d.

miqelaZe

(leqciaTa kursi)

Tbilisi - 2001
 Sesavali

bioqimia aris mecniereba, romelic cocxal organizmSi mimdinare qimiur

procesebs Seiswavlis da mis mTavar amocanas cocxali sistemebis
funqcionirebis, gamravlebis da populaciebad formirebis meqanizmebis dadgena
warmoadgens. winaT, biologiasa da qimias maTi principebiTa da kanonebiT
erTmaneTisagan damoukidebel mecnierebebad miiCnevdnen. magram dReisaTvis ukve
naTelia, rom bioqimiis gamiyofa biologiisagan SeuZlebelia, rom verc erTi
biologiuri kanoni Tu movl;ena ver aixsneba im qimiuri procesebis gareSe,
romlebic biologiur obieqtebSi mimdinareoben.
Tavisi arsiT, bioqimia warmoadgens superqimias romelSic gaerTianebulia
qimiis yvela tradiciuli dargi da miuxedavad imisa, rom ujredSi mimdinare
procesebi emyarebian cnobil qimiur Tu fizikur kanonzomierebebs, isini
gacilebiT ufro rTuli arian, vidre aracocxal sistemaSi mimdinare procesebi
da Seicaven mraval fenomens, romelTa amocnobac dRes did sirTuleebs
warmoadgens. es kanonzomierebani damyarebulia cocxali sistemebis
Taviseburebebze da maTTvis damaxasiaTebelia moqmedebis gansakuTrebuli
principebi.
pirveli gansakuTrebuli principi Tu Tavisebureba, romelic ujreds
axasiaTebs, aris makromolekulebis arseboba, es ukanaskneli ki sirTuliT da
mravalferovnebiT xasiaTdebian. magaliTad, nawlavis Cxiris (E. Coli) ujredSi
5000-mde organuli naerTia, romelTagan 3000 cilas, xolo 1000 – nukleinis mJavas
warmoadgens. rTul organizmebSi makromolekulaTa raodenoba izrdeba da
adamianebSi is ukve milionebs iTvlis. aqve unda aRiniSnos, rom cilebis
umravlesoba damaxasiaTebelia garkveuli saxeobisTvis da igive E. Coli –s 3000
cilidan arc erTi ar aris msgavsi adamianis cilisa. dReisaTvis 1200000
biologiuri saxeoba aris cnobili da advili gamosaTvlelia, rom bunebaSi
individualuri cilebis raodenoba TiTmis trilion aRwevs. aRsaniSnavia is, rom
miuxedavad aseTi mravalferovnebisa makromolekulebis agebuleba daiyvaneba
Senebis sakmaod primitiul donemde. cilebis SenebaSi monawileobas iRebs
mxolod 20 aminomJava, xolo nukleinis mJavebisa ki – 5 nukleotidi. aminomJavebi
da nukleotidebi saerToa mTeli cocxali samyarosaTvis da molekulaTa
mravalfervneba ganpirobebulia sxvadasxvaobiT mxolod maT TanmimdevrobaSi.
makromolekulebis SenebaSi monawile samSeneblo blokebs – aminomJavebs da
mononukleotidebs _ kidev erTi mniSvnelovani funqcia aqvT: isini monawileobas

2
iReben hormonebis, alkaloidebis, pigmentebis, kofermentebis da sxva organul
naerTTa sinTezSic.
zemoT Tqmulidan gamomdinareobs, rom miuxedavad cocxali ujredis
urTulesi molekuluri organizaciisa, misTvis damaxasiaTebelia sawyisi
nivTierebebis simartive, saamSeneblo masalebis erTferovneba. garda amisa,
SesaZlebelia vivaraudoT, rom dedamiwaze arsebul cocxal sistemebs saerTo
winapari hyavdaT. da bolos, zemoTTqmuli miuTiTebs imaze, rom cocxali
organizmebisaTvis damaxasiaTebelia “molekuluri ekonomia” da isini
saamSeneblod iyeneben mxolod iseT naerTebs, romlebic maT saSualebas aZleven
minimaluri danakargebiT icxovron garemos garkveul pirobebSi.
meore gansakuTrebuli principi, romelic cocxal ujreds axasiaTebs,
mdgomareobs energiis gardaqmnis TaviseburebebSi. cnobilia, rom Termodinamikis
meore kanonis Tanaxmad yoveli sistema miiswrafis mouwesrigeblobisaken,
qaosisaken, entropiis zrdisaken. cocxalma organizmebma rom SeinarCunon
molekuluri organizaciis maRali done maT esaWiroeba Tavisufali energiis
mudmivi STanTqma da misi dagroveba. amitom organizmi imyofeba garemosTan Ria
sistemur, magram arawonasworul urTierTkavSirSi. cocxali sistema garemodan
Tavisufal energias iRebs da iwvevs masSi entropiiis zrdas. organizmi
funqcionirebs garkveul temperaturaze da SeiZleba iTqvas, rom igi warmoadgens
izoTermul qimiur manqanas.
mesame gansakuTrebuli principi, romelic ansxvavebs cocxal sistemas
aracocxalisagan aris qimiuri reaqciebis Taviseburebani. ujreds SeuZlia
ifunqcios, rogorc qimiuri manqana mxolod im SemTxvevaSi, Tu qimiuri reaqciebis
mimdinareoba specifikuri da maRal efeqturi iqneba, xolo maTi siCqare _ swrafi.
bioqimiuri reaqciebi akmayofileben am moTxovnebs, radgan maTi umravlesoba
warmoebs fermentebis saSualebiT. fermentebi arian maRalspecializebuli
cilovani molekulebi, Semdgari Cveulebrivi aminomJavebisagan. isini qimiuri
reaqciebis katalizatorebs warmoadgenen da zogierTi maTgani miliardjer, da
ufro metadac, aCqarebs reaqcias. rogorc wesi, erTi fermenti erT garkveul
bioqimiur reaqcias atarebs da misi xangrZlivoba wamis meaTasebiT ganisazRvreba.
fermentebiT warmoebuli bioqimiuri reaqciebis gamosavali 100%-s aRwevs,
rac ganpirobebulia maTi maRali specifiurobiT gardasaqmneli nivTierebis
mimarT. maT urTerTobaSi isaxeba cocxalis sistemebis damaxasiaTebeli ZiriTadi
principi _ struqturuli komplementarobis (Sesabamisobis, Sesatyvisobis)
principi. am principis arsi mdgomareobs imaSi, rom makromolekulis (Cven
SemTxvevaSi fermentis) garkveuli ubani, cnobs da ikavSirebs mxolod erTi tipis

3
naerTs. makromolekulis am ubans (aqtiur centrs) dasakavSirebeli
nivTierebisadmi Sesatyvisi samganzomilebiani sivrciTi agebuleba aqvs da masTan
sxva naerTis dakavSireba gamoricxulia. es principi sakmaod gavrcelebulia
cocxal sistemebSi da gvxdeba iseTi tipis reaqciebSi, rogoric arian hormonebis
urTierTmoqmedeba receptorebTan, antisxeulebis _ antigenebTan da a.S.
cocxali sistemebisTvis damaxasiaTebeli meoTxe gansakuTrebuli principi
mdgomareos ujreduli reaqciebis TviTregulaciaSi. ujredSi aTasobiT reaqcia
mimdinareobs Tanadroulad sivrcis mcire monakveTSi. igive nawlavis CxirSi, sami
tipis substratisagan erTdroulad sinTezirdeba aTasobiT rTuli organuli
nivTiereba; mxolod wuTebi esaWiroeba garkveuli tipis cilis sinTezs (aseTi
tipis naerTebi rom xelovnurad miviRoT mecnierTa didi koleqtivis muSaobaa
saWiro Tveebis ganmavlobaSi.) rogor awarmoebs ujredi qimiuri reaqciebis aseT
efeqtur warmarTvas? upirveles yovlisa organuli naerTebis Senebisas
gamoiyeneba reaqciaTa Sualeduri produqtebi e.w. metabolitebi, romelTa
dagrovebac ujredSi garkveul fermentTa damuxruWebas, maT inhibirebas iwvevs.
magaliTad, gvaqvs reaqciaTa jaWvi A B C D I F G romlis saboloo
produqti G ujredSi didi raodenobiT dagrovda; igi iwyebs jaWvis garkveul
etapze moqmedebas da iwvevs misi momsaxure fermentis inhibirebas. dauSvaT jaWvi
damuxruWda C D doneze, maSin ujredSi grovdeba C da Tu am ukanasknels
gardaqmnis sxva gza, reaqcia iwyebs am mimarTulebiT msvlelobas, ris Sedegadac
warmoiqmneba ara G, aramed N:
A B C D

N
reaqciaTa aseTi regulacia metyvelebs imaze, rom ujredis erT nawilSi
mimdinare reaqciis siCqare SeiZleba damokidebuli iyos im naerTis
koncentraciaze, romelic ujredis sxva nawilSi warmoiqmneba. aseT naerTebs
regulatoruli metabolitebi ewodeba da maTi saSualebiT ujredi axorcielebs
naerTTa sinTezs mkacrad gansazRvruli raodenobiT reglamentirebul droSi.
mexuTe, da erT-erTi mTavari principi, romelic cocxal sisitemebs udevs
safuZvlad aris kvlavwarmoqmnis, reproduqciis principi. am princips sami
ZiriTadi maxasiaTebeli gaaCnia. pirveli _ mdgomareobs Cawerili genetikuri
informaciis kompaqturobaSi. magaliTad, informacia adamianis Sesaxeb inaxeba
dnm-Si, romlis masa 6x10-12 g tolia. meore SesaniSnavi Tviseba, romelic

4
axasiaTebs reproduqciis process aris dnm-Si Cawerili informaciis
stabiluroba. miuxedevad imisa, rom TviT dnm labilur molekulas warmoadgens,
ujredSi misi arsebobis SemTxvevaSi Sesabamisi aRmdgeneli fermentebis gamo,
iSviaTad ganicdis cvlilebebs. sakmarisia Tundac aRiniSnos is, rom Cveulebrivi
baqteriis genetikurma kodma mTeli evoluciis manZilze arc Tu iseTi didi
cvlilebebi ganicada. genetikuri informaciis stabiluroba ganpirobebulia dnm-
is ormagspiraliani jaWvis arsebobiT, romelTagan TiToeuli komplementaruli
meoresi da erT-erTis dazianebis SemTxvevaSi specialuri fermentebis
saSualebiT aRidgens Tavis struqturas meore jaWvis saSualebiT.
genetikuri informaciis gadatanis mesame Tavisebreba mdgomareobs translaciis
procesSi. am dros dnm-is molekulaSi aRwerili erTganzomilebiani informacia
Semdgari mxolod mononukleotidTa gansxvavebuli Tanmimdevrobisagan,
gadaiwereba rTul samganzomilebian cilis molekulaSi.
amrigad, Cvens mier aRwerili iyo cocxali sistemisTvis damaxasiaTebeli
kanonzomierebani, romlebsac zogadad SeiZleba sicocxlis molekuluri logika
vuwodoT. zemoTTqmuli SesaZlebelia Sejamdes Semdegnairad: cocxali ujredi
warmoadgens izoTermul Ria sistemas, romelsac TviTSenebis, TviTregulaciis da
TviTgamravlebis unari aqvs. es sistema Sedgeba didi raodenobiT erTmaneTTan
dakavSirebuli qimiuri reaqciebisagan, romlebic Cqardebian cilovani bunebis
katalizatorebiT _ fermentebiT da romlebsac TviT ujredi asinTezirebs.
ujredis qmedeba maRalekonomiuri da efeqturia.
biologiuri qimia sakmaod vrcel mecnierebas warmoadgens. igi moicavs
biologiis TiTqmis yvela disciplinas da saSualebas gvaZlevs CavwvdeT
biologiur obieqtSi momxdar movlenebs. is ras Cven gvesmis biologiuri qimiis
cnebis qveS warmoadgens im centralur mecnierebas, romelic ganmartavs
biologiis TiTqmis yvela fenomens.

5
 ujredis molekuluri komponentebi

biologiuri nivTiereba SedgenilobiT Zlier gansxvavdeba liTosferos da

atmosferos Semadgenlobisagan. im asi qimiuri elementidan, romelic dedamiwis
qerqSia aRmoCenili, ZiriTadad 32 iRebs monawileobas cocxali ujredis SenebaSi.
am ukanasknelSi yvelaze ufro xSirad gvxvdebian: naxSirbadi, wyalbadi, azoti da
Jangbadi. maTi gavrceleba cocxal sistemebSi ganpirobebulia imiT, rom jer
erTi, isini mcire zomisani arian, da Semdeg - sxvebTan SedarebiT advilad
warmoqmnian kovalentur bmas. amis gamo, maTi saSualebiT SasaZlebelia didi
raodenobiT sxvadasxva tipis maRalmolekulur naerTTa warmoqmna.
organuli biomolekula sawyiss naxSirorjangidan, wylidan da
atmosferuli azotidan iRebs. SemdgomSi am naerTebidan warmoebs e.w. Sualeduri
pruduqtebis - metabolitebis warmoqmna, romlebic dasabams aZleven
makromolekulaTa oTx did klass: cilebs, cximebs, nukleinis mJavebs da
polisaqaridebs. nukleinis mJavebis metabolitebs warmoadgenen riboza da
karbamoilfosfati, romlebidanac SemdgomSi sinTezirdebian mononukleotidebi.
es ukanasknelni erTmaneTs ukavSirdebian kovalenturi bmiT da warmoqmnian dnm-is
da rnm-is jaWvebs. cilebis SenebaSi monawileobas iReben aminomJavebi, romlebic
warmoiqmnebian -ketoaminomJavebisagan. fosfoenolpiroyurZnis da vaSlis
mJavebisagan warmoiqmnebian martivi da rTuli naxSirwylebi, xolo aminomJavidan
da malonis mJavidan – cximebi (lipidebi). Sualedur naerTTa metabolizmi
mWidrod aris dakavSirebuli erTmaneTTan da maTi urTierTgardaqmna specialuri
ciklebis saSualebiT xdeba.
cximebidan, cilebidan da nukleinis mJavebidan warmoiqmnebian zemolekuluri
kompleqsebi: membranebi, ribosomebi, ujredis SekumSvadi elementebi da sxva.
bioqimiuri organizaciis ufro rTul stadias organoidebis formireba
warmoadgens. am ukanasknelTa erToblioba warmoSobs im rTul cocxal
warmonaqmns, romelsac ujredi ewodeba.

wyali

wyali ujredis ZiriTad komponents warmoadgens da pirdapir Tu arapirdapir

monawileobas iRebs praqtikulad ujredis yvela bioqimiur procesSi. sxva
gamxsnelebTan SedarebiT mas gaaCnia maRali lRobisa da duRilis temperetura,
aorTqlebis siTbo, xvedriTi siTbotevadoba da zedapiruli daWimuloba. wylis

6
es Tvisebebi ganpirobebulia misi qimiuri TaviseburebebiT da mdgomareobs imaSi,
rom eleqtronuaryofiTi Jangbadis atomi iswrafvis miiziidos eleqtronebi
wyalbadis atomebidan, ris gamoc maT atomgulebs uCndebaT dadebiTi muxti:
+
H
O --
+
H Register
Please O
AaseTi polaruli bunebis gamo wylis molekulebi cdiloben miizidon
erTmaneTi, ris Sedegadac warmoiqmneba struqtura, sadac wylis erTi molekulis
eleqtronuaryofiTi Jangbadi orientirebuli mezobeli molekulis wyalbadTan.

H H
O H O H O
H
H

am tipis eleqtrostatikur urTierTobas wyalbadur bmas uwodeben. kovalentur

bmasTan SedarebiT igi gacilebiT sustia. wyalbadur bmas warmoqmnis aramarto
wylis molekulebi, aramed yvela is naerTi, romelsac Zlieri
eleqtronuaryofiTi atomi (mag., O, F, N) da sxva atomebTan kovalenturad
dakavSirebuli wyalbadis atomebi aqvs. wyalbaduri bmis saSualebiT xdeba
makromolekulaTa SenebaSi monawile naerTTa orientireba erTmaneTis mimarT,
rac did gavlenas axdens maT konformaciaze. wyalbaduri bma kovalenturTan
erTad warmoadgens qimiuri urTierTobis ZiriTad tipebs biologiur
makromolekulebSi.
mesame tipis ZiriTadi Zala, romelic aseve monawileobs makromolekulaTa
fiziko-qimiuri Tvisebebis CamoyalibebaSi, aris hidrofoburi urTierToba.
cnobilia, rom cximebis wyalSi gaxsnis Sedegad warmoiqmnebian micelebi,
romelTa polaruli nawili mimarTulia gareT, wylisken, xolo cximovani,
hidrofoburi _ SigniT. lipidebis aseTi orientaciis Sedegad maT hidrofobur
nawilebs Soris warmoiqmneba damatebiTi mizidulobis Zala, romelsac
hidrofoburi urTierToba ewodeba.

7
 H2O H 2O H 2O
+ - - +
- +

+
+ - + -

H 2O Register
Please H 2O H 2O

wyals, iseve rogorc sxva naerTebs axasiaTebs ionizacia. misi disociacia

SeiZleba Semdegnairad gamoisaxos:

H2O H+ + OH-

Tu wyals damatebuli aqvs mJave nivTiereba (mag, HCl), maSin xsnarSi matulobs H+-
is raodenoba, xolo Tu masSi SevitanT tutes (mag, NaOH) _ moimatebs OH-is
raodenoba. Tu H+-is ionis koncentracia [H+] tolia [OH-] _ xsnari neitraluri da
misi pH tolia 7-is. pH warmoadgens sidides, romelic asaxavs xsnarSi H+-ionTa
koncentracias. neitralur pH-ze [H+] = [OH-] da maTi koncentracia tolia 10-7 M.
pH 6-ze [H+]-is koncentracia tolia 10-6 moli/l, xolo [OH-]-isa ki 10-8 moli/l da
a.S. rac ufro maRalia xsnarSi [H+]-is ionebis koncentracia, miT ufro dabalia
xsnaris pH. magaliTad, kuWis wvenis pH tolia 1,2-3,0 da ganpirobebulia masSi HCl-
is arsebobiT; da piriqiT rac ufro naklebia xsnarSi [H+] ionebi (Sesabamisad
metia [OH-]) miT ufro maRali mniSvneloba aqvs pH (mag, kuWqveSa jirkvlis
sekretis pH tolia 7,8-8,0, xolo sisxlisa _ 7,1. igi neitraluria). pH-is sidide
biologiuri siTxeebis ZiriTad maxasiaTebels warmoadgens da amitom igi
intensiurad isazRvreba samedicino dawesebulebebSi.
zogierTi qimiuri naerTi wyalSi gaxsnisas e.w. buferul Tvisebebs amJRavnebs.
buferuli xsnarebi ewodeba iseT xsnarebs, romlebic garkveul intervalSi ar
icvlian pH-s masze tutis an mJavis damatebis Sedegad. buferul Tvisebebs
amJRavneben fosfatis, karbonatis da sxva marilebi, agreTve aminomJavebi, cilebi
da sxva organuli nivTierebebi. CamoTvlili naerTebi aniWeben sisxlis plazmas
buferul Tvisebas. Tu magaliTad 1 litr fiziologiur xsnars, romelic Sedgeba

8
0,15 M NaCl davumatebT 1 ml 10N HCl pH-is mniSvneloba Seicvleba 7-dan 2-mde. pH-is
aseTi daqveiTeba gamowveuli iqneba imiT, rom NaCl-s ar gaaCnia buferuli
Tvisebebi. Tu analogiur cdas Cavatarebs sisxlis plazmaze, misi pH umniSvnelod
Seicvleba (7,4-dan 7,2-mde). es metyvelebs sisxlis plazmis buferul Tvisebebze.

cilebi da maTi biologiuri funqcia

ujredis cxovelmoqmedebaSi monawile organul nivTierebebs Soris cilebs

wamyvani mnisvneloba eniWebaT, radgan isini awarmoeben Cawerili informaciis
realizacias. ujredis mSrali wonis 50%-s cilebi Seadgenen.

cilebis agebuleba

dReisaTvis gamoyofili da Seswavlilia aTasobiT individualuri cila. maT

SemadgenlobaSi Sedian naxSirbadi, azoti, wyalbadi da Jangbadi. garda am
elementebisa, cilebis umravlesoba Seicavs gogirds, xolo zogierTi _
fosfors, rkinas, TuTias, spilenZs. cilebs maRali molekuluri wona aqvT da
maTi daSliT miiReba dabalmolekuluri naerTebi, romlebsac aminomJavebi
ewodebaT. aminomJavebisagan aSenebulia cilovani molekulis jaWvi. cilebSi
Semaval aminomJavebSi Tavisufali karboqsilisa da aminis jgufebi Canacvlebuli
arian -naxSirbadis atomTan. aminomJavebis zogadi struqtura Semdegnairad
gamoisaxeba:

NH2 CH COOH

aminisa da karboqsilis jgufebis arseboba damaxasiaTebelia yvela

aminomJavisTvis. isini erTmaneTisagan gansxvavdebian R-gverdiTi jaWviT. cilis
sinTezisaTvis gamoiyeneba 20 tipis aminomJava, romlebic (e.w. peptiduri) bmiT
ukavSirdebian erTmaneTs. erTi aminomJavis karboqsilis jgufi peptidur bmas
warmoqmnis meore aminomJavis aminis jgufTan:

NH2 – CH – CO – NH – CH – CO –NH – CH----------------- CH – CO – NH – CH – COOH

R R R R R

9
aminomJavebis erTmaneTTan dakavSirebis Sedegad warmoiqmneba cilis polimeruli
molekula.
TiToeul cilaSi aTasobiT aminomJava SeioZleba iyos warmodgenili.
ganasxvaveben martiv da rTul cilebs. martivi cilebi mxolod aminomJavebisagan
Sedgebian, xolo rTul cilebSi aminomJavebis garda CarTulia sxva naerTebic. am
ukanasknels prosTeturi jgufi ewodeba. prosTeturi jgufis mixedviT
ganasxvaveben nukleoptoeidebs (CarTulia nukleinis mJava), lipoproteidebs
(CarTulia lipidi), metaloproteidebs (CarTulia metali). garda amisa, cilis
molekulas SesaZlebeli daukavSirdes fosformJavas naSTi (fosfoproteini),
martivi an rTuli naxSirwyali (glikoproteini) da a.S.
cilebis molekuluri wona meryeobs 10000-dan 1000000-mde. cilebi xSirad
erTiandebian zemolekuluri kompleqsebis warmoqmniT.

aminomJavebi _ cilebis struqturuli elementebi

rogorc ukve iTqva, cilis SenebaSi SesaZlebelia 20 aminomJavam miiRos

monawileoba. R radikalis polarobis mixedviT ganasxvaveben 4 tipis aminomJavas: I
– arapolaruli anu hidrofoburi aminomJavebi, II _ polaruli, aradamuxtuli
aminomJavebi, III _ dadebiTad damuxtuli aminomJavebi, IV _ uaryofiTad
damuxtuli aminomJavebi.
I jgufs anu hidrofobur aminomJavebs miekuTvnebian: alanini
valini, leicini, izoleicini, fenilalanini, prolini, triftofani, meTionini.
hidrofobur aminomJavebs axasiaTebT wyalSi susti xsnadoba.

O H2N CH C OH

O
H2N CH C OH CH2

H2N CH C OH CH CH3 CH CH3

CH3 CH3 CH3

alanini valini leicini

10
 O
O

O H2N CH C OH
H2N CH C OH
O
H2N CH C OH CH2
CH2
C OH
CH CH3

CH2
HN HN
CH3

izoleicini fenilalanini prolini triftofani

H2N CH C OH

CH2

CH2

CH3

meTionini

aminomJavebis II jgufs, romlebsac aradamuxtuli R _ radikali gaaCniaT,

miekuTvnebian: glicini, serini, treonini, cisteini, Tirozini.

O O

O H2N CH C OH H2N CH C OH

H2N CH C OH CH2 CH OH

H OH CH3

glicini serini treonini

11
 O

H2N CH C OH

CH2

H2N CH C OH

CH2

SH OH
cisteini Tirozini

III jgufs warmoadgenen e.w. mJave aminomJavebi, romelTa radikalic uaryofiT

muxts atarebs (COO-): asparaginis mJava, glutaminis mJava.

O H2N CH C OH

H2N CH C OH CH2

CH2 CH2

C O C O

OH OH
asparaginis mJava glutaminis mJava

da bolos, IV jgufSi gaerTianebuli arian aminomJavebi, romelTa radikali

dadebiTad aris damuxtuli (NH3+): lizini, arginini, histidini.

12
 O

O H2N CH C OH

H2N CH C OH CH2
O
CH2 CH2
H2N CH C OH
CH2 CH2
CH2
CH2 NH

CH2 C NH N

NH2 NH2 NH
lizini arginini histidini

neitralur pH-ze aminomJavebis karboqsilis jgufi kargavs H+-ions da imuxteba

uaryofiTad (COO-), xolo aminis jgufi ierTebs protons da imuxteba dadebiTad
(NH3+). amitom aminomJava neitralur pH-ze dipolur cviterionur formas iRebs:
+
NH 2 N H3
-
R C COOH R C COO
H Please Register H
aradisocirebuli disocirebuli
forma forma

sam aminomJavas, romlebic Seicaven aromatul birTvs, kerZod triftofans,

Tirozins da fenilalanins, ultraiisfer talRaze (280 nm) STanTqmis unari
gaaCniaT. vinaidan cilebis umravlesoba Seicavs am aminomJavebs, maTi
raodenobrivi gansazRvra advili SesaZlebelia speqtrofotometrze. garda amisa,
cilebis gansazRvra SeiZleba feradi reagentebiT, romlebic specifiurad
ukavSirdebian cilebs (aminomJavebs).
Tanamedrove meTodebis gamoyenebiT SesaZlebelia cilebis hidrolizi, masSi
Semavali aminomJavebis dayofa da analizi. aminomJavebis analizis erT-erT
gavrcelebul meTods qromatografia warmoadgens.

