasadame
mcenarisaTvis zianis miyeneba; 2) fermenti-samiznis ganmsaz-
Rvreli genebis mcenareSi amplifikacia, e.i. fermenti-samiz-
nis produqciis imdenad gadideba, rom Cveulebrivi mcena-
ris mosaspobad sakmarisi Sxami arasakmarisi aRmoCndes
fermenti-samiznis Warbi raodenobis inaqtivirebisaTvis; 3)
mcenareebSi herbicidis damSleli fermentis ganmsazRvre-
li genebis Seyvana, e.i. mcenaris iZuleba TviTon gamoimuSa-

83
Cveulebriv, herbicidebi mcenaris ujredSi sasicocxlod mniSvnelovani
fermentebidan mxolod erTis blokirebas an daTrgunvas axdenen.
167
os Sxamis mainaqtivirebeli nivTiereba. mecnierebi yvela
aRniSnul mimarTulebas warmatebiT iyeneben.
fotosinTezis procesis blokirebis (inhibirebis) ga-
momwvev yvelaze farTod gavrcelebul herbicidebs warmo-
adgenen S-triazinisa da fenilSardovanas warmoebulebi:
atrazini, simazini da diuroni. es nivTierebebi ukavSirdeba
qloroplastebis membranis SemadgenlobaSi Semaval QB-ci-
las, romelsac akodirebs qloroplastis genomis geni
psbA, Sedegad adgili aqvs qloroplastebSi eleqtronebis
transportis blokirebas da fotosinTezis inhibirebas. ge-
ni psbA gamoyofil iqna zogierTi atrazingamZle sarevelas
(mag. Amaranthus hybridus) genomidan da dadgenil iqna, rom Cve-
ulebrivi atrazin mgrZnobiare mcenaris analogiur gen-Tan
SedarebiT adgili aqvs misi produqtis (QB-cilis) an 228
mdgomareobaSi serinis Secvlas gliciniT, an 219 mdgomare-
obaSi valinis Secvlas izoleiciniT, an 225 mdgomareobaSi
fenilalaninis Secvlas ToroziniT, anu herbicidisadmi
gamZleoba dakavSirebulia psbA genSi wertilovan mutaci-
asTan, rac iwvevs cilaSi aminomJavebis Canacvlebas. aseTi
cvlileba QB-cilaSi mkveTrad amcirebs masTan herbicidis
dakavSirebas. Sedegad warmoiqmneba herbicidisadmi gamZleo-
ba.
M mutanturi 84 geni psbA CaSenebul iqna agrobaqteriul
veqtorul sistemaSi da nopalinsinTetazas promotoris
qveS gadatanil iqna Tambaqos mcenaris ujredSi, saidanac
regenerirebul iqna atarazinisadmi gamZle mcenareebi.
Semdgomi gamokvlevbiT dadginda, rom herbicidisadmi
gamZleobis genebis wyaro umaRlesi mcenareebis garda, Sei-
Zleba iyos sxva organizmebic. herbicidebTan erTad sakveb
areze kultivirebis gziT gamorCeul iqna diuronisa da ar-
tazinisadmi gamZle cianobaqteriis (Anacystis nidulans) da er-
Tujrediani eukariotuli wyalmcenaris (Chlamidomonas rein-

84
Mmutanturi anu aminomJava Canacvlebuli (serini gliciniT) psbA geni
miRebul iqna sait-specifikuri wertilovani mutaciiT.
168
hardii) mutanturi formebi. maTgan gamoyofili da klonire-
buli psbA geniT miRebul iqna transgenuri Tambaqos mcena-
re.
aromatuli aminomJavebis mainhibirebeli herbicidebi-
dan yvelaze farTod gamoyenebuls warmoadgens glifosati,
romelsac Zalian didi masStabiT awarmoebs amerikuli kon-
cerni Monsanto raindapis komerciuli saxelwodebiT. igi
efeqturi, aRmocenebis Semdgomi, mTliani moqmedebis, erT-
erTi yvelaze uvnebeli samrewvelo herbicidia. glifosati
blokirebas ukeTebs aromatuli aminomJavebis biosinTezis
erT-erT ferments, kerZod 5'-enolpiruvil-3-fosfoSikimat-
sinTetazas (SemoklebiT EPSP-sinTezata), romelsac akodi-
rebs geni aroA. aRniSnuli fermenti akatalizebs fosfoe-
nolpiruvatisa da Sikimat-3-fosfatisagan 5'-enolpiruvil-
Sikimat-3-fosfatis warmoqmnas. metabolizmis am stadiis
daTrgunva iwvevs aromatuli aminomJavebis deficits da Si-
kimatis dagrovebas, Sedegad, glofosatTan xangrZlivi kon-
taqtisas, ujredi iRupeba.
G glifosatisadmi gamZle mcenaris misaRebad mecnierebma
baqteria Salmonella thyphimirium-is ujredebSi mutagenezis
gziT miiRes Secvlili struqturis mqone (101 kodonSi pro-
lini Secvlilia seriniT), Tavisi funqcionaluri aqtiuro-
ba SenarCunebuli, magram glifosatisadmi mniSvnelovnad
gamZle fermenti EPSP–sinTetaza. Semdeg, aRniSnuli glifo-
satgamZle fermentis makodirebeli geni, izolirebul iqna
S. thyphimirium-is ujredebidan, klonirebuli baqteria E. coli-
Si da Ti-plazmidis safuZvelze Seqmnili veqtoris saSua-
lebiT Seyvanili Tambaqos mcenaris ujredebSi.
analogiuri muSaoba Catarda pomidoris, kartofilis,
Saqris Warxlis, soios, bambis, alvis xis da sxva mcenaree-
bis ujredebSi. yvela mcenare gamZle aRmoCnda glifosati-
sadmi.
sxva mkvlevarebma glifosatgamZle mcenareebi miiRes
sxva gziT. kerZod, mcenaris ujredebSi ferment EPSP–sinTe-
tazas produqciis gazrdis gziT. mecnierebma, erTi mxriv
169
EPSP-sinTetazas produqciis gadidebuli Semcvelobis mqo-
ne ujredebis seleqciiT, xolo meore mxriv, TviT EPSP-sin-
Tetazas mcenareuli genis klonirebiT, misi yvavilovani
kombostos mozaikis virusis Zlieri promotoris qveS Sem-
dgomi moTavsebiTa da aseTi rekonstruirebuli EPSP-sinTe-
zatas genis Ti-plazmidis safuZvelze Seqmnili veqtoris
saSualebiT mcenaris ujredebSi SeyvaniT, miaRwies mcena-
ris ujredebSi aRniSnuli fermentis produqciis 20_40-jer
gazrdas. Sedegad miiRes glifosatisadmi SedarebiT gamZle
(10-jer ufro gamZle sawyis mcenareebTan SedarebiT) pomi-
doris, Tambaqos da petunias mcenareebi.
herbicidebi sulfometuron meTili da qlorsulfuro-
ni (sulfonilSardovanas warmoebulebi) warmoadgenen da-
totvili jaWvis aminomJavebis (izoleicini, valini) biosin-
Tezis inhibitorebs. isini aseve didi masStabiT iwarmoeba
amerikuli kompania Du pont-is mier. aRniSnuli da sulfo-
nilSardovanas warmoebuli sxva herbicidebi gamoirCeva uC-
veulod maRali efeqturobiT, moqmedebis gamorCeviTobiT
sxvadasxva kulturebis mimarT, dabali toqsikurobiT Zu-
Zumwovrebisadmi. herbicidebi Trgunaven ferment acetolaq-
tat-sinTetazas (ALS), romelic kodirdeba ilv2 geniT da aka-
talizebs or reaqcias ujredSi, Sedegad irRveva aminomJa-
vebis valinisa da izoleicinis sinTezi.
aRniSnuli herbicidebisadmi gamZle mcenares mecnie-
rebi Rebuloben ferment acetolaqtad-sinTetazas genis
wertilovani mutaciuri cvalebadobis (prolinis SecvliT
seriniT) an mcenareSi ilv2 genis amplifikaciis gziT. herbi-
cid sulfominSardovanasadmi gamZleobis genebis donors
warmoadgens Salmonella thyphimirium, Echerichia coli, safuarebi da
sxva. magaliTad, mecnierebma baqteria Salmonella-dan gamZle-
obis genis gadatanis gziT miiRes Tambaqos transgenuri
mcenare, romelic herbicid qlorsulfuronisadmi 100-jer
ufro gamZlea, vidre sawyisi mcenare.
ganxiluli magaliTebi herbicidebisadmi gamZle mcena-
reebis miRebis pirveli da meore strategiuli mimarTule-

170
bis realizacias warmoadgens. yvela SemTxvevaSi genetikuri
inJineriis meTodebi iwvevda an fermenti-samiznis mgrZnobi-
arobis Secvlas - fermentis genSi wertilovanis mutaciis
Sedegad aminomJavebis Canacvlebas, an fermenti-samiznis
ganmsazRvreli genebis mcenareSi amplifikacias.
rezistentuli mcenaris miRebis kidev erT strategiul
mimarTulebas warmoadgens mcenarisaTvis herbicidis mole-
kulis daSlis an qimiurad modificirebis unaris miniWeba,
ris Sedegadac Sxami mcenares zians ver miayenebs.
es mimarTuleba yvelaze ukeTesia herbicidebiT niada-
gis dabinZurebisagan dacvis mizniT, magram misi realizaci-
isaTvis aucilebelia specifikuri genebi.
mecnierebma miagnes aseT genebs da amocana gadawyve-
til iqna fosfinotricinebis jgufis herbicidebisadmi (bia-
lofosi, basta da sxva) gamZle mcenareebis miRebisas.
pirvel herbicidi am jgufidan iyo bialofosi, rome-
lic Sedgeba L-alaninisa da glutaminis mJavas analogi
fosfinotricinisagan. mcenaris ujredSi moxvedrisas Sida-
ujreduli peptidazebis moqmedebiT herbicidi iSleba da
gamoiyofa fosfinotricini, romelic Seuqcevadad inaqtivi-
rebs sakvanZo ferments amoniumis asimilaciaSi – glutamin-
sinTetazas.
am jgufis herbicidebisadmi rezistentuli genebis
wyaros warmoadgens streptomicetebis ori saxeoba: Streptom-
yces hygroscopicus da Streptomyces viridochromogenes. isini gamZlea
fosfinotricinisadmi, vinaidan warmoqmnian ferment fosfi-
notricinacetiltransferazas, romelic toqsinis damRu-
pvel moqmedebas xsnis misi acetilirebis gziT. Streptomyces
hygroscopicus-Si am fermentis makodirebel gens ewoda bar, xo-
lo Streptomyces viridochromogenes-Si – pat. komerciuli transge-
nuri jiSebis miRebaSi orive geni iqna gamoyenebuli (bar –
rafsi, turnefsi, Saqria Warxali, soio, simindi ; pat – brin-
ji), anu mcenareebSi Seyvanil iqna herbicidis damSleli
fermentis ganmsazRvreli geni.
rezistentuli mcenaris miRebis es strategiul mimar-

171
Tuleba amJamad farTod gamoiyeneba. brinjis mcenaris ge-
nomSi baqteria B. subtillis-dan gamoyofili ferment proto-
porfirinogensinTetazas genis (Protox) gadataniT, miRebul
iqna difenil-eTeris rigis herbicidebisadmi gamZle tran-
sgenuri mcenareebi. am dros gamZleoba gamowveulia imiT,
rom fermentis eqspresia mcenaris ujredSi iwvevs herbici-
dis moqmedebis neitralizacias.
baqteria Alcaligenes eutrophus-dan gamoyofili ferment di-
oqsigenazas genis (tfdA) gadataniT miRebul iqna herbicide-
bis 2,4-D da 2,4,5-T-s mimarT gamZle mcenareebi. aRniSnuli
fermenti Slis am herbicidebis toqsikur nivTierebebs –
fenoqsikarboqsilnis mJavebs. oqsilinis jgufis, fotosis-
tema II-is mainhibirebeli herbicidebis mimarT gamZle bambis
mcenare miRebul iqna Klebsiella-s gvaris baqteriidan gamoyo-
fili nitrilazas genis SeyvaniT. am fermentis moqmedebiT
bromoqsinili da ioqsinili kargaven TavianT toqsikurobas.
herbicid dalaponisadmi Tambaqos mcenaris gamZleoba miR-
weul iqna baqteria Pseudomonas putida-dan gamoyofili deha-
logenazas genis gamoyenebiT, romlis eqspresiis produqtis
moqmedebis Sedegad herbicidi gardaiqmneba pro-pionmJavad.
dReisaTvis herbicidgamZle mcenareTa jiSebis qveS da-
kavebulia mTeli transgenuri jiSebis qveS dakavebuli
farTobebis ~80%. maTgan ~10%-s Seadgenen erTdroulad
herbicidebisa da mweri-mavneblebis mimarT gamZle mcenaree-
bi.

2.9.2. mweri-mavneblebisadmi gamZle transgenuri

mcenareebis miReba

qimiuri inseqticidebis uaryofiTi Tvisebebi sayovel-

Taod cnobilia: maRali toqsikuroba85 , dabali gamorCevi-

85msoflioSi pesticidebiT mowamvlis Sedegad yovelwliurad daaxloe-

biT 500 aTasi adamiani avaddeba, maT Soris 5 aTasze meti iRupeba.
172
Tobis unari 86, neli daSla niadagSi 87. maTma farTo ga-moye-
nebam, XX saukunis meore naxevarSi, gamoiwvia mravali eko-
logiuri problema, magram uaris Tqmac SeuZlebeli iyo.
sasoflo-sameurneo kulturaTa biologiuri dacvis meTo-
debis gamoyenebam problema ver moxsna. mZime mdgomareobi-
dan gamosvlis realuri imedi gaCnda genetikuri inJineriis
ganviTarebis Sedegad.
1938 wels safrangeTSi gamoyofili, sxvadasxva saxeobis
mweris sikvdilis gamomwvevi baqteria Bacillus thuringiensis-is
Seswavlis Sedegad cnobili gaxda, rom mwerebze baqteriis
damRupveli moqmedebis mizezia maT mier sporulaciis sta-
diaze gansakuTrebuli cilis sinTezi, romelic kristale-
bis saxiT grovdeba citoplazmaSi formirebadi sporebis
axlos.
B. thuringiensis-is sxvadasxva qvesaxeobis kristaluri ci-
lebis88 Seswavlam aCvena, rom yvela isini warmoadgens e.w.
protoqsinebs, romlebic mwerebis saWmlis momnelebeli
wvenis moqmedebiT gardaiqmneba aqtiur toqsinad. aseTi toq-
sini specifikurad ukavSirdeba nawlavis endoTeliumis89
ujredebis receptorebs, warmoqmnis forebs, rac iwvevs uj-
reduli homeostazis darRvevas da sabolood ujredebis
daSlas. Semdeg dazianebuli endoTeliumis gavliT, paTo-
genurobis sxva faqtorebis gamoyenebiT, paTogeni aRwevs
mweris organizmSi da iwvevs mis sikvdils.
dadginda, rom struqturiT protoqsinebi warmoadgenen

86 isini, rogorc wesi, moqmedeben aramizanmimarTulad, spoben ara marto

mavne, aramed sasargeblo organizmebsac, maT Soris iseTebsac, romlebic
warmoadgenen mavneblebisa da daavadebebis gamomwvevebis bunebriv mtrebs.
87 mravali qimiuri pesticidi didi xnis ganmavlobaSi rCeba garemoSi da
iwvevs mis arsebiT gabinZurebas.
88
B. thuringiensis-is qvesaxeobidan gamoyofili kristaluri cilebis genebi
klonirebul iqna sxvadasxva laboratoriebSi. aman gamoiwvia am jgufis
cilebis saxelwodebad mravali sinonimis gaCena: entomotoqsini, inseq-
ticiduri toqsini, endotoqsini, kristaluri proteini, delta-toqsini, δ-
toqsini, Bt- toqsini, Bt- proteini, BT - toqsini.
89
ujredebis fena, romelic qmnis sisxlisa da limfis kapilarebis
kedlebs, arteriebis, venebisa da gulis kedlebis SigniTa Sres.
173
erTjaWvian cilებs, romlებic kodirdeba erTi geniT (aRi-
niSneba bt2). aqedan daiwyo mweri-mavneblebisadmi gamZle
mcenareTa Seqmnis proeqtebis ganxorcieleba.
B. thuringiensis-is genomidan gamoyofil iqna toqsinis geni
bt2 da 35S CaMV promotoris qveS agrobaqteriuli tran-
sformaciis meTodiT integrirebul iqna Tambaqos mcenaris
genomSi. baqteriuli bt2-genis eqspresia dadasturebul iqna
rogorc transkripciis doneze, Sesabamisi m-rnm-is arsebo-
biT, ise translaciis doneze, cila-toqsinis sinTezis mi-
xedviT.
miRebuli Tambaqos transgenuri mcenare gamZle aRmoC-
nda mavneblebis mimarT. sasoflo-sameurneo kulturebis
mavneblebisagan dacvis efeqturoba naCvenebi iqna analogiu-
ri gziT miRebul, pomodoris transgenur mcenareebzec.
amasTan, mcenaris qsovilebSi sinTezirebuli baqteriuli
toqsini uzrunvelyofda inseqticiduri preparatebis gamo-
yenebiT miRebuli damcvelobiTi efeqtis analogiur efeqts.
Tambaqosa da pomidoris garda baqteriuli bt2-geni Sey-
vanil iqna mraval sasoflo-sameurneo kulturaSi, maT So-
ris kartofilSi, simindSi, bambaSi, brinjSi, soioSi, bro-
kolSi da sxva.
dReisaTvis am gziT Seqmnili simindis, soios, rafsis,
bambis mavnebel gamZle jiSebi, dakavebuli farTobebis mi-
xedviT meore adgilzea herbicidgamZle transgenuri mce-na-
reebis Semdeg.
ganxiluli meTodiT praqtikulad realizebuli ento-
mogamZle mcenareebis Seqmnis xerxis, garda muSavdeba kidev
erTi xerxi, rac gulisxmobs saWmlis momnelebeli fermen-
tebis inhibitorebis genebis Seyvanas mcenaris genomSi. vina-
idan cnobilia, rom zogierTi mcenare garkveuli mavnebli-
sagan daculia saWmlis momnelebeli fermentebis inhibito-
rebis sinTezirebis unaris gamo.
Cinuri vignidan (grZeli lobio) gamoyofili prote-
inazas genis, CaMV 35S promotoriTa da nopalinsinTetazas
genis transkripciis terminaciis saitiT (tNOS), Tambaqos

174
mcenareSi gadataniT miRweul iqna 1 mg saerTo cilaze ~ 2
mkg tripsinis inhibitoris sinTezi, ramac arsebiTad Seam-
cira mweri Heliothis virescens-is larvisagan 90 transgenuri mce-
naris dazianeba.
Llobios α-amilazas inhibitoris geni parkosanTa fito-
hemaglutinis genis promotoriT Seyvanil iqna bardas ge-
nomSi, Sedegad mcenare daculi aRmoCnda mavneblebis Callo-
sobruchus maculatus-is da Callosobruchus chinensis-is damRupveli
moqmedebisagan.
entomogamZle mcenareTa Seqmnis mizniT bolo wlebSi
ganxorcielebul axal mimarTulebas warmoadgens rnm-in-
terferenciaze91 dafuZnebuli gamZle mcenareTa miReba. ko-
lorados xoWos β-aqtinis genis ori aslis Sebrunebuli
orientaciiT kartofilis plastidur dnm-Si Seyvanis gziT
miaRwies am genis Sesabamisi orjaWviani rnm-s dagrovebas
qloroplastebSi. es ki am molekulas kartofilis rnm-in-
terferenciis sistemiT daSlisagan icavs. magram aseTi
transgenuri kartofiliT kolorados xoWos kvebisas or-
jaWviani rnm aRwevs mweris ujredebSi da iq iwvevs rnm-in-
terferencias. es ki iwvevs β-aqtinis sinTezis Sewyvetas da
mweris sikvdils.
mweri-mavneblebisadmi gamZle transgenuri mcenareebis
gamoyenebam inseqticidebis gamoyenebis mkveTri Semcireba
da mosavlianobis amaRleba gamoiwvia.

