koncefcia testi 17.
3 transkripcia dnm
1. rogor moqmedebs pre-mrnm-is 5’ da 3’ bo-
loebis cvlileba mrnm-ze, romelic tovebs
birTvs?
2. aRwereT mbrnp-ebis roli rnm splaisingSi.
3. rogor xdeba rnm-is alternatiuli
splaisingis Sedegad genebis raodenobaze
meti cilovani produqtebis warmoqmna?
17.4
koncefcia
translacia aris polipeptidis
sinTezi, romelsac rnm
warmarTavs: detalurad
Cven axla ufro detalurad ganvixilavT, rogor ga-
dadis genetikuri informacia mrnm-dan cilaSi _ trans-
laciis process. rogorc transkripciis SemTxvevaSi,
aqac yuradRebas gavamaxvilebT translaciis ZiriTad
safexurebze, romlebsac adgili aqvs prokariotebSic
da eukariotebSic da mivuTiTebT maT Soris arsebul
gansxvavebebze.
translaciis molekuluri komponentebi
translaciis procesSi, ujredi Targmnis geneti-
kur mesijs da amis mixedviT aSenebs polipeptids. me-
siji aris mrnm-is molekulis gaswvriv ganlagebuli
kodonebis wyeba, xolo mTargmnels transportuli rnm
(trnm) ewodeba. trnm-is funqciaa aminomJavebis gadata-
na aminomJavebis citoplazmuri auzidan ribosomebTan.
ujredi citoplazmas yovelTvis amaragebs ocive ami-
nomJavaTi, sxva naerTebisagan maTi sinTezis an garemom-
cveli xsnaridan STanTqmis gziT. ribosoma polipepti-
duri jaWvis mzard bolos amatebs trnm-is mier masTan
mitanil yovel aminomJavas (suraTi 17.13).
t rnm-is molekulebi identurebi ar arian. geneti-
kuri mesijis specifikur aminomJavur TanamimdevrobaSi
translirebis gasaRebi is aris, rom TiToeuli tipis
trnm-is molekula mrnm-is gansazRvrul kodons gada-
Targmnis gan­sazRvrul aminomJavaSi. ribosomasTan mo-
sul trnm-is molekulas erT bolosTan specifikuri
aminomJava aqvs dakavSirebuli. trnm-is meore boloze
aris nukleotiduri tripleti, romelsac antikodoni
ewodeba da romelic fuZeTa Sewyvilebis saSualebiT
382
mrnm
ribosoma
translacia
polipeptidi
polipeptidi aminomJavebi
trnm masTan
dakavSireb-
uli aminom-
JaviT
ribosoma
f
tr
fe gli
trnm
antikodoni
kodonebi
mrnm
S S suraTi 17.13  translacia: ZiriTadi koncefcia.
mrnm-is molekulis ribosomaSi gadaadgilebisas, kodo-
nebi, saTiTaod, translirdeba aminomJavebSi. `mTargm-
nelebs~ trnm-is molekulebi warmoadgenen, romelTa
TiToeul tips erT boloSi spe­cifikuri antikodoni aqvs,
xolo meore bo­loSi _ Sesabamisi aminomJava. rodesac
antikodoni mrnm-ze ganlagebul komplementarul kodons
ukavSirdeba, trnm Tavis aminomJavur tvirTs umatebs
mzard polipeptidur jaWvs. momdevno suraTebi gviCvenebs
translaciis zogierT detals prokario­tul ujredebSi.
ukavSirdeba komplementarul kodons mrnm-ze. magali-
Tad, warmoidgineT mrnm-is kodoni uuu, rac Seesabam-
eba aminomJava fenilalanins. trnm-s, romelic fuZeTa
Sewyvilebis saSualebiT wyalbadur bmebs warmoqmnis am
kodonTan, antikodonis saxiT aqvs aaa da Tavis meore
boloTi fenilalanins atarebs (ix. Sua trnm suraT
17.13-ze). mrnm-is molekulis gadaadgilebisas ribo-
somaSi, fenilalanini daemateba polipeptidur jaWvs
yvela SemTxvevaSi, rodesac kodoni uuu iqneba warmod-
genili translaciisaTvis. ase, kodoni kodonis Semdeg
xdeba genetikuri mesijis translacia, rameTu trnm-
ebs moaqvT aminomJavebi gansazRvruli rigis mixedviT,
xolo ribosoma aerTebs aminomJavebs polipeptidur
jaWvSi. trnm-is molekula warmoadgens mTargmnels,
 W W suraTi 17.14  transportuli rnm-is (t rnm -is) struq-
tura. SeTanxmebis mixedviT, antikodonebi iwereba 3’ _ 5’ mimar-
aminomJavas TulebiT, rom 5’ _ 3’ mimarTulebiT daweril kodonebTan Sesabam-
dakavSirebis isad ganlagdnen (ix, suraTi 17.13). fuZeTa SewyvilebisaTvis, rnm
saiti jaWvebi, dnm-is msgavsad, antiparaleluri unda iyos. magaliTad,
antikodoni 3’-aag-5’ uwyvildeba mrnm-is kodons 5’-uuc-3’.