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 cilebis pirveladi struqtura

rogorc zemoT ukve iyo naTqvami, aminomJavebi warmoqmnian jaWvs peptiduri bmis
saSualebiT (R-CO-NH-R). maTi erTmaneTTan mierTebis Semdeg warmoiqmnebian cilebi
da peptidebi. Tu makromolekulis SenebaSi monawileobas iRebs 2-dan 50-mde
aminomJava, aseT naerTs peptidi ewodeba. maTi molekuluri wona 5000-6000-s
aRemateba. umaRles organizmebSi napovnia aTasobiT peptidi, romlebic cilis
SemadgenlobaSi ar Sedian da maT damoukidebeli funqcia aqvT. zogierT maTgans
biologiuri aqtivoba aqvs da monawileobas iRebs organizmis neiro-humoralur
regulaciaSi. magaliTad, endogenuri, xuTaminomJaviani peptidi _ enkefalini
warmoadgens neirogadamcems da neirohormons, romelic tkivilgamayuCebeli
TvisebebiT xasiaTdeba.
miRebulia, rom peptids, romlis SenebaSic monawileobs Rebulobs 60-ze meti
aminomJava, cila ewodeba. cilis SemadgenlobaSi SeiZleba Sediodes aTasobiT
aminomJava. aminomJavaTa jaWvi, dakavSirebuli peptiduri bmis saSualebiT,
warmoqmnis cilis pirvelad struqturas. dReisaTvis gaSifrulia asobiT cilis
pirveladi struqtura da dadgenilia masSi Semavali aminomJavebis Tanmimdevroba.
pirveli cila, romlis pirveladi struqtura gaiSifra insulini iyo (1953 w).
aRmoCnda, rom insulini, kuWqveSa jirkvalSi sinTezirdeba proinsulinis formiT,
romelic 84 aminomJavisagan Sedgeba. sinTezis Semdeg proinsulins scildeba 33
aminomJaviani peptidi da igi gadadis aqtiur formaSi, romelic ori jaWvisagan
Sedgeba: A – 21 aminomJaviT da B – 30 aminomJaviT. A da B jaWvebi erTmaneTTan
dakavSirebuli arian gogirdis xidakebiT (disulfiduri kavSiri, S-S bma).
AA

A-jaWvi

S S

S S

S S
Please Register
B-jaWvi

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 Tavis droze did miRwevad CaiTvala hemoglobinis aminomJavuri Tanmimdevrobis
dadgena. aRmoCnda, rom hemoglobini Sedgeba 4 polipeptiduri jaWvisagan _ 2
identuri -jaWvisagan (141 aminomJava) da ori identuri -jaWvisagan (146
aminomJava). am jaWvebis aminomJavuri Tanmimdevroba waagavs erTmaneTs, maTSi
msgavsi pirveladi struqturis bevri fragmentia. aseT, msgavsi aminomJavuri
Tanmimdevrobis mqone cilebs, homologiuri cilebi ewodeba. hemoglobinis
polipeptidebs maRal homologia aqvT agreTve kunTis cila-mioglobinTan.
dReisaTvis gaSifrulia aTasobiT cilis pirveladi struqtura. gairkva, rom
cilis pirvelad struqturaSi ar gvxvdebian martivi ganmeorebadi
Tanmimdevrobani, mag. ABCDABCDABCD da a.S., erTi aminomJava miyolebiT 3-jer
metad ar meordeba, mag. AAAABBBB da a.S. cilis Seneba aris rTuli da
specifikuri.
analogiuri funqciis mqone cilebs saxeobrivi gansxvaveba gaaCniaT. magaliTad,
adamianis insulinis A jaWvi absoluturad identuria Roris, ZaRlis, kurdRlis
da kaSalotis insulinis A jaWvisa, xolo B jaWvi msgavsia mxolod spilos
analogiuri jaWvisa.
mravali cilis pirveladi struqturis dadgenam SesaZlebeli gaxada zogierTi
genetikuri daavadebis gaSifrva. magaliTad, aRmoCnda, rom avadmyofebs, romlebic
daavadebulni arian namgliseburi anemiiT, hemoglobinis -jaWvSi, me-6
mdgomareobaSi myof aminomJava glutaminis mJavis nacvlad aminomJava valini aqvT.
aseTi tipis wertilovani mutaciebi safuZvlad udevs mraval genetikur
daavadebas.

cilebis meoreuli struqtura

cnobilia, rom yovel organul molekulas erTmagi kovalenturi bmis irgvliv
trialis unari Seswevs. Tu naerTi maRalmolekuluria da misi jaWvi Sedgeba
mravali kovalenturi bmebisagan, RerZis irgvliv brunva gaZnelebulia da
molekula iRebs erTaderT, Termodinamikurad stabilur formas, konformacias.
amerikuli mecnierebis polingisa da koris gamokvlevebiT dadginda, rom
aminomJavaTa jaWvi cilis molekulaSi ganmeorebad struqturas _ -spirals
qmnis. spiraluri struqtura stabilizebulia wyalbaduri bmis saSualebiT,
romelsac warmoqmnis –CO-NH- peptiduri kavSiris wyalbadi yoveli meoTxe
aminomJavis karbonilis jgufTan. spiralis biji tolia 5,4 anqstremis da mis erT
xveulSi monawileobs 3,6 aminomJavis naSTi. aminomJavebis R radikalebi
mimarTulia spiralis gareT. spiraluri struqturis formirebaSi ar

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monawileobs aminomJava prolini. -spiralis garda gvxvdeba sworxazovani
dakecili tipis struqtura, romelic -konformaciis saxeliT aris cnobili. -
konformaciaSi myofi paraleluri peptiduri jaWvebi dakavSirebuli arian
erTmaneTTan wyalbaduri bmebiT, riTac sakmaod stabilur struqturas
warmoqmnian. aseTi tipis peptiduri konformacia gvxvdeba, magaliTad Tmis
keratinSi, abreSumis fibrionSi da sxva. -spirals da -konformacias cilebis
meoreul struqturas uwodeben.

cilebis mesameuli struqtura

polipeptidis -spirali umetes SemTxvevebSi gorgalisebr, globularul
konformacias iRebs. es ukanaskneli ganpirobebulia aminomJavaTa fiziko-qimiuri
TvisebebiT. rogorc ukve iTqva, aminomJavaTa radikalebi -spiralis warmoqmnaSi
monawileobas ar iReben da mimarTulni arian spiralis gareT. hidrofobuli
aminomJavebis radikalebi hidrofobuli urTerTobis gamo erTmaneTisken
miiswrafian, ris Sedegadac cilovan molekulas aniWeben sferosebr formas,
romlis polaruli nawilebi mimarTulia gareT, xolo arapolaruli _ SigniT.
aseTi tipis konformaciis formirebaSi, hidrofobuli Zalebis garda,
monawileobas iReben disulfiduri -S-S- kavSirebi, romlebsac warmoqmnian
aminmJava cisteinis sulfiduri (SH) jgufebi. aRweril konformacias mesameuli
struqtura ewodeba.

cila-mioglobinis mesameuli struqtura

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 mesameuli struqturis mixedviT ganasxvaveben globilarul da fibrilarul
cilebs. globularul cilebs sferosebri konformacia aqvT, xolo
fibrilarulebs _ wagrZelebuli, boWkosebri. fibrilaruli cilebiT mdidaria
SemaerTebeli qsovilebi. aseT cilebs warmoadgenen myesebSi da ZvlebSi Semavali
cila – kolageni, drekadi SemaerTebeli qsovilis cila _ elastini, kanis, Tmis
da frCxilebis cila _ -keratini da sxva. fibrilarul cilebs struqturuli
funqcia aqvT: maTi umravlesoba wyalSi uxsnadia.
fibrilaruli cilebisagan gansxvavebiT globularuli cilebi wyalSi kargad
ixsnebian; isini ujredSi dinamiur funqciebs moicaven _ fermentebis,
antisxeulebis, hormonebis umravlesoba globularul cilebs warmoadgenen.
ganasxvaveben cilebis natiur da denaturirebul mdgomareobas. natiuri
konformacia ganpirobebulia cilis mesameuli struqturiT. maRali temperaturis,
mJavebis, organuli gamxsnelebis, ultrabgeris da sxva Zlieri zemoqmedebis
Sedegad cilis mesameuli struqtura irRveva, globula iSleba uformo
polipeptidebad, ekargeba funqcia da gadadis denaturirebul mdgomareobaSi.
SedarebiT rbil pirobebSi denaturirebuli cila SeiZleba aRdges globulad,
anu moxdes misi renaturacia.

cilebis meoTxeuli struqtura

globularuli cilebis nawils meoTxeuli struqturac gaaCniaT. meoTxeuli
struqturis Camoyalibeba xdeba ori an ramdenime globularuli cilis
urTierTdakavSirebis Sedegad. aseT rTul cilovan kompleqss oligomeri
ewodeba, xolo masSi Semaval suberTeulebs ki – monomerebi. magaliTad, ferment
glutamatdehidrogenazas SenebaSi monawileobas Rebulobs 8 suberTeuli,
romelTa molekuluri wonaa – 280000.
SedarebiT kargad aris Seswavlili hemoglobinis meoTxeuli struqtura,
romelic oTxi suberTeulisagan Sedgeba. TiToeuli globula-suberTeuli
ikavSirebs rkina-porfirinis kompleqss, romelsac hemi ewodeba. hemi ierTebs
Jangbads, romlis Semdegac suberTeulebis urTierTdakavSireba Zlierdeba.
hemoglobinidan Jangbadis ganTavisuflebis Semdeg suberTeulebis
urTierTmoqmedeba sustdeba. analogiur konformaciul cvlilebebs ganicdian
fermentebi maTi substratebTan dakavSirebis Semdeg.
meoTxeuli struqturis formirebaSi da suberTeulebis dakavSirebaSi

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SedarebiT susti Zalebi arian CarTuli da amitom, rTuli oligomeruli cilis
daSla suberTeulebad advilad xdeba.

cilebis funqcia (zogadi mimoxilva)

cilebisaTvis damaxasiaTebelia funqciaTa didi sxvadasxvaoba. am

makromolekulebis yvelaze did klass fermentebi Seadgenen. dReisaTvis 3000-mde
fermentia aRwerili. fermentebs saxeobrivi specifikuroba axasiaTebT da
miuxedavad imisa, rom zogierTi fermenti erTi da igive reaqcias atarebs, maTi
aminomJavuri Semadgenloba sxvadasxva organizmebSi gansxvavdeba erTmaneTisagan.
(mag. Rorisa da xaris tripsini sxvadasxva cilebs warmoadgenen). fermentebis
umravlesobas globularuli struqtura aqvT; zogierTi fermenti erTi
suberTeulisagan Sedgeba, xolo zogierTis SemadgenlobaSi ki mravali monomeri
Sedis. TiToeul ferments aqvs aqtiuri centri – misi cilovani molekulis is
nawili, romliTac fermenti reaqcias atarebs. garda amisa, rTul oligomeruli
agebulebis fermentebs, gaaCniaT agreTve regulatoruli anu alosteruli
centric.
cilebis meore didi jgufs struqtuli cilebi warmoadgenen. maT mTavar
funqcias ujredSorisi da ujredSida sakomunikacio sistemebis formireba
Seadgens. umaRles organizmebSi erT-erTi aseTi cila kolagenia. mas
fibrilaruli konformacia gaaCnia, da am ciliT mdidaria Zvlovani da
SemaerTebeli qsovilebi. struqtuli cilebs warmoadgenen elastiuri
qsovilebis cila-elastini, kanisa da Tmis cila keratini. xrtilovan qsovilebSi
aRmoCenilia cila, romlis SemadgenlobaSi Sedis qondriotinsulfatis naSTi. es
glikoproteidi saxsrebis lorwovan sekretebs da sinovialur siTxes sapox
Tvisebebs aniWebs. struqturuli cilebiT mdidaria TviT ujredic.
cilebis erT-erT klass SekumSvadi cilebi Seadgenen. aseTebi arian, magaliTad
kunTis cilebi – aqtini da miozini. SekumSad cilebs wagrZelebuli forma aqvT.
zogierTi cila transportul funqciebs asrulebs. maT unari SeswevT
daikavSiron da sisxlis saSualebiT gadaitanon sxvadasxva tipis organuli da
araorganuli naerTebi. magaliTad, Sratis albumins gadaaqvs cximovani mJavebi,
steroidebi da sxva hidrofoburi nivTierebebi. transportuli funqcia akisria
aseve hemoglobins (gadaaqva Jangbadi)
cilebis did jgufs warmoadgenen hormonebi. hormonebis saSualebiT warmoebs
ujredSorisi urTierTobebis Camoyalibeba da ujredTa funqciuri regulacia.
cilebs warmoadgenen agreTve baqterialuri, mcenareuli da cxoveluri

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warmoSobis toqsinTa umravlesoba. magaliTad, qvewarmavlebis toqsinebi,
botulizmis gamomwvevi baqteriis (Clostridium botulinum) toqsini, difteriis toqsini,
abusalaTinis uZlieresi toqsini – ricini cilebia.
aRsanisnavia is rom, zemoTaRniSnuli cilebi, romlebsac aseTi gansxvavebuli
biologiuri aqtivoba aqvT, aSenebuli arian erTi da imave naerTebisagan –
aminomJavebisagan. am ukanasknelebs TviTon, arc biologiuri aqtivoba da arc
toqsikuroba gaaCniaT, da amdenad cilebis funqciuri sxvadasxvaoba mxolod maTi
specipikuri aminomJavuri TanmimdevrobiTaa ganpirobebuli.

antisxeulebi da imunuri reaqciebi (mokle mimoxilva)

cilebis saxeobrivi specifiuroba.

sxvadasxva cilebs Soris, romlebic adamianis Tu cxovelis organizmSi

sinTezirdeba, arseboben iseTi cilebic, romlebzec pirvelad iqna dadgenili
maTi saxeobrivi specifikuroba. am cilebs antisxeulebi anu imunoglobulinebi
ewodebaT. antisxeulebis sinTezi organizmSi maSin mimdinareobs, rodesac
sisxlSi ucxo, maRalmolekuluri naerT (antigeni) moxvdeba, da romelic
aRniSnuli saxeobisaTvis ar aris damaxasiaTebeli. am reaqcias imunuri reaqcia
ewodeba da masSi antisxeulebis warmomqmneli ujredebis B-limfocitebis garda
monawileobas iReben imunuri sistemis sxva ujredebic (T –limfocitebi,
makrofagebi da sxva). imunuri reaqciis Sedegad warmoqmnili antisxeuli
uerTdeba antigens da uvnebelyofs mas. antisxeulebi, romlebic antigenis
moxvedris Sedegad warmoiqmnebian organizmSi, sakmaod didi drois ganmavlobaSi
cirkulireben, xolo organizms am antigenebis SemoWra sisxlSi mTeli
sicocxlis ganmavlobaSi axsovs, mas ama Tu im antigenis mimarT imuniteti
uCndeba. es ukanaskneli SeiZleba xelovnuradac iyos gamowveuli, Tu organizmSi
mcire raodenobiT SeviyvanT araaqtiur denaturirebul antigens. wamyvani
mniSvneloba antisxeulebis gamomuSavebaSi eniWeba antigenebis pirvelad
struqturas, Tumca garkveuli roli SesaZlebelia mesameulma struqturamac
Seasrulos. imunuri reaqciis gamosawvevad sakmarisia ucxo antigenis 5-dan 10-mde
aminomJavis Tanmimdevroba, romelsac am antigenis epitopi ewodeba, miuxedavad
aminomJavaTa Tanmimdevrobis ganmsasazRvreli rolisa, dabalmolekuluri
peptidebis mier gamowveuli imunuri reaqcia naklebefeqturia.
antisxeulis antigenTan urTierTmoqmedebis Sedegad icvleba antigenis
konformacia da igi dezaqtivirdeba. antigen-antisxeulis kompleqsi wyalSi
cudad ixsneba da ileqeba. am reaqcias precipitaciis reaqcia ewodeba da igi

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farTod gamoiyeneba sisxlis imunologiuri statusis dasadgenad.

cilebis posttranslaciuri modifikaciebi

dReisaTvis garkveulia, rom cilebis umravlesoba aminomJavebis naSTebis garda

Seicavs arapeptiduri bunebis naerTebsac, romlebic cilovan molekulaSi arian
CarTulni. aqve unda ganvasxvavoT is cilebi, romlebic arakovalenturi
(hidrofoburi, eleqtrostatikuri da a.S.) bmiT ierTeben ama Tu im qimiur jgufs
(mag, fermentebi – metalebs, Sratis cilebi – lipidebs da sxva) im cilebisagan,
romelTa qimiuri modificireba kovalenturi bmiT warmoebs. vinaidan aseTi
kovalenturi bmis warmoqna qimiur jgufsa da cilis aminomJavas Soris cilis
ribosomebze sinTezis Semdeg xdeba, am process posttranslaciuri kovalenturi
modifikacia ewodeba.
erT-erT mniSvnelovan da gavrcelebul posstranslaciur modifikacias
cilebis fosforilireba warmoadgens. cilebis fosforilirebas, iseve rogorc
sxva tipis kovalentur modifikaciebs, fermentebi awarmoeben. dReisaTvis
aRmoCenilia daaxloebiT 1500-mde fosforilirebuli cila – fosfoproteini.
cilebis fosforilireba da Semdgom maTi defosforilireba (cilis
molekulidan fosfatis jgufis Camocileba), cvlis cilebis mesameul
struqturas, mis konformacias; da Tu es cila ferments warmoadgens – cvlis mis
katalizur aqtivobasac.
cilebis fosforilirebas didi mniSvneloba eniWeba ujredSi mimdinare
bioqimiuri procesebis regulaciaSi. fermentebi, romlebic ganicdian
fosforilirebas metabolizmis sakvanZo reaqciebSi iReben monawileobas.
hormonebis, neirogadamcemebis da sxva fiziologiurad aqtiuri Tu
farmakologiuri naerTebis moqmedeba, realizdeba ujredSi cilebis
fosforilirebis saSualebiT.
posttranslaciur kovalentur modifikacias warmoadgens agreTve cilebis
glikozilireba, romlis Sedegadac miiReba glikoproteinebi. glikozilirebis
reaqciis Sedegad cilis molekulas ukavSirdeba martivi naxSirwyali (glukoza,
galaqtoza an sxva), romelsac SemdgomSi SesaZlebelia daematos sxva
naxSirwylebic. amis gamo cilis molekulaze warmoiqmneba sakmaod grZeli
datotvili naxSirwylovani jaWvi, romelic did gavlenas axdens cilis
konformaciaze, mis fiziko-qimiur Tvisebebze. glikozilirebuli cilebis
umravlesoba ujredis plazmur membranazea lokalizebuli da monawileobas
iRebs ujredSoris kontaqtebis CamoyalibebaSi, ujredis mier hormonTa da

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neirogadamcemTa recepciaSi, antigenis blokirebaSi da sxva. ujredis garsis
zedapiruli aqtivoba mTlianad damokidebulia membranul cilebSi arsebul
glikozilis naSTebze.
mniSvnelovan posttranslaciur modifikacias warmoadgens cilebis ADP-
ribozilireba. ADP-riboza ujredis erT-erTi ZiriTadi kofermentis NAD-is
daSlis Sedegad warmoiqmneba da misi CarTva cilovan molekulaSi specialuri
fermentebis saSualebiT xdeba. sainteresoa aRiniSnos is, rom am ferments
Seicavs zogierTi mikroorganizmis toqsinic, kerZod, qoleris da difteriis
toqsinebi, romelTa moqmedebac mimarTulia cilebis ADP-ribozilirebaze.
endogenuri modifikacia mniSvnelovan rols TamaSobs dazianebuli dnm-is
aRdgebis, misi reparaciis procesSi.
ujredis gamravlebis dros warmoebs birTvuli cilebis acetilireba. erT-erT
gavrcelebul modifikacias warmoadgens amidireba, romlis drosac xdeba
glutaminisa da asparaginis mJavebis karboqsilSi NH3-amiduri jgufis Casma.
iSviaTad, magram gvxvdebian iseTi kovalenturi modifikaciebi, rogoricaa
miristilireba (cximovani mJava – miristinis mJavis cilasTan dakavSireba) da
inozitilireba (aminospirti inozitolis cilastan dakavSireba). zogierTi cila
ganicdis meTilirebas.

Ffermentebi, MmaTi funqcia da daxasiaTeba

fermentebi warmoadgenen cilebis yvelaze upro mravalricxovan da

maRalspecializebul klass. swored isini Seadgenen im samuSao aparats, romlis
saSualebiTac xdeba genTa moqmedebis realizacia. Seizleba iTqvas, rom bioqimiis
istoria fermentebis aRmoCenisa da maTi Seswavlis istorias warmoadgens.
Cveulebriv gamoyofen fermentebis moqmedebis or ZiridaT sakiTxs: fermentebis
specifikurobas da reaqciis aCqarebas. dReisaTvis orive problema gadauWrelia.
Tanamedrove enzimologia ( mecniereba, romelic fermentebis moqmedebis meqanizms

Seiswavlis) ver xsnis im kolosalur (109-1010) aCqarebebs, romlebsac fermentebi

qimiur reaqciebs aniWeben. Tundac miaxloebuli Teoriuli gaTvlebi, Tanamedrove

kvanturi meqanikisa da qimiuri kinetikis gamoyenebiT, ver wamosaxaven im
siCqareebs, riTac fermentuli reaqciebi warmoebs.

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 FfermentTa klasifikacia

fermentTa saxelwodeba warmoiqmneba Sesabamis substratze (substrati _

qimiuri naerT, romelic warmoiqmneba fermentul reaqciaSi) sufiqsi aza-s
damatebiT. magaliTad, saxelwodeba ureaza metyvelebs imaze, rom fermenti
awarmoebs Sardovanas (urea) daSlas, an citratsintaza – metyvelebs imaze, rom
fermenti axorcielebs limonmJavis (citrate) sinTezs. fermentebis
sistematizirebisaTvis msoflio saerTaSoriso komisiam SeimuSava fermentTa
nomenklatura, klasifikaciis struqtura da TiToeul ferments Sifri mianiWa.
fermentebi dayofilia 6 ZiriTad jgufad:
1. oqsidoreduqtazebi. 2. transferazebi. 3. hidrolazebi 4. liazebi 5.
izomerazebi 6. ligazebi.
oqsidoreduqtazebis jgufs warmoadgenen is fermentebi, romlebic ujredSi
awarmoeben jangva-aRdgeniT reaqciebs. transferazebi asruleben ama Tu im
funqciuri qimiuri jgufis gadatanas erTi naerTidan meoreze. hidrolazebi
axorcieleben naerTTa hidrolizs. liazebi warmoadgenen im fermentebs,
romlebic awarmoeben naerTTa mierTebas ormagi bmis adgilze. izomerazebi -
axorcieleben qimiur naerTTa izomerizacias, da bolos, ligazebi - arian
fermentebi, romlebic atareben reaqciebs AATP-is energiis xarjze.

fermentTa kofaqtorebi

fermentTa umravlesoba damoukideblad atarebs bioqimiur reaqciebs, xolo

garkveul nawils katalizis Casatareblad esaWiroeba dabalmolekuluri
nivTierebebis Tanapovniereba. aseT nivTierebebs kofaqtorebi ewodeba da isini
fermentTan uSualo kavSirSi imyofebian. arseboben kofaqtorebi, romlebic
mWidrod arian dakavSirebuli fermentis cilovan nawilTan. xSir SemTxvevebSi
es kofaqtorebi metalebs warmoadgenen (TuTia, rkina, spilenZi, manganumi da
sxv.) isini fermentul reaqciebSi Rebuloben monawileobas.
kofaqtorebTan SedarebiT ufro rTul naerTebs kofermentebi warmoadgenen.
SesaZlebelia kofermentebi fermentis molekulis SenebaSi iRebdnen

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monawileobas an fermentul reaqciebSi moqmedebdnen damoukideblad. vitaminebis
umravlesoba warmoadgens kofermentebs.
fermentuli katalizi

warmovidginoT qimiuri reaqcia, romlis drosac gardasaqmneli nivTiereba –

substrati (S) gardaiqmneba P-produqtad. qimiuri reaqcia S P mimdinareobs
imitom, rom drois yovel garkveul momentSi arsebobs S substratis
molekulebi, romlebsac SedarebiT ufro didi energia aqvT, xolo es energia
sakmarisia imisTvis, raTa daSalos an warmoqmnas qimiuri bma da gardaiqmnas P-
T. aseTi tipis S molekulebs aqtivirebul mdgomareobaSi myofi molekulebi
ewodeba. zogadad, aRwerili reaqcia SeiZleba ase gamoisaxos: S S* P. sadac S*
aris sawyisi naerTis aqtivirebuli mdgomareoba. imisTvis, rom miRweul iqnas
aqtivirebuli (gardamavali) mdgomareoba, aucilebelia S-s mieniWos garkveuli
energia. energias, romelic saWiroa sawyisi nivTierebis aqtivirebul
mdgomareobaSi gadasayvanad aqtivaciis energia ewodeba. aqtivaciis energia
Tavisufali energiis erTeulebSi gamoisaxeba _ kaloria/mol. katalizatoris
moqmedebis principi mdgomareobs swored imaSi, rom igi amcirebs aqtivaciis
energias da amiT aCqarebs qimiur reaqcias. aqtivaciis energiis SemcirebiT
aqtivirebul mdgomareobaSi myof naerTTa koncentracia izrdeba, ris gamoc
izrdeba agreTve miRebul produqtTa raodenoba da Sesabamisad _ siCqare.
fermentuli reaqciis siCqare damokidebulia moreagire nivTierebaTa
koncentraciaze. substratis koncentraciis zrdiT fermentuli reaqciis
siCqarec izrdeba. Cveulebrivi qimiuri reaqciisgan gansxvavebiT, fermentul
reaqciaSi substratis garkveuli koncentraciisas misi raodenobrivi zrda aRar
iwvevs reaqciis siCqaris matebas.

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 v

Vmax

Km [S]

am efeqts fermentis substratiT gajereba ewodeba. Tu fermenti gajerebulia

substratiT, maSin mas maqsimaluri siCqare (Vmaqs.) aqvs. es ukanaskneli
fermentebis erT-erTi damaxasiaTebeli sididea. fermentTa meore ZiriTad
maCvenebels e.w. miqaelisis konstanta (Km) warmoadgens. Km _ aris substratis
is koncentracia, romlis drosac reaqciis siCqare 1/2 Vmaqs.-is tolia. rac ufro
dabalia fermentisTvis Km, substratis miT ufro naklebi koncentraciaa saWiro
maqsimaluri siCqaris misaRwevad. dabali Km mZlavr fermentebs axasiaTebT.
fermentuli reaqcia SesaZlebelia gamovsaxoT:

S+ E ES E+P

sadac S aris substrati, P – produqti, xolo E –fermenti (enzyme-ingl)

ES warmoadgens ferment-substratul kompleqss. substrati ferments aqtiur
centrSi ukavSirdeba. fermentis aqtiuri centri rTul geometriul warmonaqmns
warmoadgens, romlis sivrciTi organizacia komplementarulia substratis
konfiguraciisa. sakuTriv qimiuri reaqcia fermentis aqtiur centrSi warmoebs.
substratis aqtiur centrTan mierTebis Semdeg warmoiqmneba ferment-
substratuli kompleqsi, romelic S-is aqtivirebul mdgomareobas warmoadgens..
fermentuli reaqcia damokidebulia sareaqcio aris pH-ze. yovel ferments
Tavisi damaxasiaTebeli pH-optimumi gaaCnia e.i. pH-is is mniSvneloba, romlis
drosac fermentuli siCqare maqsimaluria. magaliTad pepsins _ ferments,
romelic cilebis hidrolizs anxorcielebs da romelic kuWSi gamoiyofa _
mJave pH optimumi axasiaTebs (pH-1,5), xolo tripsins, romelic msgavs reaqcias

24
 atarebs, oRond TormetgojaSi gamoiyofa _ pH optimumi tute areSi aqvs (pH-7,7).
fermentuli aqtivoba sareaqcio areSi sxvadasxva fiziko-qimiuri meTodebis
gamoyenebiT isazRvreba (substratis an produqtis raodenobrivi cvlilebebis
mixedviT). fermentis ZiriTad maxasiaTebels erTeuli warmoadgens (unit-ingl.).
fermentuli aqtivobis erT erTeulad miCneulia fermentis is raodenoba,
romelic 1mkmoli substrats gardaqmnis 250-ze 1 wuTSi fermentuli reaqciis
optimalur pirobebSi. Tu es sidide gadaTvlilia 1 mg cilis raodenobaze,
maSin mas xvedriTi aqtivoba ewodeba.
fermentul reaqciaTa umravlesoba iTrguneba, inhibirdeba sxvadasxva tipis
SenaerTebiT. ganasxvaveben Seqcevad da Seuqcevad inhibitorebs. Seqcevadi
inhibitorebi, Tavis mxvriv, iyofian konkurentul da arakonkurentul
inhibitorebad. konkurentul inhibitors uwodeben naerTs, romelic substratis
nacvlad fermentis aqtiur centrs ukavSirdeba. vinaidan fermenti ver
gardaqmnis mas, igi inhibirdeba. Tu gavzrdiT substratis koncentracias, igi
gamoaZevebs inhibitors aqtiuri centridan da TviTon daukavSirdeba mas. am
SemTxvevaSi adgili aqvs substratisa da inhibitoris konkurencias fermentis
aqtiuri centris dasakaveblad, da amitom, am ukanasknels konkurentuli
inhibitori ewodeba. konkurentuli inhibitorisagan gansxvavebiT
arakonkurentuli inhibitori substratiT ar gamoZevdeba da igi ferments
aqtiur centrSi ar ukavSirdeba. rogorc konkurentuli, aseve arakonkurentuli
inhibitorebis moqmedeba fermentze Seqcevadia da igi garkveuli droisa da
zemoqmedebis Semdeg ubrundeba katalizur, aqtiur mdgomareobas. am
inhibitorebisagan gansxvavebiT Seuqcevadi inhibitorebis moqmedeba sxva xasiaTs
atarebs, da fermentis maTTan dakavSireba aqtivobis mTlian daqveiTebas iwvevs.
Seuqcevadi inhibitorebis tipiur magaliTs e.w. fosfoorganuli nivTierebebi
warmoadgenen. isini mravali fermentis aqtiur centrSi arsebul aminomJava
serins kovalenturi bmiT ukavSirdebian da Trgunaven mis funqcionirebas.
fosfoorganul naerTebs Zlieri neiroparalizuri moqmedeba axasiaTebT.
toqsikuri nivTierebebis umravlesoba fermentTa Seuqcevad inhibitorebs
warmoadgens, xolo mravali farmakologiuri nivTiereba _ Seqcevads.