90 cxovelebis individualuri ganviTarebis stadia. larva damoukideblad

cxovrobs, aRWurvilia specialuri organoebiT da jer ara aqvs srulasa-
kovani organizmis mravali nawili.
91 genis eqspresiis daTrgunvis (genis “gaCumebis”) procesi transkripciis,
translaciis doneze, m-rnm-is deadenilireba an degradacia rnm-is mcire
molekulebis saSualebiT. genis “gamorTvas” genomSi misi SenarCunebiT
uwodeben “genis nokdauns”, xolo misi sruli micilebiT – “genis nokauts”

175
 2.9.3. daavadebebisadmi gamZle transgenuri
mcenareebis miReba

M memcenareobas arsebiT zians ayeneben agreTve mcenareTa

daavadebebi, romelTa gamomwvevebs warmoadgenen virusebi,
baqteriebi da sokoebi.
gansakuTrebiT mniSvnelovania mcenareebis dacva viru-
sovani daavadebebisagan. vinaidan am daavadebebis mavneblo-
ba didia, metad rTulia diagnostika, praqtikulad ar arse-
bobs maT mimarT brZolis agroteqnikuri saSualebebi, xo-
lo tradiciuli seleqciis meTodebi nakleb efeqturia.
virusebi azianeben praqtikulad yvela kulturul mce-
nares da iwveven 300-ze met sxvadasxva daavadebas. amJamad
aRwerilia 700-ze meti fitopaTogenuri virusi. sia grZel-
deba.
virusovan infeqciebTan brZolis sirTule dakavSire-
bulia imasTan, rom virusebs, warmoadgenen ra obligatur
parazitebs, ar gaaCniaT sakuTari metabolizmi da daziane-
bisas patronis ujredebTan iseT mWidro kontaqtSi Sedian,
rom arsebiTad misi nawili xdebian, amitom maT mospobas
Tan axlavs TviT patron-ujredis mospobac.
virusebisadmi gamZle mcenareebis miRebaSi erT-erT sa-
interesi midgomas warmoadgens kapsidebis cilebis makodi-
rebeli virusuli genebis Seyvana mcenaris genomSi. dadge-
nili kanonzomierebaa, rom mcenaris ujredSi virusis gar-
sis cilebis didi raodenobiT dagrovebisas adgili aqvs
virusis reproduqciis (misi nukleinis mJavebis sinTezis)
Tu mTlian Sewyvetas ara, mis arsebiT daqveiTebas mainc92.
mcenareSi Seyvanili virusis genis eqspresia Zlieri promo-
toris qveS, iZleva virusis reproduqciis SezrudvisaTvis

92
gamZleobis amgvari meqanizmi jer ucnobia. mecnierebi varaudoben, rom
mcenaris ujredebSi virusis garsis cilis sinTezi xels uSlis am viru-
sis mimagrebas mcenaris ujredebze, an ewinaaRmdegeba virusis genetikuri
masalis (rnm an dnm) ujredSi Seyvanas. am movlenas aseve virusTaSorisi
jvaredini gamZleobis fenomensas uwodeben.
176
sakmarisi cilebis raodenobas. am gziT miRebul iqna mrava-
li virusgamZle mcenare: kartofili (foTlis sixuWuWis
virusi, S-virusi, X-virusi, Y-virusi), gogra (sazamTros
mozaikis virusi 2, yabayis yviTeli mozaikis vi-rusi), pomi-
dori (Tambaqos mozaikis virusi, pomidvris mozaikis viru-
si), kitri (kitris mozaikis viusi - CuMV), papaia (papaias
rgoluri laqianobis virusi), brinji (brinjis zolianobis
virusi), Tambaqo (Tambaqos mozaikis virusi, Tambaqos zoli-
anobis virusi, Tambaqos gravirebis virusi) da sxva.
kidev erT midgomas virusgamZle mcenareebis miRebaSi
warmoadgens virusis replikaciis meqanizmis darRveva “uSi-
naarso” rnm-s gamoyenebiT93. mcenaris ujredSi SehyavT viru-
sis kapsidis cilis genis in vitro xelovnurad sinTezirebuli
orjaWviani k-dnm, romelic transkripciisas iZleva dnm-s
“Sinaarsiani” jaWvis arakodirebad komplementarul jaWvs
(“uSinaarso” rnm-s). gamZleobis meqanizmi mdgomareobs Sem-
gegSi: ujredSi didi raodenobiT “uSinaarso” rnm-s arsebo-
bisas, igi urTierTmoqmedebs komplementarulad mcenaris
damasenianebeli virusis m-rnm-Tan da amiT xels uSlis mis
translacias rnm-interferenciis meqanizmiT orjaWviani
rnm-s daSlis gamo. am gziT miRebulia Tambaqos, kitris vi-
rusgamZle mcenareebi. orive ganxiluli midgomis uaryofi-
Ti mxarea – mcenaris dacva mxolod erTi konkretuli vi-
rusisagan.
mcenareTa virusgamZleobis amaRleba SeiZleba ribo-
nukleazisadmi 94 dabali specifiurobis genebis mcenaris ge-
nomSi SeyvaniT. saSualo donis eqspresiis manopinsinTeta-
zas genis promotoris qveS baqteria Serratia marcescens-s genis
da xaris ribonukleazas genis Tambaqos mcenaris genomSi

93
amisaTvis iyeneben genetikuri kontrolis axal tips e.w. “mic-rnm” (mRNA
– interfering complemantary RNA) kontrols. es aris “uSinaarso rnm”-iT ganxor-
cielebuli genebis aqtiurobis regulaciis tipi.
94 ribonukleazebi – fermenti-nukleazebi, romlecic akatalizeben rnm-s

degradacias.
177
agroinfeqciis meTodiT Seyvanis Sedegad, maTSi nukleazu-
ri aqtiurobis done gaizarda 7-10 jer.
dReisaTvis genetikur-inJinruli teqnologiebiT viru-
sebisagan mcenareTa inficirebis Semcirebis, virusebis gam-
ravlebis inhibirebis mravali midgoma arsebobs. miRebulia
sxvadasxva virusisadmi gamZle kartofilis, simindis, qeris
pomidvris, Tambaqos da sxva mcenareebi.
fitopaTogenuri sokoebisadmi mcenaris gamZleobis
amaRlebas mecnierebi cdiloben zogierTi mcenareuli ci-
lis eqspresiis donis gadidebiT. dadgenilia, rom paToge-
nebTan kontaqtisas mcenareebs calkeuli cilebis sinTezis
gadidebis an paTogenTan kontaqtamde ar arsebuli cilebis
eqspresiis unari aqvT. es cilebi warmoadgenen toqsikurs
da paTogenebis cxovelmyofelobis SemzRudvels, rac sabo-
lood iwvevs maT sikvdils. aseT cilebs uwodeben PR-ci-
lebs. 95 maTgan yvelaze ukeT Seswavlilia fermentebi qiti-
nazebi da β-1,3-glukonazebi, romlebic iwveven sokoebis da
zogierti saxeobis baqteriebis zrdis inhibirebas.
qitinazebis da β-1,3-glukonazebis genebi klonirebul
da promotor 35S CaMV kontrolis qveS gadatanil iqna Tam-
baqos, bambis, simindis, rafsis, pomidoris, brinjis, karto-
filis, turnefsis da sxva mcenareebis genomSi. yvela maT-
gans ganuviTarda sokovani infeqciebisadmi gamZleoba.
fungiciduri efeqtisDdabalmolekuluri cilebis –
cisteinuri cilebis, galaqturonazas inhibitorebis, MF-
jgufis cilebis da sxva genebis mcenareSi gadataniT miRe-
bul iqna rogorc sokovani, ise virusuli infeqciebisadmi
gamZle transgenuri mcenareebi.
mimdinareobs samuSaoebi sxva PR-cilebis – sokos mice-
liumze uaryofiTad momqmedi taumatinmsgavsi cilebis da
proteazebis inhibitorebis genebis mcenareebSi gadatanaze.
arsebobs sxva midgomebic. miRebulia Tambaqos, rafsis, po-

95
pathogenesis-releated proteins – paTogenezTan dakavSireuli cilebi.
178
midoris, kartofilis Rhizoctonia-sadmi gamZle da Tambaqos
Cercospora nicotiana-sadmi gamZle transgenuri mcenareebi.
baqteriuli dazianebebisadmi gamZle mcenareebis Seq-
mnis mimarTulebiT kvlevebi jer mosamzadebel fazaSia.
baqteriebis mier gamowveuli dazianebebis umetesoba dakav-
Sirebulia an peqto- da celulolitiuri kompleqsebiT
mcenareuli qsovilebis maceraciasTan 96 (tuberebis, bolqve-
bis, Zirxvenebis sxvadasxva sidample), an mcenaris gamtari
sistemis paTogenuri baqteriebis mier gamoyofili ekzopo-
lisaqaridebiT (e.w. mcenaris sistemuri Wknoba) blokirebas-
Tan.
rogorc wesi, baqteriuli Setevisagan mcenareebi Tavs
icaven e.w. hipermgrZnobiarobis reaqciis ganviTarebiT, rac
gulisxmobs ama Tu im organos dazianebuli nawilis swraf
sikvdilSi da amiT paTogenis Semdgomi ganviTarebis da da-
uzianebel nawilebSi gavrcelebis SezRudvas. baqteriebis
sawinaaRmdegi raime damcveli cilebi gamovlenili ar aris.
amJamad mecnierebi amuSaveben ramodenime imedismomcem mid-
gomas.

2.9.4. garemos araxelsayreli faqtorebisadmi

gamZle transgenuri mcenareebis miReba
garemos araxelsayreli faqtorebis (gvalva, Warbteni-
anoba, maRali da dabali temperatura, niadagis damlaSeba
da mJavianoba) eqstremaluri zemoqmedeba mcenareebze sa-
soflo-sameurneo produqciis arsebiT danakargebs iwvevs.
amitom stresuli zemoqmedebisadmi tolerantuli mcenare-
Ta miRebas didi ekonomikuri mniSvneloba aqvs.
stresisadmi mcenareTa adapturi reaqciebidan mravali
ganpirobebulia sxvadasxva genis sinqronuli urTierTmome-
debiT. genetikur-inJinruli teqnologiebisaTvis ufro mi-

96 qsovilTa ujredebis erTmaneTisagan gancalkeveba ujredSorisi

nivTierebis daSlis Sedegad.
179
sawvdomia is bioqimiuri procesebi, romlebic inducirebu-
lia uSualod stresis faqtorebiT. cnobilia, rom xan-
grZlivi wylis naklebobis stresis qveS myofi mcenareebi
agroveben rig organul dabalmolekulur naerTebs – pro-
lins, glicinbetains da sxva, romlebic warmoadgenen osmo-
regulatorebs anu osmoproteqtorebs.
stresuli faqtorebisadmi gamZle mcenareebis miRebis
saqmeSi mniSvnelovani iyo baqteriebisa da umaRlesi mcena-
reebis stresuli pasuxebis erTmaneTTan msgavseba: orive
tipis organizmis ujredebSi stresis dros adgili aqvs os-
moproteqtorebis molekulebis sinTezs, romelTa daniSnu-
lebaa osmosuri balansis damyareba citoplazmasa da gare-
mos Soris da cilebis stabilizacia.
E. coli-s genomidan gamoyofil iqna prolinis biosinTe-
zis fermentebis makodirebeli ori geni proBosm da proA da
agrobaqteriuli transformaciis meTodiT gadatanil iqna
Tambaqos mcenareSi. am baqteriuli genebis eqspresiam mcena-
ris genomSi gamoiwvia prolinis sinTezis gadideba. miRebu-
li transgenuri mcenareebis ylortebi fesviandeboda da
izrdeboda marilebis maRali koncentraciis (20 g/l) sakveb
areSi.
G gamoyofil iqna osmoproteqtor glicinbetainis sinTe-
zis makatalizebeli ferment betaindehidrogenazas geni
(BADH) da agrobaqteriuli transformaciis meTodiT gada-
tanil iqna Tambaqos mcenaris genomSi. miRebul transgenur
mcenareebs gaaCndaT marilebis maRali koncentraciisadmi
gamZleobis unari.
wylis naklebobisadmi gamZle mcenaris miReba SeiZle-
ba kidev erTi osmoproteqtoris – tregalozas gamoyenebiT.
safuara Saccharomyces cerevisiae-s genomidan gamoyofili, tre-
galozo-6-fosfatsinTetazas genis (tps1) Tambaqos plasti-
dur genomSi gadataniT 25-jer gaizarda mcenareSi trega-
lozas Semcveloba da Sesabamisad gamZleoba wylis defi-
citisadmi.

180
 wayinvebisadmi gamZleobis asamaRleblad gamoyenebul
iqna veluri kartofilidan (Solanum commersonii) da cianobaq-
teriidan (Anacystis nidulans) gamoyofili cximmJavebis biosin-
Tzis erT-erTi sakvanZo ferment delta-9-saturazas makodi-
rebeli geni. mcenaris plastidur genomSi am genis SeyvaniT
arsebiTad gaizarda ujeri cximmJavebiani lipidebis pro-
duqciis done, Sesabamisad wayinvebisadmi gamZleoba.
dadgenil iqna, rom maRali temperaturisadmi gamZleo-
ba dakavSirebulia Fad7 genTan, romelic gavlenas axdens
cximmJavebis metabolizmze. am genis inaqtivaciiT miRebuli
brinjis transgenuri mcenare normalurad izrdeboda maRa-
li temperaturis pirobebSi da SeeZlo 2 saaTamde gaeZlo
470C temperaturisaTvis.
F fizikuri faqtorebis (maRali temperatura, zedmeti ga-
naTeba, ultraiisferi sxivebis maRali dozebi) stresuli
zemoqmedebiT metabolizmis darRvevisagan (rac zRudavs
mcenaris zrdas, ganviTarebas, ujredebisa da qsovilebis
formirebas) Tavdasacavad mcenareebSi funqcionirebs fer-
menti – superoqsid-dismutaza. ujredSi igi sxvadasxva izo-
meruli formiT arsebobs: Cu/Zn-superoqsid-dismutaza –
qloroplastebSi da mcire raodenobiT citoplazmaSi, Mn-
superoqsid-dismutaza – mitoqondriebSi. am fermentebis ge-
nebi klonirebul iqna Sesabamisi m-rnm-s safuZvelze k-dnm-s
in vitro sinTezis gziT da Semdeg promotor 35S CaMV-s kon-
trolis qveS Seyvanil iqna Tambaqos mcenaris ujredebSi.
aRmoCnda, rom mcenareebi Cu/Zn-superoqsid-dismutazas genis
eqspresiiT, avlendnen fotosinTezis unars maRali ganaTe-
bis pirobebSi, xolo Cveulebriv mcenareebSi es unari
mkveTrad ecemoda.