m rnm-is msgavsad, sinTezdeba birTvSi da Semdgom unda

gadaadgildes birTvidan citoplazmaSi, sadac xor-
cieldeba translacia. prokariotul da eukariotul
wyalbaduri ujredebSic, trnm-is TiToeuli molekula mravaljera-
bmebi dad gamoiyeneba, miierTebs ra misTvis gansazRvrul ami-
nomJavas citozolSi, miitans am tvirTs ribosomasTan
da Semdeg tovebs ribosomas, rom miierTos axali ami-
nomJava.
a. rogorc suraT 17.14-zea ilustrirebuli, trnm-is
mo­lekula Sedgeba rnm-is erTi jaWvisagan, romelic,
antikodoni
daaxloebiT, mxolod 80 nukleotidis sigrZisaa (rode-
aminomJavas sac mrnm-is molekulebis umetesoba aseulobiT nukleo-
dakavSirebis
tids Seicavs). fuZeebis komplementaruli monakveTebis
saiti
arsebobis gamo, romelTac wyalbaduri bmebiT SeuZliaT,
daukavSirdnen erTmaneTs, es erTi jaWvi SesaZloa, ga-
daikecos da warmoiqmneba samganzomilebiani struqtu-
wyalbaduri ris mqone molekula. arsebuli Sewyvilebuli fuZeebis
bmebi gamosavlenad, trnm-is molekulas erT sibrtyeSi Tu
ganvixilavT, is samyuras foTols hgavs (suraTi 17.14a).
trnm ixveva da ikeceba kompaqtur samganzomilebian
struqturad, romelsac daaxloebiT L-is forma aqvs (su-
raTi 17.14b). L-is erT boloze arsebuli maryuJi Seicavs
g.
antikodons, fuZeTa triplets, romelic ukavSirdeba
mrnm-is specifikur kodons. L-formis trnm-is moleku-
b. antikodoni antikodoni
lis meore bolodan gamoSverilia misi 3’ bolo, romelic
warmoadgens aminomJavas dakavSirebis saits. amgvarad,
trnm-is struqtura mis funqcias Seesabameba.
radganac mas SeuZlia, waikiTxos nukleinis mJavas sity- genetikuri mesijis swo­­rad translirebisaTvis
va (m rnm-is kodoni) da gadaTargmnos is cilis sityvaSi saWi­­-roa amocnobis ori safexuri. pirveli, sworad
(aminomJava). unda dawyvi­l­dnen trnm da aminomJava. trnm-s, ro-
translaciis procesi martivia principSi, magram misi melic ukavSirdeba gan­­sazRvruli aminomJavas Sesabamis
bioqimia da meqanika sakmaod rTulia, gansakuTrebiT, mrnm-is kodons, ribosomasTan unda mihqondes mxolod
eukariotul ujredSi. translaciis ganxilvisas, Cven es aminomJava. TiToeul aminomJavas Sesabamis trnm-Tan
koncentrirebas movaxdenT procesis SedarebiT nak- akavSirebs specifikuri fermenti aminoacil-t rnm-
lebad rTul versiaze, rasac prokariotebSi aqvs ad- sinTetaza (suraTi 17.15). Ti­Toeuli tipis aminoacil-
gili. modiT, Tavdapirvelad ganvixiloT am ujreduli trnm-sinTetazas aqtiuri saiti Seesabameba mxolod
moqmedebis mTavari gmirebi da Semdeg vnaxoT, rogor mo- aminomJavisa da trnm-is specifikur kombinacias. arse-
qmedeben isini erToblivad polipeptidis Sesaqmnelad. bobs 20 sxvadasxva sinTetaza, yoveli aminomJavasaTvis
TiTo. yovel sinTetazas SeuZlia, daukavSirdes misTvis
transportuli rnm-is specifikuri aminomJavas makodirebel yvela sxvadasxva
trnm-s. sinTetaza akatalizebs trnm-Tan aminomJavas
struqtura da funqcia
ko­valentur dakavSirebas atf-is hidroliziT warmar-
mrnm-isa da ujreduli rnm-is sxva tipebis msgavsad, Tul pro­cesSi. Sedegad miRebuli aminoacil-trnm,
transportuli rnm-is molekulebi transkribird- romelsac ga­a qtivebul aminomJavasac uwodeben, gamoTa-
ebian dnm-is yalibebidan. eukariotul ujredSi, trnm, visufldeba fermentisagan da gadaitans Tavis aminomJa-