fermentebis moqmedebis meqanizmi, struqtura da regulacia

rogorc ukve iTqva, fermenti substrats aqtiur centrSi ierTebs. fermentis

aqtiuri centrisTvis damaxasiaTebelia maRali specifikuroba, igi Tavis

25
substrats ganasxvavebs im uamravi metabolitisagan, romelic ujredSi mis
irgvliv imyofeba. fermentis substratuli sfecifikuroba mis erT-erT ZiriTad
maxasiaTebels warmoadgens. fermentis aqtiuri centris rTul sivrciT
konfiguracias ganapirobebs mis polipeptidur jaWvSi Semavali sxvadasxva
aminomJavebi. aqtiuri centris SenebaSi SesaZlebelia monawileoba miiRon im
aminomJavebma, romlebic jaWvis dasawyisSi, SuaSi da boloSi mdebareoben.
erTmaneTisagan daSorebuli aminomJavaTa monawileoba aqtiuri centris
formirebaSi ganpirobebulia fermentis globularuli struqturiT.
ganvixiloT ferment acetilqolinesTerazis moqmedebis meqanizmi. aRniSnuli
fermenti awarmoebs acetilqolinis hidrolizs. igi substratis molekulas
qolinis naSTis mixedviT cnobs da ikavSirebs mas.

fermentTan dakavSirebuli ubani gasaxleCi bma

CH3
+
CH3 N CH 2CH2O C CH3
CH3 O

Please Register

acetilqolinesTeraza

fermentTan mierTebis Semdeg, acetilqolinis molekula iseT orientacias

ikavebs, rom SesaZlebeli xdeba fermentis aqtiur centrSi arsebuli serinis
daaxloeba acetilqoloinis karbonilis jgufTan. acetilqolinesTerazas serini
substratis molekulaSi eleqtronTa gadanawilebas iwvevs, rasac kovalenturi
bmis darRveva mosdevs. amis Semdeg, qolini da ZmarmJava (reaqciis produqtebi)
ferments cal-calke scildebian. Seuqcevadi inhibitorebi, kerZod,
fosfoorganuli neiroparalitikuri gazebi, acetilqolinesTerazis swored am
serinis naSTs ukavSirdebian.
intaqtur ujredSi mravali fermenti paralelurad muSaobs, da xSir
SemTxvevebSi, erTi fermentis produqti meore fermentis substrats warmoadgens.
sirTulis mixedviT, mravalfermentiani anu multifermentuli sistemebi SeiZleba

26
sam tipad davyoT. pirveli, da SedarebiT ufro martivi fermentuli sistemebi
citoplazmaSi arian warmodgenilni da Tavisuflad moZraoben ujredis wvenSi.
fermentuli sistemebis organizaciis ufro maRal dones fermentuli kompleqsebi
warmoadgenen. magaliTad, safuarebSi cximovani mJavebis sinTezs awarmoebs
kompleqsi, romelic 7 sxvadasxva fermentisagan Sedgeba. aseTi kompleqsebis
warmoqmna biologiurad gamarTlebulia, radgan maTi funqcionirebisas, dro
substratisa da produqtis difuziaze ar ixarjeba da erTi reaqciis Sedegad
warmoqmnili produqti iqve moixmareba meore fermentis mier.
multifermentuli sistemebis organizaciis umaRles dones qmnian. fermentTa
sistemebi, romlebic organoidebis membranebTan arian dakavSirebuli. amis
magaliTs mitoqondriuli JangviTi-fosforilirebis jaWvi warmoadgens.
multifermentuli sistemebis umravlesobas TviTregulaciis unari aqvT, rac
mdgomareobs imaSi, rom sistemis saboloo produqti pirveli fermentuli
reaqciis inhibirebas iwvevs. rogorc ki, saboloo produqtis koncerntracia
garkveul zRvrul dones miuaxlovdeba, igi iwyebs pirveli fermentis inhibirebas,
riTac gardaqmnaTa mTel jaWvs wyvets. rodesac AreaqciaTa jaWvSi:

A B C D E F

F-saboloo produqtis koncentracia Semcirdeba misi sxva reaqciebSi

moxmarebis gamo, inhibireba ixsneba da multifermentuli sistema kvlav
ganagrZobs funqcionirebas. aseTi tipis TviTregulacias, romlis drosac
xdeba sawyisi etapis saboloo produqtiT inhibireba, uku- anu
retroinhibireba ewodeba. am tipis reaqciebSi, umTavresad inhibirdeba jaWvis
pirveli fermenti. maT rTuli agebuleba aqvT da regulatoruli anu
alosteruli fermentebi ewodebaT, naerTebs ki, romlebic am fermentebze
moqmedeben - efeqtori an modulatori. regulatoruli fermentebis
umravlesoba erTi modulatoriT icvlis aqtivobas, xolo is fermentebi,
romlebic metaboluri gzebis gadakveTaze imyofebian, xSirad ramdenime
modulatoriT inhibirdebian. garda mainhibirebeli e.w. uaryofiTi
modulatorebisa, arseboben dadebiTi gamaaqtivebeli modulatorebic.
regulatorul fermentebs rTuli kinetika axasiaTebT. xSir SemTxvevebSi
isini suberTeulebisagan Sedgebian da maT meoTxeuli struqtura aqvT.
regulatorul fermentebs suberTeulTa kooperatiuloba axasiaTebT.
suberTeulTa kooperatiuli urTierTdamokidebulebis gamo, ferments

27
substratis pirveli molekulis mierTebis Semdeg uadvildeba substratis
momdevno porciebis mierTeba. es ganpirobebulia imiT, rom pirveli
katalituri aqtis Sedegad regulatoruli fermentis suberTeuli icvlis
konformacias, gadadis aqtivirebul mdgomareobaSi da gadahyavs sxva
suberTeulebic aseTive mdgomareobaSi.
regulaciis erT-erT tips fermentis molekuluri organizaciis
cvlilebebi warmoadgens. fermentebs, romlebsac erTi da igive aqtivoba aqvT,
magram gansxvavdebian erTmaneTisagan pirveladi, meoreuli an mesameuli
struqturiT izofermentebi ewodebaT. izofermentebi SeiZleba arsebobdnen
erT organizmSi, qsovilSi an ujredSi. magaliTad, ferment
laqtatdehidrogenazas gaaCnia 5 izofermenti, romlebsac erTi da igive
molekuluri wona, suberTeulTa raodenoba da katalizuri aqtivoba aqvT.
gansxvavdebian isini mxolod fermentSi Semavali suberTeulTa tipebiT.
laqtatdehidrogenaza oTxi suberTeulisagan Sedgeba: 2 M da 2 H
suberTeulebisagan. fermentis 5 izoformis arseboba ganpiribebulia
suberTeulebis dakavSirebis sxvadasxva variantebiT: 3H da 1M, 2H da 2M, 4H da
a.S. ConCxis kunTis laqtatdehidrogenaza 4M suberTeulisagan Sedgeba, xolo
gulisa ki - 4H. Tu Cven sinjaraSi am suberTeulebs SevurevT erTmaneTs, isini
yvela SesaZlo variantebiT aewyobian da 5 izoformas mogvcemen.
laqtatdehidrogenazebis izofermentebi kargad iyofian eleqtroforeziT, da
amdenad maT gansazRvras didi mniSvneloba aqvs gulis kunTis paTologiebis
diagnozisTvis.

lipidebi

lipidebi wyalSi uxsnad organul nivTierebebs warmoadgenen. isini didi

raodenobiT arian cocxal ujredSi da eqstragirdebian arapolaruli
organuli gamxsnelebiT. arsebobs lipidTa ramdenime klasi, romlebic
erTmaneTisagan cximovani mJavis agebulebiT gansxvavdebian.
lipidebs ujredSi sami ZiriTadi funqcia aqvT: 1. isini membranebis
struqturul komponentebs warmoadgenen, 2. maT sakvebi maragis funqcia aqvT,
da monawileobas Rebuloben mis gadatana-gadanawilebaSi, da bolos, 3. isini
damcvelobiTi funqciis matarebelni arian. lipidebis SemadgenlobaSi xSirad
gvxvdebian cximovani mJavebi.

cximovani mJavebi

28
 cximovani mJaveebi neitraluri cximebis, fosfigliceridebis,
glikolipidebis da qolesterinis eTerebis SemadgenlobaSi Sedian.
dReisaTvis cnobilia cximovani mJavis 70-mde tipi. maTi molekula grZeli
naxSirwyalbaduri jaWvisagan Sedgeba, romelic mTavrdeba karboqsilis
jgufiT.
CH3-CH2-CH2-CH2-CH2-CH2-CH2-CH2-CH2-CH2-CH2-CH2-CH2-CH2-CH2-COOH
CH3-(CH2)14-COOH palmitinis mJava.
CH3-CH2-CH2-CH2-CH2-CH2-CH2-CH2-CH=CH-CH2-CH2-CH2-CH2-CH2-CH2-CH2-COOH.
CH3-(CH2)7-CH=CH-(CH2)7-COOH oleinis mJava.
umaRles mcenareebSi da cxovelebSi Semavali cximovani mJavebi rigi
saerTo TvisebebiT xasiaTdebian. TiTqmis yoveli maTgani wyvil naxSirbadis
atoms Seicavs, 14-dan 22-mde. naxSirwyalbadovani jaWvi SeiZleba iyos, rogorc
najeri, aseve ujeri. cximovani mJavebi, romlebsac SedarebiT grZeli
naxSirwyalbadovani jaWvi aqvT (16-18 naxSirbadis atomi), wyalSi praqtikulad
uxsnadi arian. wyalSi micelebis warmoqmniT mxolod maTi natriumis an
kaliumis marilebi ixsnebian. amitom maT sapnis Tvisebebi aqvT.
gavrcelebul najer cximovan mJaveebs warmoadgenen miristinis, palmitinis
da stearinis mJavebi, ujers ki - oleinis, linolis da linolinis mJavebi. ori
ukanasknelis sinTezi cxovelur organizmebSi ar warmoebs.

neitraluri cximebi (acilglicerinebi)

cximovani mJaveebis glicerinis eTerebs acilglicerinebi, neitraluri

cximebi anu gliceridebi ewodebaT:

CH 2OH CH 2OR1 cximovani mJavebis

CHOH CHOR2 naxSirwyalbadovani
PleaseCH 2OH
Register PleaseCH 2OR3
Register naSTebi
glicerini acilglicerini

29
 im SemTxvevaSi, Tu eTerificirebulia glicerinis samive hidroqsilis
jgufi, maSin am naerTs triacilglicerini anu trigliceridi ewodeba, xolo
Tu cximovani mJaveebi dakavSirebulia mxolod or an erT hidroqsilTan -
Sesabamisad di- an monogliceridi ewodeba.
acilglicerinebi Tavisuflad ixsnebian qloroformSi, meTanolSi da sxva
organul gamxsnelebSi. am naerTebis mJavebSi an tuteebSi duRilisas maTi
hidrolizi glicerinamde da Tavisufal cximovan mJavamde xdeba.

fosfogliceridebi

am klasis lipidebi umTavresad membranebis SenebaSi iReben monawileobas.

acilglicerinebisagan gansxvavebiT, fosfogliceridebSi erT-erTi pirveladi
hidroqsilis jgufi eTerificirebulia ara cximovani mJaviT, aramed
fosforis mJaviT.

CH 2OR 1
CHOR2 OH
CH2 O P O
OH
fosfogliceridi (fosfatidis mJava)

fosforis mJaviT glicerinis molekulaSi SeiZleba eTerificirebuli iyos,

rogorc 1, aseve me-2 da me-3 naxSirbadis atomebi.
fosfolipidebi polarul naerTebs warmoadgenen; maT gaaCniaT uaryofiTad
damuxtuli fosformJavas naSTi. fosfogliceridebis SemadgenlobaSi
SesaZlebelia Sediodes kidev erTi, spirtuli komponenti, romelic
fosformJavas naSTs ukavSirdeba. aseTi tipis fosformJavas naerTebs
fosfodieTerebi ewodeba.

30
 O
CH2 OCR1
CH2 O C R 1
O H OCR2
CH2 O C R 2 OH H2 H2
O CH2O P O C C NH2
CH2 O P O X
-
OH O F O
fosfatidileTanolamini

X-komponentad, SeiZleba sxvadasxva tipis spirti iyos warmodgenili.

xSirad gvxvdebian eTanolamini, qolini, serini da inoziti.

O
OH OH
OH
H2N CH C OH CH N+(CH3)3 CH NH2
OH
CH2 CH2 CH2
OH

OH OH OH OH
serini qolini eTanolamini i n o z i t i

Tu fosfatidis mJavasTan SeerTebulia eTanolamini, maSin mas

fosfatidileTanolamini ewodeba. Sesabamisad miiRebian fosfatidilqolini,
fosfatidilserini da fosfatidilinozitoli. rogorc ukve aRiniSna, es
naerTebi monawileobas Rebuloben membranebis SenebaSi.
wyalSi gaxsnisas fosfogliceridebi warmoqmnian micelebs; isini
polaruli nawiliT gareT, xolo hidrofobuliT - SigniT ganlagdebian.
neitralur pH-ze fosfogliceridebis fosfati yovelTvis uaryofiTad aris
damuxtuli. fosfogliceridebis hidrolizisas ori cximovani mJava da
spirtuli X-komponenti gamoiyofa.
fosfogliceridebi agreTve specialuri fermentebiT - fosfolipazebiT
hidrolizdebian. fosfolipazebis 4 tipi arsebobs: A, B, C da D. isini
fosfogliceridebis sxvadasxva eTeruli bmis hidrolizs awarmoeben: O

31
 O
CH2 O C R 1 fosfolipaza B
O fosfolipaza A
CH2 O C R 2
fosfolipaza C
O fosfolipaza D
CH2 O P O X
-
O

fosfolipazebi A da C hormonTa membranul receptorebTan arian

SeuRlebulni. maTi moqmedebis Sedegad ujredSi Tavisufldebian araqidonis
mJava (fosfolipaza A-s SemTxvevaSi Tu R-araqidonis mJavaa),
diacilglicerini da fosfoinozitoli (fosfolipaza C-s SemTxvevaSi Tu X-
inozitia). samive naerTi ujredSida regulatorebs warmoadgenen da hormonis
moqmedebaze sapasuxod ujredis metabolizms cvlian.

sfingolipidebi da glikolipidebi

sfingolipidebi da glikolipidebi lipidTa erT-erT gavrcelebul klass

warmoadgenen. mdidaria am naerTebiT nervuli qsovili, da kerZod tvini. maT
SemadgenlobaSi Sedian cximovani mJava da sxvadasxva tipis aminospirtebi.
glikolipidebis SenebaSi, SesaZlebelia, Saqrebmac miiRon monawileoba (mag.,
galaqtozam). glikolipidebis erT-erT mniSvnelovani, rTuli Semadgenlobis
mqone warmomadgenelia gangliozidi.

steroidebi

steroidebi warmoqmoadgenen perhidrociklopentanfenantrenuli birTvis

warmoebuls:

32
 ciklopentanperhidrofenantrenuli birTvi

bunebriv steroidebs naRvlis mJavebi, mamakacis da qalis sasqeso

hormonebi, Tirkmelzeda jirkvlis hormonebi, qolesterini da sxva adamianis
organizmSi gavrcelebuli naerTebi warmoadgenen. zogierT maTgans
fiziologiuri aqtivoba axasiaTebs, xolo zogierTi - Zlier toqsikuria.
isini organizmSi mcire raodenobiT moipovebian. mdidaria qsovilebi mxolod
sterinebiT, kerZod, qolesteriniT. es ukanaskneli organizmSi, rogorc
Tavisufali, ise eTerificirebuli formiT arsebobs.
qolesterini wyalSi uxsnadia da ujredebidan eqstragirdeba mxolod
organuli gamxsnelebiT. qolesteriniT mdidaria ujredis plazmuri
membranebic.

terpenebi

terpenebi cxovelTa organizmebSi SedarebiT mcire raodenobiT arian

warmodgenili. terpenebis ZiriTadi erTeuls izopreni Seadgens.

CH2 C CH CH3
CH 3 izopreni

izoprenis monomerebisagan Semdgari naerTebiT mdidaria samkurnalo da

eTerzeTovani mcenareebi. bunebrivi kauCukic izoprenis molekulebisagan
Sedgeba. izoprenis warmoebulebi arian agreTve cximSi xsnadi vitaminebi: A. E
da K. izoprenis molekulebi monawileoben steroidebis biosinTezSic, da
amdenad, isini vitamini D-s winamorbednic arian.

prostaglandinebi

33
 prostaglandinebi lipidTa did jgufs miekuTvnebian, da cnobili arian
eikozanoidebis saxeliT. prostaglandinebi warmoiqmnebian araqidonis
mJavisagan reaqciaTa mTeli kaskadis meSveobiT. reaqciaTa am Tanmimdevrobas
araqodonis mJavis kaskadi ewodeba. araqodonis mJavis kaskadis saSualebiT,
prostaglandinebis garda, warmoiqmnebian naerTebi, romlebsac maRali
biologiuri aqtivoba aqvT. maT ricxvs miekuTvnebian leikotrienebi,
prostaciklinebi da tromboqsanebi. isini monawileobas iReben mraval
fiziologiur procesSi, iwveven gluvi kunTebis SekumSvas Tu modunebas,
aregulireben sisxlis arteriul wnevas, Trgunaven da xels uwyoben
garkveuli tipis hormonebis moqmedebas da a.S.
araqidonis mJavis kaskaduri reaqciebis siCqare da prostaglandinebis
sinTezi anTebiTi procesebis dros izrdeba. maTi intensiuri metabolizmis
gamo membranebSi zeJanguri procesebis gaaqtiveba warmoebs, rac uaryofiTad
moqmedebs ujredis funqcionirebaze. anTebis sawinaaRmdego preparatebi -
aspirini da indometacini - araqidonis kaskadis erT-erTi fermentis,
prostaglandinsinTetazis Seqcevad inhibitorebs warmoadgenen.

lipiduri micelebi, lipoproteidebi da elementaruli membrana

polaruli lipidebi, iseve rogorc sapnebi wyalSi gaxsnisas, micelebs

warmoqmnian. micelebSi lipidebis naxSirwyalbadovani, hidrofobuli jaWvebi
SigniT arian ganlagebuli da wylis molekulebs ar exebian.
acilglicerinebi da qolesterini, romlebsac polaruli nawili ar gaaCniaT
Tavad ar warmoqmnian micelebs; magram isini SesaZlebelia micelis
SemadgenlobaSi Sevidnen sakmaod rTuli struqturis formirebiT. maTgan
gansxvavebiT, polaruli lipidebi micelebs advilad warmoqmnian. garda amisa,
am ukanasknelebs SeuZliaT bimolekuluri Sris Seqmnac, gansakuTrebiT maSin,
Tu isini wyalgamyof zedapirze imyofebian.

34
 hidrofiluri nawili

hidrofoburi nawili

lipidebis bimolekuluri Sre

iseve rogorc micelebSi, bimolekulur SreSic lipidTa hidrofoburi jaWvebi

erTmaneTisken arian mimarTulni, xolo polaruliT (fosfogliceridebis
SemTxvevaSi - fosformJavas naSTebi) - wylisaken. TvisebebiT bimolekuluri
Sre biologiuri membranebis msgavsia.
biologiuri membrana rTuli struqturuli warmonaqmnia, radgan masSi,
ormagi lipiduri Sris garda ganTavsebuli arian cilebic. isini lipidur
SresTan polarobis mixedviT arian dakavSirebulni. im SemTxvevaSi, Tu
cilebs hidrofobur aminomJavaTa Tanmimdevrobis didi nawili (domeni) aqvT,
isini mTlianad ganWolaven lipidur Sres da e.w. integralur membranul
cilebs warmoqmnian. Tu membranul cilebSi polaruli aminomJavebis
raodenoba didia, maSin isini mxolod membranis zedapirTan arian
dakavSirebulni. ZiriTad Zalas, romelic membranuli makromolekulebis
urTierTdamokidebulebas gansazRvravs, hidrofoburi urTierToba
warmoadgens.

lipidebis bimolekuluri Sre integraluri ciliT

membranul cilebs lateraluri mobiluroba (gverdiTi Zvradoba)

axasiaTebT da maT membranaSi gadaadgilebis unari aqvT. garda amisa,
membranuli cilebi garkveul pirobebSi translocirdebian (gadaadgildebian)
citoplazmaSi, da piruku, - citoplazmidan adsorbirdebian membranaze.
membranaSi gamudmebiT xdeba cilebis ganaxleba, Zveli cilebis daSla da
axlebis CarTva. es yvelaferi miuTiTebs imaze, rom biologiuri membrana
dinamiur, cvlad formirebas warmoadgens, romlis biologiuri da fizikur-
qimiuri Tvisebebi ujredis sxva nawilebTan wonasworobaSi imyofe1ba.

35
 biologiuri membranis sisqe 90 anqstremis tolia. igi advilad atarebs
wyals da neitralur hidrofobur (lipofilur) naerTebs, naklebad atarebs -
polarul nivTierebebs (Saqrebs, aminomJavebs da sxva), da saerTod ar atarebs
mcire zomis ionebs (Na+,K+, Cl- da sxva). amitom es ionebi sxvadasxva raodenobiT
grovdebian membranis orive mxares, ris gamoc potencialTa sxvaobas
warmoqmnian. membranul potencials didi mniSvneloba eniWeba agznebadi
qsovilebis ujredTa plazmuri membranebis (nervuli da kunTovani) da
zogierTi organoidis (mitoqondria, endoplazmuri bade) funqcionirebaSi.
polaruli lipidebi agreTve ukavSirdebian zogierT cilas da warmoqmnian
lipoproteidebs. aseTi tipis lipoproteidebiT mdidaria sisxlis plazma,
sadac isini transportul funqciebs asruleben. maTi mTavari daniSnulebaa
gadaitanon lipidebi organizmis erTi nawilidan meoreSi. plazmis
lipoproteidebSi lipidebi cilasTan hidrofoburi urTierTobiT arian
dakavSirebulni. polaruli lipidebis garda maT SemadgenlobaSi
SesaZlebelia Sediodnen cximovani mJavebi, qolesterini da sxva neitraluri
lipidebi. plazmis lipoproteidebi erTmaneTisagan lipiduri SemadgenlobiT,
da Sesabamisad, simkvriviT gansxvavdebian.

Saqrebi

Saqrebad anu naxSirwylebad iwodebian polioqsialdehidebi,

polioqsiketonebi da maTi warmoebulni. naxSirwylebis empiriuli formula
gamoixateba (CH2O)n, sadac n 3 da masze metia. naxSirwylebi SesaZlebelia
davyoT monosaqaridebad, oligosaqaridebad da polisaqaridebad.
monosaqaridebi anu martivi Saqrebi erTi polioqsialdehiduri an
polioqsiketonuri erTeulisagan Sedgebian. martivi Saqrebidan yvelaze ufro
gavrcelebuls eqvsnaxSirbadiani Saqari D-glukoza warmoadgens. am
ukanasknelidan warmoiqmnebian danarCeni Saqrebi. D-glukoza organizmis
ZiriTadi sawvavi nivTierebaa. oligosaqaridebi warmoadgenen iseT Saqrebs,
romelTa SemadgenlobaSic 2-dan 6-mde monosaqariduli erTeuli Sedis. isini
erTmaneTTan glikoziduri kavSiriT arian dakavSirebuli. rTuli Saqrebi anu
polisaqaridebi ewodebaT iseT Saqrebs, romelTa jaWvi didi raodenobis
monosaqariduli erTeulebisagan Sedgeba. polisaqariduli jaWvi SesaZlebelia
iyos rogorc sworxazovani, aseve datotvili. polisaqaridebi erTi an ori

36
tipis monosaqariduli erTeulisagan Sedgebian, da amitom, nukleinis mJavebisa
da cilebisagan gansxvavebiT, isini ar SeiZleba informaciul
makromolekulebs warmoadgendnen. magram datotvili jaWvis formirebis gamo,
polisaqaridebma SeiZleba miiRon gacilebiT ufro mravalferovani
konfiguracia, vidre cilebma an nukleinis mJaveebma. amitom olisaqaridTa
datotvili jaWvebi monawileobas iReben iseT informaciul procesebSi,
rogorebic arian ujredSorisi urTierTobani - kontaqtebis Camoyalibeba,
adhezia, urTierTSecnoba, imunuri reaqciebi da a.S.

monosaqaridebi

monosaqaridebis molekulaSi sami an meti naxSirbadis atomi Sedis. maT

aradajaWvul narSirbadovan jaWvSi naxSirbadis erTi atomi dakavSirebulia
karbonilis jangbadTan, xolo danarCeni - hidroqsilis jgufTan. Tu
karbonilis jgufi jaWvis boloSi mdebareobs, monosaqaridi warmoadgens
aldehids da mas aldoza ewodeba, xolo Tu es jgufi molekulis Sua
nawilSia - igi ketozas warmoadgens.

HC O CH2OH

H OH O

CH2OH CH2OH

gliceraldehidi dioqsiacetoni
(aldotrioza) (ketotrioza)

umartivesi, sami naxSirbadisagan Semdgari monosaqaridebi, triozebi

gliceraldehidi da dioqsiacetoni arian. gliceraldehidi aldotriozas,
xolo dioqsiacetoni - ketotriozas warmoadgens.
naxSirbadovani ConCxis gagrZelebisas miiRebian tetrozebi (4 naxSirbadiani
naxSirwylebi), pentozebi (5 naxSirbadiani), heqsozebi (6 naxSirbadiani),
heptozebi (7 naxSirbadiani) da a.S. TiToeuli maTgani Sedgeba aldo- da keto-
formebisagan. magaliTad, glukoza 6 naxSirbadiani aldoheqsozaa, xolo
fruqtoza - 6 naxSirbadiani ketoheqsoza. monosaqaridebis fiziko-qimiurma
analizma cxadyo, rom maTi umravlesoba Ria struqturis mqone naerTs ki ar
warmoadgens, aramed warmoqmnis daxurul birTvs, romelsac aldoheqsozebis

37
SemTxvevaSi piranozuli,xolo ketoheqsozebis SemTxvevaSi - furanozuli
struqtura axasiaTebT. am struqturebis gamosaxva xouersis proeqciuli
formulebis saSualebiT xdeba.
monosaqaridebs optikuri aqtoba aqvT, da amitom maT, SesaZlebelia, D- da
L-forma hqondeT. monosaqaridebis hidroqsilis jgufi SeiZleba acilirdes an
meTilirdes. garda amisa, SesaZlebelia karbonilis jgufis aRdgena
wyalbadiT, ris Sedegadac miiRebian Saqarspirtebi. magaliTad, glukozis
wyalbadiT aRdgenisas miiReba sorbiti, xolo manozisas - maniti.
Saqarspirtebi ujredSi sustad metabolizirdebian. maT tkbili gemo aqvT da
amitom isini diabetiT daavadebulTa racionSi ixmarebian. bunebrivi
Saqarspirtebidan aRsaniSnavia glicerini da inoziti. isini lipidebis
SemadgenlobaSi Sedian. garda amisa, inozitis warmoebulni ujredSida
regulatorTa funqcias asruleben.

38
 CHO CHO CHO

H OH HO H H OH

HO H HO H HO H

H OH H OH HO H

H OH H OH H OH

C H 2O H C H 2O H C H 2O H

D -g l u k o z a D -m a n o z a D -g a l a q t o z a

H
H
HO O
HO
HO H

H OH
H OH

D -g l u k o z a (f u r a n o z u l i f o r m a )

H OH

H O
HO
HO H
H OH

H OH

D - g l u k o z a ( -p i r a n o z u l i f o r m a )
H
H
OH OH H OH.