2.9.5. maRalmosavliani transgenuri mcenareebis

miReba
genetikur-inJinruli meTodebiT maRalmosavliani ji-
Sebis Seqmnis problemis gadawyvetis samuSaoebi aqtiurad
mimdinareobs Semdegi momarTulebebiT: mcenareSi fotosin-
181
Tezuri aqtiurobis amaRleba da calkeuli nivTierebebis
sinTezis gazrda.
fotosinTezuri aqtiurobis amaRlebis samuSaoebi tar-
deba C4-tipis mcenareebis genebis C3-tipis mcenareebSi gada-
tanis mimarTulebiT. simindis C4-tipis mcenaris foTlis me-
zofilis ujredebSi atmosferuli CO2 -is fiqsaciaSi mo-na-
wile ZiriTadi fermentis – fosfoenolpi-ruvat-karboqsi-
lazas makodirebeli geni klonirebul da agrobaqteriuli
transformaciis meTodiT gadatanil iqna brinjis C3-tipis
mcenareSi. transgenuri mcenareebis analizma aCvena, rom fe-
rmentis aqtiuroba brinjis mcenareSi 2-3-jer metia, vidre
simindSi, ramac gamowvia fotosinTetikuri aqtiurobis da
mosavlianobis gazrda.
garda fotosinTezis procesSi uSualod monawile ge-
nebis klonirebisa da gamoyenebisa, identificirebul iqna
ujredSi qloroplastebis raodenobis makontrolebeli ge-
nebi. maTi gamoyeneba aseve gamoiwvevs fotosinTezis unaris
gazrdas. kidev sxva gzas warmoadgens yovel qloroplas-
tSi qlorofilis raodenobis gazrda. miRebul iqna qlo-
rofil a da b-s specifikurad damakavSirebeli modificire-
buli cilebi da naCvenebi iqna, rom maTi eqspresia trans-
genur mcenareebSi arsebiTad zrdis biomasas.
sxva mimarTulebas warmoadgens transgenur mcenareeb-
Si metabolizmis cvlilebis samuSaoebi. naxSirwylebis me-
tabolizmis regulaciaSi sakvanZo fermentis simindis saqa-
rozofosfatsinTetazas genis (SPS-geni) sxva mcenareebis
genomSi Seyvana iwvevda naxSirwylebis cvlis cvlilebas
da mcenareTa produqtiulobis amaRlebas. amJamad aseTi
transgenuri mcenareebi miRebulia pomidorSi, kartofilSi,
rafsSi, bambaSi.
dReisaTvis savele gamocdebi warmatebiT gaiara da sa-
soflo-sameurneo warmoebaSi iyeneben mosavlianobisa da
produqciis xarisxis amaRlebis ganmsazRvreli sxvadasxva
genebiT transformirebul simindis Svid da xorblis erT
transgenur jiSs.

182
2.9.6. gaumjobesebuli gemuri da sasaqonlo
Tvisebebis mqone transgenuri mcenareebis
miReba
transportirebisa da Senaxvis procesSi Mmemcenareobis
zogierT produqtis Senaxvisunarianoba yovelTvis iyo mni-
Svnelovani amocana da mis gadasawyvetad mimarTavdnen an
mosavlis adre aRebas (moumwifebeli nayofebis krefa) an
nayofebis damuSavebas eTilenis97 gamoyofis xelSemSleli
qimiuri preparatebiT. genetikur-inJinruli meTodebiT
problemis gadawyvetisas gamoyenebul iqna ramodenime xer-
xi.
erT-erTi mdgomareobs am genebis i-rnm-s matricidan k-
dnm-s miRebasa da konstruqciis SeqmnaSi, romelic mcenaris
ujredSi eqspresiisas iZleva „uSinaarso“ rnm-s. am gziT mi-
Rebul farTod cnobil pomidoris jiS „FLAVR SAVR“-is na-
yofebSi fermentis raodenoba Seadgenda mxolod 10%-s Cve-
ulebrivi jiSebis nayofebTan SedarebiT.
Semdeg xerxs warmoadgens eTilenis producirebis
uunaro mcenareebis miReba. amisaTvis Sewavlil iqna eTile-
nis sinTezis gza da dadgenil iqna, rom mis winamorbeds
warmoadgens S-adenozinmeTionini. ferment АСС-sinTetazas
moqmedebiT is gardaiqmneba 1-aminociklopropan-1-karbonis
mJavad (АСС), romelic Semdeg АСС-oqsidaziT ixliCeba nax-
SiorJangad, amiakad, WianWvela mJavad da eTilenad. Seqmnil
iqna genetikuri konstruqcia, romelic eqspresiisas iZleo-
da АСС-sinTetazas da АСС-oqsidazas „uSinaarso“ rnm-s.
mcenareSi maTi Seyvanisas adgili hqonda gacilebiT mcire
raodenobis eTilenis dagrovebas nayofebSi. garda amisa,
cnobilia tombaqteriebs gaaCniaT fermenti АСС-dezaminaza,
romelic АСС-s gardaqmnis α-ketobutiratad. aRniSnuli
fermentis baqteriuli geni 35S CaMV promotoris kontro-

97
nayofebis darbileba xdeba gamoyofili eTileniT gaaqtiurebuli mcena-
reuli celulazebis da poligalaqturonazas genebis moqmedebiT, romle-
bic moumwifebel nayofebSi ar aris aqtiuri.
183
lis qveS gadatanil iqna pomidoris genomSi da aseve mii-
Res mcenareebi nela mwifadi nayofebiT. es midgoma gamoiye-
nes sxva kulturebis mimarTac. am gziT miRebuli nesvis
jiSi ukve gamoiyeneba, xolo bananis transgenuri mcenaree-
bi gadian savele gamocdas.
Senaxvisunarianobis garda xil-bostneuli produqciis
mwarmoeblebisaTvis arsebiT faqtors warmoadgens gasayidi
saqonlis gemo da garegnoba. nayofebisaTvis ufro tkbili
gemos micemis erT-erT variants warmoadgens tropikuli
mcenaris dioskoreofilum kaminsis (Dioscoreophyllum cummin-
sii) nayofis cila monelinis genis gamoyeneba. es cila 3000-
jer ufro tkbilia Saqarze da 100000-jer – saqarozaze, mi-
si utilizeba xdeba bunebrivad, naxSirwyalbadovani cvlis
gziT, amitom potenciurad misaRebia diabetiT daavadebul-
Ta produqtebis warmoebaSi. magram cila Zalian aramdgra-
dia Termiulad, misi meoTxeuli struqtura advilad iSle-
ba organuli mJavebis sust xsnarebSi, ris Sedegadac kar-
gavs tkbil gemos. nedli saxiT gamoyenebuli mcenareebis
genomSi aRniSnuli cilis Seyvanis mizniT mecnierebma cila
monelinis makodirebeli geni agrobaqteriuli kointegraci-
uli veqtoruli sistemis saSualebiT gadaitanes pomidoris
genomSi Е8 cilis promotoris qveS, xolo salaTis genomSi
35S CaMV promotoris qveS. miRebuli transgenuri mcenaree-
bis foTlebi (salaTis) da mwife nayofebi (pomidoris) Sei-
cavda cila monelins.
mimdinareobs kvlevebi gemuri Tvisebebis gasumjobeseb-
lad sxva tkbili cilebis – taumatinebis gamoyenebis Sesa-
xeb. am mxriv gansakuTrebiT sayuradReboa afrikaSi mozar-
di tropikuli mcenaris taumatokokusis (Thaumatococcus dani-
ellii) gamoyeneba.
Kmemcenareobis kidev erT problemas warmoadgens meqa-
nikuri zemoqmedebis da Senaxvisas mcenareuli qsovilebis
feris Secvla, rac mkveTrad aqveiTebs produqtis sasaqon-
lo Rirebulebas. es xdeba polifenoloqsidazebiT katali-
zebuli monofenolebis da orTo-difenolebis orTo-qinone-
bad gardaqmnis Sedegad. polifenoloqsidazebis genebi lo-
184
kalizebulia birTvSi, maTi produqtebi ganlgebulia moto-
qondriebis da plastidebis membranebSi. kartofilis poli-
fenoloqsidazis „uSinaarso“ orientaciis k-dnm-s 35S CaMV
promotoris qveS kartofilis mcenareSi Seyvanis Sedegad
miRebul iqna transgenuri mcenare, romlis tuberebi gaci-
lebiT naklebad Savdeboda gaWrisas.
garegnoba gansakeTrebiT mniSvnelovania dekoraciul
da yvavilovan mcenareebSi. მraval farulTeslovan mcena-
reSi gvirgvinis Seferiloba ganpirobebulia specifikuri
pigmentebis – anTocianinebis sinTeziT. am pigmentebis saer-
To winamorbeds warmoadgens 4,2′,4′,6′-tetra-hidroqsil-hal-
koni, romelic warmoiqmneba ferment halkonsinTetazas sa-
SualebiT da aris yviTeli feris nivTiereba. Tu tetrahid-
roqsilhalkoni halkonizomerazas moqmedebiT gardaiqmneba
naringeninSi, xolo Semdeg flavanon-3-hidroqsilazas moq-
medebiT – dihidrokempferolSi, Seferiloba qreba, vinaidan
orive es nivTiereba uferoa. samagierod dihidrokempfero-
li sxvadasxva mcenaris ujredSi ferment dihidroflava-
nol-4-reduqtazas moqmedebiT SeiZleba gardaiqmnas, an
delfinidinis warmoebul lurji feris nivTierebad, an ci-
anidinis warmoebul wiTeli feris nivTierebad. amdenad
ferment dihidroflavanol-4-reduqtazas genis sxvadasxva
mcenareSi Seyvanisas SesaZlebelia sxvadasxva feris pro-
duqtis miReba. simindis genomidan dihidroflavanol-4-re-
duqtazas genis petunias genomSi gadataniT mecnierebma mii-
Res petuniasaTvis ucxo feris gvirgvinis furclebiani yva-
vili, ris gamoc transgenurma mcenarem garkveuli popula-
roba moipova.
genetikur-inJinruli gziT miRebul iqna TeTri, nazi-
vardisferi da nazi-cisferi qrizanTemas, titas, mixakis
mcenareebi. amisaTvis mcenareTa genomSi Seyvanil iqna hal-
konsinTetazas m-rnm-is k-dnm 35S CaMV promotoris qveS ro-
gorc pirdapiri, ise „uSinaarso“ orientaciiT. orive Sem-
TxvevaSi iTrguneboda fermentis sinTezi, magram sxvada-
sxva efeqturobiT. k-dnm-s pirdapiri orientaciiT CaSenebi-
sas mcirdeboda halkonsinTetazas produqcia, Sedegad mci-
185
rdeboda gvirgvinis furclebis Seferilobis intensioba.
k-dnm-s „uSinaarso“ orientaciiT CaSenebisas mTlian iTrgu-
neboda halkonsinTetazas sinTezi, Sedegad gvirgvinis fur-
clebis Seferiloba iyo absoluturad TeTri.
dekoraciul mcenareebSi genetikuri inJineriis miRwe-
vis kidev erT naTel magaliTs warmoadgens e.w. cisferi
vardebis miRebis istoria. vardis gakulturebidan TiTqmis
5 aTasi wlis manZilze veravin SeZlo cisferi vardebis mi-
Reba. is gaxda adamianis miuRwevlobis erTgvari sinonimi,
meyvavileebis Rirsebis saqme. belgiisa da didi britaneTis
mebaReobis sazogadoebam maRali jildoc ki daniSna cisfe-
ri vardis miRebisaTvis.
XX saukunis bolosaTvis dadginda, rom cisferi vardi
ar SeiZleba iyos, radganac mcenaris ujredebs ar gaaCniaT
anTocianuri pigmentis dihidrokempferolis anTocianur
pigment delfinidinad gardaqmnis fermentebi. magram Sem-
degSi dadginda, rom sxva mcenareebs gaaCniaT delfinidi-
nis ojaxis anTocianuri pigmentebis sinTezis ganmsazRvre-
li genebi, rac gaxda am mimarTulebiT genetikur-inJinruli
samuSaoebis warmoebis safuZveli.
1986 wels avstraliaSi Seiqmna bioteqnologiuri fir-
ma Calgene Pacific, romlis ZiriTadi komerciuli mizani iyo
swored cisferi vardebis Seqmna. 1994 wlidan firmas ewoda
Florigene. warmatebisken pirveli nabiji iyo 1991 wels petuni-
idan klonirebuli pigment delfinidinis geni. 90-iani wle-
bis SuaSi mkvlevarebma miiRes pirveli transgenuri mcenare
delfinidinis genis maRali eqspresiiT, magram mizani ver
iqna miRweuli, vinaidan samuSaoebi tardeboda vardis wi-
Tel pigmentian jiSze. gvirgvinis furclebSi warmoiqmne-
boda pigmentebis cianidinisa da delfinidinis narevi, rac
mas aniWebda mimzidvel muq-mewamul fers, magram ara cis-
fers.
sxva dekoraciul yvavilovan mcenareebze transforma-
ciuli samuSaoebis Catarebis dros firmis mkvlevarebma pe-
tunias delfinidinis geni gadaitanes TeTryvavilovan mi-
xakSi, romelSic mutirebuli iyo dihidroflavonilreduq-
186
tazas geni da miiRes mixakis mcenareebi cisferi da iisfe-
ri yvavilebiT. ris Sedegadac mecnierebma daiwyes mutire-
buli dihidroflavonilreduqtazas genis mqone vardis mce-
nareebis Zebna, magram aseTi ver ipoves. maSin gadawyvetil
iqna ganexorcielebinaT aRniSnuli genis „gamorTva“ vardis
ujredebSi misi Sebrinebuli orientaciis Tanmimdevrobebis
SeyvaniT.
samuSao dasrulebuli da problema gadawyvetili iq-
na aRniSnuli firmis iaponiis filialSi Suntory Flowers, sa-
dac ias mcenaridan klonirebuli delfinidinis genis, zam-
baxis mcenaridan klonirebuli dihidroflavanolreduqta-
zas genis da Sebrinebuli orientaciis vardis dihidrof-
lavanolreduqtazas genis vardis mcenareSi gadataniT 2005
wels miRebul iqna namdvili cisferi vardi.

2.9.7. gaumjobesebuli kve biTi da samrewvelo

Rirebulebis mqone transgenuri mcenareebis
miReba
adamianisa da cxovelebisaTvis ZiriTad sakveb nivTi-
erebas mcenareul ujredebSi warmoadgens naxSirwylebi
(saxamebeli, mono- da disaqaridebi), romlebic TiTqmis yve-
la mcenaris ujredSi erTnairia da aris mniSvnelovani ra-
odenobiT. cilebis raodenoba da miT ufro maTi aminomJa-
vuri Semadgenloba ar Seesabameba kvebis mrewvelobisa da
mecxoveleobis moTxovnebs, radganac mcenareuli warmoSo-
bis cilebSi arasakmarisadaa srulfasovani kvebisaTvis au-
cilebeli Seucvleli aminomJavebi.
cilebiT yvelaze mdidar organos mcenareSi warmoad-
gens Teslebi, sadac grovdeba e.w. samarago cilebi. dadgin-
da, rom Teslebis samarago cilebSi ama Tu im Seu-cvleli
aminomJavis Semcveloba saxeobrivi niSania da kontroldeba
Sesabamisi genebis erTobliobiT. es gaxda gaumjobesebuli
Sedgenilobis samarago cilebis mqone mcenareTa miRebis
strategiuli midgomebis SemuSavebis safuZveli.
erT-erTi midgoma aris sxvadasxva saxeobis mcenareTa
samarago cilebis genebis gaerTianeba erT mcenareSi (anu
187
aminomJavuri Sedgenilobis mixedviT srulfasovani sxva
mcenareebis axali cilebis genebis Seyvana ujredSi). aseTi
midgomis principuli SesaZlebloba demonstrirebul iqna
jer kidev gasuli saukunis 80-ian wlebSi, roca parkosanTa
samarago cila fazeolinis geni gadaitanes Tambaqos ujre-
debSi, xolo simindis samarago cila zeinis geni – mzesum-
ziras mcenareSi.
meore midgoma gulisxmobs ama Tu im mcenaris sakuTa-
ri cilebis produqciis gazrdas maTi genebis ufro mZlav-
ri promotoris qveS dayenebiT.
mesame midgoma iTvaliswinebs samarago cilebis struq-
turuli genebis Secvlas maTSi Seucvleli aminomJavebis
axali kodonebis CarTvis gziT, anu samaragi cilebis ami-
nomJavuri Sedgenilobis gaumjobesebas. am mizniT in vitro mu-
tagenezis gziT samarago cilis genis TanmimdevrobaSi Seh-
yavT Sesabamisi Seucvleli aminomJavas kodonebi. polipep-
tidur jaWvSi aseTi cvlilebis Setanis saukeTeso ad-gils
warmoadgens C-bolos e.w. hipervariabeluri ubani. Tumca
aseTi genebis eqspresiaSi jer kidev arsebobs garkveuli
problemebi. samarago cilebis aminomJavuri Sedgenilobis
gaumjobeseba SeiZleba aseve sxva gziTac, kerZod, Seucvle-
li amonomJavebis saerTo raodenobis gazrdiT. Seucvleli
aminimJavebis nawili (izoleicini, treonini, meTionini da
lizini) sinTezirdeba aspartatidan da procesis ZiriTad
fermentebs warmoadgenen aspartatkinaza da dihidrodipiko-
linatsintaza. orive fermentis aqtiuroba regulirdeba
liziniT ukukavSiris principiT. Corynebacterium (dapA)-dan ga-
moyofili dihidrodipikolinatsintazas geni da Escherichia
(lysCM4)-dan gamoyofili aspartatkinazas geni klonirebul
da mierTebul iqna bardas rbfk-as98 mcire subnawilakis ge-
nis liderul TanmimdevrobasTan da Semdeg orivesTan da-
kavSirebul iqna parkosanTa cila fazeolinis genis promo-

98
ribulozo-1,5-bifosfat-karboqsilaza – fotosinTezis sakvanZo fermenti.