383
aminomJava aminoacil-trnm (a) organzomilebiani struqtura. yvela trnm-isaTvis
sinTetaza (fermenti) damaxasiaTebelia Sewyvilebul fuZeTa oTxi ubnisa da
sami maryuJis arseboba, aseve 3’ boloze aminomJavas da-
kavSirebis saitis fuZeTa Tanamimdevroba. antikodonuri
1 tripleti unikaluria TiToeuli tipis trnm-isaTvis.
aqtiuri saiti ukavSirdeba
aminomJavasa da atf-s. (varskvlavebiT aRniSnulia qimiurad modificirebuli
adenozini
fuZeebi, rac axasiaTebs trnm-s.)
atf (b) samganzomilebiani struqtura
(g) trnm-isaTvis am wignSi gamoyenebuli simbolo.
2
atf kargavs or P
jgufs da ukavSirdeba ami­
nomJavas amf-is saxiT.
buli fuZe u SeiZleba Seuwyvildes mrnm-is kodonis
mesame poziciaze myof (3’ bolosTan) a-s an g-s. fuZeTa
pirofosfati adenozini
Sewyvilebis wesebis aseT Sesustebas qanaoba ewodeba.
qanaoba xsnis, ratom SeiZleba, rom mocemuli aminom-
Javas Sesabamisi sinonimuri kodonebi erTmaneTisagan
fosfatebi gansxvavdebodnen, Cveulebriv, mesame fuZis mixedviT,
magram ara sxva fuZeebiT.
trnm
ribosomebi
3
Sesabamisi trnm ribosomebi, cilis sinTezis procesSi, trnm-is an­
kovalenturad
ukavSirdeba tikodonebis mrnm-is kodonebTan specifikur Sewyvi­
aminomJavas, lebas uwyoben xels. ribosoma sakmaod didia, rom
Caanacvlebs
ra amf-s. eleqtronul mikroskopSi gamoCndes da is ori suber-
adenozini Teulisagan _ didi da patara suberTeulebisagan _
amf Sedgeba (suraTi 17.16). ribosomuli suberTeulebi age-
bulia cilebisa da rnm-is molekulebisagan, romelTac
4 ribosomuli rnm, an rrnm, ewodebaT. eukariotebSi, su-
fermenti gamoyofs gaaq-
tivebul aminomJavas. berTeulebi birTvakSi iqmneba. qromosomul dnm-ze ar-
sebuli ribosomuli rnm-is genebi transkribirdeba, rnm
ganicdis procesings da aiwyoba citoplazmidan Semot-
anil cilebTan erTad. Sedegad miRebuli ribosomuli
aminoacil trnm (`gaaq-
tivebuli aminomJava~) suberTeulebi birTvis forebis gavliT gadadian ci-
toplazmaSi. prokariotebsa da eukariotebSic, didi da
patara suberTeulebi funqcionaluri ribosomis Camo-
S S suraTi 17.15  aminoacil t rnm sinTetaza specifi- sayalibeblad mxolod m rnm -is molekulasTan dakav-
kur aminomJavebs akavSirebs trnm-Tan. trnm-isa da Sirebis Semdeg erTdeba. ribosomis masis daaxloebiT
aminomJavas dakavSireba endergonuli procesia, romelic ori mesamedi r rnm -ia. radganac ujredebis umetesoba-
atf -is xarjze xorcieldeba. atf kargavs fosfatis or Si aTasobiT ribosomaa, r rnm rnm -is yvelaze gavrcele-
jgufs da gardaiqmneba amf-ad (adenozin monofosfatad). bul tips warmoadgens.
prokariotebisa da eukariotebis ribosomebi Za­lian
vas ribosomaze, mzard polipeptidur jaWvze. msgavsia struqturisa da funqciebis mixedviT. Tumca,
amocnobis meore sa­f exurs warmoadgens trnm-is eukariotebSi ribosomebi odnav didia da prokari-
anti­kodonsa da mrnm-is kodons Soris swori Sewyvileba. otebis ribosomebisagan garkveulwilad gansxvavdeba
aminomJavebis Sesabamisi TiToeuli mrnm-is kodonisaT- molekuluri SemadgenlobiT. es gansxvavebebi samedi-
vis TiTo trnm rom arsebobdes, trnm-ebis raodenoba cino TvalsazrisiT mniSvnelovania. garkveul antibio-
61 unda iyos (ix. suraTi 17.5). sinamdvileSi, arsebobs tikur wamlebs SeuZliaT prokariotuli ribosomebis
mxolod 45 trnm, rac miuTiTebs, rom zogierT trnm-s inaqtivireba, eukariotuli ribosomebis mier cilis
erTze met kodonTan dakavSireba unda SeeZlos. es mov- sinTezis unaris inhibirebis gareSe. aseTi wamlebi, maT
lena imitomaa SesaZlebeli, rom kodonis mesame fuZisa Soris tetraciklini da streptomicini, baqteriul in-
da t rnm-is antikodonis Sesabamisi fuZis Sewyvilebis feqciebTan sabrZolvelad gamoiyeneba.
wesebi ar aris iseTi mkacri, rogorc dnm-isa da m rnm-is ribosomebis struqtura asaxavs mis funqcias, rom
kodonebisaTvis. magaliTad, trnm-is 5’ boloze moTavse- erTmaneTTan `miiyvanos~ mrnm da aminomJavebis matare-