39
 CH2OH CHO

O CHO CHO H NH2

H OH H OH CH2 HO H

H OH H OH H OH HO H

H OH H OH H OH H OH

CH2OH CH2OH CH2OH CH2OH

H
D-fruqtoza OH D-riboza
dezoqsiriboza galaqtozamini

specifikuri fermentebis moqmedebis Sedegad aldozebis aldehidebis

jgufi SesaZlebelia daiJangos Sesabamisi karbonis mJaveebis warmoqmniT,
romlebsac Saqris mJavebi ewodeba. biologiuri funqciis mixedviT
mniSvnelovania uronis mJavebi, romlebic aldoheqsozebidan, kerZod
glukozidan da galaqtozidan warmoiqmnebian. magaliTad, glukozis daJangvis
Sedegad miRebuli glukuronis mJava, mravali polisaqaridis ZiriTad
komponents warmoadgens. Saqris mJavebs miekuTvneba agreTve askorbinis mJava-
vitamini C. mis struqturaSi ormagi bmis arsebobis gamo, igi aramdgrad
naerTs warmoadgens da advilad iJangeba.
ujredSi sakmaod gavrcelebulnia Saqrebi, romlebsac romelime
naxSirbadis atomTan erTi hidroqsilis jgufi akliaT. maT dezoqsiSaqrebi
ewodebaT. dezoqsiriboza dnm-is naxSirwylovan komponents warmoadgens. Tavis
tvinis metabolizmSi didi mniSvneloba eniWeba galaqtozis dezoqsiformas -
fukozas. igi monawileobas Rebulobs mexsierebis procesebSi.
Saqrebs, romelTa hidroqsilis jgufi Canacvlebulia aminojgufiT,
aminoSaqrebi ewodebaT. magaliTad, glukozis aminirebuli forma -
glukozamini, monawileobas iRebs mravali polisaqaridis SenebaSi, xolo
galaqtozis aminirebuli forma - galaqtozamini, xrtilis polisaqarid-
qondroitinsulfatis mTavar Semadgenel komponents warmoadgens.

disaqaridebi

disaqaridebidan yvelaze ufro gavrcelebuli maltoza, laqtoza da

saqaroza arian. maltoza saxameblidan warmoiqmneba ferment amilazis

40
moqmedebis Sedegad. igi glukozis ori molekulisgan Sedgeba. disaqaridi
laqtoza, romelic mxolod rZeSia aRmoCenili galaqtozisa da glukozisagan
warmoiqmneba, xolo lerwmis Saqris anu saqarozis SemadgenlobaSi,
glukozisa da fruqtozis naSTebi Sedian. es ukanaskneli, bunebaSi erT-erT
yvelaze ufro gavrcelebul Saqars warmoadgens.

polisaqaridebi

polisaqaridebis did jgufs e.w. samarago polisaqaridebi warmoqmnian,

romelTa Soris mniSvnelovani funqcia saxamebels (mcenareebSi) da glikogens
(cxovelebSi) aqvT. isini sakmaod didi granulebis saxiT grovdebian
citoplazmaSi, saidanac organizmis moTxovnilebisamebr Tavisufldebian
maTSi Semavali monosaqariduli formiT.
saxamebeli ori formiT arsebobs: -amilozisa da amilopeqtinis formiT.
isini erTmaneTisagan monosaqariduli jaWvis agebulebiT gansxvavdebian; -
amilozis jaWvi sworxazovania, xolo amilopeqtinis ki - datotvili.
saxamebeli adamianis saWmlis momnelebel sistemaSi ferment amilazas
saSualebiT gadamuSavdeba, romelic mas monosaqaridul erTeulebamde
(glukozamde) Slis. glikogeni umTavresad RviZlSi da kunTebSi grovdeba.
mas, iseve rogorc amilopeqtins datotvili jaWvi aqvs.
mcenareul samyaroSi farTod gavrcelebulia polisaqaridi - celuloza,
romelic struqturuli polisaqaridia. mis ganmeorebad erTeuls ori
glukozis naSTisagan Semdgari disaqaridi celobioza warmoadgens. am
polisaqaridis glikozidur bmis hidrolizs fermenti celulaza awarmoebs,
romelic ZuZumwovarTa saWmlis momnelebel sistemaSi ar moipoveba
(gamonakliss mcoxnavi cxovelebi warmoadgenen, romelTa kuWSi arseboben
celulazis gamomuSavebeli mikroorganizmebi).

Saqrebi da ujredis garsi

umaRles cxovelTa ujredis garsi (plazmuri membrana) rbili, dinamiuri

struqturuli warmonaqmnia. is bimolekuluri lipiduri Srisa da sxvadasxva
tipis cilebisagan Sedgeba, romlebic sakmaod rTul, datotvil saqaridul
jaWvs Seicaven. miuxedavad imisa, rom plazmur membranaSi arsebuli
biomolekulebis struqtura nawilobriv dadgenilia, bevri ram dReisaTvis
cnobili ar aris. kerZod, daudgenelia is, Tu rogor xdeba ujredebis

41
urTierTSecnoba da urTierTdakavSireba, ra ganapirobebs ujredebis
erTmaneTTan mimagrebas da maT populaciebad Camoyalibebas. ase magaliTad,
Tu gancalkevebul Tirkmlis ujredebs davTesavT sakveb areze, isini ecdebian
gaerTianebas da asociaciebis warmoqmnas. aseve, Tu erTmaneTSi avurevT
Tirkmlis da RviZlis disociirebul danawevrebul kulturebs, ujredis
TiToeuli tipi Seecdeba "Tavisianebis" moZebnas da maTTan populaciis
Camoyalibebas. garda amisa, normaluri ujredebi arasodes qaotur
dajgufebebs ar warmoqmnian, isini erT fenad lagdebian da wyveten
gamravlebas. ujredTa urTierTSecnobas da maT dakavSirebas ujredTa garsze
mdebare datotvili Tu dautotavi saqariduli jaWvebis mTeli speqtri
anxorcielebs, romlebic membranul cilebTan kovalenturad arian
dakavSirebuli, da am cilebis saSualebiT - ujredSida metabolizmTanac. Tu
es ukanaskneli ujredSi Seuqcevadad icvleba, maSin ujreds ekargeba
kontaqturi Tvisebebi, igi scildeba Tavis populacias da sisxlis nakadSi
Tavisuflad moZraobas iwyebs. da Tu aseT ujreds onkologiurad
gadagvarebuli ujredi warmoadgens, avTvisebiani procesebi vrceldeba mTel
organizmSi (metastazebi).
ujredTa garsSi warmodgenilni arian agreTve gansakuTrebuli tipis mJave
bunebis polisaqaridebi, romlebmac mJave mukopolisaqaridebis saxelwodeba
miiRes. maT blanti konsistencia aqvT da amitom isini ujredebis adheziaSi,
maT "cementirebaSi" iReben monawileobas. erT-erT aseT mukopolisaqarids
hialuronis mJava warmoadgens. igi TiTqmis yvela tipis ujredis garsis
SemadgenlobaSi Sedis da didi raodenobiT aris warmodgenili brolisebr
sxeulSi da sinovialur siTxeSi. meore tipis mukopolisaqarids qondroitoni
warmoadgens. am naerTis gogirdmJava eTerebi, qondroitinsulfatebi, Zvlebis,
xrtilis, Tvalis rqovana garsis da sxva SemaerTebeli qsovilebis
SemadgenlobaSi gvxvdebian. zogierTi qsovilebis (RviZli, filtvebi,
arteriebi) ujredSoris siTxeSi msgavsi struqturis kidev erTi
mukopolisaqaridi - heparinia warmodgenili.

42
 nukleotidebi da polinukleotidebi

nukleotidebis mTavar funqcias warmoadgens maTi monawileoba nukleinis

mJavebis SenebaSi. garda amisa, nukleotidebi ujredis metabolizmSi, mis
energetikaSi da regulaciaSi iReben monawileobas.
nukleotidebi Sedgebian azotovani fuZisagan, naxSirwylovani
komponentisagan (pentoza) da fosformJavas naSTisagan. azotovani fuZe
SesaZlebelia pirimidinisa da purinis warmoebulebiT iyos warmodgenili.
nukleotidebis SemadgenlobaSi ZiriTadad gvxvdebian pirimidinis sami fuZe:
uracili, Timini da citozini, da purinis ori fuZe: adenini da guanini.

NH2 O

N N
N NH

N N
H N H N NH2

adenini guanini

NH2 O O

N NH NH

N O N O N O
H H H

citozini Timini uracili

ujredSi pirimidinisa da purinis fuZeebs specialuri fermentebis

saSualebiT xuTnaxSirbadiani naxSirwyali riboza uerTdeba. igi ukavSirdeba
glikoziduri bmiT pirimidinis 1 da purinis me-9 azots. warmoqmnil naerTebs
nukleozidebi ewodebaT. nukleozidebis SemadgenlobaSi ribozis nacvlad
SeiZleba Sediodes dezoqsiriboza.

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 nukleozidebs aseve specialuri fermentebis saSualebiT ukavSirdeba
fosformJavas naSTi, ris Sedegadac warmoiqmneba nukleotidi. fosformJava
naxSirwylis molekulis erT-erT hidroqsils (2',3' an 5' mdgomareobaSi)
ukavSirdeba. nukleotidi SesaZlebelia Seicavdes erT, or an sam fosforis
mJavas naSTs, romlebic erTmaneTTan fosfodieTeruli bmiT arian
dakavSirebuli:

NH2

N
N

N
O O O N

-
O P O P O P O 5
O
4 H H 1
O- O- O-
3 2
H H
OH OH

adenozintrifosforis mJava

nukleotidebi ujredSi ZiriTadad nukleoziddifosfatebisa da

nukleozidtrifosfatebis saxiT arian warmodgenilni. am naerTebSi,
Sesabamisad ori da sami fosformJavas naSTi ribozis me-5 naxSirbadTan aris
dakavSirebuli. magaliTad, adenozin-5'-trifosfatSi(ATP) fuZe-purini
(adenini) dakavSirebulia ribozis 1 naxSirbadTan, xolo fosfatebi me-5
naxSirbadTan. nukleotidebis Semoklebuli dasaxelebaa:
ATP - adenozintrifosforis mJava
ADP - adenozindifosforis mJava
AMP - adenozinmonofosforis mJava
GTP - guanozintrifosforis mJava
CTPP- citidintrifosforis mJava
UTP - uridintrifosforis mJava
TTP - Timidintrifosforis mJava

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 nukleozid-5'-difosfatebis da nukleozid-5'-trifosfatebis fosformJavas
naSTebi advilad hidrolizdebian Sesabamis mono- da
difosfonukleotidebamde.magaliTad. nukleoziddi- da
nukleozidtrifosfatebis fosfodieTeruli bmis hidrolizis Sedegad
gamoiyofa didi energia (7 kkal/molze), romelic sxva fermentebis mier
moixmareba. amitom am bmas makroerguls uwodeben.
ATP ADP + araorganuli fosfati (Pi) + 7 kkal/moli
ADP A AMP + araorganuli fosfati (Pi) + 7 kkal/moli
ATP ujredis mTavar da universalur energetikul naerTs warmoadgens.
warmoadgenen ra energiis wyaros, misi fosfaturi jgufebi ujredis sxvadasxva
procesebSi iReben monawileobas. ATP SemakavSirebel rgols warmoadgens im
procesebs Soris, romlebic energiis warmoqmniTa da daxarjviT
mimdinareoben. ADP da AMP, romlebic ATP-s defosforilirebiT
warmoiqmnebian, specialuri sunTqvis sistemebis saSualebiT kvlav
fosforilirdebian. ATP-s saSualebiT ujredSi energiis ganawileba warmoebs.
zogierT reaqciebSi ATP-s magivrad sxva nukleozidtrifosfatebi
gamoiyenebian (mag. GTP). energetikuli funqciebis garda, nukleoziddi- da
trifosfatebs sxva daniSnulebac aqvT; kerZod, isini monawileoben sxvadasxva
naerTTa sinTezSi. magaliTad, uridindifosfati monosaqaridTa naSTebs
aaqtivebs polisaqaridebis sinTezisas, xolo citidintrifosfati - lipidTa
zogierT komponents.
nukleozidtrifosfatebis erT-erT wamyvan funqcias dnm-is da rnm-is
sinTezSi maTi monawileoba warmoadgens. isini am makromolekulebis
"saamSeneblo blokebad" gamoiyenebian. dnm-is da rnm-is sinTezis dros es
nivTierebebi kargaven or bolokidura fosfatebs da
nukleozidmonofosfatebis saxiT nukleinis mJaveebis SenebaSi iReben
monawileobas.
aRwerili mononukleotidebis garda ujredSi gvxvdebian am naerTTa sxva
tipebic, romlebsac aseve mniSvnelovani funqcia akisriaT. erT-erT aseTi
tipis naerTs nikotinamidadenindinukleotidi warmoadgens, romlis azotovani
fuZe aseve ribozis naSTTan aris dakavSirebuli. am naerTis mokle
saxelwodebaa NAD-i. NAD-Si azotovan fuZes nikotinis mJavis amidi
warmoadgens.

45
 COOH CONH2

N N
Please Register
nikotinis mJava nikotinis mJavis amidi

nikotinis mJava vitamin PP-s saxelwodebiT aris cnobili. NAD-is msgavs

naerTs flavinadenindinukleotidi (SemoklebiT FAD) warmoadgens. FAD-is
SemadgenlobaSi Sedis riboflavini (vitamini B2). NAD-i da FAD-i
monawileobas Rebuloben Jangva-aRdgeniT reaqciebSi kofermentebis funqciiT.
mniSvnelovan naerTs, romelic mraval metabolur procesebSi iRebs
monawileobas warmoadgens, kofermenti A (SemoklebiT - CoA). mis SemadgenlobaSi
Sedis adenini, panToTenis mJava da -aminoeTanTioli. CoA-s molekula
mTavrdeba sulfihidrilis (SH-) jgufiT, romelTanac rTuleTerul bmas
warmoqmnian karbonis mJaveebi. am naerTs ujredSi gadaaqvs cximovani, Zmris da
sxva organuli mJaveebi, romlebsac karboqsilis jgufebi aqvT. CoA-sa da
karbonis mJaveebs Soris kavSiri makroergulia da igi tolfasia ATP-is
fosfodieTeruli bmisa.
polinukleotidebi warmoadgenen naerTebs, romlebic sxvadasxva tipis
mononukleotidTa jaWvebisagan Sedgebian. Tu polinukleotidi aSenebulia
dezoqsinukleotidebisagan e.i. maT SemadgenlobaSi naxSirwyalbadovani
komponenti warmodgenilia dezoqsiriboziT, maT dezoqsiribonukleinis
mJavebi (dnm) ewodebaT; xolo polinukleotidebs, romlis SemadgenlobaSic
Sedis riboza - ribonukleinis mJavebi. dnm-s da rnm-s msgavsi fiziko-qimiuri
Tvisebebi aqvT, orive Sedgeba mononukleotiduri jaWvisagan, dakavSirebuls
erTmaneTTan dezoqsiribozis (dnm-Si) an ribozis (rnm-Si) naSTebiT.
miuxedavad imisa, rom dnm-is molekula pirvelad 1869 wels gamoyves
ujredidan, misi qimiuri struqturis dadgena mxolod wina saukunis 50-ian
wlebSi moxerxda. dReisaTvis cnobilia, rom yoveli organizmis dnm-i erTi
da igive mononukleotidebisagan Sedgeba. dnm-is SenebaSi monawileobas iRebs
4 mononukleotiduri erTeuli: dezoqsiAMP, dezoqsiGMP, dezoqsiTMP da
dezoqsiCMP (SemoklebiT dAMP, dGMP,dTMP,dCMP).
nukleotidebi erTmaneTTan dakavSirebulni arian sxvadasxva
TanmimdevrobiT. nukleotidebis jaWvi warmoiqmneba kovalenturi bmis

46
saSualebiT; erTi nukleotidis dezoqsiriboza meore ribozis naSTs
ukavSirdeba 3’5’'-fosfodieTeruli xidakebiT:
NH2

N
N

N
N
adenini
HO O
O
H H
NH
H H
O H

O P O- N O

Timini
O
O O
H H
N
H H NH
O H

N
O P O- N NH2

O- guanini
O
H H

H H
O H

O P O-

O-
da a.S.

(dnm-is SemTxvevaSi naxSirwyali dezoqsiribozaa, xolo rnm –is riboza)

dnm-is preparatebi, gamoyofili sxvadasxva organizmebidan gansxvavdebian

erTmaneTisagan, rogorc mononukleotiduri SemadgenlobiT, ise maTi
TanmimdevrobiT. dnm-s didi molekuluri wona gaaCnia da misi udidesi nawili

47
ujredis birTvSia lokalizebuli. dnm-i aRmoCenilia agreTve
mitoqondriebSic.
dnm-is qimiuri analizis safuZvelze dadgenil iqna Semdegi
kanonzomierebani:
1. dnm-is nukleotiduri Semadgenloba damaxasiaTebelia yoveli biologiuri
saxeobisTvis.
2. erTi organizmis sxvadasxva qsovilSi da ujredSi arsebuli dnm-is
Semadgenloba identuria da ar icvleba gamravlebis, ganviTarebis, zrdis,
diferenciaciis da sxva procesebSi. masze ar moqmedeben asaki, kveba da sxva
fiziologiuri faqtorebi.
3. monaTesave organizmebs msgavsi Semadgenlobis dnm-i aqvT.
4. yoveli organizmis dnm-Si purinis fuZeebis (adenini + guanini) raodenoba
tolia pirimidinis (Timini + citozini) raodenobisa. garda amisa adeninis
raodenoba tolia Timinisa, xolo guaninis - citozinisa.
am kanonzomierebaTa safuZvelze ingliselma mecnierebma uotsonma da
krikma SeimuSaves dnm-is struqturuli modeli, romlis safuZvelsac
warmoadgens ormagi marjvniv daxveuli spirali. spirali Sedgeba ori
antiparaleluri nukleotiduri jaWvisagan, romelTa fuZeebi erTmaneTTan
wyalbaduri bmiT arian dakavSirebuli. sivrciTi SezRudulobis gamo ormag
jaWvSi adenini ukavSirdeba mxolod Timins, xolo guanini - citozins. Tu
ormagi spirali daiSala or jaWvad, TviTeul maTganze moxdeba paraleluri
jaWvis aSeneba mxolod komplementaruli principiT, rac gulisxmobs imas,
rom adeninis sawinaaRmdegod yovelTvis dadgeba Timini, xolo guaninisa -
citozini, da piriqiT.
ujredSi aRmoCenilia sami tipis ribonukleinis mJavebi: informaciuli anu
matriculi rnm-i (m-rnm), ribosomuli rnm-i (r-rnm) da transportuli rnm-i (t-
rnm). samive tipi xasiaTdeba garkveuli molekuluri woniT da nukleotiduri
TamnmimdevrobiT. rnm-is jaWvebi (iSviaTi gamonaklisis garda) ormag spirals
ar warmoqmnian. TiToeuli rnm-i warmodgenilia ramdenime molekuluri
saxeobiT. magaliTad, cnobilia ribosomuli rnm-is sami molekuluri
varianti, transportuli rnm-is - 60 varianti, xolo informaciuli rnm-is
molekuluri formebi aTasebs aRemateba.
informaciuli rnm-is jaWvi Sedgeba oTxi tipis fuZisagan - adeninisagan,
guaninisagan, citozinisagan da uracilisagan. rnm-is es forma birTvSi
transkripciis procesSi sinTezirdeba. am dros dnm-is erT-erT jaWvze
komplementaruli rnm-is awyoba warmoebs specialuri fermentebis saSualebiT.

48
transkripciis Sedegad dnm-Si Cawerili informacia "gadaiwereba" rnm-is
jaWvze. transkripciis damTavrebis Semdeg i-rnm gadadis citoplazmaSi, sadac
ribosomebze gamoiyeneba matricis saxiT. i-rnm-is nukleotiduri
Tanmimdevroba gansazRvravs Sesatyvisi cilis aminomJavur Tanmimdevrobas. i-
rnm-is sami nukleotidi- tripleti (kodoni) Seesabameba erT aminomJavas da
tripletebis morigeoba ganapirobebs aminomJavaTa Tanmimdevrobas. TviTeuli
cila sinTezirdeba i-rnm-is erT molekulaze.
transportuli rnm-is molekulebi i-rnm-Tan SedarebiT mcire zomis arian.
maT mTavar funqcias cilis sinTezis procesSi aminomJavaTa gadatana
warmoadgens. TiToeul aminomJavas Seesatyviseba TiTo t-rnm-i. yvela tipis t-
rnm-i bolovdeba c-c-a (citozini-cotozini-adenini) TanmimdevrobiT.
ukanasknels (adenins) aqvs Tavisufali hidroqsilis jgufi, romelic
eTerificirebulia Sesabamisi aminomJavis karboqsilis jgufiT. t-rnm-Tan
aminomJavis dakavSireba specifiuri fermentebiT warmoebs. warmoqmnil
produqts aminoacil-t-rnm-i ewodeba. aminoacil-t-rnm gadaitaneba
ribosomebze, sadac peptiduri jaWvis sinTezi mimdinareobs. t-rnm-s aqvs
meore specifiuri ubani, Semdgari sami nukleotiduri Tanmimdevrobisagan -
igi komplementarulia masTan dakavSirebuli aminomJavis Sesatyvisi
tripletisa, romelic i-rnm-Sia lokalizebuli. t-rnm-is am specifiur
nukleotidur Tanmimdevrobas antikodoni ewodeba.
ribosomuli rnm-i ribosomebis SenebaSi iRebs minawileobas. miuxedavad
imisa, rom ribosomuli rnm-i ujredis rnm-is mniSvnelivan nawils Seadgens,
misi funqcia bolomde garkveuli ar aris. ribosomebi Sedgebian 50% rnm-isa
da 50% cilisagan. isini ujredis endoplazmur badeze arian ganlagebulni.
ribosomebi ori suberTeuliT arian warmodgenilni: didi suberTeuliT,
romlis molekuluri wonaa 2 milioni, da mcire suberTeuliT - molekuluri
woniT 1 milioni.

virusebi

virusebi cilebisa da nukleinis mJavebis erToblivi funqcionirebis erT-

erT magaliTs warmoadgenen. isini warmoqmnian zemolekulur kompleqsebs,
romelTa SemadgenlobaSic garkveuli TanmimdevrobiT Calagebuli nukleinis
mJava da cilebi Sedian. izolirebul mdgomareobaSi virusi ver mravldeba.
magram sakmarisia virusuli nawilaki (virioni) moxvdes "maspinZlis" ujredSi,
rom mas saSualeba eZleva sakuTari replikaciis daregulirebisa. virusuli

49
nukleinis mJava, romelic virionis infeqciur komponents warmoadgens, ikavebs
"maspinZlis" ujredis biosinTezur aparats, ris saSualebiTac awarmoebs
sakuTari cilebis sinTezs.
arsebobs dnm- da rnm-Semcveli virusebi. rnm-Semcveli virusebis
SemTxvevaSi ujredis ribosomebi upiratesobas virusul rnm-s aniWeben, ris
gamoc es ukanaskneli gamoiyeneba, rogorc matrica virusuli cilebis
sinTezisaTvis. dnm-is Semcvel virusebis SemTxvevaSi "maspinZlis" ujredSi
virusuli dnm-is transkripcia mimdinareobs da warmoqmnili i-rnm-dan
warmoebs virusuli cilebis sinTezi.

50
 metabolizmi

aqamde Cvens mier ganxiluli iyvnen is ZiriTadi bioqimiuri naerTebi,

romlebic cocxal sistemebSi moipovebian da romlebic ujredTa SenebaSi
iReben monawileobas. aRsaniSnavia is, rom es naerTebi umoqmedo statikur
mdgomareobaSi ar imyofebian. isini warmoiqmnebian, arseboben da iSlebian
garkveuli drois ganmavlobaSi da maTi "sicocxlis" xangrZlivoba
damokidebulia ujredis funqcionur mdgomareobaze. intensiurad momuSave,
dinamiur ujredSi daSlis, kataboluri procesebi sWarboben sinTetur,
anabolur reaqciebs. da piriqiT, mosvenebul mdgomareobaSi ufro intensiurad
aRdgeniTi, sinTeturi reaqciebi mimdinareoben.
dReisaTvis 2000-mde fermenti, romlebic bioqimiur gardaqmnebs
anxorcieleben da 4000 reaqcia aris cnobili. am reaqciaTa safuZvelze
Semdgaria metaboluri ruqa, romelic cocxal ujredSi mimdinare bioqimiur
procesebs asaxavs. ujredSi mimdinare dinamiuri procesebi qaotur xasiaTs ar
atareben. metabolizmi maRalkoordinirebul da mizanmimarTul process
warmoadgens, romelSic mravali bioqimiuri reaqcia Tanadroulad
mimdinareobs da sadac sakvebi nivTierebebis da plastiuri masalis daSla
Serwymulia biosinTetur reaqciebTan.

ATP da ujredis bioenergetika

Tavisufali energiis wyaros heterotrofuli ujredebisTvis, romelsac

isini qimiuri formis saxiT iReben, sakveb produqtTa daSla anu katabolizmi
warmoadgens. am dros producirebul energias ujredi Semdegi miznebisTvis
moixmars: 1. dabalmolekuluri nivTierebebidan warmoqmnis biomolekulebs, 2.
atarebs meqanikur samuSaos 3. gadaaqvs nivTierebebi membranis erTi nawilidan
meoreSi koncentraciuli gradientis sawinaaRmdegod, 4. uzrunvelyofs swori
informaciis gadacemas.
mTavar damakavSirebel rgols ujredul reaqciebs Soris, agreTve energiis
moxmarebisa da gamoyofis procesebs Soris ATP warmoadgens.
maRalenergetikuli ujreduli sawvavis daSlis procesSi
gamoTavisuflebuli Tavisufali energia ATP-s warmoqmnaSi ixarjeba ADP-dan
da araorganuli fosfatidan (Pi):
ADP + Pi ATP

51
 warmoqmnili ATP garkveuli bioqimiuri reaqciebis Catarebisas iSleba:
ATP ADP + Pi
am dros gamoyofili energia (7 kkal/molze) ujreduli procesebis
Casatareblad moixmareba. amrigad, ATP ujredSi qimiuri energiis gadamtanad
gvevlineba da mas metabolur procesebSi centraluri adgili ukavia. ATP-s
hidrolizisas Tavisufali energiis didi raodenibiT gamoyofa
ganpirobebulia nukleotidebSi arsebuli fosfodieTeruli bmis
TaviseburebebiT. Tu ATP-s da ADP-s bolokidura fosfatebis hidrolizis
procesSi 7 kkal/mol gamoiyofa, AMP-s fosfatis Camocilebisas - mxolod 3,4
kkal/mol. amitom, ATP-s molekulaSi arsebul or kidura qimiur bmas

makroerguls uwodeben da aRniSnaven mas ~ niSniT. adenozini - P ~ P ~ P

fosfatebis makroerguli bma SesaZlebelia warmoiqmnas agreTve maTi
dakavSirebisas karboqsilis, amidis da sxva qimiur jgufebTanac. zogierT
aseT naerTs 2-jer meti energiac ki aqvT, vidre ATP-s. magaliTad,
fosfoenolpiruvatis fosfatis hidrolizis dros gamoiyofa 14,8 kkal/molze.
ATP-s, rogorc energiis gadamtani naerTis roli, SesaZlebelia
Semdegnairad warmovidginoT: metaboluri procesebis Sedegad warmoqmnili
naerTi, romelsac makroerguli bma aqvs, gadascems Tavis energias ADP-s:

X ~P+ ADP ADP P (ATP)

ATP Tavis mxriv gadaitans makroergul fosfats sxva naerTze da
gaaqtiurebs mas:

ATP + Y Y ~P P + ADP
fermentebi, romlebic awarmoeben ATP-s molekulidan fosfatis jgufis
gadatanas ama Tu im naerTze (aqceptorze) - kinazebi ewodebaT.
bioqimiur naerTTa gaaqtivebis garda ATP ujredis or did procesSi iRebs
monawileobas: meqanikur muSaobaSi da osmosur wonasworobaSi. ujredis
meqanikuri muSaoba yvelaze ufro mkafiod Cans kunTovan qsovilSi, sadac
warmodgenilia mZlavri SekumSvadi specialuri struqtura - miofibrilebi. es
ukanasknelni ATP-s hidrolizis xarjze icvlian konformacias, riTac
ujredis formis cvlilebebs ganapirobeben. vinaidan danarCeni tipis
ujredebSi miofibrilebis msgavsi struqturebi ar arian aRmoCenili, maTSi
sivrciTi organizaciis formirebas mikromilakebi da mikrofilamentebi
awarmoeben. isini ganapirobeben cilebis, nukleinis mJavebis, organoidebis da
sxva nawilakebis transports ujredSi, maT gadaadgilebas (translokacias) da
koncentrirebas. mikromilakebis da mikrofilamentebis SemadgenlobaSi Sedian

52
e.w. SekumSvadi cilebi: aqtini, miozini, tubulini da sxva, romelTa mTavar
funqcias konformaciuli cvlilebebi warmoadgens. es ukanaskneli ki
mTlianad damokidebulia ATP-is hidrolizis energiaze.
meore procesi, sadac ATP-s energiis ZiriTadi fondi ixarjeba, ujredis
osmosi warmoadgens. cnobilia, rom ujredSi mimdinare osmosuri procesebis
mimdinareoba damokidebulia masSi organuli da araorganuli ionebis
raodenobiT. ujredSi ionTa Tavisufali difuzia SezRudulia morfologiuri
barierebiT, romlebic apirobeben am ionTa araTanabar gadanawilebas. aseTi
araTanabari gadanawileba warmoqmnis ionTa koncentraciul gradientebs
ujredis sxvadasxva nawilSi, riTac formirdeba ujredis osmosuri
potenciali. ionTa koncentraciuli gradientebis SesanarCuneblad ujredis
membranebSi (plazmur, organoidebis da sxva) arseboben specialuri saqaCavebi,
romlebsac es ionebi gadaaqvT membranis erTi mxridan meoreze, maTi
gradientis sawinaaRmdegod. am saqaCavebs ATP-azebi (adenozintrifosfatazebi)
ewodebaT, radgan ionTa membranaze gadatana ATP-s hidrolizis energiis
xarjze warmoebs.ujredSi ramdenime tipis aseTi saqaCavebi arseboben: Na,K-
ATPaza, Ca-ATP-aza, H-ATPaza, da sxva. process, romlis drosac xdeba ionTa
gadatana membranaze, maTi koncentraciuli gradientis sawinaaRmdegod, ionTa
aqtiuri transporti ewodeba.
miuxedavad ATP-s intensiuri hidrolizisa, misi koncentracia ujredSi
mudmivi rCeba. es metyvelebs ujredSi am nukleotidis mZlavr masinTezirebeli
sistemebis arsebobaze, romlebSic wamyvani da centraluri roli glikolizs
da mitoqondriebSi mimdinare JangviT-fosforilirebas ganekuTvneba.

katabolizmi

glikolizi

glukoza organizmis ZiriTad "sawvav" nivTierebas warmoadgens. igi

organizmSi an Tavisufali an glikogenis formiT inaxeba da saWiroebis
SemTxvevaSi bioqimiur reaqciebSi ixarjeba. glukozis daSla glikolizis
meSveobiT iwyeba. glikolizis procesSi glukozis 6-naxSirbadiani molekula
or 3-naxSirbadian piroyurZnis mJavamde iSleba. glukozis piroyurZnis
mJavamde daSla 10 Tanmimdevruli fermentuli reaqciis saSualebiT warmoebs.
am dros gamoTavisuflebuli energia ATP-s saxiT grovdeba.