188
tori da transkripciis terminatori. orive genetikuri
konstruqcia agrobaqteriuli transformaciis gziT Seiyva-
nes soios da rafsis genomSi. miRebul trans-genuri mcena-
reebis ujredebSi lizinis Semcveloba iyo 100-jer meti,
vidre sawyis mcenareebSi da rac mTavaria, soios samaragi
cilaSi lizinis Semcveloba gaizarda 5-jer, xolo rafsis
TeslebSi – 2-jer. aseT soios didi perspeqtivebi gaaCnia
mecxoveleobaSi iafi kobinirebuli sakvebis dasamzadeb-
lad.
sasoflo-sameurneo mcenareTa kvebiTi Rirebulebis
gaumjobesebis kidev erT magaliTs warmoadgens e.w. “oqros
brinjis” Seqmna. 1999 wels brinjis mcenaris ujredebi tran-
sformirebul iqna genetikuri konstruqciiT, romelic Sei-
cavda: 1) brinjis glutelinis genis promotors, abusa-la-
Tinis katalazas genis pirvel introns, bardas rbfk-as
mcire subnawilakis genis liderul Tanmimdevrobas, baqte-
ria Erwinia uredovora-s karotindesaturazas gens crtI da no-pa-
linsinTetazas genis terminators; 2) glutelinis genis
promotors, abusalaTinis katalazas genis pirvel introns,
nargizisfitoinsintazas gens da nopalinsinTetazas genis
terminators. aseTi mcenareebi Teslis endospermSi yovel 1
gramze asinTezebdnen da agrovebdnen 1.6 mkg karotinoidebs.
marcvlebs hqondaT yviTeli feri, saidanac warmoiSva maTi
pirvandeli saxelwodeba. Tumca unda aRiniSnos, rom gada-
taniTi mniSvnelobiT es brinji marTlac aris oqros mniS-
vnelobis. vinaidan Seicavs provitamin A-s, romelic Cveu-
lebriv deficituria aziis calkeuli qveynebis Raribi mo-
saxleobis sakvebSi, sadac farTodaa gavrcelebuli provi-
tamin A-s naklebobasTan dakavSirebuli, Seuqcevadi sibr-
mavis gamomwvevi, Tvalis daavadeba. Semdgom wlebSi axali
genetikuri manipulaciebiT es brinji kidev ufro gaumjo-
besebuli iqna.
miRebul iqna kartofilis pirveli mcenareebi saxameb-
lis gaumjobesebuli xarisxobrivi maxasiaTeblebiT. Cveu-
lebriv saxamebeli Sedgeba amilopeqtinisa da amilozasa-
gan. miT ufro xarisxiania saxamebeli, rac met amilopeq-
189
tins Seicavs. Sebrunebuli orientaciis amilozas genis
Zlieri promotoris qveS kartofilis genomSi SeyvaniT mi-
Rebul mcenareSi amilozas sinTezi daeca nulamde. “uSina-
arso” rnm komplementarulad ukavSirdeba amilozas norma-
luri genis rnm-s da xels uSlis mis translacias.
kvebis da parfiumeriul mrewvelobaSi farTod gamoi-
yeneba mcenareuli cximebi. maTi gamoyenebis xasiaTi da mo-
culoba didadaa damokidebuli zeTSi Semavali cximmJave-
bis raodenobriv da xarisxobriv Sedgenilobaze.
ZiriTad cximzeTovan mcenareebs warmoadgens soio, pa-
lmebi – Cocos nucifera, Elaeis guineensis, Orbignya barbosiana, rafsi
da mzesumzira.
sxvadasxva mcenaris zeTebis cximmJavebis Sed-geniloba
erTnairi ar aris, magram umetesi zeTovani kultura iZleva
zeTebs palmitinis, stearinis, oleinis, lonolinis, lino-
leinis mJaviT. sxva cximmJavebi (kaprili, kaprilinis, miris-
tinis da sxva) naklebad warmoiqmneba.
imis mixedviT, Tu romeli cximmJava monawileobs zeTis
SedgenilobaSi, maT sxvadasxva gamoyeneba aqvT – sakvebi,
kosmetikuri an teqnikuri. amitom mrewvelobis sxvadasxva
dargisaTvis sxvadasxva mcenareebis moyvanaa saWiro. nawili
maTgan sakmaod Znelad kultivirebadia (mag. merqnianebi) iz-
rdeba mxolod garkveul klimatur pirobebSi. amitom bio-
teqnologiuri firmebis pirveladi amocanaa cximmJavebis
Sevlili Sedgenilobis mcenareebis miReba. am sferoSi li-
der amerikul korporacia Calgene-Si Seqmnilia rafsis ramo-
denime jiSi samrewvelo gamoyenebisaTvis. isini erTmaneTi-
sagan gansxvavdebian zeTSi konkretuli cximmJavas Semcve-
lobiT da Sesabamisad maT sxvadasxva gamoyeneba aqvT. maga-
liTad, jiSi 40% stearinis mJavas SemcvelobiT gamoiyeneba
margarinisa da Sokoladis zeTis warmoebaSi, 40% da 60%
laurinis mJavas SemcvelobiT – kosmetikuri da teqnikuri
gamoyenebis detergentebis warmoebaSi, 80% oleinis mJavas
SemcvelobiT – kvebis produqtebis, sacxebi saSualebebis,
melnis warmoebaSi da a.S.

190
 Seqmnilia rafsis jiSi unikaluri cvilis – xoxobas
SemcvelobiT, romelic arc erT mcenares ara aqvs da veSa-
pebze nadirobis akrZalvis Semdeg warmoadgens praqtiku-
lad erTaderT wyaros samedicino saSualebebis, avtomobi-
lebis da TviTmfrinavebis gansakuTrebuli sacxebi masa-
lis, transformatorebis zeTebis da plastmasebis warmoe-
baSi. ramdeni da romeli genebia gadatanili am mcenaris Se-
saqmnelad ucnobia – informacia ar vrceldeba.
soios mcenareSi igive soios ferment delta-12 desa-
turazas genis damatebiTi aslebis Seyvanis Sedegad zeTSi
Semcirda linolis da linolenis mJavas Semcveloba da ga-
izarda oleinis mJavas Semcveloba. aseTi zeTi iTvleba uf-
ro Zvirfasad, vinaidan naklebad iSleba gacxelebisas da
ar sqeldeba Senaxvisas.
Mmimdinareobs kvlevebi bambis boWkos teqnologiuri ma-
xasiaTeblebis gaumjobesebis mimarTulebiT. am mizniT bam-
bis mcenaris plastidur genomSi Seyvanil iqna bunebrivad
degradirebadi polimeris – poli-β-hidroqsilbutiratis
sinTezisaTvis aucilebeli genebi. transgenuri mcenare Sei-
cavda 30%-mde polimerul danamats. kidev erT gzas bambis
boWkos gasaumjibeseblad warmoadgens bambis mcenareSi qi-
tinis da qitinazis msgavsi monomerebis makodirebeli gene-
bis Seyvana da maTi celulozas boWkos struqturaSi Car-
Tva, romlis realizacia mimdinareobs belgiaSi 2010 wli-
dan.
sasoflo-sameurneo warmoebisaTvis mniSvnelovani prob-
lemis genetikur-inJinruli gadawyvetis magaliTs warmo-
adegens heterozisis SemTxveva. cnobilia, rom mravali sa-
soflo-sameurneo kulturis (simindi, kitri, pomidori, xo-
rbali da sxva) saTesle masalas (e.w. hibriduli Teslebi)
Rebuloben specialur meTesleobis meurneobebSi ori sxva-
dasxva jiSis an xazis Sejvarebis Sedegad. aseTi Sejvare-
biT miRebuli F1 Taobis mcenareebis Teslebi iZlevian gaci-
lebiT maRalmosavlian heterozisul mcenareebs. magram ase-
Ti Teslebis samrewvelo masStabiT miReba gaZnelebulia
TviTdamtvervis SesaZleblobiT.
191
 XX saukunis 20-iani wlebidan problemis gadasawyvetad
iyeneben citiplazmatur mamrobiT sterilobas (cms), rac
warmoiqmneba zogierT mcenareSi birTvuli da citoplazma-
turi genomebis Seusabamobis Sedegad. Tumca meTodis gamo-
yenebam simindis kulturaSi aCvena, rom igi periodulad iZ-
leva “Cavardnebs”, anu Teslebis garkveuli (zogjer mniS-
vnelovani) raodenoba ar aris hibriduli, an miiReba mosa-
lodnelze naklebi raodenobis Teslebi.
garantirebuli sicocxlis uunaro mtvrianebiani mcena-
reebis misaRebad yvelaze warmatebuli aRmoCnda genetikuri
konstruqciebi baqteria Bacillus amyloliquefaciens-is genebis ga-
moyenebiT. am gziT miRebuli damtvervis kontrolis siste-
miani rafsis mcenare, aseve warmoadgens saukeTesos biou-
safrTxoebis TvalsazrisiTac.
aq gamoyenebul genetikur konstruqciaSi seleqtiur
markerad warmodgenilia ara tradiciuli kanamicinrezis-
tentobis geni aramed Streptomyces-is gvaris aqtinomicetidan
gamoyofili herbicid fosfinotricinis gamZleobis geni,
romelsac hqvia bar da akodirebs ferment fosfinotricin-N-
acetiltransferazas.
amiT ori kargi saqme gakeTda: mcenarem axali sasargeb-
lo Tviseba (herbicidgamZleoba) SeiZina da moixsna sakama-
To sakiTxi antibiotikrezistentuli genebis gavrcelebis
Sesaxeb. konstruqciaSi ZiriTad gens warmoadgens baqteria
Bacillus amyloliquefaciens-dan gamoyofili, Tambaqos qsovilspe-
cifikur genis ТА29 promotorTan dakavSirebuli ribonuk-
leazas (rnmaza)99 geni, romelic aqtiuria mxolod mtvria-
nas qsovilebSi.
aseTi mcenare normaluria da mis yvela ujredSi, gar-
da mtvris marcvlis kedlis ujredebisa, baqteriuli ribo-
nukleaza ar sinTezirdeba da rnm ar iSleba. orive genis
Semcvelma konstruqciam miiRo saxelwodeba – barnase. magram

99
rnm-s degradaciis makatalizebeli fermenti-nukleaza.

192
im SemTxvevaSi, roca pirveli Taobis hibridebma unda war-
moqmnan Teslebi, aucilebelia fertilurobis aRdgena, ami-
tom meTesleobaSi rafsis heterozisuli hibridebis misa-
Rebad Rebuloben xazebs, romelTa genomSi Seyvanilia gene-
tikuri konstruqcia saxelwodebiT barstar. igi barnase kon-
struqciisagan gansxvavdeba mxolod imiT, rom masSi ribo-
nukleazas makodirebeli Tanmimdevrobis magivrad Seicavs
igive baqteria Bacillus amyloliquefaciens-dan gamoyofil ribo-
nukleazas inhibitoris genis Tanmimdevrobas. barnase trans-
genuri mcenaris barstar transgenuri mcenaris mtvris mar-
cvliT damtverianebisas genetikuri konstruqciebi erT ge--
nomSi gaerTiandeba da heterozisuli mcenaris mtvrianas
qsovilebis ujredebSi ribonukleazas aqtiuroba daiTrgu-
neba.
amJamad barnase/barstar meTods farTod iyeneben rafsis
jiSebidan komerciuli Teslebis misaRebad.

2.9.8. samedicino preparatebis producenti

transgenuri mcenareebis miReba
bolo wlebSi aqtiurad mimdinareobs genetikur-inJin-
ruli samuSaoebi uSualod sakvebSi gamoyenebadi an cilo-
vani produqtidan gamoyofili da Semdgom gawmendili sa-
xiT gamoyenebadi samedicino daniSnulebis cilebis eqspre-
sirebadi transgenuri mcenareebis misaRebad.
pirveli Sedegebi samkurnalo cilebis sinTezirebadi
mcenareebis Seqmnis saqmeSi miRebul iqna gasuli saukunis
bolos imunoglobulin IgG1-s eqspresirebadi Tambaqos tra-
nsgenuri mcenaris SeqmniT.
dReisaTvis miRebulia mravali transgenuri mcenare,
romlebic asinTezeben samkurnalo daniSnulebis sxvadasxva
cilebs, maT Soris veterinariaSi gamoyenebul somatotro-
pins (Tambaqo), herpersis virusis sawinaaRmdego antisxeu-
lebs (Tambaqo, simindi, soio), B da C hepatitis samkurnalo
interferonebs (brinji, turnefsi), antimikrobuli moqmede-

193
bis laqtoferins (kartofili), trombozebis samkurnalo
girudins (rafsi), protein C-s (Tambaqo) da sxva.
calke interess iwvevs saWmeli vaqcinebis misaRebad
aucilebeli imunizaciis antigenebis maproducirebeli tra-
nsgenuri mcenareebis miReba.
pirveli saWmeli vaqcina iyo B hepatitis sawinaaRmdego
vaqcina, romelic miRebul iqna jer Tambaqos mcenareSi,
Semdeg – kartofilSi. Tagvebis sakvebSi aseTi kartofilis
damatebiT izrdeboda imuniteti aRniSnuli daavadebis ga-
momwvevi virusis winaaRmdeg. SemdegSi miRebul iqna hepati-
tis sawinaaRmdego vaqcina salaTSi da xanWkolaSi.
amJamad ukve Seqmnilia da klinikuri gamocda gaiara
transgenuri mcenareebidan miRebulma rigma vaqcinam, maT
Soris gastroenteritis (Tambaqo, kartofili), qoleris vib-
rionis (kartofili), citomegalovirusis (kartofili, Tam-
baqo), papilomis virusis (kartofili) da sxva sawinaaRmde-
go vaqcinam. umetes SemTxvevaSi am cilebis eqspresireba-di
saWmeli transgenuri mcenareebiT kvebisas viTardeboda spe-
cifikuri imunuri reaqcia.
iaponiaSi tardeba klinikuri gamocda antialergiuli
transgenuri brinjis monawileobiT. muSavdeba ganviTarebad
qveynebSi farTod gavrcelebuli bavSvTa nawlavebis infeq-
ciuri daavadebebis gamomwvevebis enterobaqteriebisa da
vibrionebis sawinaaRmdego „saWmeli vaqcinebis“ Semcveli
transgenuri mcenareebis (banani da sxva) Seqmnis teqnolo-
giebi.
dReisaTvis samedicino daniSnulebis saWmeli mcenareu-
li cilebis, vaqcinebis, antisxeulebis miReba warmoadgens
genetikuri inJineriis perspeqtiul mimarTulebas.

194
 2.9.9. transgenuri mcenareebis gavrcelebis
Tanamedrove mdgomareoba
pirveli genetikurad modificirebuli (transgenuri)
jiSebi warmoebaSi gaCnda 1992 wels. ganvlil periodSi,
tradiciuli seleqciis meTodebiT Seqmnil jiSebTan Seda-
rebiT, am mcenareebma aCvenes maRali efeqturoba, ris gamoc
maT qveS dakavebuli farTobebi Zalian swrafad gaizarda.
ISAAA100 -s monacemebiT (cxrili6), Tu 1996 wels am kulture-

tran sgen uri mcen areebi s qveS dakavebuli

farTobebi heqtarSi da akrSi (1996-2017 w . w . )
cx r ili 6
wel i h eqtari ( mln) akri (mln)
1996 1.7 4.2
1997 11.0 27.2
1998 27.8 68.7
1999 3 9.9 98.6
2000 44.2 109.2
2001 52.6 13 0.0
2002 58.7 145.0
2003 67.7 167.3
2004 81.0 200.2
2005 90.0 222.4
2006 102.0 252.0
2007 114.3 282.4
2008 125.0 3 08.9
2009 13 4.0 3 3 1.1
2010 148.0 3 65.7
2011 160.0 3 95.4
2012 170.3 420.8
2013 175.2 43 2.9
2014 181.5 448.5
2015 179.7 444.0
2016 185.1 457.4
2 0 17 189.8 469.0
j ami 23 39.5 5780.0
bis qveS dakavebuli farTobebi Seadgenda 1.7 mln. ha-s (4.2
mln. akrs), 2016 wlisaTvis igi gaizarda 185.1 mln. ha-mde

100
ISAAA - International Service for the Acquisition of Agri-biotech Applications – agrobi-
oteqnologiuri aplikaciebis Sesyidvis saerTaSoriso samsaxuri.

195
(457.4 mln.akri), rac kultivirebadi nayofieri miwebis 12%-
Seadgenda, xolo 2017 wlisaTvis man miaRwia 189.8 mln. ha-s
(469.0 mln. akrs), anu kidev gaizarda 3%-iT (4.7 mln. ha).
transgenuri mcenareebis warmoebaSi lideria aSS. amJa-
mad aseTi mcenareebis warmoebaSi danergvis maRali tempe-
biT gamoirCevian ganviTarebadi qveynebi. 2012 wlidan dawye-
buli (sur. 60) transgenuri mcenareebis qveS dakavebuli
farTobebis mixedviT ganviTerebadi qveynebi (100.6 mln. ha –
2017 w.) uswreben industriulad ganviTarebul qveynebs (89.2
mln. ha – 2017 w.). daaxloebiT 18 milioni fermeridan 90%-ze
meti modis ganviTarebadi qveynebis mcire meurneobebze.