384
 (a)moqmedi ribosomis kompiute­ruli modeli. es aris
baqteriuli ribosomis modeli, romelic mis mTlian
transkripcia
dnm formas gviCvenebs. eukariotuli ribosoma, daaxloebiT,
msgavsia. ribosomuli suberTeuli aris ribosomuli rnm-is
mrnm molekulebisa da cilebis erToblioba.
ribosoma
translacia tunelis (b)dakavSirebis saitebis maCvenebe­li sqematuri modeli.
polipeptidi gasasvleli ribosomas aqvs mrnm-is dakavSirebis saiti da trnm-is
mzardi dakavSirebis sami saiti, A, P da E saitebi. es sqematuri
polipeptidi ribosoma Semdgom diagramebSic Segvxvdeba.
trnm molekulebi
(g)sqematuri modeli mrnm-iTa da trnm-iT. trnm Tavsdeba
dakavSirebis saitze, rodesac misi antikodoni uwyvil-
deba mrnm-is kodons. P saitTan dakavSirebulia mzard
polipeptidTan bmuli trnm. A saitTan aris trnm, romelsac
didi
polipeptidur jaWvze dasamatebeli Semdegi aminomJava
suberTeuli moaqvs. gantvirTuli trnm ribosomas tovebs E saitidan.
E P
A
WW suraTi 17.16  moqmedi ribosomis anatomia
mcire
suberTeuli

beli trnm-ebi. mrnm-is dakavSirebis saitis garda,

TiToeul ribosomas aqvs trnm-Tan dakavSirebis
mrnm
sami saiti (ix. suraTi 17.16). P saiti (peptidil-
trnm-saiti) ikavSirebs trnm-s, romelic mzar­di
P saiti Asaiti
(peptidil- trnm-is (aminoacil - trnm-is polipeptiduri jaWvis matarebelia. A saiti (ami-
dakavSirebis saiti) dakavSirebis saiti) noacil-trnm saiti) ikavSirebs trnm-s, romelic
jaWvze dasamatebel momdevno aminomJavas atarebs.
E saiti gantvirTuli (aminomJavisagan ganTavisuflebuli)
(gasasvleli trnm-ebi ribosomas toveben E saitis (gasvlis
saiti) didi
E P A saiti, ingl. exit-dan) saSualebiT. ribosoma gana-
suberTeuli
Tavsebs trnm-sa da mrnm-s erTmaneTTan axlo man-
Zilze da awvdis axal aminomJavas mzardi polipep-
mrnm-is
dakavSirebis mcire tidis karboqsi-boloze dasamateblad. Semdgom, is
saiti suberTeuli akatalizebs peptiduri bmis warmoqmnas. rodesac
polopeptidi grZeldeba, is gadis ribosomis didi
suberTeulis gasasvlel gvirabSi. polipeptidis
sinTezis dasrulebis Semdeg, is gamoTavisufldeba
citozolSi gasasvleli gvirabidan.
ribosomis struqturis genetikuri da bioqi-
miuri kvleva oTxi aTeuli weli mimdinareobda da
amino bolo mzardi misi kulminaciaa baqteriuli ribosomis detal-
polipeptidi uri struqtura, romlis modelic warmodgenilia
Semdegi aminomJAva, 17.1 suraTze. uaxlesi monacemebi mxars uWers
romelic unda hipoTezas, rom ribosomis struqturasa da fun-
SeuerTdes poli- qcionirebaSi mTavari roli akisria rnm-s, da ara
peptidur jaWvs
cilas. cilebi, romlebic ribosomaze, ZiriTadad,
garedan arian ganTavsebuli, xels uwyoben rrnm-is
E trnm konformaciul cvlilebebs, rodesac isini trans-
mrnm laciis procesSi katalizs axorcieleben. rnm
3’
warmoadgens ribosomis ori suberTeulis sazRvr-
isa da A da P saitebis mTavar Semadgenels da is
kodoni
5’ akatalizebs peptiduri bmis warmoqmnas. amrigad,
ribosoma SeiZleba, ganvixiloT, rogorc erTi
uzarmazari ribozomi!

385
 ribosomis
1 didi
suberTeuli

P saiti
Met Met

iniciatori trnm gtf gdf

mrnm

start kodoni
ribosomis
patara
suberTeuli translaciis iniciaciis
mrnm-is dakavSirebis saiti
kompleqsi
2

1 ribosomis patara suberTeuli ukavSirdeba m rnm -is molekulas. prokariotul ujredSi, am suberTeulze arsebuli m rnm -is da-

kavSirebis saiti amoicnobs specifikur nukleotidur Tanamimdevrobas mrnm-ze start kodonis gverdze, aRmavali mimarTulebiT.
iniciatori trnm, Tavisi uac antikodoniT, uwyvildeba start kodons aug-s. am trnm-s gadaaqvs aminomJava meTionini (Met).

2 ribosomis didi suberTeulis mierTeba asrulebs iniciaciis kompleqss. translaciis yvela kom­p onentis erTad Sesakrebad

saWiroa cilebi, romelTac iniciaciis faqtorebs uwodeben (ar aris naCvenebi). gtf iZleva am asociaciisaTvis saWiro ener-
gias. iniciatori trnm P saitzea. A saiti misawvdomia momdevno aminomJavas matarebeli trnm-isaTvis.