53
 glikolizis produqti - piroyurZnis mJava (piruvati) SesaZlebelia sami
gziT moixmaros. aerobul organizmebSi, glikolizi glukozis daSlis
mxolod pirvel stadias warmoadgens, radgan warmoqmnili piruvati SemdgomSi
mTlianad iSleba CO2-mde da H2O-mde. piruvatis daSla, am SemTxvevaSi,
grZeldeba JangviTi dekarboqsilirebis reaqciiT, romlis drosac mas
waerTmeva CO2. darCenili 2-naxSirbadiani ZmarmJavas naSTi, aqtivirebuli
kofermenti A-Ti (CoA) erTveba Semdgom gardaqmnebSi, romelsac limonmJava
anu krebsis cikli ewodeba.
glukozis daSlis meore gzas warmoadgens piruvatis aRdgena rZis mJavamde
(laqtati). adamianis organizmSi es procesi ZiriTadad ConCxis kunTebSi
mimdinareobs maTi intensiuri muSaobisas. am dros Jangbadis moxmareba didia,
da amitom, misi ujredSida koncentracia - qveiTdeba. Seqmnili anaerobuli
pirobebis gamo, piruvati iwyebs gadasvlas laqtatSi. am process anaerobuli
glikolizi ewodeba. analogiuri procesi zogierT mikroorganizmebSic
mimdinareobs, sadac igi rZemJava duRiliT aris cnobili.

glukoza
glikolizi

2 piruvati

anaerobuli anaerobuli
pirobebi pirobebi
O2
aerobuli
pirobebi
2 eTanoli + 2CO2 2 laqtati
2 H2 O
spirtuli 2 acetil-CoA anaerobuli
duRili glikolizi
O2

limonmJavas
cikli

4 CO2 + 4 H2O

54
 piruvatis gardaqmnis mesame gzas spirtuli duRili warmoadgens. am gzis
saSualebiT zogierTi mikroorganizmebi, da maT Soris safuarebi,
piruvatidan warmoqmnian eTanols. anaerobuli duRili evoluciurad erT-erT
uZveles biologiur process warmoadgens.
glikolizis procesSi SeiZleba ori stadia gamoiyos. pirvel stadiaSi
gaerTianebulia pirveli xuTi raeqcia, romlebsac pirobiTad SesaZlebelia
glikolizis mosamzadebali etapi vuwodoT. am reaqciebis Sedegad glukoza
fosforilirdeba jer naxSirbadis me-6 atomTan, xolo Semdeg - 1 atomTan.
warmoqmnili fruqtozo-1,6-difosfati iSleba gliceraldehid-3-fosfatamde.

glukoza
ATP heqsokinaza
glukozo-6-fosfati
fosfoglukoizomeraza
fruqtozo-6-fosfati
ATP fosfofruqtokinaza
fruqtozo-1,6-difosfati
aldolaza
gliceraldehid-3-fosfati + dihidroacetonfosfati
triozofosfatizomeraza
gliceraldehid-3-fosfati (2 molekula)
gliceraldehidfosfatdehidrogenaza
3-fosfogliceroilfosfati (2 molekula)
2ATP fosfogliceratkinaza
3-fosfoglicerati (2 molekula)
fosfogliceratmutaza
2-fosfoglicerati (2 molekula)
enolaza
fosfoenolpiruvati (2 molekula)
G 2ATP piruvatkinaza
piruvati (2 molekula)

55
 amrigad, glikolizis pirvel, mosamzadebel stadiaze xdeba 6-naxSirbadiani
glukozis daSla 2 molekula samnaxSirbadian naxSirwylamde -
gliceraldehid-3-fosfatamde, risTvisac 2 molekula ATP-is energia ixarjeba.
am stadiaze SesaZlebelia daiSalos sxva heqsozebic: D-galaqtoza, D-
fruqtoza, D-manoza (mas Semdeg, rac moxdeba maTi fosforilireba).
glikolizis mosamzadebel stadiaSi monawile fermentebi sakmaod kargadaa
Seswavlili. isini gamoyofili arian ujredidan da maTi moqmedebis
meqanizmebi cnobilia. pirvel reaqcias atarebs fermenti heqsokinaza. igi
regulatorul, alosterul ferments warmoadgens da inhibirdeba sakuTari
produqtiT, glukozo-6-fosfatiT. RviZlSi warmodgenilia am fermentis sxva
forma, romelsac glukokinaza ewodeba. heqsokinazisagan gansxvavebiT, es
ukanaskneli specifikuria mxolod D-glukozis mimarT da ar inhibirdeba
glukozo-6-fosfatiT. RviZlis glukokinaza mxolod mas Semdeg Sedis
reaqciaSi, rodesac glukozis koncentracia sisxlSi izrdeba (magaliTad,
saWmlis monelebisas). am dros, glukokinazas glukozis Warbi raodenoba
gadahyavs glukozo-6--fosfatSi, romlisganac SemdgomSi glikogeni
sinTezirdeba. Saqriani diabetiT daavadebul adaminebSi am fermentis aqtioba
daqveiTebulia, da amitom glikogenis maragis dagroveba RviZlSi ver
xerxdeba.
glikolizis me-2 reaqciaSi xdeba glukozis izomerizacia, romelsac
fermenti fosfoglukoizomeraza anxorcielebs. Sedegad glukozo-6-fosfati
gardaiqmneba fruqtozo-6-fosfatad. me-3 reaqciis, iseve rogorc pirveli
reaqciis dros ATP ixarjeba naxSirwylis fosforilirebisTvis. fruqtozo-
1,6-difosfats warmoqmnis fermenti fosfofruqtokinaza. es ukanaskneli,
heqsokinazis msgavs rTul regulatorul ferments warmoadgens da
gansazRvravs mTeli glikolizis procesis siCqares. sakmarisia intensiuri
muSaobis Sedegad qsovilSi (mag. ConCxis kunTebSi) Semcirdes ATP-is
koncentracia, an moimatos misi daSlis produqtTa (ADP, AMP) raodenobam,
fosfofruqtokinazis aqtivoba, da Sesabamisad mTeli glikolizis siCqare,
izrdeba. xolo ATP-is zrdis SemTxvevaSi, piriqiT - fermenti inhibirdeba. me-4
reaqciaSi fermenti aldolazas meSveobiT fruqtozo-1,6-difosfati ixliCeba
2 samnaxSirbadian naerTad gliceraldehid-3-fosfatad da
dihidroacetonfoafatad, xolo es ukanaskneli triozofosfatizomerazis
mier gardaiqmneba gliceraldehid-3-fosfatad. amrigad, glikolizis pirvel
"mosamzadebel" stadiaze adgili aqvs heqsozis daxleCas or molekula

56
samnaxSirbadian gliceraldehid-3-fosfatamde, risTvisac ixarjeba ori ATP-is
tolfasi energia - 14 kkal/molze.
glikolizis meore stadia, romelic gliceraldehid-3-fosfatis gardaqmniT
iwyeba, aseve 5 reaqciisagan Sedgeba. am stadiaze, jer erTi aRdgeba wina
stadiaze daxarjuli ATP-is ori molekula, da Semdeg - damatebiT
sinTezirdeba ori axali molekula. gliceraldehid-3-fosfatis gardaqmnas 3-
fosfogliceroilfosfatad awarmoebs fermenti
gliceraldehidfosfatdehidrogenaza. am reaqciis Sedegad xdeba
araorganuli fosfatis mierTeba gliceraldehid-3-fosfatis aldehidis
jgufze. warmoqmnil anhidrids acilfosfati (COO-H2PO3)Oewodeba da misi
hidrolizis Sedegad didi energia (11,8 kkal/molze) gamoiyofa. am reaqciis
koferments nikotinamidadenindinukleotidi (NAD) warmoadgens. rogorc ukve
iTqva, mis SemadgenlobaSi Sedis nikotinis mJavis amidi (vitamini PP). reaqciis
msvlelobisas NAD-Si Semavali nikotinamidi daJanguli formidan (NAD+)
aRdgenilSi gadadis (NADH), radgan substratidan (gliceraldehid-3-
fosfatidan) erT protons (H+) ierTebs. momdevno, fosfogliceratkinazis
mier warmoebul reaqciaSi, fosfogliceroilfosfatis acilfosfaturi bmis
hidrolizi xdeba da gamonTavisuflebuli energiis xarjze fosfaturi jgufi
ADP-ze gadaitaneba. Sedegad warmoiqmneba ATP.
am reaqciiT warmoqmnili 3-fosfoglicerati ferment
fosfogliceratmutazis mier gardaiqmneba 2-fosfogliceratad, romelsac
SemdgomSi ferment enolazis saSualebiT waerTmeva wyali. warmoqmnili
produqti - fosfoenolpiruvati - zemaRal energetikul naerTs warmoadgens,
radgan misi fosfaturi jgufis hidrolizis dros gamoyofili energia ( 14,8
kkal/mol) 2-jer aRemateba ATP-s kidura fosfatebis hidrolizis energias. da
bolos, fosfoenolpiruvatis fosfati gadaisroleba ADP-ze ferment
piruvatkinazas daxmarebiT. reaqciis Sedegad warmoiqmneba ATP da piruvati
(piroyurZnis mJava).
piroyurZnis mJavas didi mniSvneloba eniWeba ujredis metabolizmSi, igi
mravali daSlis Tu biosinTeturi gzebis gadakveTaze imyofeba. im
SemTxvevaSi, Tu glikolizi aerobul pirobebSi mimdinareobs da ujreds
Jangbadis sakmarisi fondi aqvs, piruvati krebsis ciklSi erTvis da
agrZelebs daSlas - daJangvas. gliceraldehid-3-fosfatdehidrogenazuli
reaqciis Sedegad warmoqmnili NADH aseve ganagrZobs daJangvas
mitoqondriebSi sawyis formamde - NAD+-mde. anaerobul pirobenSi, rodesac

57
ujredi Jangbadis deficits ganicdis (mag. intensiurad momuSave ConCxis
kunTebSi), NADH-is daJangva TviT piruvatis saSualebiT xdeba da es
ukanaskneli rZis mJavad (laqtati) gardaiqmneba:

piruvati + NADH + H+ L-laqtati + NAD+

am reaqcias fermenti laqtatdehidrogenaza warmarTavs. es fermenti didi

raodenobiT aris aRmoCenili kunTovan qsovilSi, mas xuTi izoforma aqvs,
romelTagan erTi - H4, specifikuria gulis kunTisTvis. piruvatis laqtatad
gardaqmna intensiurad warmoebs jangbadis ukmarisobisas da iwvevs didi
raodenobiT rZis mJavis dagrovebas. amis gamo qsovilebSi pH-is davardnas,
SemJavianebas aqvs adgili. swored laqtatis dagroveba iwvevs adamianSi
"daRlili kunTebis" SegrZnebis fenomens.
amrigad, glikolizis procesSi glukozis molekulis orad iSleba.
glukozis molekulis dayofisas amoiReba 14 kkal/molze energia, romelic 2
molekula ATP-Si Seinaxeba. mis mimdinareobas Jangbadi ar esaWiroeba, igi
anaerobul pirobebSi xdeba. glikolizis procesi citoplazmaSi warmoebs da
masSi SesaZlebelia CaerTon sxva Saqrebic: polisaqaridebidan - glikogeni da
saxamebeli, disaqaridebidan - maltoza, laqtoza, da saqaroza, xolo
monosaqaridebidan - fruqtoza, manoza da galaqtoza.

glikogenolizi

glikogeni glukozis naSTebisagan Semdgar polisaqaridul jaWvs

warmoadgens. igi maragdeba ujredebSi (umTavresad RviZlSi) da organizmis
energetikuli moTxovnilebis mixedviT iSleba glukozad. glikogeni
hidrolizdeba fosforolizis saSualebiT, da am process fermenti -
glikogenfosforilaza warmarTavs. es ukanaskneli bolokidura glukozis
fosforilirebas iwvevs da warmoqmnili glukozo-1-fosfati Camocildeba
glikogenis jaWvs:

(glukoza)n + Pi (glukoza)n-1 + D-glukozo-1-fosfati

vinaidan, glikolizSi monawileobas Rebulobs ara glukozo-1-fosfati,

aramed glukozo-6-fosfati, adgili aqvs am ukanasknelis sinTezs ferment

58
fosfoglukomutazis saSualebiT. am reaqciis Semdeg glukozo-6-fosfati
erTvis glikolizis procesSi. ConCxis kunTebSi glikogenfosforilaza
warmodgenilia ori formiT: dabalaqtiuri fosforilaza b-Ti da
maRalaqtiuri fosforilaza a-Ti. dabalaqtiuri fosforilaza b mxolod mas
Semdeg gadadis aqtiur formaSi (fosforilaza a-Si), rodesac moxdeba misi
cilovani molekulis posttranslaciuri kovalenturi modifikacia, kerZod
fosforilireba. fosforilirebul mdgomareobaSi myofi fermenti (e.i.
fosforilaza a) gacilebiT ufro swrafad Slis glikogens, vidre -
arafosforilirebuli. glikogenfosforilazis fosforilireba-
defosforilirebas da Sesabamisad a da b formebis urTierTgardaqmnas
awarmoebs ori fermenti: fosforilazis kinaza da fosforilazis fosfataza.
pirveli iwvevs glikogenfosforilazis fosforilirebas, xolo meore - mis
defosforilirebas.
fosforilazis kinaza, erT-erTi tipiuri warmomadgenelia regulatorul
fermentTa im didi klasisa, romlebsac proteinkinazebi ewodebaT. isini
awarmoeben sxvadasxva cilebis (maT Soris fermentebis) kovalenturi
modifikaciis centralur reaqcias - fosforilirebas. maTi moqmedebis
Sedegad ATP-s bolokidura fosfati gadaitaneba cilovan molekulaze da
ukavSirdeba am cilovani molekulaSi Semaval aminomJava serinis naSTs:
proteinkinaza

cila + ATP cila-P + ADP

cilis mier uaryofiTad damuxtuli fosfatis jgufis mierTeba scvlis mis

mesameul struqturas, konformacias. Tu aseT cilas fermentuli aqtivoba
aqvs, icvleba misi aqtivobac: izrdeba an mcirdeba.
aseTi modificirebuli cilidan fosfatis jgufis Camoxsnas, meore
fermenti - proteinfosfataza awarmoebs:

proteinfosfataza

cila-P cila + Pi

am reaqciis Sedegad cilis molekula Tavisufldeba fosfatis jgufisagan

da ubrundeba sawyis mdgomareobas.
fosforilirebis da defosforilirebis reaqciebiT ujredSi asobiT
cilisa da fermentis regulacia xdeba. vinaidan, mafosforilirebeli

59
fermentebis, proteinkinazebis umravlesoba, dabalmolekuluri naerTebiT
aqtivirdeba, romlebic xSirad ujredSida regulatorebs warmoadgenen, da
romelTa koncentraciac ujredSi hormonebis, mediatorebis da sxva
fiziologiurad aqtiuri nivTierebebis zegavleniT icvleba, gasagebi xdeba
Tu ra didi mniSvneloba eniWebaT am fermentebs metabolizmis regulaciaSi.
Cvens mier ganxilul SemTxvevaSi, fermenti fosforilazis kinaza
aqtivirdeba Ca-is ionebiT. aqedan gamomdinare - rac ufro maRalia ujredSi
Ca-is koncentracia, miT ufro aqtiuria fosforilazis kinaza da miT ufro
intensiurad warmoebs glikogenis daSla. aseTi tipis regulacias didi
mniSvneloba eniWeba kunTovan qsovilSi, radgan masSi, kalciumis
koncentraciis cvlilebebi dakavSirebulia kunTis SekumSvasTan.
amrigad, glikogenis daSlis regulacia SesaZlebelia Semdegi sqemiT
gamoisaxos:
glikogeni

fosforilaza a-P (aqtiuri)

glukozo-1-fosfati
fosforilazis kinaza
fosforilaza b (pasiuri)

unda aRiniSnos, rom glikogenis daSlis regulacia amiT ar mTavdeba; TviT

fermenti fosforilazis kinaza ganicdis fosforilirebas sxva tipis
proteinkinaziT da SesaZlebelia iyos aqtiur da araaqtiur mdgomareobebSi.
ukanasknel ferments cAMP-damokidebuli proteinkinaza ewodeba da misi
kontroli hormonebisa da mediatorebis saSualebiT xdeba. am reaqciaTa
meSveobiT glikolizi fiziologiurad aqtiuri nivTierebebiT regulirdeba.
garda zemoTaRwerili meqanizmebisa, glikolizi sxvadasxva tipis
metabolitebiTac ganicdis cvlilebebs. glikolizis pirvel regulirebad
ferments heqsokinaza warmoadgens. kunTebSi es fermenti sakuTari produqtiT,
glukozo-6-fosfatiT inhibirdeba. amitom, sakmarisia kunTebSi am naerTis
koncentraciam moimatos, rom igi gamoiwvevs heqsokinazis damuxruWebas, da
Sesabamisad, mTeli glikolizis procesis Senelebas. rogorc zemoT ukve iyo
naTqvami, RviZlSi am fermentis garda arsebobs analogiuri aqtivobis mqone -
glukokinaza, romelic glukozo-6-fosfatiT ar inhibirdeba. amitom RviZlSi,

60
kunTebisagan gansxvavebiT glukozis Warbi raodenoba moixmareba glikogenis
sinTezisaTvis.
glikolizi regulirdeba ara marto glukozis miwodebis doneze, aramed is
ganicdis cvlilebebs im metabolitebiTac, romlebic TviT am procesSi
warmoiqmnebian. am mxriv, mniSvnelovania ori sakvanZo fermentis -
fosfofruqtokinazis da piruvatkinazis regulacia. pirveli aqtivirdeba AMP-
iT, ADP-iT, fruqtozo-1,6,-difosfatiT da araorganuli fosfatiT, e.i.
naerTebiT, romlebic ujredSi energetikuli masalis simcireze metyveleben;
da inhibirdeba ATP-iT da citratiT - metabolitebiT, romlebic ujredis
maRalenergetikul dones gamoxataven. aseve regulirdeba ATP-iT fermenti--
piruvatkinaza.
da bolos, aucilebelia aRiniSnos, rom zogierT mikroorganizmebSi
glukozis daSla mimdinareobs eTanolamde. am process spirtuli duRili
ewodeba. aseTi tipis mikroorganizmebs aqvT fermenti piruvatdekarboqsilaza,
romelic glikolizis Sedegad warmoqmnili piruvatis dekarboqsilirebas
awarmoebs. Sedegad warmoiqmneba acetaldehidi, romelic ferment
alkoholdehidrogenazas saSualebiT eTanolad gardaiqmneba:

CH3 CH3 CH 3

C O C O CH 2OH

COOH H Register
Please
piruvati acetaldehidi eTanoli

limonmJavas cikli

wina TavebSi Cvens mier ganxiluli iyo glikolizis procesi. igi

anaerobul pirobebSi mimdinareobs da glukozis katabolizmis evoluciurad
uZveles gzas warmoadgens. am procesis sabolo stadiamde warmarTvis
Sedegad ujredSi grovdeba rZis mJava an eTanoli. radgan organizmebis
umravlesobas funqcionireba uxdebaT Jangbadis garemoSi, aerobul pirobebSi,
glukozis daSla aseTi tipis organizmebSi daJangvis reaqciebiT grZeldeba
da am dros gamoyofili energia gacilebiT aRemateba anaerobul pirobebSi
generirebul energias.

61
 glukozis aerobuli daJangva piroyurZnis mJavidan iwyeba da igi sam
stadias gadis. pirvel stadiaze piruvatidan acetil-CoA warmoiqmneba,
romelic meore stadiaze iSleba limonmJavas ciklis saSualebiT. mesame
stadiaze daSlili produqtebidan amoiRebian eleqtronebi, romlebic
mitoqondriebSi arsebuli specialuri satransporto jaWvis gavlis Semdeg
generireben energias. es energia, saboloo jamSi, ATP-is molekulebis qimiur
bmaSi Seinaxeba.
acetil-CoA-s warmoqmna. glikolizis Sedegad miRebuli piruvati ganicdis
dekarboqsilirebas, kargavs erT naxSirbadis atoms. warmoqmnili acetilis
jgufi (ZmarmJavas naSTi) uerTdeba koferment A-s (CoA) da sinTezirdeba
acetil-CoA. CoA rTul naerTs warmoadgens, romlis SemadgenlobaSic Sedian
ADP, panToTenis mJava (vitamini B3) da TioeTilaminis naSTi. misi molekula
bolovdeba TioeTilaminis sulfihidrilis-SH jgufiT, romelsac uerTdeba
ZmarmJavas karboqsili:

CH3 C S CoA

O O

gogirdsa da naxSirbads Soris arsebuli bma makroergulia, misi hidrolizis

SemTxvevaSi ATP-s fosfatebis tolfasi energia (7 kkal/molze) gamoiyofa.
amitom xSirad ZmarmJavis naSTis mierTebas CoA-Tan acetatis gaaqtivebas
uwodeben. gaaqtivebuli acetati Sedis biologiuri daJangvis meore stadiaSi
- limonmJavas ciklSi. aqve unda aRiniSnos, rom aqtivirebuli acetati anu
acetil-CoA warmoiqmneba agreTve cximebis da aminomJavebis katabolizmis
Sedegadac. amrigad, igi im erT-erT centralur naerTTagania, romlebic
metaboluri gzebis gadakveTaze imyofebian.
piruvatis dekarboqsilirebas da acetil-CoA-s warmoqmnas awarmoebs
rTuli multifermentuli - piruvatdehidrogenazuli kompleqsi, romelic
sami fermentisagan Sedgeba: TviT piruvatdehidrogenazisagan, dihidrolipo-
ilacetiltransferazisagan da dihidrolipoildehidrogenazisagan. am
kompleqsis mier warmoebuli jamuri reaqcia Semdeg saxes atarebs:

piruvati + CoA + NAD+ acetil-CoA + NADH + H+ + CO2

62
 am reaqciaSi monawileobas iRebs 5 kofermenti: Tiaminpirofosfati, FAD,
CoA, NAD da lipois mJava, romelTa SemadgenlobaSic 4 adamianisTvis
aucilebeli vitaminia (Tiamini-B1-TiaminpirofosfatSi, riboflavini-B2-FAD-Si,
panToTenis mJava-B3-CoA-Si da nikotinamidi--NAD-Si). piruvatdehidrogenazuli
reaqciis Sedegad warmoqmnili acetil-CoA limonmJavas ciklSi
metabolizirdeba.
limonmJavas cikli. limonmJavas cikli, glikolizisagan gansxvavebiT,
daxurul cikls warmoadgens. igi pirvelad 1937 wels iyo aRwerili krebsis
mier da amitom mas xSirad krebsis ciklsac uwodeben. misi reaqciebi
mitoqondriebis Sida membranebze mimdinareoben da maTi msvleloba 8
stadiaSi xdeba.
I stadias warmoadgens acetil-CoA-s kondensacia (dakavSireba)
oqsaloacetatTan (mJaunZmarmJavasTan), romelsac fermenti citratsinTaza
akatalizirebs:
COOH

COOH CoA CH2

CO COH COOH
+ CH3CO SCoA
citratsinTaza
CH2 CH2

COOH C OOH
oqsaloacetati citrati

igi rTul alosterul ferments warmoadgens, regulirdeba sxvadasxva

metabolitebiT da kontrols uwevs mTeli krebsis ciklis mimdinareobas
(aseTi tipis fermentebs malimitirebeli fermentebi ewodebaT).
II stadias Seadgens izomerizaciis reaqcia, romelsac warmarTavs fermenti
akonitaza, da romlis drosac limonis mJava (citrati) gardaiqmneba izo-
limonis mJavad (izocotratad):

63
 COOH COOH

CH2 CH2

COH COOH CH2 COOH

akonitaza
CH2 CHOH

C OOH C OOH

citrati izocitrati
III stadiaze xdeba warmoqmnili izocitratis dehidrireba da miiReba -
ketoglutaris mJava. am reaqcias atarebs fermenti - izocitratdehidrogenaza:
COOH COOH

CO2
CH2 NAD+ NADH + H+ CH2

CH2 COOH CH2

izocitratdehidrogenaza
CHOH CO

C OOH C OOH
izocitrati -ketoglutarati

IV stadiaze -ketoglutaratis JangviTi dekarboqsilireba mimdinareobs,

romelsac -ketoglutaratdehodrogenazuli kompleqsi awarmoebs. am dros
warmoiqmneba qarvis mJava (suqcinati), romlis karboqsilis jgufi CoA-Ti
aris aqtivirebuli:
COOH COOH
CoA

CO2
CH2 NAD+ NADH + H+ CH2

CH2 CH2
-ketoglutaratdehidrogenaza
CO CO

C OOH S CoA
-ketoglutarati suqcinil-CoA

64
 es reaqcia hgavs zemoT aRweril piruvatdehidrogenazul reaqcias, romlis
drosac warmoiqmneboda acetil-CoA. -ketoglutaratdehidrogenaza aseve sami
fermentisagan Sedgeba, romlebic Sesabamis kofermentebTan erTad
monawileoben reaqciaSi. reaqciis Sedegad warmoqmnili suqcinil-CoA, acetil-
CoA-s msgavsad maRalenergetikul naerTs warmoadgens.
V stadia warmodgenilia suqcinil-CoA-sinTetazuri reaqciiT, romlis
drosac suqcinil-CoA-s energia gadadis nukleozidtrifosfatis, kerZod GTP-s
fosfodieTerul bmaSi. am dros xdeba GDP-s aRdgena GTP-mde.
COOH COOH
GTP
GDP + Pi HSCoA
CH2 CH2

CH2 suqcinil-CoA-sinTetaza CH2

CO COOH

S CoA

suqcinati
suqcinil-CoA

reaqciis Sedegad warmoqmnili GTP-s bolokidura fosfati gadaisroleba

ADP-ze ferment nukleoziddifosfatkinazas meSveobiT:

GTP + ADP GDP + ATP

da sabolood ATP miiReba.
VI stadiaze suqcinatis dehidrireba warmoebs, romlis Sedegadac
warmoiqmneba fumaris mJava (fumarati). am reaqcias fermenti
suqcinatdehidrogenaza asrulebs. mis prostetul jgufs FAD-i warmoadgens,
romelic reaqciis msvlelobisas ikavSirebs or wyalbadis atoms da aRdgeba
FADH2-mde.