200

180

160

140

120

100

80

60

40

20

1996 2002 2009 2017

Total Industrial Developing

wyaro: ISAAA, 2017

s ur. 60. tr ans g e nur i m ce nar e e bis f ar To bi s zr d a

ind us tr iul d a g anv i Tar e bad q v e y ne bS i
(Total – s a e rTo farTo bi; Industrial – i n dustriulad ganviTa r e b u l i q v e y n e b i ;
Developing – g a n v i T a r e a d i q v e y n e b i )

2017 wlis bolos transgenuri mcenareebis qveS yve-

laze meti farTobi dakavebuli iyo Semdeg qveynebSi:

196
 qveyan a farTobi , kul tura
№
mln . ha
simindi, soio, bamba, rafsi,
1 aSS 75,0 Saqris Warxali, ionja, papa-
ia, gogra, kartofili
2 brazilia 50,2 soio, simindi, bamba
3 argentina 23,6 soio, simindi, bamba
rafsi, simindi, soio, Saqris
4 kanada 13,1
Warxali
5 indoeTi 11,4 bamba
6 paragvai 3,0 soio, simindi, bamba
7 pakistani 3,0 bamba
8 CineTi 2,8 bamba, papaia, wiwaka
saamxreT afri-
9 2,7 simindi, soio, bamba
kis respublika
10 bolivia 1,3 soio, simindi

ZiriTad transgenur kulturebs warmoadgenen soio, si-

mindi, bamba da rafsi (sur. 61). aseve mniSvnelovani moTxov-
100

90

80

70

60

50

40

30

20

10

1996 2001 2007 2013 2017

soio simindi bamba rafsi

sur. 61. ZiriTadi t ransgenuri kulturebi msofli oSi (mln. ha)

wyaro: ISAAA, 2017

197
nilebaa kartofilis mimarT. 2017 wlis monacemebiT trans-
genur soios ekava mTeli soios qveS dakavebuli farTobe-
bis 77% (94.1 mln. ha), transgenur siminds – simindis far-
Tobebis 32% (59.7 mln. ha), transgenur bambas – mTeli bam-
bis plantaciebis 80% (24.1 mln. ha), xolo transgenur rafs
– 30% (10.2 mln. ha).
transgenuri mcenareebidan yvelaze ufro gavrcelebu-
lia herbicidgamZle, mavne-mwerebisadmi gamZle da orive
Tvisebis matarebeli mcenareebi (sur. 62).
120

100

80

60

40

20

1996 2001 2007 2013 2017

sur. 62. t ransgenuri mcenareebis gavrceleba genetikur-inJinruli
Tvisebebis mixedviT (mln. ha)
( h erb i ci d g amZl e mwereb i s ad mi g amZl e ori ve T vi s eb a)
wyaro: ISAAA, 2017

2017 wlis monacemebiT transgenuri mcenareebi mohyavT

yvela kontinentze, antarqtidis garda (sur. 63). 2017 wels
es kulturebis mohyavdaT 24 qveyanaSi 101, sadac cxovrobs
dedamiwis mosaxleobis ~ 60%, anu ~ 4 mlrd. adamiani. am
qveynebidan 18 gadavida mega-agrobioteqnologiuri qveynebis
ricxvSi, anu qveynebisa, sadac transgenuri mcenareebis qveS
dakavebuli farTobebi aRemateba 50000 heqtars (sur. 63). me-

101 rekorduli iyo 2015 weli – 28 qveyana.

198
 №4 kanada №22 p ortu- №1 5 espaneTi №26 C exeTi №25 slovake- №27 burkina №7 p akistani
1 3 .1 mln. h a g alia 0.1 mln. ha ar a Ti f as o ara 3 .0 mln. h a
ra f si, simindi < 0.05 mln. ha a ra №28 r umine-
s oio, Warxali s i mindi s i mindi T i ara b a m ba

№1 aSS №8 C ineTi
75 .0 mln. ha 2.8 m ln. ha
s i mindi, soio,
b a m ba, rafsi, b a m ba, papaia,
W a rxali.i onja, w i waka
p a paia, gogra,
k a rtofili
№5 i ndoeTi
№23 b angla- 1 1 .4 mln. ha
d eSi b a m ba
< 0.05 mln. ha
b a drijani

№1 4 mianmari
№1 7 meqsika 0.3 mln. h a
0.1 mln. ha b a m ba
b a m ba, soio

№1 3 filipi-
№20 h ondu- n ebi
r asi 0.8 m ln. ha
< 0.05 mln. ha s i mindi
s i mindi

№1 2 avstra-
№24kostarika li a
< 0.05 mln. ha 0.9 m ln. ha
b a m ba, soio, b a m ba, rafsi
a n anasi

№1 8 kolum- №1 6 sudani
bia 0.2 m ln. ha
0.1 mln. ha b a m ba
b a m ba, simindi

№1 1 bolivia №1 9 vietnami
1 .3 mln. ha < 0.05 mln. ha
s oio s i mindi

№21 Cile №6 p ar agvai №3 argentina №1 0 urugvai №2 brazilia №9 s . afrika

< 0.05 mln. ha 3 .0 mln. h a 23 .6 mln. ha 1 .1 mln. ha 5 0.2 mln. ha 2.7 m ln. ha
s oio. simindi, s oio. simindi, s oio. simindi, s i mindi, soio.
s i mindi b a m ba b a m ba s oio. simindi b a m ba b a m ba

1 8 m ega-qveyana, r omlebsac t ransgenuri mcenareebi m ohyavT 50000 heqtarze met f arTobze ..

s ur. 63. tr ansgenur i m ce nar e e bis g av r ce l e bis g e o g r af ia d a

m war m o e be l i q v e y ane bi 2 01 7 we l s

urneobis am sferoSi dasaqmebulia daaxloebiT 18 milioni

fermeri.
2016 wels komerciuli warmoebisaTvis daSvebuli iyo
29 kulturis transgenuri mcenareebis 404 jiSi, maT Soris,
356 – sasursaTo kulturebis, 3 – xe-mcenaris, 22 – xexilo-
vani kulturebis da 23 – dekoraciuli mcenareebis (sur. 64).
SemdgomSi mosalodnelia am monacemebis gaumjobeseba.
G genetikuri transformaciis gziT miRebul yvelaze gav-
199
 tran s g en u ri
ji S i

wyaro: ISAAA, 2016

s ur . 64 . 2 01 6 wl is bo l o s s am r e wv e l o g am o y e ne bis aTv is
d aS v e bul i tr ans g e nur i m ce nar e e bis jiS e bi
(404 t ransgenuri jiSi, maT Soris: 3 56 – sasursaTo kul t ur a; 3 – xe-
mc en are; 22 – xexi l ovan i ; 23 – dekorac i ul i )

rcelebul transgenur mcenareTa jiSebis mokle daxasiaTe-

ba motanilia cxrilebSi 7 – 11.
transgenuri mcenareebis qveS farTobebis swrafi zrda
TavisTavad ukve warmoadgens Cveulebriv jiSebze maTi upi-
ratesobis naTel dadasturebas. cifrebSi es naCvenebia
ISAAA-s monacemebSi. 1996 wlidan 2016 wlis CaTvliT trans-
genuri mcenareebis kultivirebam moitana: sasoflo-sameur-
neo produqciis realizaciidan miRebuli Semosavlis gaz-
rda 150 mlrd. amerikuli dolaris odenobiT; 583,5 mln. kg
sxvadasxva pesticidis ekonomia, ramac arsebiTad Seamcira
araxelsayreli zemoqmedeba garemoze; mxolod 2011 wels
atmosferoSi СО2-is gamofrqveva sasoflo-sameurneo teqni-
kis naklebi gamoyenebis gamo, Semcirda 23,2 mlrd.kg-iT, rac
200
praqtikaSi gamoyen ebul tran sgen ur mcen areTa ji Sebi s
mokle daxasi aTeba
(ojaxi: gogrisebrTa, papaiasebrTa, parkosanTa, nacarqaTamasebrTa)
cx r ili 7
kultu- S e yvanil i cil e bi
g e nis wy a r o S e Ze nil i niS ani
ra (g e ne bi)
ne s v i dezamina 1-amino- Pseudomonas momwifebis vadebis dag-
(Cucumis ciklopropan-1-ami- morganii rZeleba mcenareul hor-
melo) nokarbonis mJava mon eTilenis winamor-be-
dis degradaciis Sede-gad
y abay i kitris mozaikis vi-kitris mozai- sazamTros mozaikis viru-
(Cucur- rusis garsis pro- kis kukumovi- sisa da kitris mozaikis
bita pepo teini rusi virusisadmi gamZleoba
var. gi-
raumon- cukinis yviTeli cukinis yvi- sazamTros mozaikis vi-
tia) mozaikis virusis Teli mozai- rusisa da cukinis yviTe-
garsis proteini kis virusi li mozaikis virusi sad-
mi gamZleoba
p ap aia papaias rgoluri papaias rgo- papaias rgoluri laqia-
(nesv is laqianobis viru- luri laqia- nobis virusisa da sxva vi-
xe) sis garsis prote- nobis virusi rusebisadmi gamZleoba
(Carica ini
papaya)
fosfinotricin Streptomyces fosfinotricinuli her-
N-acetiltransfe- hygroscopicus bicidebisadmi (amoniumis
raza (bar) glufozinati) gamZleoba
5-enolpi-Sikomat-3- Agrobacterium herbicid glifosatisadmi
fosfatsintazas tumefaciens CP4 gamZleoba
s o io mutanturi geni
(Glycine TeslebSi cximmJavebis
max) delta-12 desatura- Glycine max Semcvelobis modifikacia
zas geni (oleinis mJavas maRali
Semcveloba)
TeslebSi cximmJavebis
delta-12 desatura- Glycine max Semcvelobis modifikacia
zas geni (linolenis mJavas dabali
Semcveloba)
fosfinotricin Streptomyces fosfinotricinuli her-
S aqr is N-acetiltransfe- hygroscopicus bicidebisadmi (amoniumis
W ar x a- raza (bar) glufozinati) gamZleoba
li
(Beta 5-enolpi-Sikomat-3- herbicid glifosatisadmi
vulgaris) fosfatsintazas Escherichia coli gamZleoba
mutanturi geni
(aroA)

201
praqtikaSi gamoyen ebul tran sgen ur mcen areTa ji Sebi s
mokle daxasi aTeba
(ojaxi: marcvlovanTa)
cx r ili 8
kultu- S e yvanil i cil e bi
g e nis wy a r o S e Ze nil i niS ani
ra (g e ne bi)
Ddelta-endotoqsi- Bacillus thu- simindis fesvis Wiasadmi
ni ringiensis gamZleoba
Acetollaqtat-sin- Arabidopsis imidazolinuri herbicide-
Tetaza thaliana bisadmi gamZleoba
delta-endotoqsini Bacillus thurin- simindis mavne mwerisadmi
+ giensis gamZleoba
glifosat oqsid- + +
reduqtaza Ochrobactrum Gglifosatisadmi gamZleo-
anthropi ba
5-enolpiruvil-Si- herbicid glifosatisad-
kimat-3-fosfatsin- Zea mays mi gamZleoba
s imindi tazas geni CaM V
(Zea promotoris qveS
mays) imidizolinuri herbicide-
acetolaqtat-sin- Nicotiana taba- bisadmi gamZleoba (imaze-
taza cum tapiri)

fosfinotricin N- Streptomyces fosfinotricinuli her-bi-

acetiltransfe-ra- hygroscopicus cidebisadmi (amoniumis
za (bar) .+ glufozinati) gamZleoba
+ Bacillus amyloli- +
rnmaza quefaciens damtvervis sistemis kon-
troli: mamrobiTi steri-
loba
delta-endotoqsiniBacillus thuringi- simindis mavne mwerisadmi
+ ensis gamZleoba
fosfinotricin N- + +
acetiltransfe-ra- Streptomyces fosfinotricinuli her-bi-
za (bar) hygroscopicus cidebisadmi (amoniumis
glufozinati) gamZleoba
fosfinotricin N- Streptomyces fosfinotricinuli her-bi-
acetiltransfe-ra- hygroscopicus cidebisadmi (amoniumis
za (bar) glufozinati) gamZleoba
fosfinotricin N- Streptomyces fosfinotricinuli her-bi-
br inji acetiltransfe-ra- viridochromoge- cidebisadmi (amoniumis
(Oryza za nes glufozinati) gamZleoba
sativa) acetolaqtat-sin- Arabidopsis imidizolinuri herbicide-
taza thaliana bisadmi gamZleoba

202
praqtikaSi gamoyen ebul tran sgen ur mcen areTa ji Sebi s
mokle daxasi aTeba
(ojaxi: kombostosebrTa)
cx r ili 9
S e yvanili cilebi
kultura g e nis wy a r o S e Ze nil i niS ani
(g e ne bi)
fosfinotricin N- Streptomyces fosfinotricinuli her-
acetiltransfe-ra- hygroscopicus bicidebisadmi (amoniumis
za (bar) glufozinati) gamZleoba
kaliforniuli cximmJavebis Semcvelobis
dafnis xe Um- modificireba TeslebSi,
Tioesteraza bellularia californi- gansakuTrebiT laurinis
ca da miristinis mJavebis
maRali done
Klebsiella pneu- oqsilinuri herbicidebi-
nitrolaza moniae subsp.oza- sadmi (bromoqsinilis da
r af s i enae ioqsinilis CaTvliT) gam-
ar genti-
Zleoba
nul i glifosat oqsido- Ochrobactrum Hherbicid glifosatisa-
(Brassica
reduqtaza anthropi dmi gamZleoba
napus)
fosfinotricin N- Streptomyces fosfinotricinuli her-
acetiltransfe-ra- hygroscopicus bicidebisadmi (amoniumis
za (bar) + glufozinati) gamZleoba
+ Bacillus amyloli- +
rnmaza quefaciens damtvervis sistemis kon-
troli: mamrobiTi steri-
loba
fosfinotricinuli her-
fosfinotricin N- Streptomyces bicidebisadmi (amoniumis
acetiltransfe-ra- hygroscopicus glufozinati) gamZleoba
za (bar) + +
+ Bacillus amyloli- damtvervis sistemis kon-
rnmazas inhibitori quefaciens troli: fertilobis aR-
dgena
r af s i fosfinotricin N- Streptomyces fosfinotricinuli her-
p o lonu- acetiltransfe-ra- hygroscopicus bicidebisadmi (amoniumis
r i (tur - za (bar) glufozinati) gamZleoba
ne f s i)
(Brassica glifosat oqsido- Ochrobactrum Hherbicid glifosatisa-
rapa) reduqtaza anthropi dmi gamZleoba

203
praqtikaSi gamoyen ebul tran sgen ur mcen areTa ji Sebi s
mokle daxasi aTeba
(ojaxi: balbisebrTa, selisebrTa, mixakisebrTa)
cx r ili 1 0
kul tu- S e yvanil i cil e bi
g e nis wy a r o S e Ze nil i niS ani
ra (g e ne bi)
proteinaze bis inhi- Solanum tuberosum bambis mavne-mwerebisad-
bitor i + mi gamZleoba
+ Bacillus thuringien -
delta-endotoqsini sis

sulfonilSardovanas
acetolaqtat-sin- Nicotiana tabacum herbicidebisadmi (tria-
Tetaza sulfuroni da meTsul-
bam ba furonmeTili) gamZleo-
(Gossypi- ba
um hirsu- Klebsiella pneumoniae oqsilinuri herbicide-
tum) nitralaza subsp.ozaenae bisadmi (bromoqsinilis
da ioqsinilis CaTvl-
iT) gamZleoba
Bacillus thuringien - bambis mavne-mwerebisad-
delta-endotoqsini sis mi gamZleoba
+ + +
nitralaza Klebsiella pneumo- oqsilinuri herbicide-
niae subsp.ozaenae bisadmi (bromoqsinilis
da ioqsinilis CaTvl-
iT) gamZleoba
fosfinotricin N- Streptomyces fosfinotricinuli
acetiltransfe-ra- hygroscopicus herbicidebisadmi (amo-
za (bar) niumis glufozinati)
gamZleoba
5-enolpi-Sikomat-3- Hherbicid glifosati-
fosfatsintazas Escherichia coli sadmi gamZleoba
mutanturi geni
(aroA)
s e li (Li- sulfonilSardovanas
num usita- acetolaqtat-sin- Nicotiana tabacum herbicidebisadmi (tria-
tissimum) Tetaza sulfuroni da meTsul-
furonmeTili) gamZleo-
ba

204
 sulfonilSardovanas
herbicidebisadmi (tria-
acetolaqtat-sin- Nicotiana tabacum sulfuroni da meTsul-
Tetaza + furonmeTili) gamZleo-
+ Dianthus caryop- ba
aminiciklopropan- hyllus +
sintazas geni uin- moWrili yvavilebis Se-
naarso orientaci- naxvis vadis dagrZe-le-
iT ba eTilenis Semcirebu-
li dagrovebis gziT

m ix aki
(Dianthus sulfonilSardovanas
caryop- acetolaqtat-sin- Nicotiana tabacum herbicidebisadmi (tria-
hyllus) Tetaza + sulfuroni da meTsul-
+ Petunia hybrida furonmeTili) gamZleo-
dihidroflavanol- ba
reduqtaza +
gvirgvinis feris Sec-
vla

sulfonilSardovanas
acetolaqtat-sin- Nicotiana tabacum herbicidebisadmi (tria-
Tetaza + sulfuroni da meTsul-
+ Petunia hybrida furonmeTili) gamZleo-
flavinoid-3′,5′- hid- ba
roqsil aza +
gvirgvinis feris Sec-
vla

sulfonilSardovanas
acetolaqtat-sin- Nicotiana tabacum herbicidebisadmi (tria-
Tetaza + sulfuroni da meTsul-
+ Viola sp. furonmeTili) gamZleo-
flavinoid-3′,5′- hid- ba
roqsil aza +
gvirgvinis feris Sec-
vla

205
praqtikaSi gamoyen ebul tran sgen ur mcen areTa ji Sebi s
mokle daxasi aTeba
(ojaxi: ZaRlyurZenasebrTa)
cx r ili 1 1
kul tu- S e yvanil i cil e bi
g e nis wy a r o S e Ze nil i niS ani
ra (g e ne bi)
aminiciklopropan- momwifebis vadis dag-
sintazas geni uin- Lycopersicon escu- rZeleba eTilenis Sem-
naarso orientaci- lentum cirebuli dagrovebis
iT gziT
momwifebis vadis dag-
1-aminociklopro- rZeleba mcenareul ho-
Pseudomonas morga- rmon eTilenis winamor-
pan - 1- aminokarbo-
nis mJavas dezamina- nii bedis degradaciis Se-
p o m id o - za degad
r i (Lyco- momwifebis vadis dag-
persicon 5-adenozilmeTio- baqteriofagi rZeleba mcenareul ho-
esculen- nin hidrolaza ТЗ rmon eTilenis winamor-
tum) Е. соli bedis degradaciis Se-
degad
poligalaqturona- Senaxvis dagrZelebuli
zas geni uinnaarsoLycopersicon escu- periodi: nayofebi ar
orientaciiT lentum rbildeba poligalaq-
turonazas mcire rao-
denobis gamo.
Tam baqo Klebsiella pneu- oqsilinuri herbicide-
(Nicotiana nitralaza moniae bisadmi (bromoqsinilis
tabacum) subsp.ozaenae da ioqsinilis CaTvl-
iT) gamZleoba
delta-endotoqsini Bacillus kolorados xoWosadmi
thuringiensis gamZleoba
helikaza kartofilis kartofilis foTlis
+ foTlis sixu- sixuWuWis virusisadmi
kartofi-
delta-endotoqsini WuWis virusi gamZleoba
l i (Sola-
+ +
num tube- Bacillus thuringi- kolorados xoWosadmi
rosu)
ensis gamZleoba
kartofilis Y vi- kartofilis Y kartofilis Y virusi-
rusis garsis pro- virusi sadmi gamZleoba
teini + +
+ Bacillus thuringi- kolorados xoWosadmi
delta-endotoqsini ensis gamZleoba

206
gzidan 12 mln. avtomobilis moxsnis eqvivalenturia; daax-
loebiT 18 milioni fermeris da maTi ojaxis wevrebis (60
milionamde adamianis) keTildReobis amaRleba da sxva.