SS suraTi 17.17  translaciis iniciacia

polipeptidis aSeneba radganac gansazRvravs kodonTa wakiTxvis CarCos mrnm-

translacia, polipeptiduri jaWvis sinTezi, Sei-
Zleba daiyos sam stadiad (transkripciis etapebis
analogiurad): iniciacia, elongacia da terminacia. sa-
mive etapi saWiroebs cilovan `faqtorebs~, romlebic
exmarebian mrnm-s, t rnm-sa da ribosomebs translaciis
procesSi. jaWvis iniciaciisa da elongaciis garkveu-
li aspeqtebisaTvis aseve saWiroa energia. is moipoveba
gtf -is (guanozin trifosfatis), atf-is monaTesave
molekulis, hidrolizis saSualebiT.
ribosomis asociacia da translaciis
iniciacia
translaciis iniciaciis safexuri erTmaneTTan
aerTebs mrnm-s, polipeptidis pirveli aminomJavis
matarebel trnm-sa da ribosomis or suberTeuls (sura-
Ti 17.17). pirvelad, ribosomis patara suberTeuli uka-
vSirdeba mrnm-sa da specifikur iniciatorul trnm-s,
romelic aminomJava meTioninis matarebelia. patara su-
berTeuli Semdeg gadaadgildeba mrnm-is gaswvriv daR-
mavali mimarTulebiT, anu axdens mis skanirebas, sanam ar
miaRwevs start kodonamde, aug, romelic translaciis
dawyebis signals warmoadgens. es mniSvnelovani faqtia,
isaTvis. kompleqsTan ukve asocirebuli iniciatoruli
trnm amis Semdeg ukve wyalbadur bmebs amyarebs start
kodonTan.
m rnm-is, iniciatoruli trnm-isa da ribosomis pa-
tara suberTeulis kavSirs emateba ribosomis didi su-
berTeulic, riTac sruldeba translaciis iniciaciis
kompleqsis Camoyalibeba. yvela am komponentis erTad
Sesakrebad saWiroa cilebi, romelTac iniciaciis faq-
torebs uwodeben. iniciaciis kompleqsis Sesaqmnelad,
ujredi energiasac xarjavs, gtf-is molekulebis saxiT.
iniciaciis procesis dasrulebisas, iniciatoruli trnm
ganlagebulia ribosomis P saitze, xolo vakanturi A
saiti mzad aris Semdegi aminoacil trnm-isaTvis. yur-
adReba miaqcieT, rom polipeptidis sinTezi yovelTvis
erTi mimarTulebiT xorcieldeba, sawyisi meTioninidan
amino-boloze, aseve uwodeben N-bolos, saboloo ami-
nomJavisaken karboqsi-boloze, aseve uwodeben C-bolos
(ix. suraTi 5.18).
polipeptiduri jaWvis elongacia
translaciis elongaciis stadiaze, aminomJavebi,
saTiTaod, ematebian winamdebare aminomJavas. TiToeul

386
 polipeptidis
kodonis amocnoba. Semosuli
amino bolo
transkripcia dnm aminoacil trnm-is antikodoni
uwyvildeba mrnm-ze komplemen-
mrnm
tarul kodons A saitze. gtf-is
ribosoma
translacia 1 hidrolizi zrdis am safexuris
sizustesa da efeqturobas.
polipeptidi

mrnm

ribosoma gamzade-
saiti saiti
buli Semdegi amino-
acil trnm-isTvis gtf

2 gdf

2
3
gdf
translokacia. ribosoma peptiduri bmis warmoqmna.
axdens A saitze ganlagebuli didi suberTeulis rrnm-is
gtf
trnm-is translokacias P molekula akatalizebs
saitze. P saitze mdebare peptiduri bmis warmoqm-
Tavisufali (carieli) trnm nas A saitze mdebare axal
gadadis E saitze, saidanac is aminomJavasa da P saitze
gamoiyofa. mrnm gadaadgil- ganlagebuli mzardi
deba masTan dakavSirebuli polipeptidis karboqsi
trnm-ebiT da Semdgom kodons bolos Soris. am safexuris
miitans translaciisaTvis A Sedegad, polipeptidi aR-
saitze. moCndeba A saitze mdebare
trnm-Tan dakavSirebuli.

S S suraTi 17.18  translaciis elongaciis cikli. diagramaze ar aris naCvenebi cilebi, romelTac elongaciis
faqtorebi ewodebaT. gtf-is hidrolizi mniSvnelovan rols asrulebs elongaciis procesSi.