65
 COOH COOH

FAD FADH2
CH2 CH

CH
CH2 suqcinatdehidrogenaza

COOH COOH

suqcinati fumarati

FAD-is garda suqcinatdehidrogenaza Seicavs rkinis atomebs, romlebic

reaqciis Sedegad valentobas icvlian, Fe(II) gadadis Fe(III)-Si.
VII stadiaze warmoebs fumaratis hidratireba da warmoiqmneba vaSlis
mJava(malati). reaqcias fermenti fumarathidrataza akatalizirebs:
COOH COOH
H2O

CH COH

CH fumarathidrataza CH2

COOH COOH
fumarati malati
VIII, ciklis ukanasknel stadiaze vaSlis mJavidan warmoiqmneba
mJaunZmarmJava (oqsaloacetati), romelsac fermenti malatdehidrogenaza
atarebs. am dros xdeba NAD+-is aRdgena NADH-mde:
COOH COOH

NAD+ NADH + H+
COH CO

CH2 malatdehidrogenaza CH2

COOH COOH
malati oqsaloacetati

am reaqciiT mTavrdeba limonmJavas ciklis erTi wre. amrigad, limonmJavas

ciklSi Tavdapirvelad Sedis ornaxSirbadiani naerTi - acetati, romelic

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gaaqtivebulia CoA-Ti. ZmarmJavas naSTi ukavSirdeba mJaunZmarmJavas limonis
mJavis warmoqmniT. ciklis msvlelobisas orive naxSirbadi Tavisufldeba ori
molekula naxSirorJangis saxiT da ciklis boloSi oqsaloacetati
regenerirebulia.
ciklSi monawileobas iRebs 4 dehidrogenaza, romelTa reaqciebis Sedegad
xdeba 4 wyvili wkalbadis atomis Camocileba metabolitebidan. aqedan sami
wyvili xmardeba NAD+-is aRdgenas NADH-mde, xolo erTi - FAD-is aRdgenas
FADH2-mde. 4 wyvili eleqtronebisa, romlebic protonis kofermentebTan
mierTebis Semdeg rCebian, gadaecemian eleqtronTa gadatanis jaWvs. am jaWvis
saSualebiT, isini aRadgenen Jangbadis molekulas (O2) or molekula wylamde.
ciklis procesSi warmoebs erTi molekula makroerguli nivTierebis (GTP)
sinTezi.
limonmJavas ciklis (sunTqvis jaWvTan erTad) saboloo gantoleba
SesaZlebelia Semdegnairad gamoisaxos:

CH3-COOH + 3O2 ________ 2CO2 + 2H2O

aRsaniSnavia is, rom am erTi SexedviT ubralo reaqciis Casatareblad,

ujredisaTvis saWiro gaxda mTeli limonmJavas ciklis Seqmna. saqme imaSi
mdgomareobs, rom ZmarmJavis meTilis jgufis dasaJangavad aucilebelia
aqtivaciis didi energia, rac TavisTavad miuRebelia ujredisaTvis. amitom,
arCeulia ZmarmJavis daJangvis meore gza, romlis drosac adgili aqvs wvis
energiis ara erTjerad, didi raodenobiT gamoTavisuflebas, aramed mis
etapobriv gadanawilebas.
piroyurZnis mJavis gardaqmna acetil-CoA-mde, iseve rogorc mTeli
limonmJavas cikli, ujredis centralur metabolur gzas warmoadgens, da
amitom sxvadasxva tipis nerTebiT regulirdeba. magaliTad,
piruvatdehidrogenazuli kompleqsi acetil-CoA-Ti da NADH-iT inhibirdeba.
orive naerTi am kompleqsis produqtebs warmoadgens da maTi ujredSi
dagroveba fermentis SeCerebas iwvevs. garda amisa, piruvatdehidrogenaza Ca-is
ionebiT aqtivirdeba. es ukanasknelni, mitoqondrialur JangviT-
fosforilirebis sistemebTan mWidro kavSirSi imyofebian. Ca2+-is
koncentraciis zrda ujredSi energetikuli procesebis Semcirebaze
metyvelebs, da amitom am ionebiT piruvatdehidrogenazis aqtivacias didi
mniSvneloba eniWeba ATP-s raodenobis regulaciaSi.

67
 TviT limonmJavas ciklis fermentebidan regulirdeba citratsinTaza, mis
Sekavebas suqcinil-CoA iwvevs. es ukanaskneli, limonmJavas ciklis momdevno
stadiebze warmoiqmneba da misi didi raodenobiT dagroveba ciklis
intensiur muSaobaze metyvelebs. fermenti izocitratdehidrogenaza, romelic
izolimonis mJavidan -ketoglutarats warmoqmnis, ADP-Ti aqtivirdeba. ADP-s
Warbi raodenoba ujredSi energetikuli deficitis pirobebSi warmoiqmneba, da
amitom am naerTiT krebsis ciklis erT-erTi wamyvani fermentis stimulireba
mTeli ciklis gaZlierebas, da Sesabamisad, ATP-s producirebas iwvevs.
normalur pirobebSi glikolizis siCqare SeTanxmebulia limonmJava
ciklis siCqaresTan: ujredSi imdeni glukoza iSleba piruvatamde, ramdenic
saWiroa misi ciklSi CasarTavad. arc piruvati da arc acetil-CoA didi
raodenobiT ar grovdebian. maTi koncentracia sakmaod dabal doneze
meryeobs da isini dinamiur wonasworobaSi imyofebian krebsis ciklis sxva
metabolitebTan. am metabolitebSi, ATP-s da NADH-is garda, romlebic
glikolizisa da sunTqvis stadiebis saerTo gamosaval produqtebs Seadgenen,
wamyvani mniSvneloba limonmJavas eniWeba. es ukanaskneli, warmoadgens ra
krebsis ciklis pirvel produqts, iwvevs glikolizis erT-erTi ZiriTadi
fermentis - fosfofruqtokinazis - inhibirebas. amitom, bunebrivia, rom
citratis dagroveba ujredSi gamoiwvevs rogorc glikolizis, aseve krebsis
ciklis SeCerebas.
limonmJavas cikli mravali organuli nivTierebis daSlis saboloo
stadias warmoadgens. naxSirwylebis, cilebis da cximebis katabolizmis
produqtebi limonmJavas ciklis metabolitebi arian. garda amisa, piriqiT,
ciklis zogierTi naerTi moixmareba biosinTezur reaqciebSi. aseTebi arian -
ketoglutaris mJava, suqcinati, oqsaloacetati. miuxedavaT imisa, rom es
metabolitebi didi raodenibiT moixmarebian anabolur reaqciebSi (mag.
aminomJavaTa sinTezSi) maTi koncentraciebi mudmivi rCeba saTadarigo
sistemebis amuSavebis gamo. erT-erT aseT saTadarigo ferments
piruvatkarboqsilaza warmoadgens, romelic piruvatidan warmoqmnis
oqsaloacetats, Tu es ukanaskneli didi raodenobiT ixarjeba aminomJavaTa
sinTezSi. aseT fermentTa saSualebiT xdeba limonmJavas ciklis Sevseba
metabolitebiT da maT anapreloturi sistemebi ewodebaT.
mcenareebSi arsebobs limonmJavas ciklis modifikacia, romelsac
glioqsalaturi cikli ewodeba.

68
 glukozis katabolizmis meoradi gzebi - pentozofosfaturi gza.

miuxedavad imisa, rom glukozis daSla cxovelur organizmebSi,

ZiriTadad, piruvatamde mimdinareobs, xolo es ukanaskneli ki - bolomde
iJangeba limonmJava ciklSi, arseboben meoradi gzebi, romelTa saSualebiTac
glukozidan warmoiqmnebian sxva, specialuri daniSnulebis produqtebi. erT-
erT aseT, glukozis meoradi metabolizmis gzas, pentozofosfaturi anu
fosfoglukonaturi gza warmoadgens. Tu glikolizis SemTxvevaSi glukozis
daJangva mimdinareobs ATP-s sinTezisTvis, pentozofosfaturi reaqciebis
Sedegad miiReba ori mniSvnelovani metaboliti:
nikotinamiddinukleotidfosfati (NADP) da ribozo-5-fosfati. NADP, iseve
rogorc NAD, monawileobas iRebs dehidrogenazul reaqciebSi da aRdgeba
NADPH-mde. isini msgavsi funqciebis naerTebi arian. aqve unda aRiniSnos, rom
NADP (iseve, rogorc NAD) ATP-s msgavsad energiis matarebels warmoadgens,
oRond im SemTxvevaSi, Tu nukleozodtrifosfatis energia inaxeba mis
fosfodieTerul bmebSi, NADP energiis utilizacias awarmoebs Tavisi
aRdgeniTi potencialiT. NADP-s es funqcia did mniSvnelobas iRebs im
qsovilebSi, romlebSic warmoebs cximovani mJavebis da steroidebis
biosinTezi da maSinac, roca qsovili maTi winamorbedebis deficits
ganicdis. aseT SemTxvevebs adgili aqvT sarZeve jirkvlebSi, cximovan
qsovilebSi, Tirkmelzeda jirkvalSi, RviZlSi da sxv. cximovani mJavebis
biosinTezis dros NADPH-is saSualebiT xdeba ormagi bmebis aRdgena, rac
aucilebelia am naerTTa normaluri metabolizmisTvis. im qsovilebSi,
romlebSic cximovani mJaveebis metabolizmi sustia (mag. kunTebSi)
pentozofosfaturi gza praqtikulad ar arsebobs. am gzis meore
daniSnulebaa ujreds miawodos pentozebi, da kerZod, D-riboza. es
ukanaskneli ki nukleinis mJavebis biosinTezSi gamoiyeneba.
pentozofosfaturi gza citoplazmaSi mimdinareobs.
glukozis pentozofosfaturi daJangvis pirvel reaqcias warmoadgens
glukozo-6-fosfatis dehidrireba, romlis Sedegadac miiReba 6-
fosfoglukono- -laqtoni. am reaqcias glukozo-6-fosfatdehidrogenaza
atarebs.
II stadiaze fosfoglukonat- -laqtoni gadadis 6-fosfoglukonatSi ferment
laqtonazis saSualebiT. 6-fosfoglukonatis JangviTi dekarboqsilirebis
Sedegad miiReba D-ribulozo-5-fosfati. am reaqcias 6-glukonatdehidrogenaza

69
atarebs. da bolos, ferment fosfopentozoizomerazis saSualebiT
warmoiqmneba D-ribozo-5-fosfati.
zemoT aRwerili dehidrogenazebis kofermentebs NADP warmoadgens,
romelic NADPH-mde aRdgeba. amrigad, erTi molekula glukozis
pentozofosfaturi gziT daJangvisas ori molekula NADPH miiReba. reaqciebis
Sedegad miRebul saboloo produqts 5-naxSirbadiani ribozo-5-fosfati
warmoadgens.
rogorc ukve iTqva, glukozis katabolizmis pentozofosfaturi gza
sxvadasxva intensivobiT mimdinareobs sxvadasxva qsovilebSi da
damokidebulia am qsovilebSi cximebis gardaqmnaTa Taviseburebebze.
eriTrocitebSi, magaliTad, NADPH membranebSi Semaval ujer cximovan mJavebs
Jangbadis zemoqmedebidan icavs. amitom iseTi memkvidreobiTi daavadebebis
dros, rodesac genetikuri defeqti pentozofosfaturi gzis fermentebSi
gvxvdeba, adgili aqvs eriTrocitebis membranebis destruqcias, maT hemolizs.

eleqtronTa gadatana da JangviTi fosforilireba

JangviTi-fosforilireba aerobul ujredebSi sunTqvis terminalur

stadias warmoadgens. naxSirwylebis, aminomJavebis da cximovani mJavebis
daSlis produqtebi, sabolood sunTqviT jaWvSi katabolizirdebian. am
procesSi eleqtronebi organuli substratebidan Jangbadze gadaitanebian da
gamoTavisuflebuli energia gamoiyeneba AATP-is sinTezisaTvis.
limonmJavas ciklis erTi bruniT Semotrialebisas specifiuri
dehidrogenazebi iwveven wyvili wyalbadis atomebis amoRebas izocitratidan,
-ketoglutaratidan, suqcinatidan da malatidan. wyalbadis atomebi
eleqtronebs gadascemen eleqtronTa gadacemis jaWvs, xolo protonebi (H+-
ionis formiT) gadadian Sidamitoqondrialur sivrceSi eleqtronebi, jaWvis
saSualebiT gadaecema jer garkveul naerTebs, xolo Semdeg - Jangbads,
romelic aerobul ujredebSi eleqtronTa saboloo aqceptors warmoadgens.
ikavSirebs ra Jangbadis atomi eleqtronebis wyvils, ierTebs agreTve H+ -
ionebs, romlebic wina stadiebze gamoTavisufldnen jer dehidrogenazebis,
xolo Semdeg sunTqviTi jaWvis mier. reaqciis Sedegad miiReba wylis (H2O)
molekula

70
 sunTqviTi jaWvi

eleqtronebis gadacema sunTqviT jaWvSi warmoebs. es ukanaskneli rigi

cilebisagan Sedgeba, romlebsac mWidrod dakavSirebuli prostetuli jgufi
aqvT; maT eleqtronebis mierTeba da gacema SeuZliaT. sunTqviT jaWvSi cilebi
garkveuli TanmimdevrobiT arian ganlagebulni. yoveli maTgani eleqtronebs
winamdebare cilidan ierTebs da gadascems momdevno cilas. sawyis etapze
eleqtronebs didi Tavisufali energia aqvT, romelic jaWvSi moZraobisas
mcirdeba. am energiis mniSvnelovani nawili Seinaxeba ATP-is molekulaSi.
yoveli ori eleqtronis gadacemisas 3 molekula ATP sinTezirdeba. ATP-is
sinTezi sunTqviTi jaWvis specialur ubnebSi (3 ubani) warmoebs da maT
fosforilirebis ubnebi ewodebaT.
s u n T q v i T i j a W v i

NADP, -ketoglutarati, izocitrati piruvati, malati da sxva.

NADH

NADH-dehidrogenaza
(FMN, Fe-S) ADP
ATP
ubiqinoni

citoqromi b ADP
ATP
citoqromi c1

citoqromi c

citoqromi aa3
(citoqromoqsidaza) ADP
A ATP
O2

71
 specifiuri dehidrogenazebi, romlebic piruvatisa da sxva organuli
naerTebis daJangvas axorcieleben mitoqondrialur matriqsSi arian
ganlagebulni. matriqsi garSemortymulia ori tipis: gare- da
Sidamitoqondrialuri membranebiT. am ukanasknelze lokalizebulia
eleqtronTa gadamtani jaWvi da fermentebi, romlebic ATP-is sinTezs
awarmoeben. garemitoqondrialuri membrana ionebis da metabolitebis
umravlesobas advilad atarebs, xolo Sida ki - praqtikulad gaumtaria.
nivTierebebi Sidamitoqondrialur membranas mxolod specialuri
transportuli sistemebis saSualebiT gadian.
cnobilia, rom yoveli naerTi, romelsac eleqtronis gacemis an mierTebis
unari aqvs xasiaTdeba Jangva-aRdgeniTi potencialiT, da mas standartuli
Jangva-aRdgeniTi potenciali ewodeba. es sidide voltebSi izomeba. sunTqviTi
jaWvi iwyeba NADH-iT, romelsac momdevno naerTebTan SedarebiT, ufro
maRali Jangva-aRdgeniTi potenciali aqvs da igi tolia - 0,32 V. amitom
pirvel stadiaze NADH gadascems 2e- da ijangeba: NADH NAD+ + H+ + 2e- [aseT
naerTebSi (-) niSani imaze miuTiTebs, rom mocemul nivTierebas axasiaTebs
dabali Tvisoba (swrafva) eleqtronis mimarT. rac ufro uaryofiTia Jangva-
aRdgeniTi potenciali, miT ufro sustia am naerTis swrafva eleqtronebis
mimarT da miT ufro advilad kargavs igi eleqtronebs]. jaWvis saboloo
stadiaze myofi Jangbadis Jangva-aRdgeniTi potenciali tolia + 0,82 V da igi
Tavisuflad ierTebs eleqtronebs:

1/2 O2+ 2H++ 2 e- H2O

eleqtronebis pirvel donorsa da saboloo aqceptors Soris sunTqviTi jaWvis

arseboba aucilebelia eleqtronTa energiis TandaTanobiTi SemcirebisTvis,
raTa igi gamoyenebul iyos ATP-is sinTezSi. sunTqviT jaWvSi eleqtronTa
energiis SedarebiT mniSvnelovani dacema sam punqtSi xdeba. potencialis I
vardnas adgili aqvs maSin, rodesac NADH-dehidrogenazidan eleqtroni
gadaecema ubiqinons, meored - eleqtronis energia Tavisufldeba misi
citoqrom b-dan citoqrom c1-ze gadasvlisas da mesamed - citoqrom a-dan
Jangbadze gadatanisas. samive ubanze warmoebs energiis Senaxva ATP-is
molekulaSi da amitom am ubnebs, xSirad, fosforilirebis punqtebs
uwodeben.

72
 eleqtronTa wyvilebis umravlesoba sunTqviT jaWvs NADH-is an NADPH-is
saSualebiT miewodeba. isini NAD-damokidebuli dehidrogenazebis kofermentebs
warmoadgenen. zogierTi maTgani Cvens mier ukve ganxilulo iyo glikolizisa
da limonmJavas ciklis aRwerisas. am fermentebis garda ujredSi sxva
dehidrogenazebic arseboben, romlebic analogiur reaqcias atareben: iwveven
substratis daJangvas e.i. misi molekulidan ori wyalbadis atomis amoRebas.

erTi wyalbadis atomi hidrid-ionis formiT (:H-) gadaitaneba NAD+ an NADP+-

ze, xolo meore ki Tavisufldeba protonis (H+) formiT. TviTeul hidrid-

ions gaaCnia ori aRdgeniTi ekvivalenti da misi mierTeba NAD-is molekulas
Semdegnairad scvlis:
H

H
CONH2 CONH2

+
N N

R Please Register R
NAD+ NADH

dehidrogenazebis umravlesobas wyalbadis atomebi gadaaqvT NAD+-is

molekulaze da amitom, SeiZleba iTqvas, rom es nukleotidi asrulebs
aRdgeniTi ekvivalentebis koleqtorul, Semagrovebel funqcias.
eleqtronTa gadatana sunTqviT jaWvze TviT dehodrogenazebis molekulaSi
iwyeba. am ferments, prostetuli jgufis saxiT, mWidrod dakavSirebuli
flavinmononukleotidi (FMN) aqvs, romlis SemadgenlobaSic Sedis
riboflavini. NADH-is ori aRdgeniTi ekvivalenti gadaecema FMN-s da miiReba
FMN-H2. garda FMN-isa NADH-dehidrogenaza Seicavs rkina-gogirdovan centrs
(Fe-S), romelic FMN-is Semdeg iRebs or aRdgeniT ekvivalents da maT
ubiqinons gadascems. swored am dros warmoebs aRdgeniTi potencialis
pirveli sagrZnobi dacema. ubiqinoni warmoadgens cximSi xsnad qinons,
romelsac grZeli izoprenoiduli gverdiTi jaWvi aqvs. misi aRdgeniTi
eqvivalenti, SemdgomSi, citoqromebs gadaecema.
citoqromebi rkinaSemcvel, wiTeli an yavisferi Seferilobis mqone cilebs
warmoadgenen. isini garkveuli TanmimdevrobiT moqmedeben da eleqtronebi
ubiqinonidan Jangbadze gadaqvT. citoqromebi hemproteinebs ganekuTvnebian, maT

73
SemadgenlobaSi Sedis rkinaporfirinis jgufi anu hemi. igi Tavisi
agebulebiT hemoglobinSi Semaval rkina-porfirinis kompleqss waagavs.
arsebobs citoqromebis sami klasi: a, b da c. TviTeul maTganSi Semavali
rkina daJangul formaSi [Fe(III)] imyofeba, romelic eleqtronebis mierTebis
Semdeg zeJangul formaSi [Fe(II)] gadadis. ubiqinonidan eleqtronebs ierTebs
citoqromi b, romelic maT Semdgom sxva citoqromebs gadascems: b c1 c
aa3. bolo stadiaze myof citoqroms, xSirad, citoqromoqsidazas uwodeben,
radgan mas eleqtronebi pirdapir Jangbadze gadaaqvs. citoqromebs sxvadasxva
Jangva-aRdgeniTi potenciali aqvT: yvelaze maRali - b-s, xolo dabali - aa3-s.
swored maTi potencialTa sxvaoba ganapirobebs eleqtronTa dinebas b-dan aa3-
mde.
sunTqviTi jaWvis bolo stadias, rogorc ukve iTqva, Jangbadis mier
eleqtronebis mierTeba warmoadgens. am dros Jangbadis molekula mTlianad
aRdgeba wylamde, risTvisac eleqtronTa ori wyvilia saWiro:

O2 + 4 H+ + 4 e- 2 H2O

im SemTxvevaSi, Tu moxda Jangbadis molekulis arasruli aRdgena da man

miierTa ara oTxi, aramed ori eleqtroni, warmoiqmneba wyalbadis zeJangi -
-
H2O2, xolo Tu erTi eleqtroni - warmoiqmneba superoqsiduri radikali (:O2 ).
orive naerTi ujredisTvis, da gansakuTrebiT membranebisTvis Zlier
toqsikuria, radganac moqmedeben fosfolipidebSi Semaval ujer cximovan
mJavebze, iwveven maT daJangvas da destruqcias. superoqsiduri radikali
mravali paTologiuri mdgomareobisas warmoiqmneba, magram mas
gansakuTrebuli mniSvneloba eniWeba anTebiTi procesebisas, romlis drosac
imunuri ujredebis mier producirebuli zeJanguri produqtebis raodenoba
maRal dones aRwevs. superoqsiduri radikalebis gauvnebelyofa ujredSi
ferment superoqsiddismutazis saSualebiT xdeba:

-
2O2 + 2H+ H2O2 + O2

sunTqviTi jaWvis Tanmimdevrobis dadgenaSi, misi calkeuli etapebis

garkvevaSi didi roli iTamaSes specifiurma inhibitorebma. bevr maTgans
SerCeviTi moqmedeba axasiaTebT. sunTqviTi jaWvis pirvel ubans amuxruWebs

74
rotenoni, erT-erTi Zlieri mcenareuli Sxami, meore ubans - toqsikuri
antibiotiki - antimicini A, xolo mesames kaliumis cianidi.

ATP-is sinTezi

rogorc zemoT avRniSSneT ATP-is sinTezi sunTqviTi jaWvis sam punqtSi

anu monakveTSi warmoebs. maT xSirad SeuRlebis punqtebsac uwodeben, radgan
am punqtebSi warmoebs eleqtronTa transportis SeuRleba fosforilirebis
procesTan, ATP-is sinTezTan. NADH-dan Jangbadamde erTi eleqtronuli
wyvilis gadatanisas Tavisufldeba daaxloebiT 52 kkal/moli siTbo,
romlisganac mxolod 22 kkal/moli Seinaxeba 3 molekula AATP-Si
(gavixsenoT, rom erTi molekula ATP-is sinTezisTvis ADP da araorganuli
fosfatidan saWiroa 7,3 kkal/moli). energiis danarCeni nawili mitoqondriebis
mier gamoiyeneba Ca, ATP-s, ADP-s da sxva naerTTa transportisTvis, agreTve
gamoiyofa siTbos saxiT.
ATP-is masinTezirebeli (ATP-sinTetaza) sistema Sidamitoqondrialur
membranaSia moTavsebuli. igi rTul membranul kompleqss warmoqmnis da
Sedgeba ori tipis cilebisagan. erTi maTgani, kerZod Fo, mTlianad CaZirulia
mitoqondrialur membranaSi (indeqsi o miuTiTebs fermentis am nawilis
mgrZnobiarobaze toqsikuri antibiotikis-oligomicinis mimarT), xolo meore,
F1, dakavSirebulia Fo-Tan da mimarTulia Sidamitoqondrialur sivrceSi.
orive komponenti warmoqmnis kompleqss, romelsac ATP-sinTetaza ewodeba.
miuxedavad imisa, rom ATP-sinTetazis agebuleba da misi suberTeulebis
fizikur-qimiuri Tvisebebi sakmaod kargad aris Seswavlili, am fermentis
moqmedebis molekuluri meqanizmebi jer-jerobiT gaurkvevelia. amitom,
sakiTxi imis Sesaxeb, Tu rogor xdeba eleqtronTa transportirebisas
gamoTavisuflebuli energiis gardaqmna ATP-Si Caweril qimiuri bmis enegiad,
Ria rCeba. Tanamedrove Sexedulebebis Tanaxmad, romlebic ingliseli
mecnieris mitCelis qemiosmotur hipoTezas eyrdnoba, eleqtronTa
transportis Tanadroulad mimdinareobs wyalbadis ionTa (H+) gadaqaCva
Sidamotoqondrialuri sivrcidan membranaTaSoris sivrceSi. H+-is aseTi
transportirebis Sedegad warmoiqmneba transmembranuli potenciali,
ganpirobebuli wyalbadis ionTa koncentraciuli gradientiT. swored am
gradientis energia (osmosuri energia) gamoiyeneba ATP-is sinTezisaTvis.

75
 mitCelis Teoriis Tanaxmad, sunTqvis jaWvSi eleqtronTa gadanacvlebis
dros gamoTavisuflebuli energia ixarjeba protonTa transportze maTi
koncentraciuli gradientis sawinaaRmdegod e.i. protonTa saqaCavis
muSaobaze. membranaTaSoris sivrceSi gadasuli protonebi, koncentraciuli
gradientis arsebobis gamo eswrafvian SigniT Sesvlas specialuri arxebis
saSualebiT. qemiosmosuri hipoTezis Tanaxmad es arxebi ganlagebulia ATP-
sinTetazur kompleqsSi, sadac gradientis energia gardaiqmneba ATP-is qimiur
energiad.
amrigad, 1 molekula glukozis biologiuri daJangvisas CO2-mde da H2O-mde,
glikolizis, limonmJava ciklis da sunTqviTi jaWvis gavliT miiReba 38
molekula ATP, rac biologiuri sistemebis ekonomiurobaze da efeqturobaze
migviTiTebs.

mikrosomuli daJangva

mikrosomebi warmoadgenen centrifugirebis Sedegad miRebul subujredul

fraqcias, romelSic warmodgenili arian endoplazmuri badis fragmentebi,
xSirad ribosomebiTurT. mikrosomebis membranebi Seicaven fermentebs,
romlebsac SeuZliaT awarmoon sxvadasxva tipis organuli naerTebis
daJangva, maT molekulaSi gangbadis atomis CarTva.
aseT fermentebs oqsigenazebi ewodebaT. miuxedavad imisa, rom ujredSi
arsebuli Jangbadis 90% mitoqondriebSi citoqromoqsidazis mier moixmareba,
mikrosomalur oqsigenazebs sakmaod mniSvnelovani funqcia akisriaT.
oqsigenazebis ori tipi arsebobs: mono- da dioqsigenazebi.
isini organul naerTebs, Sesabamisad erTi da or Jangbadis atoms
ukavSireben. monooqsigenazebs xSirad hidroqsilazebsac uwodeben, radgan
isini Jangbadis CarTvas hidroqsilis jgufis formiT axdenen.
monooqsigenazebi monawileobas iReben kateqolaminebis biosinTezSi.
oqsigenazebis moqmedebis Sedegad lipofiluri nivTiereba SedarebiT
polaruli da wyalSi advilad xsnadi xdeba. amitom am naerTis daSla da
misi organizmidan gamoyvana advildeba. amrigad, oqsigenazebis erT-erT
mniSvnelovan funqcias, maTi detoqsikaciuri moqmedeba warmoadgens.
mikrosomuli daJangvis ZiriTadi elementia citoqromi P-450, romelic mravali
organuli nivTierebebis hidroqsilirebas awarmoebs. detoqsikaciuri
daJangvis sistemebiT mdidaria RviZlis ujredebi.