2.10. biousafrTxoebis problemebi da maTi

gadawyveta

2.10.1. genetikurad modificirebuli mcenareebis

SesaZlo uaryofiTi efeqtebi adamianis
janmrTelobaze, maTi Sefasebis meTodebi
da Tavidan acilebis xerxebi

miuxedavad genetikurad modificirebuli mcenareebis

swrafi gamravlebisa da didi mniSvnelobisa adamianisa da
garemosaTvis, dRemde arsebobs adamianis janmrTelobisa da
garemosaTvis genetikurad modificirebuli mcenareebis di-
di safrTxis - atomuri eleqtrosadguris katastrofis Se-
degebis tolfasi safrTxis Sesaxeb azri (genetikurad mo-
dificirebuli organizmebis mowinaaRmdegeebi).
is, rom adamianebis nawili skeptikurad, eWvis TvaliT
uyurebs mecnierebis am siaxleebs kargia, vinaidan winaaR-
mdegoba warmoadgens ganviTarebis aucilebel faqtors.
mTa-varia obieqturi da samarTliani argumentebiT dadgin-
des WeSmariteba.
arainformirebuli adamianebis mier damoukidebeli
logikuri daskvnebis gakeTebis mizniT, aucilebelia aRniS-
nul organizmebTan dakavSirebuli adamianis janmrTelobi-
saTvis potenciuri riskebis analizi, maTi Sefasebis meTo-
debi da metad Sromatevadi, xangrZlivi, mravalmxrivi da
mizanmimarTuli eqsperimentebis safuZvelze SemuSavebuli
usafrTxoebis RonisZiebebi.
adamiani Zalian xSirad midis riskze, iZulebiT ri-
skze. roca adamiani Ria cecxlidan Rumelze gadavida, gaC-
nda damwvrobis, sacxovreblis dawvis riski; Rumelidan ga-
zis gamaTboblebze gadasvliT warmoiqmna sadarbazoebis,
207
korpusebis afeTqebis riskebi; Tburi eleqtrosadgurebidan
atomurze gadasvliT miviReT Cernobili; nakelis da sxva
organuli sasuqebis qimiuriT Secvlis Sedegad daiwyo or-
ganizmis mowamvla; nior-arayidan antibiotikebze gadasvlam
arnaxuli alergiebi, mowamvlebi, axali problemebi gamo-
iwvia; etlidan avtomobilSi, matarebelSi da TviTmfrinav-
Si gadajdomam sikvdilianobis riskebi mravaljer gazarda
da a.S. am yvelaferze adamiani keTildReobisaTvis, siamov-
nebisaTvis, progresisaTvis midis.
bioteqnologia axalia adamianisaTvis da amitom isic
aucilebeli riskis jgufia: genetikurad modificirebuli
mcenareebi mravldebian, ujvardebian, cvlian genetikur ma-
salas.
risks Sefaseba, raodenobrivi da obieqturi gazomva
da saTanado daskvnebis gakeTeba sWirdeba. riskia magali-
Tad, asteroiduli safrTxe. magram misi dedamiwaze Camo-
var-dnis albaTobis, masis, parametrebis codna, miuxedavad
dedamiwaze arsebuli 40-60 km diametriani Rrmulebisa, ada-
mians ar aiZulebs mTeli resursebi mimarTos misgan Tavda-
sacavad. jer sakmarisia mcire xarjebi avtomaturi tele-
skopebisaTvis. asea am SemTxvevaSic.
„sruli garantia“, „absoluturi usafrTxoeba“, ro-
gorc transgenuri mcenareebis mowinaaRmdegeebi moiTxoven
bunebaSi ar arsebobs. saSiSia yvelaferi, absoluturad yve-
laferi. oRond safrTxe aris Cveulebrivi da cnobili, an
piriqiT. uCveulo da ucxo axali safrTxis warmoqmnisas
ucnobia misi sazRvrebi, zianis xarisxi.
genetikurad modificirebuli organizmebis usafrTxo-
ebaze pirvelebi SeSfoTdnen mecnierebi. jer kidev ameri-ke-
lebma robert polakma da pol bergma 1971 wels, rekombi-
nantuli dnm-s erTi mikrobidan meoreSi gadatanis pirveli
praqtikuli SesaZleblobis miRebisTanave, misi usafrTxo-
ebis gadasawyvetad moiwvies saerTaSoriso konferencia. ga-
dawyvites, rom samuSaoebi hibridul dnm-Tan saSiSia, Zali-
an saSiSia. magram riskis obieqturi Sefaseba ver SeZles. im
dros aseTi Sromebi erTeuli iyo.
208
 aseTi gamowvevebi iyo adamianis winaSe odnav adrec.
1945 wels, roca mzad iyo gamocdisaTvis msoflioSi pirve-
li atomuri bombi, daisva sakiTxi: xom ar Seva haeris azo-
ti jaWvur reaqciaSi, romelic moicavs mTel atmosferos
da Sedegad mTeli dedamiwa ar gadaiqceva erT saerTo er-
Tjerad bombad. Teoretikosebis erT nawili ambobda „ki“,
meore - „ara“. amerikeli fizikosis gregori breitis das-
kvnam - „ara“, gza gauxsna pirvel sacdel afeTqebebs da
Semdgom atomur eras.
1975 wels asilomarSi (aSS) Seikriba msoflios wam-
yvani mecnierebi. genetikur-inJinruli samuSaoebi aRiarebu-
li iqna gansakuTrebulad saSiSad da SemuSavebul iqna Za-
lian mkacri usafrTxoebis zomebi. SezRudul iqna gamo-
kvlevebi obieqtebis da Semsruleblebis mixedviT. Tumca
TandaTanobiT, mas Semdeg, rac genetikuri gamokvlevebis
realuri ziani aravis mier iqna damtkicebuli, zianis ris-
kebi Semcirda da SezRudvebi kvlevebze etapobrivad arse-
biTad Semcirda.
dRemde arc erTi genetikuri katastrofa ar momxdara.
am wlebSi miRebul iqna da moqmedebs aTeulobiT saerTaSo-
riso SeTanxmeba da kanoni, romlebic aregulireben geneti-
kurad modificirebuli organizmebis gamoyenebis sxvadas-
xva aspeqtebs.
dReisaTvis damuSavebulia adamianis janmrTelobasa
da garemosaTvis transgenuri mcenareebis usafrTxoebis Se-
fasebis efeqturi sistema. is moicavs savaraudo genetikuri
modifikaciebis dagegmvis dawyebidan transgenuri mcenaree-
bis saxelmwifo registraciis etapamde (rac mis sameurneod
gamoyenebis uflebas iZleva) gamoyenebul mTel rig midgo-
mebsa da meTodebs.
adamianis janmrTelobaze genetikurad modificirebu-
li mcenareebis SesaZlo uaryofiTi efeqtebis riskis Sefa-
sebisas daculia Semdegi ZiriTadi principebi:
1. transgenuri mcenaris usafrTxoebis eqspertiza
xorcieldeba mecnierulad dasabuTebuli saxiT. misi Cata-
rebisas iyeneben samecniero-teqnikur literaturaSi gamoq-
209
veynebul da specializirebuli monacemTa bazaSi arsebul
informacias, eqspertebis daskvnebs, eronuli da saerTaSo-
riso organizaciebis mier SemuSavebul meTodur miTiTe-
bebs da a.S., anu riskis Sefasebas aqvs mecnieruli safuZve-
li da ara raime varaudi. riski izomeba meTodurad koreq-
tuli cdebiT;
2. riskis Sefaseba transgenuri mcenaris yoveli vari-
antisaTvis xdeba idividualurad;
3. riskis Sefaseba meordeba axali informaciis gamov-
lenisas;
4. genetikurad modificirebul organizmebTan an maT
Semcvel produqtebTan dakavSirebuli riskebi ganixileba
arsebuli intaqturi recipientuli organizmebis riskebTan
konteqstSi, vinaidan adamianis janmrTelobasa da garemo-
saTvis potenciurad saSiSi SeiZleba iyos tradiciuli se-
leqciiT miRebuli organizmebic. genetikuri modifikaciis
efeqtis gamosavlenad aucilebelia aseTi organizmis Seda-
reba sawyis organizmTan.
transgenuri mcenaris gamoyenebasTan dakavSirebul
adamianis janmrTelobis potenciur riskebs Soris mniSvne-
lovania Semdegi:
1. recipientis organizmisaTvis axali cilebis – tra-
nsgenebis produqtebis sinTezi, romlebic SeiZleba iyos
toqsikuri da/an alergiuli;
2. CaSenebuli ucxo dnm-s gavleniT cocxali organiz-
mebis calkeuli genebis aqtiurobis Secvla, romlis Sede-
gad SeiZleba moxdes am organizmebidan miRebul kvebis
produqtebSi samomxmareblo Tvisebebis gauareseba, mag. to-
qsikuri da alergenuli nivTierebebis usafrTxoebis norma-
ze meti raodenobiT dagroveba;
3. transgenebis horizontaluri gadacema sxva orga-
nizmebisaTvis, kerZod antibiotikebisadmi gamZleobis mar-
keruli genebis gadacema gmo-dan saWmlis momnelebeli
traqtis mikroorganizmebisaTvis.
genetikurad modificirebuli kvebis produqtebis usa-
frTxoebis Sefasebis strategia dafuZnebulia „arsebiTi eq-
210
vivalentobis“ principze, romlis mixedviT fasdeba ara kve-
bis axali produqtebis, rogorc aseTis usafrTxoebis do-
ne, aramed misi gansxvaveba, usafrTxo gamoyenebis mraval-
wliani istoriis mqone, tradiciuli kvebis analogebTan Se-
darebiT.
kvebis produqtebis usafrTxoebasa da kvebiT Rirebu-
lebaze momqmedi genetikurad modificirebul organizmebis
axali niSnebis identifikaciisaTvis detalurad analizde-
ba transgenezis genis donori organizmis Sesaxeb informa-
cia, agreTve genetikuri modifikaciis xasiaTi. Semdeg xdeba
genetikurad modificirebuli organizmisa da sawyisi ara-
modoficirebuli organizmebis SedarebiTi analizi. amisaT-
vis erTmaneTs adareben agronomiul maCveneblebs, CaSenebu-
li genis/genebis produqtebs, sakvanZo qimiuri komponente-
bis Sedgenilobas, ZiriTad metabolitebs, nedleulis gada-
muSavebis Sedegebs.
axali produqti (jiSi) SeiZleba iyos: 1) eqvivalentu-
ri sawyis analogTan arsebiTi niSnebiT; 2) eqvivalenturi
sawyis analogTan erTi (ramodenime) niSnis garda; 3) araeq-
vivalenturi sawyis analogTan arsebiTi niSnebiT.
me-2 da me-3 SemTxvevaSi tardeba sawyisi analogisgan
gansxvavebuli gmo-s niSnebis usafrTxoebis detaluri ana-
lizi Semdegi maCveneblebis mixedviT: potenciuri toqsiku-
roba, potenciuri alergiuloba, antibiotikebisadmi gamZle-
obis genebis saWmlis momnelebeli traqtis mikroorganiz-
mebSi gadatanis SesaZlebloba, kvebiTi Rirebulebisa da
sakvebi nivTierebebis SeTvisebis potenciuri gauaresebis
albaToba.
potenciuri toqsikurobis Sefaseba xdeba Semdegnai-
rad. Tu axali produqti warmoadgens mcenareuli sakvebis
cnobil komponents, romelsac xangrZlivi usafrTxo gamo-
yenebis istoria aqvs, maSin toqsikurobis Sefaseba aucile-
beli ar aris. sxva SemTxvevaSi tardeba: 1) potenciuri toq-
sinebis koncentraciis gansazRvra mcenaris saWmel nawil-
Si; 2) aRniSnuli produqtis xvedriTi wilis dadgena mosax-
leobis garkveul jgufSi; 3) Tu cilebia, aminomJavuri Tan-
211
mimdevrobis Sedareba cnobil toqsinebTan da kvebiT anta-
gonistebTan; 4) axali nivTierebebis Termuli damuSavebi-
sadmi stabilurobis Sefaseba; 5) potenciuri toqsinebis
daSlis siCqaris gansazRvra modelur sistemebSi; 6) in vitro
kulturaSi axali nivTierebebis toqsikurobis donis ana-
lizi; 7) toqsikurobis analizi laboratoriuli da Sinau-
ri cxovelebis (Tagvebi, wiwilebi, Tevzebi, mewveli Zroxe-
bi) iZulebiTi kvebis eqsperimentebSi xangrZlivi (1-2 weli,
Cveulebrivi koncentraciiT) an mokle (2 kviramde, maRali
koncentraciiT) drois ganmavlobaSi.
transgenebis potenciuri alergiulobis SefasebisaT-
vis gamoiyeneba janmrTelobis msoflio organizaciisa da
FAO-s eqspertebis mier SemuSavebuli sqema, romlis mixed-
viT nebismieri transgenuri jiSi, sanam sameurneo gamoyene-
bis uflebas moipovebdes, gadis mravalmxriv da mraval-
wlian guldasmiT Semowmebas usafrTxoebaze.
unda aRiniSnos, rom amJamad kultivirebul transge-
nur jiSebSi gadatanili genebis produqtebSi naklebadaa
iseTebi, romlebic seriozul eWvs iwvevdnen potencialur
toqsikurobasa da alergiulobaze.
erT SemTxvevaSi adgili aqvs mcenareTa sakuTari gene-
bis manipulacias ucxo genebis monawileobis gareSe (mag.
soio da rafsi zeTis gaumjobesebuli SedgenilobiT, pomi-
dori da nesvi nayofebis gaxangrZlivebuli momwifebis va-
diTa da SenaxviT). sxva SemTxvevaSi transgenebi akodireben
fermentebs, romlebic analogiuria rogorc mcenareebSi,
ise mikroorganizmebSi (glifosatisadmi gamZle transgenu-
ri mcenareebi).
sulfonilSardovanas, imidazolinis da zogierTi sxva
herbicidisadmi tolerantuli jiSebis misaRebad gamoyene-
buli iqna ferment acetolaqtatsinTetazas mcenareuli ge-
nebi, romlebsac usafrTxo gamoyenebis xangrZlivi istoria
aqvT. igive SeiZleba iTqvas virusebisadmi tolerantul
transgenur jiSebzec. kartofilis nebismieri Cveulebrivi
jiSis tuberebSi yovelTvis aris virusuli cilebi asjer
meti SemcvelobiT, vidre transgenur jiSebSi.
212
 analogiuri magaliTebi Zalian bevria. mravalferovani
transgenuri jiSebidan mxolod erTeulebia, romlebSic ge-
netikuri modifikaciis Sedgad warmoqmna axali, saxeobis
Cveulebrivi jiSebisaTvis aradamaxasiaTebeli naerTebi.
esenia fermentebi fosfinotricinacetil-transferaza da
neomicinfosfotransferaza, romlebic Sesabamisad herbi-
cid amoniumis glufozinatis da antibiotikebis kanamici-
nis, neomicinis da geneticinis dezaqtivacias iwveven. miuxe-
davad amisa, yvela produqti, maT Soris usafrTxo gamoye-
nebis xangrZlivi istoriis mqonec, xangrZliv guldasmiT
Semowmebas gadis.
kvebis produqtebSi samomxmareblo Tvisebebis gauare-
sebis (kvebiTi Rirebulebisa da sakvebi nivTierebebis SeT-
visebis potenciuri gauaresebis albaTobis) SefasebisaTvis,
praqtikaSi Cveulebriv, didi raodenobiT miRebul transge-
nur formebs Soris, tradiciuli seleqciis gziT, gamoarCe-
ven nimuSebs arsebiTi mutaciis gareSe, Semdeg guldasmiT
aanalizeben am formebSi sakvebi nivTierebebis - cilebis,
cximebis, naxSirwylebis, mineraluri elementebis, vitamine-
bis da a.S., aseve adamianis janmrTelobisaTvis potenciu-
rad saSiSi nivTierebebis Semcvelobas.
imisaTvis, rom transgenuri jiSi daSvebul iqnas same-
urneo gamoyenebisaTvis, is arsebiTad ar unda gansxvavdebo-
des sawyisi jiSisagan, garda transgenezis niSnisa an gene-
tikuri modifikaciiT gaTvaliswinebuli niSnebisa, anu da-
culi unda iyos „arsebiTi eqvivalentobis“ principi. maga-
liTad, raundapisadmi gamZle soios 1400 analitikuri eq-
sperimentidan yvelam daadastura transgenuri da sawyisi
jiSebis sruli identoba rogorc sakvebi (cilebi, cximebi,
boWko, nacris elementebi, naxSirwylebi, kaloriuloba,
marcvlis tenianoba, gadamuSavebuli marcvlis sakvebi Tvi-
sebebi da a.S.), ise antisakvebi (cximmJavebi, aminomJavebi,
tripsinis, leqtinebis inibitorebi da sxva) niSnebis mixed-
viT.
adamianis janmrTelobis Semdeg potenciur araxelsay-
rel faqtors warmoadgens transgenebis (pirvel rigSi an-
213
tibiotikebisadmi gamZleobis genebis) horizontaluri gada-
cema genetikurad modificirebuli organizmidan adamianisa
da cxovelis saWmlis momnelebeli traqtis mikroflori-
saTvis.
rogorc wesi, nebismieri transgenuri konstruqciis
SedgenilobaSi, garda sakuTvriv transgenisa da misi regu-
latoruli elementebisa, Sedis transformirebuli ujre-
debis gamorCevisaTvis aucilebeli e.w. seleqtiuri (anu
markeruli) geni.
Cveulebriv am mizniT iyeneben garkveuli antibiotike-
bisadmi (kanamicini, ampicilini, streptomicini da sxva) gam-
Zleobis genebs.
antibiotikebisadmi gamZleobis seleqtiuri genebis ar-
seboba dReisaTvis ar warmoadgens safrTes adamianis jan-
mrTelobisaTvis, vinaidan, jer erTi, am antibiotikebma, ro-
gorc samedicino preparetebma, ukve dakarges TavianTi mniS-
vneloba, mikroorganizmebis mier maTdami gamZleobis una-
ris SeZanis gamo; meore, genmodificirebuli orgnizmidan
miRebuli kvebis produqtebis dnm-dan saWmlis momnelebeli
traqtis mikroorganizmebSi seleqtiuri genebis gadatanis
albaToba Zalian dabalia (10-17). amisaTvis saWiroa ganxor-
cieldes ramodenime Zalian nakleb savaraudo movlena: se-
leqtiuri genis Semcveli dnm-s monakveTi ar unda dazian-
des saWmlis monelebis procesSi, aucilebelia seleqtiuri
genis an mis mimdebare raionebis dnm-s da saWmlis momnele-
beli traqtis daavadebis gamomwvevi mikroorganizmis qro-
mosomis an plazmidis dnm-s Soris homologia, xolo imi-
saTvis, rom seleqtiuri geni eqspresirebdes masSi gadata-
nis Semdeg, aucilebelia es ukanaskneli integrirdes Sesa-
bamisi prokariotuli promotoris qveS. aseTis realizaci-
is SesaZlebloba da Sesabamisad, aRniSnuli riskis realo-
bis seriozuli ganxilva, didi absurdulobis gamo, mxo-
lod gauTviTcnobierebel adamianebTan SeiZleba.
miuxedavad amisa, vinaidan adamianebis SeSfoTeba am
faqtis mimarT maRalia, mecnierebma daiwyes alternatiuli
seleqtiuri sistemebis SemuSaveba (ix. gv. 142).
214
 bolo dros sul ufro xSirad seleqtiur genebad iye-
neben aratoqsikuri Saqrebisadmi (qsiloza, manoza, deoqsi-
glukoza) gamZleobis genebs, fotohormonebis eqspresiis ma-
iducirebel genebs da sxva. farTod daiwyes markerul ge-
nebad e.w. reportiori genebis (luciferazas, β-glukuroni-
dazas, mwvane fluorescenciis cilis genebi da sxva) gamo-
yeneba.
aseve SemuSavebulia seleqtiuri proceduris Semdeg
transformantebidan seleqtiuri genebis mocilebis an
umarkero transgenuri mcenareebis miRebis meTodebi.
amrigad, savaraudo riskebis fonze, genmodificirebu-
li sasoflo-sameurneo kulturebis qveS dakavebuli fa-
rTobebis raodenoba msoflioSi yovelwliurad matulobs.
risks Sefaseba, raodenobrivi da obieqturi gazomva da sa-
Tanado daskvnebi da qmedebebi sWirdeba. „sruli garantia“,
„absoluturi usafrTxoeba“, rogorc am organizmebisagan
kritikosebi moiTxoven, bunebaSi ar arsebobs. saSiSia yve-
laferi. potenciurad saSiSia genetikurad modificirebuli
organizmi. magram dReisaTvis gmo-s mwarmoebel qveynebSi
damuSavebuli adamianis janmrTelobisaTvis transgenuri
mcenaris usafrTxoebis Sefasebisa da dacvis sistemebis
analizi gviCvenebs, rom gmo-s gamoyenebasTan dakavSirebuli
adamianis janmrTelobis potenciuri riskebis (transgenebis
produqtebis toqsikurioba da/an alergiuloba, am organiz-
mebidan miRebuli kvebis produqtebSi samomxmareblo Tvise-
bebis gauareseba da transgenebis horizontaluri gadacema
sxva organizmebisaTvis) realizaciis SesaZlebloba, daxve-
wili, obieqturi da mizanmimarTuli meTodebisa da xerxe-
bis gamoyenebis gamo, praqtikulad SeuZlebelia da saSiS-
roebis albaToba arafriT aRemateba tradeciuli teqnolo-
giebiT Seqmnili organizmebidan mosalodneli riskebis al-
baTobas.