damatebaSi monawileobs ramodenime cila, romelTac
elongaciis faqtorebs uwodeben. axali aminomJavas dam-
ateba sam-safexuriani cikliT xorcieldeba, rac aRwer-
ilia suraT 17.18-ze. energiis xarjva xdeba pirvel da
mesame safexurebze. kodonis amocnoba saWiroebs gtf-is
ori molekulis hidrolizs, rac am safexuris sizustesa
da efeqturobas zrdis. kidev erTi gtf hidrolizdeba
translokaciis safexuris energiiT uzrunvelsayofad.
m rnm gadaadgildeba ribosomaSi mxolod erTi
mimarTulebiT, 5’ boloTi win; es Seesabameba ribosomis
gadaadgilebas m rnm -ze 5’ _ 3’ mimarTulebiT. mniS-
vnelovania, rom ribosoma da m rnm erTmaneTis mimarT
moZraoben erTi mimarTulebiT da gadaadgileba
kodonobiT xdeba (anu, SeiZleba iTqvas, rom moZraobis
biji aris kodoni). prokariotebSi elongaciis cikls
wamis meaTedze naklebi dro sWirdeba da jaWvze yove-
li aminomJavas damatebisas Tavidan meordeba, sanam ar
dasruldeba polipeptidis sinTezi.
translaciis terminacia
translaciis saboloo stadia terminaciaa (suraTi
17.19). elongacia grZeldeba, sanam mrnm-ze ganlage-
buli stop kodoni miaRwevs ribosomis A saits. fuZeTa
tripletebi uag, uaa da uga ar kodireben aminomJavebs,
aramed moqmedeben, rogorc translaciis dasrulebis
signalebi. cila, romelsac gamoTavisuflebis (ril-

387
 1 2 3
gamoTavisuflebis Tavisufali
faqtori polipeptidi

stop kodoni (uag, uaa an uga)

1 rodesac ribosoma miaRwevs stop kodons m rnm -ze, ribosomis A saiti, t rnm -is nacvlad, Rebulobs cilas, saxelad gamoTavisu-
flebis faqtori.
2 gamoTavisuflebis faqtori axdens P saitze mdebare t rnm -sa da polipeptiduri jaWvis bolo aminomJavas Soris arsebul bmis
hidrolizs. amrigad, polipeptidi gamoTavisufldeba ribosomidan.
3 ori ribosomuli suberTeuli da ansamblis sxva komponentebi disocirdebian.

SS suraTi 17.19  translaciis terminacia

izing) faqtori ewodeba, pirdapir ukavSirdeba stop poliribosomebi

kodons A saitze. gamoTavisuflebis faqtori iwvevs
polipeptidur jaWvTan, aminomJavis nacvlad, wylis
molekulis mierTebas. es reaqcia iwvevs dasrulebuli
polipeptidis hidrolizs P saitze ganlagebuli trnm-
isagan; da polipeptidi gamoiyofa ribosomis didi su-
berTeulis gasasvleli gvirabidan (ix. suraTi 17.16a).
amis Semdeg, translaciuri ansamblis danarCeni nawile-
bic iSleba.
mzardi
polipeptidi
Semomavali
ribosomuli
suberTeuli
erT ribosomas SeuZlia saSualo zomis polipepti-
dis sinTezi wuTze nakleb droSi. Tumca, Cveulebriv,
erTi mrnm erTdroulad polipeptidis mravali aslis
Sesaqmnelad gamoiyeneba, radganac ramdenime ribosomas
erTdrulad SeuZlia erTi mrnm-dan mesijis transli-
reba. mas Semdeg, rac ribosoma gaivlis start kodons,
mrnm-s SeiZleba daukavSirdes meore ribosoma; amgvarad,
m rnm-is gaswvriv SeiZleba imyofebodes ramodenime ri-
bosoma. ribosomebis aseT rigs poliribosomebs (an
awyobili
polisomebs) uwodeben da maTi danaxva eleqtronuli
polipeptidi mikroskopiTaa SesaZlebeli (suraTi 17.20). poliribo-
somebi nanaxia rogorc proka­riotul, ise eukariotul
ujredebSi. isini ujreds saSualebas aZleven, Zalian
swrafad daasinTezos po­lipeptidis mravali asli.

poliribosoma
mrnm-is funqcionaluri cilis warmoqmnis
mrnm-is
dasawyisi
a. (5’ bolo) bolo dasruleba da mizanmimarTuli gadatana
(3’ bolo)
translaciis procesi xSirad ar aris sakmarisi
b. funqcionaluri cilis Sesaqmnelad. am nawilSi, Tqven
gaecnobiT modifikaciebs, romelTac polipeptiduri
jaWvebi ganicdian translaciis procesis Semdeg da

ribosomebi (a) mrnm-is molekula, Cveulebriv, erTdroulad translirdeba

mrnm ramodenime ribosomis mier, romelTa klasterebsac poliribo-
somebs uwodeben.

(b) am mikrofotoze Cans didi poliribosoma prokariotul uj­

S S suraTi 17.20   poliribosomebi redSi (TEM- transmisiuli eleq­tronuli mikroskopi).