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 O oqsidaciuri stresi

ujredSi gamudmebiT mimdinareobs Jangva-aRdgeniTi reaqciebi, romlebic

safuZvlad udevs qimiuri energiis warmoqmnas, rTuli cikluri aRnagobis mqone
nivTierebis biosinTezs da maT degradacias. yovel ujredul sistemas Tavisi,
misTvis damaxasiaTebeli Jangva-aRdgeniTi, e.w. redoqs-statusi aqvs, romelic
damokidebulia JangviTi da aRdgeniTi reaqciebis intensivobiT, daJanguli da
aRdgenili naerTebis TanafardobiT. ZiriTad nivTierebebs, romlebic ujredis
redoqs-statuss gansazRvraven arian wyvilebi: glutaTioni(daJanguli) da
glutaTioni(aRdgenili); NAD+(daJanguli) da NADH(aRdgenili); NADP(daJanguli)
da NADPH(aRdgenili). am wyvilebs Soris wamyvani mniSvneloba glutaTions
mieniWeba, romelic ujredis ZiriTad redoqs-bufers warmoadgens. Tu intensiuri
JangviTi reaqciebis Sedegad am sistemis SesaZlebloba amowurulia, ujredis
redoqs-statusi icvleba da sistema paTologiur mdgomareobaSi iwyebs gadasvlas.
ujredSi JangviTi procesebis ZiriTad generators TviT Jangbadi warmoadgens.
Jangbadis eleqtronuli konfiguracia isea awyobili, rom mas saSualeba eZleva
miierTos zedmeti eleqtronebi, ris gamoc igi Zlieri damJangveli xdeba.
mitoqondriaSi mimdinare sunTqviTi procesebis dros Jangbadis molekula
ierTebs ra wyalbadis 4 atoms (4 protons da 4 eleqtrons) wylamde aRdgeba. am
procesis kontroli substratebis saSualebiT xorcieldeba, da amitom wylis
warmoqmna limitirebulia fermentebis (dehidrogenazebis) kinetikuri
maxasiaTeblebiT. miuxedavad amisa, garkveuli tipis zemoqmedebiT, ZiriTadad
sunTqviTi jaWvis susti mimdinareobis dros, SesaZlebelia moxdes Jangbadis
arasruli aRdgena. Tu Jangbadis molekulam miierTa mxolod ori eleqtroni
warmoiqmneba wyalbadis zeJangi – H2O2, xolo Tu erTi eleqtroni –
superoqsiduri anionradikali (O2-). Jangbadis es naerTebi Zlier agresiuli arian
da azianeben ujredSi arsebul makromolekulebs.
sunTqviTi jaWvis normaluri msvleloba:
O2 + 4H+ + 4e - 2H2O (wyali)
sunTqviTi jaWvis Senelebuli msvleloba:
O2 + 2H+ + 4e - H2O2 (wyalbadis zeJangi)
sunTqviTi jaWvis damuxruWebuli msvleloba:
O2 + e - O2- (superoqsiduri anionradikali)

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 wyalbadis zeJangi SesaZlebelia warmoiqmnas agreTve monooqsigenazebis
moqmedebis Sedegadac. magaliTad, kateqolaminebis (adrenalinis, dofaminis,
noradrenalinis) damSleli fermentis-monooqsigenazis aqtivobis Sedegad miiReba
wyalbadis zeJangi. superoqsidanionradikali SesaZlebelia warmoiqmnas ferment
qsantinoqsidazis moqmedebiT, romelic purinuli nukleotidebis katabolizmis
saboloo produqts – qsantins gardaqmnis Sardis mJavad. am radikalis kidev erT
mZlavr wyaros warmoadgens citoqromi P-450, romelic qsenobiotikebis (adamianis
organizmSi moxvedrili ucxo nivTierebebis, da maT Soris eTilis spirtis)
daSlas ganapirobebs. superoqsidis generacias agreTve fermenti NADPH oqsidaza
awarmoebs. am fermentis gaaqtiveba ujredis hormonebiT zemoqmedebiT
mimdinareobs.
ujredi reaqciuli jangbadis gauvnebelyofisTvis rig fermentebs
gamoimuSavebs, romelTa Sorisac yvelaze mniSvnelovania superoqsiddismutaza.
misi zemoqmedebiT superoqsiduri anion-radikali SedarebiT sust wyalbadis
zeJangad gadaiyvaneba:

2 O2- + 2 H+ H2O2 + O2

warmoqmnili wyalbadis zeJangi ferment katalazas saSualebiT neitralizdeba:

2H2O2 2 H2O + O2

am reaqciis garda wyalbadis zeJangis gauvnebelyofa ujredis ZiriTadi

aRmdgeneli komponentis – glutaTionis saSualebiT mimdinareobs.
aseTi mZlavri dacviTi antioqsidanturi meqanizmebis arsebobis gamo
reaqciuli Jangbadis Semcveli naerTebi mitoqondriebSi SedarebiT mcire
raodenobiT warmoiqmneba. magram, sakmarisia sakvebis deficitis, radiaciis,
toqsinebis, an sxva paTogenuri zemoqmedebis Sedegad sunTqviTi jaWvis
msvleloba Seferxdes adgili aqvs Jangbadis arasrul aRdgenas, ris gamoc didi
raodenobiT warmoiqmneba Jangbadis Semcveli Tavisufali radikalebi. isini
upirveles yovlisa TviT mitoqondriebs azianeben, riTic kidev ufro amZimeben
darRveul metabolizms. bunebrivia, rom Tu gamauvnebelyofeli sistemebis
moqmedeba daqveiTebulia, mitoqondriebis dazianeba gardauvalia. amaze miuTiTebs
Mmravali tipis neirodegeneraciuli paTologiebis dros mimdinare molekuluri
procesebis analizi. magaliTad, amiotropuli lateraluri sklerozis ojaxuri
forma viTardeba Ca/Zn superoqsiddismutazis genSi momxdari mutaciis Sedegad.

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katalazur da glutaTioperoqsidazur (glutaTionis aRmdgeneli fermenti)
sistemebSi momxdari cvlilebebi SesaZlebelia gaxdes alcheimeris,
hantingtonis, parkinsonis daavadebebis, spinocereberuli ataqsiis da sxva
neiropaTologiebis ganviTarebis mizezi.
ujredis intoqsikaciis, daqveiTebuli metabolizmis da Rrma iSemiuri
procesebis mimdinareobis fonze superoqsiduri anionradikalidan warmoiqmneba
hidroqsilis radikali (OH.). am naerTis JangviTi potenciali gacilebiT aRemateba
superoqsidanionis JangviT potencials, ris gamoc igi intensiurad azianebs
biologiur makromolekulebs. hidroqsilis radikali miiReba fentonis reaqciiT
rkinis Tanaobisas:

H2O2 + Fe2+ OH. + OH- + Fe3+

dadgenilia, rom ujredis nekrozuli gadagvareba iwyeba hidroqsilis radikalis

warmoqmnis Semdeg, ris gamoc am naerTis metabolizmi gansakuTrebul yuradRebas
ipyrobs. aqve unda aRiniSnos, rom mitoqondriebi sakmaod didi raodenobiT
rkinas Seicaven citoqromebSi da NADH-dehidrogenazebSi Semavali kompleqsebis
saxiT. am cilovani molekulebis swrafi daSla an maTi susti biosinTezi
Tavisufali rkinis Warbi raodenobiT warmoqmnis mizezi xdeba, rac hidroqsilis
radikalis formirebis winapirobas warmoadgens. amitom, Tavisufali rkinis
acilebas, rogorc mitoqondriebidan, ise ujredis sxva regionebidan
damcvelobiTi mniSvneloba aqvs. mTavar cilas, romlis saSualebiTac
Tavisufali rkinis helatireba, misi dakavSireba, deponireba, da Sesabamisad
gauvnebelyofa xdeba – feritini warmoadgens.
hidroqsilis radikalis erT-erT mZlavr wyaros ujredSi azotis oqsidi
(NO) warmoadgens. rogorc cnobilia, azotis oqsidi e.w. “ujreduli Suamavlebis”
sistemas miekuTvneba anu ujredis zogierTi hormoniT gaRizianebis SemTxvevaSi
sinTezirdeba, gamodis ujredidan da axlomdebare ujredebze iwyebs moqmedebas.
azotis oqsidi fiziologiurad aqtiuri naerTebis ricxvs miekuTvneba da mis
moqmedebis yvelaze mkafio gamoxatulebas sisxlZarRvTa gafarToeba –
vazodilatacia warmoadgens. dasaSvebia, rom azotis oqsidis warmoqmnas ujredSi
kompensaciuri xasiaTi hqondes, radgan misi intensiuri sinTezi iSemiuri
procesebis dros mimdinareobs. am dros ujredSi Ca-is ionebi grovdeba,
romlebic NO-s masinTezirebeli fermentis NO-sinTazis aqtivatorebs
warmoadgenen. es ukanaskneli kalciumis Tanaobisas aminomJava argininidan iwyebs
NO-s warmoqmnas.

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 azotis oqsidi TviT warmoadgens anionradikals, magram misi redoqs-
potenciali SedarebiT sustia. superoqsiduri anionradikalis Tanaobisas igi
jer peroqsinitrilis radikals, xolo Semdgom hidroqsilis radikals
warmoqmnis:

NO. + O2- ONOO-

ONOO- + H+ ONOOH
ONOOH OH. + NO2.

am reaqciis gamo, azotis oqsidi, romelic fiziologiur koncentraciebSi

normalur metabolur reaqciebSi iRebs monawileobas, maRal koncentraciebSi
toqsikuri xdeba da ujreduli komponentebis ngrevas ganapirobebs. magaliTad
makrofagebi, ucxo ujredis destruqcias azotis oqsidis da
superoqsidanionradikalis sekreciiT iwveven. amitom, azotis oqsidis didi
raodenobiT warmosaqmnelad imunur ujredebs damatebiT NO-sinTazis
inducirebadi geni aqvT.
ujredis JangviTi potencialis matebas, misi homeostazis wanacvlebas
JangviTi procesebisken “oqsidaciuri stresi” ewodeba. JangviT da aRmdgenel
sistemebSi arsebuli balansis darRveva SesaZlebelia ganpirobebuli iyos
reaqciuli Jangbadis intensiuri warmoqmniT an antioqsidaciuri sistemebis
deficitiT. sxvadasxva tipis egzogenurma stresma SesaZlebelia ujredSi
oqsidaciuri talRa gamoiwvios. magaliTad, radiaciis zegavleniT mimdinare
wylis hidrolizi warmoqmnis hidroqsilis radikals. Jangbadis moxmarebis mateba
(hiperoqsia) aseve iwvevs didi raodenobiT zeJanguri produqtebis warmoqmnas.
makrofagebSi, rogorc ukve iTqva, oqsidaciuri talRa imunuri pasuxis Sedegad
viTardeba. sxvadasxva tipis qsenobiotikebma, gansakuTrebiT antibiotikebma,
citoqromi P-450 (CYP-is) gaaqtivebis gamo aseve SesaZlebelia oqsidaciuri stresi
gamoiwvion.
im SemTxvevaSi, Tu oqsidaciuris stresi ujredis sacicocxlo funqciebze
moqmedebs, magaliTad dnm-is molekulaSi iwvevs cvlilebebs, igi Seuqcevadi, da
amitom sasikvdilo xdeba; xolo im SemTxvevaSi, Tu oqsidaciuri stresi xanmokle
xasiaTs atarebs, igi gardamavalia da Seqcevad procesad gvevlineba. amitom
antioqsidanturi sistemebis xangrZlivi daqveiTeba an qronikuli anTebiTi
procesebi ujredis iZulebiT sikvdils – nekrozs ganapirobeben.

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 cximovani mJavebis daJangva

triacilglicerinebi anu trigliceridebi cxovelTa organizmisTvis erT-

erTi kaloriuli sakvebia (maTi daJangvis Sedegad 9 kkal/molze energia
gamoiyofa). isini, agreTve, organizmis ZiriTad samarago nivTierebebs
warmoadgenen da didi raodenobiT grovdebian sxvadasxva tipis qsovilebSi.
RviZlSi, gulisa da ConCxis kunTebSi mosvenebis dros triacilglicerinebi
saWiro energiis TiTqmis naxevars iZlevian.
triacilglicerinebis energiis 95% moTavsebulia cximovan mJavaTa sam
naSTSi da mxolod 5% - glicerinis molekulaSi. Aam naerTTa daSla
mitoqondriebSi mimdinareobs da maTi daSlis produqtebi, glukozis daSlis
produqtebis msgavsad, limonmJavas ciklSi erTvian.
cximovan qsovilebSi trigliceridebis daSla hormon-damokidebuli
procesia. Ffermenti trigliceridlipaza, romelic triglicerididan
digliceridsa da cximovan mJavas warmoqmnis, fofsforilirebiT aqtivirdeba. Mmis
fosforilirebas cAMP-damokidebuli proteinkinaza anxorcielebs, romelic
Tavis mxriv cAMP-iT stimulirdeba: es ukanaskneli ki, ujredSi hormonis
zegavleniT warmoiqmneba. mTliani lipolizis (romelSic triglicerildlipazis
garda monawileobas iReben mono- da diglicerildlipazebic) Sedegad
warmoqmnili cximovani mJavebi sisxlis saSualebiT sxva organoebTan
gadaitanebian.

cximovani mJavebis transporti mitoqondriebSi

ujredis citoplazmaSi cximovani mJavebi SesaZlebelia ori gziT moxvdnen.

Pirvel SemTxvevaSi, isini transportirdebian sisxlis albuminis saSualebiT
ujredamde, scildebian mas, Tavisuflad gadian plazmur membranas da Sedian
citoplazmaSi. Mmeore SemTxvevaSi cximovani mJavebis wyaros warmoadgenen TviT
citoplazmaSi arsebuli triacilglicerinebi, romlebic ferment lipazas
saSualebiT iSlebian.
Citoplazmidan cximovani mJavebi mitoqondrialur matriqsSi mxolod
samstadiani gardaqmnis Semdeg xvdebian. Ppirvel stadiaze maT, acetil-CoA-
sinTetazuri reaqciis Sedegad, CoA uerTdeba:

RCOOH + CoA-SH R-CO-S-CoA + AMP + Ppi

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 warmoqmnili naerTi, acil-CoA, romelic cximovani mJavis gaaqtivebul formas
warmoadgens, mitoqondrialur membranaSi ver gadis da amitom cximovan mJavas
CoA-s nacvlad uerTdeba karnitini - naerTi, romelic aminomJava lizinis
gardaqmnis produqts warmoadgens:

acil-CoA+karnitini —› acil-karnitini + CoA

cximovan mJavasTan warmoqmnili karnitinis eTeri, romelsac acil-karnitini

ewodeba, advilad gadis mitoqondrialur membranas da mitoqondriaSi kvlav
Cainacvlebs CoA-Ti:

acil-karnitini + CoA —› acil-CoA + karnitini

Sidamitoqondrialur sivrceSi warmoqmnili acil-CoA ukve ganicdis daSlas.

cximovani mJavebis aseTi rTuli gadatanis reaqciebi, romlebsac sami
fermenti emsaxureba, saWiroa imisTvis, rom gaiTiSos erTmaneTisgan CoA-is ori
fondi (puli): citoplazmuri fondi, romelic ZiriTadad sinTezur reaqciebs (mag.
cximovani mJavebis sinTezs) emsaxureba da mitoqondriuli CoA-is fondi, romlis
saSualebiTac JangviTi degragaciis - daSlis reaqciebi (mag. piruvatis, cximovani
mJavebis, aminomJavebis) mimdinareoben.
Aacilkarnitinis warmomqmneli fermenti, karnitinacetiltransferaza
regulatorul ferments warmoadgens. igi gansazRvravs cximovani mJavebis
transports da Sesabamisad, maTi daJangvis intensiobas mitoqondriebSi

cximovani mJavebis daJangva mitoqondriebSi

cximovani mJavebis daJangva or stadias moicavs: pirvel stadiaze maT

ornaxSirbadiani fragmenti - acetil-CoA - CamoeWreba. TiToeuli aseTi cikli
oTxfermentiani reaqciisagan Sedgeba da saboloo produqts ori naxSirbadiT
Semoklebuli cximovani mJavis fragmenti warmoadgens. magaliTad, 16-naxSirbadiani
palmitinis mJavis Semoklebisas miiReba 14-naxSirbadiani miristinis mJava.
darCenili fragmenti kvlav mokldeba ori naxSirbadiT da es procesi grZeldeba
manamde, sanam ar warmoiqmneba 2-naxSirbadiani ZmarmJavas naSTi:

C16-palmitinis mJava —› C14-miristinis mJava —› C12-laurilis mJava —› ……….—›

—› C2-ZmarmJava

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 daJangvis am reaqciebSi monawileobas Rebuloben mxolod cximovani mJavebis
CoA eTerebi. Aamitom cximovani mJavebis daSlis I stadiis saboloo reaqcia aseT
saxes miiRebs (palmitinis mJavis daSlis SemTxvevaSi):

acetil-CoA acetil-CoA acetil-CoA

palmitil-CoA — —› miristil-CoA — —› lauril-CoA — —›……..—› acetil-CoA

e.i. erTi molekula 16-naxSirbadiani palmitinis mJavidan miiReba rva 2-

naxSirbadiani ZmarmJavas naSTi.
rogorc ukve iTqva, TiToeul aseT reaqcias axorcielebs oTxi fermenti,
romelTagan ori dehidrogenazas warmoadgens. maTi moqmedebis Sedegad aRdgeba 1
NAD+-i NADH-mde da 1 FAD-i FADH2-mde. Ees ukanasknelni, gadascemen ra
eleqtronebs sunTqviT jaWvs, ATP-is sinTezs iwveven. Aadvili gamosaTvlelia, rom
1 molekula palmitinis mJavis daSlis Sedegad miiReba 8 molekula acetil-CoA
da 35 molekula ATP.
Cvens mier aRweril reaqciaTa Tanmimdevrobas adgili aqvs im SemTxvevebSi,.
Tu cximovani mJava luwi raodenobis naxSirwyalbadis naSTebisagan Sedgeba. kenti
naxSirbadiani cximovani mJavebis daJangvis saboloo stadiaze warmoiqmneba 3-
naxSirbadiani propionil-CoA, romelic jer gardaiqmneba meTilmalonil-CoA-T,
xolo Semdeg – suqcinil-CoA-T. es ukanaskneli krebsis ciklis metabolits
warmoadgens da am ciklis saSualebiT agrZelebs daJangvas.

Ppropionil-CoA —› meTilmalonil-CoA —› suqcinil-CoA

meTilmalonil-CoA-s suqcinil-CoA-d gardaqmnis fermenti

meTilmalonilmutaza, romelic am reaqciisTvis kofermentad iyenebs vitamin B12-
is
warmoebuls – dezoqsiadenozilciankobalamins. vitamini B12 erTaderTi vitaminia,
romlis SemadgenlobaSic Sedis metali - kobalti. igi sakmaod rTuli
aRnagobisaa da mis centralur birTvs porfirinis msgavsi struqtura Seadgens.
vitamini B12 mxolod mikroorganizmebSi sinTezirdeba. Mmisi ukmarisoba
adamianebSi iwvevs avTvisebiani, makrocituli (megaloblasturi) anemiis
ganviTarebas.
meTilmalonil-CoA manawileobas iRebs agreTve aminomJavebis - meTioninis,

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valinis da izoleicinis cvlaSi, e.i. reaqciebSi, romlebSic meTilis jgufis
gadatana an Canacvleba xdeba.

COOH COOH
CH CH3 CH2
C O CH 2
S Register
Please CoA CO Register
Please S CoA
meTilmalonil-CoA suqcinil-CoA

amrigad, cximovani mJavebi, romlebic luwi raodenoba naxSirbadis

atomebisagan Sedgebian, iSlebian acetil-CoA-mde, xolo kenti-CH2-raodenobis
mqone cximovani mJavebi ki - acetil-CoA-mde da propionil-CoA-mde. es ukanaskneli
warmoqmnis jer meTilmalonil-CoA-s, xolo Semdeg suqcinil-CoA-s, romelic
uSualod iJangeba krebsis ciklSi. ujeri cximovani mJavebi, ori damatebiTi
fermentuli reaqciis saSualebiT, gardaiqmnebian jer najerad, xolo Semdgom
iSlebian aRwerili reaqciebis meSveobiT.
cximovani mJavebis daJangvis meore stadiaze acetil-CoA iSleba limonmJavas
ciklis saSualebiT da am dros gamoyofili energia ATP-is formiT Seinaxeba. 1
molekula 16-naxSirbadiani palmitinis mJavis daJangvisas 131 molekula ATP-i
sinTezirdeba ADP-dan, rac gacilebiT aRemateba sxva produqtebidan miRebul
energias (gavixsenoT, rom 1 molekula glukozis daJangvisas miiReba mxolod 38
molekula ATP).

Kketonuri sxeulebi da maTi metabolizmi

RviZlSi, cximovani mJavebis daJangvis Sedegad miRebuli acetil-CoA

SesaZlebelia ori gziT gardaiqmnas: daiSalos limonmJava ciklis saSualebiT, an
warmoqmnas acetoacetati, oqsibutirati da acetoni. es ukanasknelni, ketonuri
sxeulebis saxelwodebiT arian cnobilni.

Aacetil-CoA + acetil-CoA acetoacetil-CoA + CoA

acetil-CoA + acetil-CoA acetoacetati + 2CoA
acetoacetati + H+ acetoni + CO2
acetoacetati + NADH + H+ oqsibuTirati + NAD+

ketonuri sxeulebis warmoqmnis biologiuri arsi mdgomareobs imaSi, rom

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organizmi Warbi raodenobis energetikul masalas ekonomiurad iyenebs da
Tanabrad anawilebs mas periferiul organoebSi. Aaq isini bolomde, CO2-mde da
H2O-mde iJangebian, riTac xels uwyoben am organoebis kvebas.
normaSi ketonuri sxeulebis raodenoba sisxlSi mcirea. Ddiabetis an
SimSilis SemTxvevebSi maTi koncentracia Zlier izrdeba. Ees ganpirobebulia
imiT, rom am SemTxvevaSi, cximovani mJavebis daJangvis siCqare sWarbobs ketonuri
sxeulebis moxmarebis siCqares periferiuli organoebis mier. glukozis
SeuTviseblobis (diabeti) an misi simciris (SimSili) gamo, da energetikuli
deficitis Sesavsebad, RviZlSi cximovani mJavebi intensiurad iJangebian. ketonuri
sxeulebis Warbi raodenobiT SeTviseba ki periferiul organoebs ar SeuZliaT.

aminomJavebis daJangva da Sardovanas cikli

adamianis organizmSi metaboluri energiis udidesi nawili (90%)

trigliceridebisa da naxSirwylebis daJangvis Sedegad gamomuSavdeba. Eenergiis
danarCeni (10-15% racionis mixedviT) miiReba aminomJavaTa daJangvis dros. Ees
ukanaskneli Zlierdeba maSin, rodesac racioni mxolod cilovani sakvebisagan
Sedgeba. Ggarda amisa, cilebis daSlis procesi SesaZlebelia gaZlierdes
SimSilisa da garkveuli paTologiebis SemTxvevebSic, rodesac cximovani da
naxSirwylovani sakvebis SeTviseba organizmis mier ver xerxdeba.

transaminazebi

endogenuri da egzogenuri cilebis daSlis produqtebs aminomJavebi

warmoadgenen. aminomJavaTa daSla, maTTvis damaxasiaTebeli aminojgufis
CamocilebiT iwyeba. am process JangviTi dezaminireba ewodeba. Tu es
aminojgufebi cilebisa da aminomJavebis Semdgomi sinTezisaTvis ar gamoiyenebian,
isini Tavs erT sabiloo produqtSi iyrian da am produqtis saxiT gamoiyofian
organizmidan. adamiansa da cxovelebis umravlesobaSi aseT produqts Sardovana
warmoadgens.
aminomJavebisagan -aminojgufis Camocilebas axorcieleben fermentebi,
romlebsac transaminazebi anu aminotransferazebi ewodebaT. transaminazul
reaqciebSi aminomJavebis -aminojgufi gadaitaneba -ketoglutaris mJavis -
naxSirbadis atomze, ris Sedegadac miiReba sawyisi aminomJavis ketoanalogi da
aminomJava – L-glutaminis mJava:

85
 COOH COOH
COOH C O CHNH2
COOH
CH2 CH 2
R CH
CH 2 R C O
CH2
NH 2
COOH COOH
Please Register
L- aminomJava -ketoglutaris ketomJava L-glutaminis
mJava mJava

magaliTad, alaninis gadaaminirebisas warmoiqmneba ketomJava, piroyurZnis mJava

(piruvati), asparaginis mJavisagan – MmJaunZmarmJava, Tirozinisagan –
hidroqsifenilpiruvati da a.S.
transaminazuli reaqciebis Sedegad aminomJavaTa aminojgufebi erTiandebian
glutamatSi (glutaminis mJavaSi). es aminomJava aminis jgufebis koleqtors
warmoadgens. amrigad, cilebSi monawile ocive aminomJavis katabolizmi mTavrdeba
erT aminomJavaSi, glutamatSi, romelic maTi gadaaminirebis saboloo produqts
warmoadgens.
transaminazebs ganasxaveben substratis – sawyisi aminomJavis mixedviT.
magaliTad, alaninis gadaaminirebas -ketoglutaratze awarmoebs fermenti-
alanintransaminaza anu alaninaminotransferaza, aspartatisa (asparaginis
mJavaisa) – aspartatransaminoza anu aspartataminotransferaza a.S.

transaminazebis prostetul jgufs piridoqsalfosfati – vitamini B6-is

warmoebuli – warmoadgens. piridoqsalfosfati mWidrod aris dakavSirebuli

fermentis molekulasTan. am naerTis saSualebiT xdeba fermentis mier
donoruli aminomJavidan aminojgufis mierTeba da misi -ketoglutaratze
gadatana. transaminazuli reaqciebis Sedegad gamoTavisuflebuli ketomJavebi, an
iSlebian limonmJavas ciklSi, (mag. piruvati, oqsaloacetati) an monawileobas
iReben sxva biosinTezur reaqciebSi.
transaminazebi Zlier da swraf, didi brunvaTa ricxvis mqone fermentebs
warmoadgenen da amitom isini warmatebiT gamoiyenebian daavadebaTa
diagnostikisaTvis. magaliTad, alaninaminotransferaza da
aspartataminotrasferaza ganisazRvreba gulisa da RviZlis paTologiebis dros.
rodesac trombozebisa Tu adgilobrivi anoqsiis Sedegad warmoiqmneba
koronaruli infarqtis kera, miokardiumis dazianebuli ujredebidan sxvadasxva
fermentebis da maT Soris transaminazebis gadasvla xdeba sisxlSi; amitom, maTi

86
aqtivobis gansazRvra sisxlis SratSi miokardiumis infarqtis erT-erTi
informaciuli sadiagnostiko saSualebaa da metyvelebs am paTologiis xarisxze
da siRrmeze. am fermentebis garda aRniSnul SemTxvevebSi xSirad isazRvrebian
kreatinkinaza da laqtatdehidrogenaza (H4-izofermenti).
transaminazebis gansazRvras didi mniSvneloba aqvs hepatotoqsikologiur
analizSi da mas iyeneben zogierTi profdaavadebaTa da intoqsikaciebis
diagnostikaSi. mag. maTi saSualebiT warmoebs organul gamxsnelebTan momuSave
personalis profilaqtikuri gamokvleva. RviZlis ujredebi mdidaria
transaminazebiT da maTi sisxlSi gadasvla metyvelebs hepatotoqsikuri
nivTierebebis Sedegad gamowveul RviZlis qsovilis degeneraciul movlenebze.
Aamrigad, transaminazebis moqmedebis Segegad warmoebs aminomJavebis -
aminojgufis gadatana -ketoglutaratze, romlis drosac glutaminis mJava
warmoiqmneba.

glutamatdehidrogenaza

aminomJavebis gadaaminirebis Sedegad miRebuli glutaminis mJava Semdgom

reaqciebSi aTavisuflebs aminis jgufs amoniumis ionis (NH4+) saxiT. glutaminis
mJava, im erTaderT aminomJavas warmoadgens, romelzec aseTi tipis reaqciis
Catareba aris SesaZlebeli. misi es Tviseba ganpirobebulia aminuri jgufebis
koleqtoruli funqciiT, razec zemoT iyo msjeloba. glutaminis mJavidan
amoniakis gamoTavisuflebas fermenti glutamatdehidrogenaza awarmoebs:

glutamati + NAD+ + H2O -getoglutaris mJava + NADH + H+ + NH4+

es fermenti mxolod mitoqondrialur matriqsSia lokalizebuli da mas

rTuli, suberTeulebisagan Semdgari struqtura aqvs. mas aloqsteruli
regulacia axasiaTebs: igi aqtivdeba ADP-iT da inhibirdeba GTP-iT im
SemTxvevebSi, rodesac ujredebi (magaliTad, RviZlis) energetikul deficits
ganicdian da Tu maTSi ADP-is koncentracia maRalia, fermenti aqtivdeba da
iwyebs glutaminis mJavis intensiur daSlas. warmoqmnili -ketoglutarati
limonmJavas ciklSi iJangeba da avsebs energetikul deficits, xolo amoniaki –
organizmis mier gamoidevneba da piriqiT, Tu GTP-is koncentracia maRalia (GTP
krebsis ciklSi monawile erT-erTi fermentis uSualo produqts warmoadgens), da
Tu krebsis cikli intensiurad mimdinareobs, maSin glutamatdehidrogenazuli

87
reaqcia muxruWdeba da cikls mcire raodenobiT -ketoglutarati miewodeba.
JangviTi dezaminirebis reaqciebi maRali siZlieriT RviZlSi mimdinareobs,
radgan es ukanaskneli, amoniakis organizmidan gamodevnis ZiriTad organos
warmoadgens. miuxedavad amisa, glutamatdehidrogenazuli reaqcia, RviZlis garda
metnaklebad, yvela tipis ujredSia warmodgenili.
amoniaki toqsikur nivTierebas warmoadgens. miuxedavad imisa, rom
neitralur pH-ze (pH 7,4) amoniaki TiTqmis mTlianad warmodgenilia amoniumis
ionebiT (NH4+), romlebic sustad gadian plazmur membranebSi da amitom ver
moqmedeben ujredis mgrZnobiare struqturebze. neitraluri amoniaki (NH3),
romelic mxolod 1% Seadgens – membranebSi Tavisuflad ganvladi, da
Sesabamisad, Zlier toqsikuria. gansakuTrebul mgrZnobiarobas neitraluri
amoniakis mimarT amJRavneben tvinis ujredebi. mis mcire koncentraciamac ki
SeiZleba gamoiwvios koma. amoniakis aseTi toqsikuri moqmedeba gamowveulia
glutamatdehidrogenazuli reaqciis SemobrunebiT glutamatis sinTezisaken.
Gglutamati tvinSi mTavar aRmgznebel neirogadamcems warmoadgens da misi zrda
nervul struqturebSi aRgznebadobis matebas iwvevs.
toqsikuri amoniakis acilebas tvini da zogierTi sxva periferiuli qsovili
glutaminis saSualebiT axdens. am organoebSi, TviT glutaminis mJava ikavSirebs
amoniaks da warmoqmnis glutamins. Ees ukanaskneli, glutaminis mJavis amidirebul
modifikacias warmoadgens. igi sisxlis saSualebiT gadaitaneba RviZlSi, sadac
isev iSleba glutaminis mJavad da amoniakad.