215
2.10.2. genetikurad modificirebuli mcenareebis
SesaZlo uaryofiTi efeqtebi garemoze, maTi
Sefasebis meTodebi da Tavidan acilebis gzebi

genetikurad modificirebuli mcenareebis garemoze Se-

saZlo uaryofiTi moqmedebis faqtorebidan (riskebidan) mni-
Svnelovania Semdegi:
1. biologiur Tanasazogadoebaze damangreveli moqme-
deba da Zvirfasi biologiuri resursebis dakargva, genmo-
dificirebuli mcenareebidan gadatanili genebiT adgilob-
rivi jiSebis dabinZurebis Sedegad.
bunebaSi zedmeti araferia, nebismieri bunebrivi Tana-
sazogadoebis farglebSi calkeul saxeobebs Soris gar-
kveuli balansi arsebobs. cocxali organizmebi erTmaneT-
Tan mWidro kontaqtSi da urTierTdamokidebulebaSi imyo-
febian. adamianis Carevis gareSe biologiuri mravalferov-
nebis Secvlis albaToba umniSvneloa. drois garkveul mo-
nakveTSi, magalTad klimaturi faqtorebis cvlilebiT, ro-
melime saxeobis ricxovnebis mateba, maSinaTve rTavs zrdis
SemzRudvel meqnizmebs da saxeobebs Soris balansi isev
aRdgeba.
amitom, roca laparakoben biologiur Tanasazogado-
ebaze transgenebis damangrevel moqmedebaze, mxedvelobaSi
aqvT Semdegi: calkeuli transgenuri niSnebis, pirvel rig-
Si garemosadmi adaptaciis unaris mimniWebeli - sicivisad-
mi, sicxisadmi, gvalvisadmi, damlaSebisadmi gamZleobis
niSnebis kulturuli jiSebidan maT velur saxeobebze ga-
datanisas, SesaZlebelia am ukanasknelebma SeiZinon damate-
biTi upiratesobebi arsebobisaTvis brZolaSi. es ki iwvevs
saxeobebs Soris bunebaSi arsebuli balansis darRvevas. Se-
degi SeiZleba iyos damangreveli - erTi saxeobis raodeno-
bis mateba gamoiwvevs sxva saxeobis/saxeobebis raodenobis
Semcirebas an srul gadaSenebas.
2. axali parazitebis, pirvel rigSi sarevelebis Seq-
mna da arsebulebis mavneblobis unaris gaZliereba trans-

216
genuri mcenareebis saSualebiT an transgenebis sxva saxeo-
bebSi gadatanis Sedegad.
sarevelebi warmoadgenen mcenareTa jgufs garkveuli
adaptaciuri niSnebiT (Teslebis sxvadasxva pirobebSi aR-
moceneba, Teslebis xangrZlivi sicocxlisunarianoba, yvavi-
lobis fazis swrafi gavla, mravalTeslianoba, mtvris mar-
cvlebis advili gavrceleba, maRali konkurentunarianoba,
advili vegetatiuri gamravleba da sxva), romlebic (ra
Tqma unda yvela niSani erTad arc erT saxeobas ara aqvs)
maT sxvadasxva garemoSi, maT Soris kulturuli mcenaree-
bis farTobebze, advilad gavrcelebaSi exmareba, miuxeda-
vad amis sawinaaRmdegod adamianis mudmivi Carevisa. geneti-
kurad modificirebuli organizmebis saziano moqmedeba sa-
revelebTan brZolaSi da Sesabamisad SesaZlo riski mdgo-
mareobs, arsebul sarevelebSi agresiulobis gazrdis po-
tenciur SesaZleblobaSi (e.w. supersarevelebis miRebaSi),
am mcenareebis mier transgenebidan axali adaptaciis niSne-
bis, pirvel rigSi sxvadasxva stresuli faqtorebisadmi
gamZleobis genebis SeZenaSi. garda amisa, aseTi transgene-
bis wyalobiT saSiS sarevelebad SeiZleba gadaiqces zogi-
erTi, Tavisi bunebiT gareuli formebisagan naklebad gan-
sxvavebuli kulturuli mcenare (mag. rafsi, xanWkola da
sxva). aseve, arsebobs transgenebis kulturuli saxeobebi-
dan maT gareul formebSi gadatanis da Sesabamisad, maT sa-
revelebad gadaqcevis saSiSroeba.
3. transgenuri mcenareebis mier gamoyofili (sinTezi-
rebuli) toqsikuri nivTierebebis uaryofiTi (damRupveli)
moqmedeba am mcenareebze mcxovreb an am mcenareebiT mkve-
bav, transgenuri niSnebis „arasamizne“ organizmebze (mag.
futkari, sxva sasargeblo an daculi saxeobebi).
inseqticiduri Tvisebebis (t-genis gamo) mqone trans-
genuri jiSebis Seqmnam maSinaTve gaaCina kiTxva xom ar iqo-
niebs es uaryofiT gavlenas biologiur mravalferivnebaze,
„samizne“ organizmebis (mag. kolorados xoWo) garda, xom
ar daziandeba (ganadgurdeba) sxva, maT Soris sasargeblo
organizmebic. sabednierod,Bt-proteinebs maRali gamorCevi-
217
Tobis unari aqvT. miuxedavad amisa, usafrTxoebis Sefasebi-
sas detalurad unda iqnes awonili genmodificirebuli
organizmebis aramiznobrivi moqmedebis negatiuri efeqtebi.
4. toqsikuri nivTierebebis (mag. herbicidebis) uaryofi-
Ti moqmedeba ekosistemaze, tolerantuli transgenuri mce-
nareebis qveS gadidebuli dozebis gamoyenebis gamo.
herbicidgamZle transgenuri pirveli mcenareebis gamo-
CenisTanave gaCnda saSiSi qimikatebis, mag. herbicidebis ara-
sruli daSlis toqsikuri nivTierebebis garemoze uaryofi-
Ti moqmedebis saSiSroeba. vinaidan adgili eqneba am nivTie-
rebebis gazrdili raodenobiT moxvedras bunebaSi.
aRniSnuli SesaZlo riskebis dasadgenad SemuSavebu-
lia specialuri meTodika, romelic maTi usafrTxoebis
kompleqsuri, yovelmxrivi Sefasebis Catarebis saSualebas
iZleva. es meTodika gamoyenebulia yvela qveyanaSi, sadac
genetikurad modificirebuli organizmebi mohyavT. misi Zi-
riTadi wesebi dafiqsirebulia rig saerTaSoriso SeTanxme-
bebSi, rac mas savaldebulos xdis mierTebuli qveynebisaT-
vis.
genmodificirebuli orgazizmebis garemoSi gavrcele-
bis SesaZlo ekologiuri riskebis Sefasebisas:
1. pirvel rigSi mxedvelobaSi Rebuloben recipientu-
li da donoruli organizmebis biologiur Taviseburebs:
a) adgili sistematikaSi, gamravlebisa da gavrcelebis
xerxebi, garemoSi gadarCena;
b) geografiuli gavrceleba, bunebrivi gavrcelebis
adgilis aRwera;
g) toqsikuroba.
2. gansakuTrebuli yuradReba eqceva genetikur-inJin-
ruli modifikaciis Taviseburebebs:
a) recipientuli organizmis genomSi CaSenebuli dnm-s
fragmentis aRwera (zoma da wyaro, yoveli Semadgeneli na-
wilis, maT Soris regulatoruli elementebis savaraudo
funqcia);
b) dnm-s CaSenebuli fragmentis struqtura da funqci-

218
onaluri Sesabamisoba, maTSi cnobili funqcionalurad sa-
SiSi Tanmimdevrobebis arseboba, CaSenebulis lokalizacia,
transgenebis aslebis raodenoba.
3. srulyofilad ganixileba gmo-s biologiuri Tavi-
seburebebi da garemosTan misi urTierTmoqmedebis xasiaTi,
kerZod:
a) gmo-s genetikuri stabiluroba, transgen(eb)is eq-
spresiis xarisxi da done;
b) transgen(eb)is mier kodirebuli cil(eb)is aqtiuro-
ba da Tvisebebi;
g) genetikuri informaciis gadatanis unarianoba (po-
tenciur garemoSi genmodificirebul organizmebTan hibri-
dizaciis unaris mqone gareuli da kulturuli mcenareebis
arseboba, genmodificirebuli organizmebidan aseT organi-
zmebSi transgenebis gadatanis albaToba);
d) genmodificirebuli organizmis konkurentuli
upiratesobis albaToba intaqtur recipientul organiz-
mTan SedarebiT, potenciur garemoSi genmodificirebuli
organizmebis populaciis raodenobis mkveTri zrdis alba-
Toba;
e) genmodificirebuli organizmis samizne da arasa-
mizne organizmebTan urTierTmoqmedebis SesaZlo meqanizme-
bi da Sedegebi.
garemosaTvis genmodificirebuli organizmis usafrT-
xoebis Sesaxeb saboloo daskvnebi miiReba yvela zemoT aR-
niSnuli monacemebisa da potenciuri garemos Taviseburebe-
bis (nakveTis geografiuli ganlageba, misi siaxlove nak-
rZalebTan, aRkveTilebTan da sxva bunebis damcav terito-
riebTan, zomebi da damuSavebis done, klimaturi, geologi-
uri, niadagobrivi maxasiaTeblebi, flora, fauna) gaTvalis-
winebiT.
meTodika kargad muSaobs praqtikaSi. dRemde arsebi-
Tad ar arsebobs genmodififirebuli organizmebis mier ga-
remoze negatiuri moqmedebis arc erTi SemTxveva.
aRniSnulidan gamomdinare, SeiZleba vTqvaT rom, gene-
tikurad modificirebuli mcenareebis, rogorc Zalian ma-
219
Rali teqnologiis, gamoyenebis potenciuri saSiSroeba
udavod arsebobs, magram TiToeuli transgenuri mcenaree-
bis biousafrTxoebis ganxiluli meTodikiT detaluri Se-
fasebisa da kontrolis gamo:
1. genmodificirebuli mcenareebidan gadatanili gene-
biT adgilobrivi jiSebis dabinZurebis Sedegad biologiur
Tanasazogadoebaze damangreveli moqmedebis da Zvirfasi
biologiuri organizmebis dakargvis, bunebaSi saxeobebs
Soris arsebuli balansis darRvevis saSiSroeba ar arse-
bobs;
2. transgenuri mcenareebis saSualebiT axali parazi-
tebis, maT Soris supersarevelebis Seqmna, an arsebuli pa-
razitebis mavneblobis unaris gaZlierebis riskebi ganeit-
ralebulia;
3. transgenuri niSnebis „arasamizne“ organizmebze
uaryofiTi (damRupveli) moqmedebis albaToba ar arsebobs;
4. toqsikuri nivTierebebis (mag. herbicidebis) uaryo-
fiTi moqmedeba ekosistemaze, tolerantuli transgenuri
mcenareebis gamoyenebis dros mcirdeba, vinaidan am kombina-
ciiT sarevelebze kontroli ufro maRalia, vidre Cve-
ulebriv praqtikaSi.