388
 1 ribosoma 5

4 4
mrnm
3 6
sasignalo 2
peptidi
endoplazmuri
badis membrana

signalis
amomcnobi
nawilaki
(san) sasignalo peptidi cila
mocilda

san
cila
receptori

endoplazmuri translokaciis
badis sanaTuri kompleqsi

S S suraTi 17.21  cilebis eb-isken mizanmimarTuli gadatanis sasignalo meqanizmi. polipeptidi, romelic gamiznu-
lia endomembranuli sistemis an ujredidan sekreciisaTvis, iwyeba sasignalo peptidiT, aminomJavebis rigiT, romelic
mimarTavs mas eb-isaken. es suraTi gviCvenebs sekretoruli cilis sinTezs da mis imavdroul imports eb-Si. eb-sa da Semdeg
goljis aparatSi, cila ganicdis Semdgom procesings. sabolood, satransporto vezikula gadaitans mas plazmur mem-
branasTan, ujredidan gamosayofad (ix. suraTi 7.10).

aseve zogierT im meqanizms, romelTa saSualebiT xdeba
dasrulebuli cilebis mizanmimarTuli gadatana ujre-
dis specifikuri ubnebis, saitebisaken.
cilis daxveva da post-translaciuri
modifikaciebi
sinTezis procesSi, polipeptiduri jaWvi spon-
tanurad iwyebs daxvevasa da dakecvas da warmoiqmneba
specifikuri konformaciis funqcionaluri molekula:
sam-ganzomilebiani molekula meoradi da mesameuli
struqturiT (ix. suraTi 5.20). geni gansazRvravs pirve-
lad struqturas, xolo pirveladi struqtura, Tavis
mxriv, gansazRvravs konformacias. xSir SemTxvevaSi,
Caperonuli cila (Caperonini) exmareba polipeptids
sworad daxvevaSi (ix. suraTi 5.23).
sanam cila Tavisi GgansakuTrebuli samuSaos Ses-
rulebas SeZlebs ujredSi, SesaZloa, saWiro iyos
damatebiTi safexurebi _ post-translaciuri modi-
fikaciebi. garkveuli aminomJavebi ganicdian qimiur
modifikaciebs Saqrebis, lipidebis, fosfatis jgufebi-
sa an sxva danamatebis mierTebiT. fermentebma Sesa-
Zloa, erTi an meti aminomJava moaSoron polipeptiduri
jaWvis aminobolos. zog SemTxvevaSi, erTi polipepti-
duri jaWvi, SesaZloa, fermentulad or an met nawilad
gaixliCos. magaliTad, cila insulini Tavdapirvelad
erTi polipeptiduri jaWvis saxiT sinTezdeba, magram
aqtiuri mxolod mas Semdeg xdeba, rac fermenti jaWvis
centralur nawils amoWris da cila ori, erTmaneTTan
disulfiduri xidakebiT dakavSirebuli polipepti-
duri jaWvebisagan Sedgenili rCeba. sxva SemTxvevebSi,
damoukideblad dasinTezebuli ori an meti polipep-
tidebi, SesaZloa, SeerTdes da meoTxeuli struqturis
mqone cilis suberTeulebad iqces.
polipeptidebis mizanmimarTuli gadatana
specifikuri adgilebisaken
cilis sinTezSi aqtiuri eukariotuli ujredebis
eleqtronul mikrifotografiebze Cans ribosomebis (da
poliribosomebis) ori populacia: Tavisufali da daka-
vSirebuli (bmuli) (ix. suraTi 6.11). Tavisufali ribo-
somebi citozolSia gabneuli da umetesad asinTezeben
cilebs, romlebic ixsneba citozolSi da aqve funq-
cionirebs. amis sapirispirod, bmuli ribosomebi daka-
vSirebuli arian endoplazmuri badis (eb) citozolur
mxaresTan an birTvis garsTan. dakavSirebuli ribosome-
bi asinTezeben endomembranuli sistemis (birTvis gar-
si, eb, goljis aparati, lizosomebi, vakuolebi da
plazmuri membrana) cilebs da aseve cilebs, romlebic
unda gamoiyos ujredidan, rogoricaa insulini. Ta-
visTavad, ribosomebi identuria da SeuZliaT statusis
Secvla Tavisuflidan bmulisaken.
ra gansazRvravs, nebismier konkretul dros ribo-

389
soma Tavisufali iqneba citozolSi, Tu xorklian eb-
sTan iqneba bmuli? polipeptidis sinTezi yovelTvis
citozolSi iwyeba, sadac Tavisufali ribosoma mrnm-is
molekulis translacias iwyebs. procesi aqvec dasrul-
deba, Tu mzardi polipeptidi Tavad ar ukarnaxebs ri-
bosomas, daukavSirdes eb-s. endomembranuli sistemis
an sekreciisaTvis gamiznuli cilebis polipeptidebi
moniSnulia sasignali peptidiT, romelic mimarTavs
cilas eb-saken (suraTi 17.21). sasignalo peptidi, daax-
loebiT 20 aminomJavasagan Sedgenili Tanamimdevroba,
ganlagebulia polipeptidis wamyvan (amino) boloSi,
an mis siaxloves, da ribosomidan mis gamosvlisTanave
amoicnoba cila-rnm-uli kompleqsis, saxelwodebiT _
signalis amomcnobi nawilakis (san-ingl. SRP), mier.
es nawilaki moqmedebs, rogorc adaptori, romelsac mi-
aqvs ribosoma eb-is membranaSi CaSenebul receptorul
cilasTan. es receptori warmoadgens multicilovani
translokaciuri kompleqsis nawils. polipeptidis sin-
Tezi aq grZeldeba da mzardi polipeptidi cilovani
foris saSualebiT gaivlis membranas da aRwevs eb-is
RruSi (lumenSi). sasignalo peptidi, Cveulebriv, fer-
mentulad moSordeba polipeptids. dasrulebuli po­
li­peptidis danarCeni nawili, Tuki is sekretoruli
cila unda iyos, gamoTavisufldeba eb-is RruSi arse-
bul xsnarSi (rogorc suraT 17.21-ze); da piriqiT, Tuki
polipeptidi membranuli cila unda iyos, is rCeba eb-is
membranaSi nawilobriv CaSenebuli.
sxva tipis sasignalo peptidebi gamoiyeneba polipep-
tidebis mizanmimarTulad gadasatanad mitoqondriebis,
qloroplastebis, birTvis SigTavsisa da sxva organ-
elebisaken, romlebic ar ekuTvnian endomembranul si­
s­t emas. am SemTxvevebSi, mniSvnelovani gansxvaveba is
aris, rom translacia citozolSive mTavrdeba da poli-
peptidi Semdgom importirdeba organelaSi. translo-
kaciis meqanizmebi aseve varirebs, magram yvela aqamde
Seswavlil SemTxvevaSi, `safosto indeqss~, romelic
mimarTavs cilebs sekreciisa Tu ujreduli dislokaci-
isaken, raime tipis sasignalo peptidi warmoadgens. pro-
kariotebi aseve iyeneben sasignalo Tanamimdevrobebs
cilebis mizanmimarTulad gadasatanad sekreciisaTvis.
koncefcia 17.5
rnm mraval rols asrulebs
ujredSi: mimoxilva
rogorc Cven ukve vnaxeT, cilis sinTezis (da eb-
isaken mizanmimarTuli gadatanis) ujredul manqanaSi
dominirebdnen sxvadasxva tipis rnm-ebi. mrnm-is garda,
aq Sedis trnm, rrnm da, eukariotebSi, mbrnm da san rnm
(cxrili 17.1). rnm-is kidev erTi tipi _ mcire birT-
vakuli rnm (mbkrnm, ingl. snoRNA) xels uwyobs pre-
rrnm transkriptebis procesings birTvakSi. es pro-
cesi aucilebelia ribosomis warmoqmnisaTvis. rnm-is
am mcire molekulebs mravalferovani funqciebi aqvT:
struqturulidan informaciulamde da katalizuramde.
Tanamedrove kvlevebma aseve gamoavlina mcire, erT an
or-jaWviani rnm-is molekulebis arseboba, romlebic
moulodnelad mniSvnelovan rols asruleben genebis
rnm-is warmomadgenlebi
17.1
cxrili
eukariotul ujredSi
rnm-is warmomadgenlebi funqciebi
matricul rnm-s (mrnm) gadaaqvs informacia dnm-dan
ribosomebamde, romelic gansazRvavs
cilebSi aminomJavebis Tanmimdevrobas.
satrasnporto rnm-i (srnm) moqmedebs rogorc molekula-adap-
tori cilebis sinTezisas; gardaqmnis
mrnm-is kodonebs amino mJavebad.
ribosomuli rnm-i (rrnm) asrulebs ribosomebSi katalizur
(ribozimul) da struqturul rols.
pirveladi transkripti mrnm-sTvis, srnm-sTvis an rrnm-
saTvis `winamorbedis~ rols TamaSobs
iqamde,sanam ar Sewyvets arsebobas
Serwymis an, piriqiT, daSlis Sedegad.
zogi rnm introni moqmedebs rogorc
ribozimi, da katalizirebs sakuTar
daSlas.
mcire birTvuli rnm-i (mbrnm) TamaSobs struqturul da katalizur
rols splaisozomebSi, romlebic war-
moadgenen cilis da rnm-is kompleqss,

17.4
koncefcia testi
1. romeli ori procesi warmoadgens ga­rants,
rom mzard polipeptidur jaWvs daemateba
swori (saWiro) aminomJava?
2. aRwereT, ra sargebloba SeiZleba mouta-
nos ujreds poliribosomebis warmoqmnam?
3. aRwereT, rogor transportirdeba endo-
membranul sistemaSi sasekrecio polipep-
tidi?
san (rnm)
mcire birTvuli rnm (mcbrnm)
mcire interferirebuli
rnm (mirnm) da mikro-rnm (mirnm)
romelic Tavis mxriv aerTianebs
pirvelad transcripts.
warmoadgens signalis amomcnobi
nawilakis komponents (san), cilis da
rnm-is kompleqsi, romelic amoicnobs
sasignalo peptidebs polipeptidebSi,
endoplazmuri badisaTvis.
monawileobas iRebs birTvSi mimdin-
are pre-rnm-is transkripciaSi ribo-
somuli suberTeulis formirebisas.
CarTulia genis eqspresiis
regulaciaSi.

390