2.10.3. genetkurad modificirebuli mcenareebis

usafrTxoebis kontrolis Tanamedrove
sistema

pirveli genmodificirebuli mcenareebis warmoebaSi

danergvidan 25 welze meti gavida. am xnis ganmavlobaSi ma-
Ti rogorc farTobebi, ise saxeobaTa mravalferovneba da
warmoebuli produqciis moculoba arsebiTad gaizarda mi-
uxedavad amisa, dRemde ar wydeba transgenuri mcenareebis
mowinaaRmdegeTa protestebi, maT gamoyenebasTan dakavSire-
buli SesaZlo potenciuri riskebis Sesaxeb. am potenciuri
riskebis gaTvaliswinebiT transgenuri mcenareebis mwarmoe-
bel qveynebSi Seqmnilia genetikurad modificirebuli or-
ganizmebis usafrTxoebis kontrolis Tanamed-
220
rove sistemebi.
genetikurad modificirebuli mcenareebis gamoyenebas-
Tan dakavSirebul adamianis janmrTelobasa da garemoze
potenciur riskebi ganxilulia qveTavebSi 2.10.1. da 2.10.2.
genmodificirebuli mcenareebis adamianis janmrTelo-
basa da garemosaTvis usafrTxoebis kontrolis sistemis
Seqmnisas didi mniSvneloba aqvs am organizmebis miRebisa
da praqtikaSi gamoyenebis riskebis Sefasebis fundamenturi
principebis SemuSavebas, rac gulisxmobs riskebis mecni-
erul da ara varaudebis doneze gamovlenas, maT Sefasebas
TiToeuli (individualuri) transgenuri organizmisaTvis,
am sferoSi yvela xelmisawvdomi informaciis ganxilvas da
sxva.
imisaTvis, rom Seiqmnas saTanado warmodgena genmodi-
ficirebuli mcenareebis usafrTxoebis Tanamedrove kont-
rolis sistemaze, mizanSewonilia igi ganxiluli iqnas aSS-
s, rogorc genmodificirebuli organizmebis miRebis da da-
nergvis yvelaze didi gamocdilebis mqone qveynis magaliT-
ze. am qveyanaSi yvela genmodificirebuli mcenaris usaf-
rTxoebas erTmaneTisagan damoukideblad amowmebs sami sa-
xelmwifo organo. kvebis produqtebisa da samkurnalo sa-
Sualebebis kontrolis komisia, romelic pasuxismgebelia
mosaxleobis kvebis usafrTzoebaze, genmodificirebul
produqtebs swavlobs 3-5 wlis ganmavlobaSi gaZlierebuli
reglamentiT; garemos dacvis saagento akontrolebs gmo-
dan biocenozis cocxal organizmebSi herbicidebis, mavneb-
lebis da daavadebebisadmi gamZleobis genebis gavrcelebas;
soflis meurneobis saministro pasuxismgebelia imaze, rom
nebismierma axalma jiSma ar moaxdinos mavne gavlena sxva
mcenareebze da mTlianad sasoflo-sameurneo procesze.
garda amisa arsebobs postregistraciuli monitoringi, ro-
melic akontrolebs registrirebuli, anu nebadarTuli
transgenuri mcenaris danergvisa da gamoyenebis process.
kvebis produqtebisa da samkurnalo saSualebebis kon-
trolis komisiis mier genmodificirebul produqtebis au-
cilebeli kvleva moicavs sam etaps: 1) arsebiTi eqvivalen-
221
tobis dadgena; 2) axali cilis Tvisebebis dadgena; 3) ga-
mocda cxovelebze (Tagvebi, wiwilebi, Tevzebi, mewveli
Zroxebi).
genmodificirebuli produqtebis arsebiTi eqvivalen-
tobis dadgena ganxilulia qveTavSi 2.10.1.
meore etapze tardeba mediko-genetikuri da medikobio-
logiuri Sefaseba. mediko-genetikuri Sefaseba, romelic
dafuZnebulia polimerazul jaWvur reaqciasa da dnm-s seq-
venirebaze, afasebs gadatanili dnm-konstruqcias, Tanmx-
leb markerul genebs, maT sakuTar mutabelobas, mutagenur
aqtiurobas garemomcvel genomze. mediko-biologiuri Sefa-
sebis dros specialuri kvlevebiT dgindeba CaSenebuli
konstruqciis gavlena imunur statusze, axali genmodifi-
cirebuli organizmis mutagenuri, kancerogenuri, genotoq-
sikuri, neirotoqsikuri moqmedeba.
mesame etapze produqtis qronikuli toqsikuroba xan-
grZlivi drois ganmavlobaSi fasdeba laboratoriul cxo-
velebze (Tagvebi, wiwilebi, Tevzebi, mewveli Zroxebi), ra-
cionSi gamosakvlevi produqtis maqsimalurad SesaZlebeli
dozis damatebiT. tardeba dinamikuri dakvirveba bioqimiur,
fiziologiur, morfologiur parametrebze, maT Soris dgi-
ndeba reproduqciuli funqciebi.
am samuSaoebis paralelurad garemos dacvis saagento
atarebs Tavis gamokvlevebs. igi amowmebs Semdeg savaraudo
riskebs:
1. gadatanili genebi SesaZlebelia damtverianebis
gziT gadaeces axlonaTesaur gareul formebs da maTi hib-
riduli Taoba SeiZens axal Tvisebebsa da sxva mcenareeb-
Tan konkurenciis unars (invazia);
2. transgenuri mcenareebi SeiZleba gaxdenen sareve-
lebi da SeiZleba gamodevnon irgvliv arsebuli sxva mce-
nareebi;
3. genmodificirebuli mcenareebis produqtebi moxvde-
bian veluri bunebis wrebrunvaSi baqteriebis saSualebiT
da a.S.

222
 4. genmodificirebuli mcenareebis produqtebi moxvde-
bian balaxmWameli cxovelebis (mag. Zroxebis) saWmlis mom-
nelebeli traqtis mikrofloraSi.
TvalsaCinoebisaTvis sainteresoa aseTi kvlevebis pra-
qtikuli cdebis ramdenime magaliTi: 1) invaziis Sefasebis
mizniT erTmaneTs Seadares rafsis sami jiSi: sakontrolo,
misi kanamicingamZle analogi da herbicidgamZle (basta)
analogi. maT narevs zrdidnen izolirebul farTobebze
sxvadasxva adgilebSi da sxvadasxva klimatur pirobebSi.
Teslebi momwifebis Semdeg niadagSi Tavisuflad cvivoda,
sarevelebis winaaRmdeg brZola ar Catarebula. Sedegebi
aseTi miiRes: genmodificirebul analogebSi invazia sta-
tistikurad ar aris meti, vidre Cveulebriv rafsSi. dad-
ginda didi sxvaoba Teslebis gadarCenis mxriv - ufro Zli-
eri sakontrolo aRmoCnda; 2) ekosistemaze genmodificire-
buli organizmis gavlenis Sefasebis mizniT Tambaqos (pro-
teazebis mainhibirebeli genis gadatanis Sedegad foTlebs
hqonda inseqticiduri Tvisebebi) Camocvenili foTlebi Ca-
bares niadagSi. 5-7 dRis Semdeg foTlebSi proteazebis in-
hibitorebis Semcveloba iyo 0.05%, xolo 14 dRis Semdeg sa-
erTod ar iyo. mwerebis raodenoba ar Secvlila, nematode-
bi gaizarda. Sedegad entomofaunis cvlileba Sefasda ara-
kritikulad; 3) transgenuri mcenareebis stabilurobis Se-
fasebis, anu Seyvanili genebis sxva organizmebSi horizon-
taluri gadatanis albaTobis dadgenis mizniT herbicidgam-
Zle (basta) Saqris Warxlis foTlebi Cabares niadagSi. dak-
virvebiT dadginda, rom mcenareuli dnm didxans inaxeba ni-
adagSi da transgenuri dnm ar gadadis niadagis mikroflo-
raSi. baqteriaACinetobacter calcoaceticus-iT genebis e.w. horizon-
taluri gadatanis arsebobis gasarkvevad dadginda SesaZlo
transformaciis sixSire: transformantebi ar gamovlinda
maTi laboratoriuli gamokvleviT 10-13 xarisxamde, rac
mindvris pirobebSi 10-16-is tolia, anu erTi SesaZlo Sem-
Txveva 10 aTas trilionSi. aseTi transformaciis sixSire
aRniSnul cdaSi miCneul iqna usafrTxod; 4) cxovelebis
saWmlis momnelebeli traqtis mikroflo-raSi transgenebis
223
moxvedris albaTobis dadgenis eqsperimentiT gairkva, rom
genmodificirebuli sakvebiT cxoelis gamokvebisas, kuWis
baqteriebis genomSi modificirebuli monakveTis gadatanis
albaToba Seadgens 2.10-17. ZuZumwov-rebis genomSi rekombi-
nantuli dnm-s monakveTis CaSeneba principulad SeuZlebe-
lia, vinaidan maT ujredebs gaaCniaT aseTi CaSenebisgan
Tavdacvis mZlavri meqanizmi.
rac Seexeba mesame Semmowmebels - soflis meurneobis
saministros, isic paralelur reJimSi asrulebs Tavis fun-
qcias. Tu rogor keTdeba es, amis sailustraciod isev mo-
viyvan cdebis magaliTs: 1) transgenebis sxva mcenareebSi ga-
datanis sixSiris Seswavlisas, kerZod, herbicid bastasadmi
gamZlebis genis (bar-geni) genmodificirebuli rafsidan mis
Cveulebriv analogze gadatanis dasadgenad izolirebul 1
ha (100X100 m) farTobze daTeses Cveulebrivi rafsi da cen-
trSi wriulad 9m diametris farTobze daTeses genmodifi-
cirebuli analogi. jvaredini damtvervis xelSewyobisaTvis
ganalages futkris skebi. damtvervis Semdeg Catarda gamo-
naskvnili Teslebis analizi. hibriduli mcenareebis raode-
noba wridan 1 metris dacilebiT iyo - 1.4%, 3 metris man-
Zilze - 0.4%, 12 metris manZilze - 0.01%, xolo 47 metris
manZize - 0.0003%, anu sami mcenare milioni mcenaridan, rac
umniSvneloa; 2) igive mizniT erTmaneTis gverdiT darges
Cveulebrivi kartofili da misi genmodificirebuli analo-
gi. jvaredini damtverva maTi sazRvridan 1 metrSi Seadgen-
da 24%-s, 10 metrSi - 0.017%-s, xolo 20 metrSi hibriduli
mcenareebi ar aRmoCnda. daskvna dadebiTia.
analogiuri Semowmeba aqvs gavlili dReisaTvis war-
moebaSi rekomendirebul genmodificirebul jiSebs (2016
wlis monacemebiT 29 saxeobis 404 jiSi).
ra Tqma unda yvela genmodificirebuli mcenare usaf-
rTxo ar aris. maTi udidesi nawilis wundeba xdeba TviT
Semqmnelebis mier, kidev didi nawili ver gadis registra-
ciis tests, darCenilebidan bevri ver uZlebs postregis-
traciul monitorings da mxolod erTeulebi aRweven war-
moebamde - namdvilad usafrTxo da uvnebeli adamianisa da
224
garemosaTvis.
sanam arsebobs genetikur-inJinruli saqmianobis SesaZ-
lo riskebi adamianisa da garemos usafrTxoebaze, maTi re-
gulireba unda moxdes saxelmwifoebriv doneze zemoT gan-
xiluli sistemis Sesabamisad.
genmodificirebuli organizmebis SesaZlo uaryofiTi
moqmedebebis mimarTulebiT adamianebis mudmivi fiqri, sif-
rTxile gadawyvetilebebSi da brZola usafrTxoebisaTvis
yovelTvis aucilebelia, vinaidan mis gareSe am saqmeSi Sem-
dgomi ganviTareba ver iqneba.
G

225
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 s a r C e v i
Sesavali ...................................................................................................................... 3
Tavi 1. Uujreduli inJineria ...........................................................................6
1 .1 u jredebis g enetikuri konstruirebis u jredul-inJinruli
meTodebi ................................................................................................................... 8
1 .1 .1. m cenaris u jredebis konstruireba izolirebuli
protoplastebis SerwymiT ..............................................................................8
1 .1 .2. m cenaris u jredebis konstruireba subprotoplastebis
SerwymiT................................................................................................................... 2 0
1 .1 .3. m cenaris u j redebis konstruireba u jreduli
organelebis SeyvaniT .......................................................................................2 1
1 .1 .4. m cenaris u jredebis g enetikuri cvalebadoba in vitro
manipulaciebiT .................................................................................................... 2 2
1 .1 .5. m cenaris u j redebis m utaciuri cvalebadoba in vitro
kultivirebisas .................................................................................................... 2 4
1 .1 .6. S ecvlili (konstruirebuli) u jredebis g amorCeva da
Senaxva ........................................................................................................................2 5
1.2. ujreduli asociaciebi mikroorganizmebTan ............................... 2 6
1 .2 .1. u j reduli a sociaciebis S eqmnis a rsi, ZiriTadi
mizani da tipebi ................................................................................................. 2 6
1.2.2. u j redul populaciebSi m ikroorganizmebis S eyvanis
kvleviTi muSaobis Sedegebi ........................................................................ 2 8
1 .3. u j redu li seleqciiis ra ob a , Ta viseb u reb eb i da
Sedeg ebi ..................................................................................................................... 31
1.3.1 . ujreduli seleqciis zogadi sqema .................................................. 31
1.3.2 . ujreduli seleqciis obieqtebi ........................................................34
1 .3.3. in vitro u j redu li ku ltu reb is m eTodeb i
mcenareTa seleqciaSi ...................................................................................... 42
1 .3.3.1. in vitro u j reduli ku lturebis da mxmare m eTodebi
mcenareTa seleqciaSi .......................................................................................43
1 .3.3.1.1. in vitro g anayofiereba (progamuri S euTavseblobis
daZleva) ....................................................................................................................43
1 .3.3.1.2. Teslkvirtebis, naskvebisa d a m o umwifebeli hibriduli
Canasaxebis kultura (postgamuri SeuTavseblobis daZleva) ........ 44
1 .3.3.1.3. letaluri hibridebis q sovilebidan m cenareTa
regeneracia (morfogenezi hibriduli qsovilebis kulturaSi) ... 47
1.3.3.1.4. haploidebis miReba................................................................................ 47

228
1 .3.3.1.5. ax ali jiSebis, hibridebis, x azebis kl onuri
mikrogamavleba („xelovnuri Teslebis“ Seqmnis CaTvliT) .............. 50
1 .3.3.1.6. m cenareuli g e nofondis kr ioSenaxva (ujredebisa d a
qsovilebis banki) ................................................................................................. 52
1 .3.3.2. u j redul seleqciaSi g enetikuri m ravalferovnebis
Seqmnis meTodebi ................................................................................................. 54
1.3.3.2.1. somaklonuri cvalebadoba................................................................. 54
1.3.3.2.2. somaturi hibridizacia ....................................................................... 58
1.3.3.2.3. in vitro inducirebuli mutacia........................................................... 66
1.3.4. gamorCevis meTodebi ujredul seleqciaSi ............................... 7 2
1 .3.5. u j reduli seleqciis u piratesobebi, S emzRudveli
pirobebi da faqtorebi ................................................................................... 7 6
1.3.6.. ujreduli seleqciis S edegebi da perspeqtivebi ..................... 7 7
1.3.6.1. somaklonuri cvalebadoba ................................................................... 7 7
1.3.6.2 somaturi hibridizacia........................................................................... 8 0
1.3.6..3. in vitro inducirebuli mutacia ........................................................... 8 3
Tavi 2. genetikuri inJineria........................................................................ 8 6
2 .1 .GmcenareTa g enetikuri inJineriis ra oba da teqnologiis
etapebi....................................................................................................................... 8 6
2.2. mcenareebSi genebis gadatanis meTodebi ....................................... 9 0
2.3. genebis gadatana veqtorebis gamoyenebiT.................................... 9 1
2.3.1. veqtorisadmi wayenebuli moTxovnebi................................................. 9 1
2.3.2. mcenareTa genetikuri inJineriis veqtorebi ................................... 9 4
2.3.2.1. Aagrobaqteriis veqtorebi..................................................................... 9 4
2.3.2.1.1. mcenareTa bunebrivi transformacia agrobaqteriebiT ........ 9 4
2.3.2.1.2. veqtorebi agrobaqteriis plazmidebis safuZvelze ............1 1 3
2.3.2.2. veqtorebi fitovirusebis safuZvelze.......................................... 1 2 0
2 .3.2.3. v e qtorebi m cenaris q l oroplasturi d a m ი toqondriuli
dnm-s gamoyenebiT ................................................................................................. 1 2 3
2 .3.2.4. v eqtorebi m obiluri e lementebis (transpozonebis)
safuZvelze ............................................................................................................. 1 2 4
2.3.3. veqtorebis saSualebiT genebis gadatanis meTodebi ............... 1 2 6
2.4. genebis pirdapiri gadatanis meTodebi ......................................... 1 30
2.5. g enebis gadatanis kombinirebuli meTodebi .............................. 1 38
2.6. mcenaris ujredis transformaciis calkeuli sakiTxebi.... 1 39
2.6.1. m cenareSi ucxo genis eqspresiis regulreba............................... 1 39
2.6.2. markeruli (reportiori, seleqtiuri) genebis gamoyeneba ..... 1 42

229
2 .6.3. tr ansformirebuli m cenareebi m arkeruli s eleqtiuri ge-
nebis gareSe............................................................................................................1 47
2.6.4. Tambaqos transformacia „foTlis rgolebis“ meTodiT ........1 48
2.6.5. mcenaris plastomuri genomis transformacia ............................1 50
2.6.6. polimerazuli jaWvuri reaqcia.........................................................1 56
2.7. mcenaris transformaciis dadastureba .........................................1 6 3
2.8. mcenaris genomSi ucxo genis eqspresia (funqcionireba) ...1 6 3
2.9. genetikuri inJineriis miRwevebi .......................................................1 65
2 .9 .1. he rbicidebisadmi g amZle tr ansgenuri m cenareebis
miReba .........................................................................................................................1 66
2 .9 .2. m weri-mavneblebisadmi g amZle tr ansgenuri m cenareebis
miReba .........................................................................................................................1 7 2
2 .9 .3. d a av ad e be bis ad m i g am Zl e tr ans g e nur i m ce nar e e bis
miReba .........................................................................................................................1 7 6
2 .9 .4. g aremos araxelsayreli f aqtorebisadmi g amZle
transgenuri mcenareebis miReba....................................................................1 7 9
2.9.5. maRalmosavliani transgenuri mcenareebis miReba ..................... 1 8 1
2 .9 .6. g a umjobesebuli g emuri d a s asaqonlo Tvisebebis
mqone transgenuri mcenareebis miReba .......................................................1 8 3
2 .9 .7. g a umjobesebuli kvebiTi d a s amrewvelo R irebulebis
mqone transgenuri mcenareebis miReba ........................................................1 8 7
2 .9 .8. s amedicino p reparatebis p roducenti tr ansgenuri
mcenareebis miReba ...............................................................................................1 9 3
2 .9 .9. tr ansgenuri m cenareebis g avrcelebis Ta namedrove
mdgomareoba ............................................................................................................1 9 5
2.10. biousafrTxoebis problemebi da maTi gadawyveta ............. 2 07
2 .1 0.1. g e netikurad m odificirebuli o rganizmebis S esaZlo
uaryofiTi e feqtebi adamianis janmrTelobaze, m aTi S efasebis
meTodebi da Tavidan acilebis xerxebi ................................................... 2 07
2 .1 0.2. g e netikurad m odificirebuli o rganizmebis S esaZlo
uaryofiTi e feqtebi g aremoze, m aTi Sefasebis m e Todebi d a Ta-
vidan acilebis gzebi.........................................................................................2 1 6
2 .1 0.3. g enetkurad m odificirebuli m cenareebis usafrTxoebis
kontrolis Tanamedrove sistema................................................................. 2 2 0
gamoyenebuli literatura.......................................................................... 2 2 6

230
 vaxtang qobalia

mcenareTa genetikuri konstruirebis

meTodebi

VAKHTANG KOBALIA
Methods of plant genetic construction

teqredaqtori m. Tofuri a

wigni daibeWda akaki wereTlis saxelmwifo

universitetis stambaSi

xelmowerilia dasabeWdad 24.01.2019

pirobiTi nabeWdi Tabaxi 14.4

fasi saxel Sekrulebo

231