COOH
CHNH2
CH2
CH2

PleaseCONH
Register
2

glutamini

intensiurad momuSave ConCxis kunTebidan amoniaki RviZls aminomJava

alaninis saxiT gadaecema. kunTebSi gamomuSavebuli glutaminis mJava
transaminirebis reaqciis Sedegad gadascems aminis jgufs piruvats. es
ukanaskneli, kunTebSi intensiurad mimdinare glikolizis gamo aq didi
raodenobiT grovdeba. piruvatis aminirebis Sedegad warmoqmnili alanini,
sisxlis saSualebiT gadaecema RviZls, sadac isev transaminazuli reaqciis
Sedegad viRebT glutaminis mJavas. am process did mniSvneloba eniWeba qsovilTa

88
koordinirebuli muSaobisTvis, radgan, jer erTi, intensiurad momuSave
kunTebidan xdeba amoniakis CamoSoreba, da meorec - RviZlis ujredebs glukozis
resinTezisTvis piroyurZnis mJava miewodeba.
Aamrigad, qsovilebSi cilebis katabolizmis Sedegad warmoqmnili
toqsikuri amoniaki inertuli aminomJavebis (glutamini, alanini) saSualebiT
gadaecema RviZls, sadac transaminazuli reaqciebiT kvlav warmoiqmneba
glutaminis mJava

Sardovanas cikli

cxovelur samyaroSi aminuri azotis gamoyofa sami naerTis saSualebiT

xdeba: Tavisufali amoniakis, Sardis mJavasa da Sardovanas saxiT. organizmebs,
romlebic aminur azots amoniakis saxiT gamoyofen, amonioteluri organizmebi
ewodebaT, xolo iseTebs romlebic Sardis mJavas aTavisufleben – urikoteluri
organizmebi. pirvelebis tipiuri warmomadgenlebi Tevzebi arian, xolo meoresi –
frinvelebi da qvewarmavlebi. ZuZumwovrebis umravlesoba da adamiani ureotelur
organizmebs ganekuTvnebian, radgan aminur azots Sardovanas saxiT gamoyofen.
Sardovana warmoiqmneba cikluri procesis Segegad, romelic ZiriTadad
RviZlSi mimdinareobs.

NH2 CO NH2 NH2 CO H2PO3

Sardovana karbamoilfosfati

+
glutamatdehidrogenazuli reaqciis Sedegad warmoqmnili NH4 uerTdeba
naxSirorJangs, romelic JangviTi reaqciebis erT-erT saboloo produqts
warmoadgens. am reaqciaze ATP-is ori molekula ixarjeba. warmoqmnili
maRalenergetikuli naerTi – karbamoilfosfati – uerTdeba ornitins, ris
Sedegadac miiReba citrulini. es ukanaskneli mitoqondriebidan citoplazmaSi
gamodis, sadac ukavSirdeba asparaginis mJavas. amis Sedegad warmoiqmneba
argininosuqcinati, romelic Semdgom iSleba argininad da fumaris mJavad. ciklis
saboloo stadiaze arginins scildeba Sardovana da warmoqmnili ornitini kvlav
transportirdeba mitoqondriebSi, Semdgom ciklSi monawileobis misaRebad:
aRsaniSnavia is, rom Sardovanas warmoqmnisas, xdeba erTi ioni HCO3 –isa da ori
ioniNH4-is dakavSireba, ris Sedegadac organizmi metabilizmis ori narCeni
produqtisagan Tavisufldeba: amoniakisa da bikarbonatisagan. Sardovana
RviZlidan sisxlSi gadadis, xolo am ukanasknelidan ki Tirkmelebis saSualebiT

89
idevneba organizmidan Sardis saxiT.
Sardovanas ciklis saboloo gantolebaa:

-
2NH4 + HCO3 + 3ATP + H2O Sardovana + 2ADP + AMP + PPi + 2Pi + H+

Sardovana

arginini ornitini
karbamoilfosfati
fumaris
mJava

argininosuqcinati citrulini
asparaginis mJava

Sardovanas cikli

aminomJavebis dekarboqsilireba

aminomJavebis katabolizmis erT-erT mniSvnelovan reaqcias maTi

dekarboqsilireba warmoadgens. am reaqciebis Sedegad aminomJavas Camoscildeba
karboqsilis jgufi da igi naxSirorJangis saxiT Tavisufldeba.
dekarboqsilirebis reaqciebs fermentebi – dekarboqsilazebi atareben, romelTa
SenebaSic, prostetuli jgufis saxiT monawileobs piridoqsalfosfati (vitamini
B6). dekarboqsilazebis moqmedebis Sedegad warmoiqmnebian naerTebi, romlebsac
biogenuri aminebi ewodebaT. am naerTTa zogierT warmomadgenels Zlieri
fiziologiuri da farmakologiuri efeqti aqvs: isini hormonebisa da
neirogadamcemebis rols asruleben da monawileobas iReben qsovilTa
regulaciaSi.
ganasxvaveben 4 tipis dekarboqsilazur reaqcias, romelTa Soris yvelaze
mniSvnelovania -- -dekarboqsilireba:

90
 R-CHNH2-COOH R-CH2NH2 + CO2

dekarboqsilazebis moqmedebis Sedegad glutaminis mJavidan miiReba gama-

aminoerbos mJava (gaem) dioqsifenilalaninidan (warmoiqmneba aminomJava Tirozinis
daJangviT) – dofamini, triftofanidan – triftamini, oqsitriftofanidan
(warmoiqmneba aminomJava triftofanis daJangviT) – serotonini, histidinidan –
histamini:

CH2NH2
CH 2 CH2 CH 2NH2 HO CH2CH 2NH2
HO
CH2
COOH HO NH
gaem dofamini serotonini

CH2CH 2NH2

NH N

histamini
Ggaem-i centraluri Semakavebeli neirogadamcemia. misi Seyvana organizmSi
refleqsebis daqveiTebasa da umaRlesi nervuli moqmedebis Sesustebas iwvevs.
trankvilizatorebis - benzodiazepinebis sedaciuri efeqti gaem-is receptorebis
gaaqtivebiT aris ganpirobebuli. Ggaem-i da misi analogebi, xSirad dadebiT
efeqts iZlevian epilefsiis mkurnalobisas.
dofamini centraluri da periferiuli neirogadamcemia. tvinis nervuli
gzebi, romlebic dofamins Seicaven, monawileoben iseT daavadebaTa paTogenezSi,
rogorebic aris Sizofrenia, parkinsonis daavadeba, depresia da sxva. dofaminis
antagonistebs neirolefsiuri efeqti aqvT.
serotoninic da histaminic centralur da periferiul mediatorebs
warmoadgenen. garda amisa, histamins lokaluri moqmedebac gaaCnia. igi didi
raodenobiT gamoiyofa anTebiTi procesebis adgilze, afarTovebs sisxlZarRvebs,
da amiT xels uwyobs leikocitebis SeRwevas anTebis kerasTan. histamini, agreTve
monawileobas iRebs kuWis mier marilmJavis gamoyofis procesSi. antihistaminuri
preparatebi (mag. dimedroli) farTod gamoiyenebian qsovilTa sensibilizaciis
(mgrZnobiarobis) dasaTrgunavad, agreTve kuWis paTologiebis samkurnalod
(cimetidini).

91
 aminomJavaTa naxSirbadovani ConCxis katabolizmi

organizmSi arsebuli cilebi ZiriTadad 20 aminomJavisagan Sedgeba, amitom

aminomJavaTa naxSirbadovani ConCxis daSlis 20 gza arsebobs. aminomJavaTa
daSlis Sedegad gamoyofili energia organizmis moTxovnilebis mxolod 10%
akmayofilebs. aminomJavebis daSlis Sedegad 5 produqti warmoiqmneba, romelic
krebsis ciklis saSualebiT agrZelebs daJangvas. 10 aminimJavisagan, kerZod,
alaninisagan, treoninisagan, glicinisagan, serinisagan, cisteinisagan,
fenilalaninisagan, Tirozinisagan, leicinisagan, lizinisagan da
triftofanisagan warmoiqmneba: acetil-CoA; argininisagan, histidinisagan,
glutaminis mJavisagan da prolinisagan -- -ketoglutarati, meTioninisa da
valinisagan – suqcinil-CoA, asparaginis mJavidan – mJaunZmarmJava. fenilalaninisa
da Tirozinisagan acetil-CoA-is garda warmoiqmneba agreTve fumaris mJava.

zogierTi memkvidreobiTi daavadebebi, dakavSirebuli aminomJavaTa

katabolizmTan

dReisaTvis aRmoCenilia, rom memkvidreobiTi paTologiebis nawili

ganpirobebulia aminomJavaTa cvlaSi monawile fermentebis fizikur-qimiuri da
biologiuri Tvisebebis darRvevebiT. memkvidreobiT daavadebebs safuZvlad
fermentis makodirebel genSi momxdari mutaciuri cvlilebebi udevs, romelic
fermentis molekuluri organizaciisa da Sesabamisad, katalizuri aqtivobis
cvlilebas iwveven. xSirad, es cvlilebebi ganpirobebulia cilis molekulaSi
mxolod ramdenime aminomJavis SecvlasTan. mutaciuri cilis anomaliuri
fermentuli aqtivoba ujredSi garkveuli metabolitebis dagrovebas iwvevs. aseTi
metabolitebi qsovilTa funqcionirebaze moqmedeben, aqveiTeben maT normalur
ganviTarebas.
memkvidreobiTi daavadebebidan yvelaze metad gavrcelebilia darRvevebi,
romlebic fenilalanin-Tirozinis gardamqmnel fermentebs moicaven. es
ukanasknelni, mgrZnobiare arian genetikuri cvlilebebis mimarT. erT-erT aseT,
sakmaod gavrcelebul daavadebas, fenilketonuria warmoadgens. fenilketonuriis
dros adgili aqvs ferment fenilalaninhidroqsilazis mutaciur cvlilebebs,
ris gamoc, fenilalaninidan Tirozinis sinTezi veRar xerxdeba. am
avadmyofobisaTvis damaxasiaTebelia bavSvebSi gonebrivi ganviTarebis daqveiTeba.
mis simptoms SardSi didi raodenobiT fenilpiroyurZnis mJavis gamoyofa

92
warmoadgens.
xSir SemTxvevebSi mutaciur cvlilebebs ganicdis geni, romelic
fenilalaninis metabolizmis kidev erT ferments – homogentizinis mJavis
oqsidazas aprogramirebs. am ukanasknelis defeqtis gamo, viTardeba daavadeba,
romelic alkaptonuriiT aris cnobili da romelic xasiaTdeba SardSi
homogentizinis mJavis didi raodenobiT (es mJava haerze moxvedrisas Savdeba).
fenilalaninis gardaqmnis gzebSi monawile fermentis mutaciis Sedegad
viTardeba agreTve albinizmi. es genetikuri cvlileba ganpirobebulia ferment
Tirozinhidroqsilazis defeqtiT, romelic veRar awarmoebs Tirozinis daJangvas
dioqsifenilalaninSi. amis Sedegad pigmenti melanini ar warmoiqmneba.

Aanabolizmi

naxSirwylebis biosinTezi cxovelTa organizmSi

wina TavebSi Cvens mier ganxiluli iyo kataboluri reaqciebi, romelTa
saSualebiTac xdeba ZiriTadi sakvebi nivTierebebis, cximebis, naxSirwylebis da
cilebis daSla, maTi CarTva krebsis ciklSi da daSlili produqtebidan ATP-is
miRebis gzebi. es ukanaskneli biosinTezuri, anaboluri reaqciebis Casatareblad
gamoyeneba.
anabolizmi da katabolizmi ujredSi Tanadroulad mimdinareobs.
katabolizmis Sedegad gamoyofili energia biosinTetur reaqciebs xmardeba da
amiT ujredebSi mimdinare procesebs Soris dinamiuri wonasworoba myardeba.
unda aRiniSnos, rom makromolekulebis biosinTezis da daSlis gzebi
araidenturia. isini regulirdebian sxvadasxva fermentebiT da xSirad,
gamoyofili arian erTmaneTisagan morfologiuri, subujreduli barierebiT.

glukoneogenezi
cxovelTa organizmSi biosinTezuri procesebis centralur rgols
warmoadgenen reaqciebi, romlebic glukozis sinTezs awarmoeben. D-glukoza,
mravali qsovilis da organos, maT Soris nervuli sistemis, Tirkmelebis,
sakvercxeebis, eriTrocitebis, embrionis erTaderT energetikul wyaros
warmoadgens. magaliTad, adamianis tvini mxolod dRe-Ramis ganmavlobaSi 120 gr
glukozas moixmars. amitom, organizmSi glukoza gamudmebiT sinTezirdeba
aranaxSirwyalbadovani komponentebisagan da am process glukoneogenezi (axali
Saqris warmoqmna-laT.) ewodeba. cxovelebSi glukoneogenezi, upiratesad, RviZlSi

93
mimdinareobs (SedarebiT ufro sustad es procesi, Tirkmelebis qerqovan
nivTierebaSic warmoebs).
N ZiriTad nivTierebas, romlisganac glukoza sinTezirdeba, piroyurZnis mJava
warmoadgens. es ukanaskneli ki – mravali nivTierebis daSlis (TviT glukozisac)
saboloo produqtia. glikolizis im aTi reaqciidan, romlis saSualebiTac
glukozidan piruvati warmoiqmneba, Svidi glukoneogenezSi gamoiyeneba, radgan
isini advilad Seqcevadi arian. glikolizis sami reaqcia Seuqcevadia da amitom,
glukozis sinTezi sxva fermentuli sistemebis saSualebiT warmoebs. Tavis mxriv,
glukoneogenezis es fermentuli reaqciebic Seuqcevadia, rac imaze metyvelebs,
rom glukozis, rogorc daSlis, aseve sinTezis gzebi, praqtikulad pirdapiri
mimarTulebiT mimdinareobs. ufro metic, isini damoukideblad regulirdebian.
amrigad, piruvatidan glukozis sinTezSi glikolizis ukureaqciebi
gamoiyenebian, garda sami SemovliTi gzisa, romlebic glukoneogenezs mxolod
pirdapiri mimarTulebiT amuSaveben. pirvel SemovliT gzas fosfoenolpiruvatis
sinTezi warmoadgens. glikolizis procesSi fosfoenolpiruvatis piruvatamde
gardaqmna ferment piruvatkinazas saSualebiT warmoebs. piruvatkinazis meSveobiT
ukureaqciis Catareba anu piruvatidan fosfoenolpiruvatis sinTezi SeuZlebelia,
radgan amisaTvis aucilebelia daixarjos sakmaod didi aqtivaciis energia.
amitom, piruvatidan jer warmoiqmneba mJaunZmarmJava, anapreloturi fermentis
piruvatkarboqsilazis saSualebiT, xolo Semdeg ki – oqsaloacetatidan –
malati malatdehidrogenazis meSveobiT. warmoqmnili malati gamodis
mitoqondriebidan, gadadis citoplazmaSi da isev enolpiruvatkarboqsikinazis
saSualebiT, oqsaloacetatidan warmoiqmneba fosfoenolpiruvati, romelic
agrZelebs “aRmaval” gardaqmnas glukozisaken, glikolizis fermentebis
SeqceviTi reaqciebiT manamde, sanam ar warmoiqmneba fruqtozo-1,6-difosfati. am
ukanasknelis sinTezs glikolizSi fermenti fosfofruqtokinaza axorcielebs,
romlis ukureaqciis Catareba praqtikulad SeuZlebelia. amitom, fruqtozo-6-
fosfatis warmoqmnas fruqtozo-1,6-fosfatidan, glukoneogenezis SemTxvevaSi,
sxva fermenti, kerZod, fruqtozodifosfataza awarmoebs. da bolos, mesame
reaqcias, romelsac SemovliTi gza esaWiroeba, warmoadgens glukozis warmoqmna
glukozo-6-fosfatidan. glukoneogenezSi, glikolizisagan gansxvavebiT, am
reaqcias ara heqsokinaza, aramed glukozo-6-fosfataza warmarTavs.
glukoneogenezis Sedegad warmoqmnili glukoza an sisxlis saSualebiT
gadaecema sxva organoebs, an glikogenis saxiT Seinaxeba TviT RviZlSi.
glikolizi da glukoneogenezi reciprikulad regulirdebian. centralur
metabolits, romelic am or process aregulirebs, acetil-CoA warmoadgens. es

94
naerTi ferment piruvatkarboqsilazas dadebiTi modulatoria da piruvatidan
fosfoenolpiruvatis sinTezs aCqarebs, e.i. zrdis glukoneogenezis pirveli
stadiis siCqares. amasTanave, igi ferment piruvatkinazas uaryofiTi modulatoria
da anelebs fosfoenolpiruvatidan piruvatis warmoqmnas, e.i. akavebs glikolizis
me-10 reaqciis siCqares. amdenad, acetil-CoA-s RviZlis ujredebSi dagroveba
aZlierebs glukoneogenezs da ainhibirebs glikolizs.
garda amisa, radgan glukozis sinTezSi monawiloobas Rebulobs krebsis
ciklis erT-erTi metaboliti – mJaunZmarmJava – yoveli naerTi, romelic krebsis
ciklis metabolitebamde iSleba, gazrdis glukozis warmoqmnis siCqares. aseT
naerTebs ki – rodorc cnobilia, cximebis, naxSirwylebis da aminomJavebis
ZiriTadi tipebi warmoadgenen. amrigad, urTierTsawinaaRmdegod mimarTuli
glikolizisa da glukoneogenezis reaqciebis msvleloba SesaZlebelia Semdegi
sqemiT gamovsaxoT:

glukoza
glukozo-6-fosfataza heqsokinaza
glukozo-6-fosfati
fosfoglukoizomeraza
fruqtozo-6-fosfati
fruqtozodifosfataza fosfofruqtokinaza
fruqtozo-1,6-difosfati
aldolaza
gliceraldehid-3-fosfati + dihidroacetonfosfati
triozofosfatizomeraza
gliceraldehid-3-fosfati
gliceraldehidfosfatdehidrogenaza

glukoneogenezi 3-fosfogliceroilfosfati
fosfogliceratkinaza
3-fosfoglicerati
fosfogliceratmutaza
2-fosfoglicerati

enolaza glikolizi
oqsaloacetati fosfoenolpiruvati
malati piruvatkinaza
oqsaloacetati piruvati

95
 glikogenis sinTezi

cxovelebSi glikogenis sinTezi TiTqmis yvela tipis qsovilebSi

mimdinareobs, magram gansakuTrebuli intensivobiT es procesi RviZlSi da
ConCxis kunTebSi xasiaTdeba. glikogenis sinTezi iwyeba glukozidan glukozo-6-
fosfatis warmoqmniT, romelic momdevno reaqciaSi glukozo-1-fosfatad
gardaiqmneba. es orive reaqcia saerToa glikogenis daSlisa da misi
sinTezisaTvis da warimarTeba mTliani procesis mimdinareobis mimarTulebiT.
glikogenis sinTezis sakvanZo reaqcias, romelic glikogenolizis procesSi
ar gvxvdeba, glukozo-1-fosfatidan uridindifosfoglukozis warmoqmnaa:

UTP + glukozo-1-fosfati UDP-glukoza + Ppi

UDP-glukozidan glukozis naSTi ferment glikogensinTetazis meSveobiT

glikogenis molekulas uerTdeba:

UDP-glukoza + (glukoza)n UDP + (glukoza)n+1

glikogensinTetaza-fermenti, romelic warmoqmnis glikogens da

glikogenfosforilaza-fermenti, romelic Slis glikogens, reciprokulad
regulirdebian. Tu glikogenfosforilazis aqtiur formas warmoadgens
fosforilirebuli fermenti (fosforilaza a), glikogensinTetazis aqtiuri
forma arafosforilirebuli fermentia.

glikogenfosforilaza glikogenfosforilaza-P
(pasiuri) (aqtiuri)

proteinkinaza

glikogensinTetaza glikogensinTetaza-P
(aqtiuri) (pasiuri)

Tu gaviTvaliswinebT imas, rom proteinkinazebi hormonebiT regulirdebian,

gasagebi gaxdeba, rom ujredSi glikogenis daSla da misi warmoqmna mxolod

96
erTi mimarTulebiT SeiZleba mimdinareobdes.

naxSirwylebis cvlis darRvevebi

NnaxSirwylebis cvlis daavadebaTa erT-erTi ZiriTadi gamovlineba aris

sisxlSi glukozis raodenobis meryeoba. ansxvaveben hiperglikemias –
mdgomareobas, rodesac sisxlSi glukozis raodenoba momatebulia da
hipoglikemias – mdgomareobas, rodesac glukozis raodenoba sisxlSi mcirdeba.
hiperglikemia endokrinul sistemaTa darRvevis erT-erT tipiur simptoms
warmoadgens.
Saqriani diabeti. glikolizisa da glikogenolizis regulaciaSi wamyvan
rols asrulebs insulini, romlis ukmarisobis dros viTardeba daavadeba –
Saqriani diabeti. am dros glukozis koncentracia sisxlSi (hiperglikemia) da
SardSi (glukozuria) izrdeba, xolo glikogenis maragi RviZlSi mcirdeba. amave
dros, kunTovani qsovili kargavs glukozis utilizaciis unars. insulinis
Seyvana iwvevs metaboluri darRvevebis koreqcias: normalizdeba glukozis
STanTqma kunTis ujredebis mier, aRdgeba wonasworoba glikolizsa da
glukoneogenezs Soris. insulini glikolizSi monawile fermentebis –
fosfofruktokinazis, heqsokinazis, piruvatkinazis – induqciasa da
glukoneogenezSi manawile fermentebis – represias, maTi sinTezis daTrgunva
iwvevs
Saqriani diabetis garda hiperglikemia SeiZleba gamowveuli iyos zogierTi
sxva endokrinuli jirkvlis funqciis darRvevis Sedegad ase, magaliTad,
hiperglikemia gamoiwveva Tirkmelzeda jirkvlis avTvisebiani gadagvarebis,
farisebri jirkvlis hiperfunqciis Sedegad.
hipoglikemia. hipoglikemia SesaZlebilia gamowveul iyos hipofizaruli
kaxeqsiis, adisonis daavadebis, hipoTireozis Sedegad. glukozis koncentraciis
Zlier daqveiTebas sisxlSi iwvevs kuWqveSa jirkvlis adenomebi, -ujredebis mier
insulinis didi raodenobiT gamomuSavebis gamo. Hhipoglikemia viTardeba agreTve,
SimSilis, xangrZlivi fizikuri datvirTvis, -ganglioblokatorebis miRebis
Sedegad.
glukozuria. SardSi glukozis Warbi raodenoba, xangrZlivi drois
ganvalobaSi, metyvelebs kuWqveSa jirkvlis paTologiur cvlilebebze (Saqriani
diabeti, mwvave pankreatiti da a.S.). SedarebiT ufro iSviaTad gvxvdeba
Tirkmlovani warmoSobis glukozuria, romelic ganpirobebulia Tirkmlis
milakebidan glukozis rezorbciis ukmarisobiT. glukozuria, rogorc droebiTi

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movlena, SesaZlebelia warmoiqmnas zogierTi mwvave infeqciuri da nervuli
daavadebebis, intoqsikaciis Sedegadac.
naxSirwylovani cvlilebebi hipoqsiuri mdgomareobisas. hipoqsiuri
mdgomareobisas piruvatis daJangvis siCqare xSirad CamorCeba glikolizis
intensivobas. amitom, sisxlSi piruvati da laqtati grovdeba, ris gamoc, sisxlSi
pH-is Secvla (SemJaveba, acidozi) xdeba.
hipoqsiuri mdgomareoba SesaZlebelia gamowveuli iyos sisxlis mimoqcevis
an sasunTqi sistemebis darRvevebiT, simaRlis avadmyofobiT, anemiiT, Jangva-
aRdgeniTi fermentebis daqveiTebiT zogierTi infeqciisa da intoqsikaciis
Sedegad, hipo-da avitaminozebis dros. laqtatisa da piruvatis mateba sisxlSi
SesaZlebelia agreTve gamowveul iyos RviZlis parenqimis dazianebiT (hepatitis
gviani stadiebi, cirozi da sxva) am dros glukoneogenezis Sekaveba xdeba.
glikogenozebi. glikogenis cvlis darRvevasTan aris dakavSirebuli
zogierTi memkvidreobiTi daavadebebic, romlebsac zogadad glikogenozebi
ewodebaT. es daavadebebi warmoiqmnebian glikogenis sinTezsa da daSlaSi
monawile fermentebis susti aqtivobis an maTi ukmarisobis gamo.
aseTi tipis daavadebaTa Soris SedarebiT xSiria I tipis glikogenozi anu
girkes daavadeba, romelic ganpirobebulia ferment glukozo-6-fosfatazis
sinTezis genetikuri defeqtiT.
cnobilia glikogenozebis 9 tipi.

L lipidebis biosinTezi

lipidebis da maT Soris triacilglicerinebis anu trigliceridebis

biosinTezi cxovelTa organizmSi maRali intensiobiT mimdinareobs, rac
ganpirobebulia imiT, rom isini organizmis ZiriTad samarago fonds
warmoadgenen. am naerTebSi dagrovili energia sakmarisia nivTierebaTa cvlis
SesanarCuneblad 8 kviris ganmavlobaSi da isini adamianis masis 15-20%-s
Seadgenen. im SemTxvevebSi, rodesac adamiani didi raodenobiT iRebs
naxSirwylovan sakvebs da organizms ukve aRar ZaluZs misi Senaxva glikogenis
formiT, adgili aqvs triacilglicerinebis sinTezs, romlis didi raodenobac
SesaZlebelia dagrovdes cximovan qsovilebSi. neitraluri lipidebisagan
mono – , di – da triacilglicerinebisagan gansxavebiT polaruli lipidebi ar
maragdebian organizmSi, magram monawileoben ra membranebis SenebaSi, maT
gaaCniaT swrafi ganaxlebis periodi (magaliTad, RviZlis ujredebis plazmur
membranebSi arsebuli fosfolipidebis ganaxlebis periodia-2-3 dRe) da amitom

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aqtiurad sinTezirdebian organizmSi.

cximovan mJavaTa sinTezi

malonil-CoA-warmoqmna. adamianisa da cxovelTa organizmSi arsebuli

cximovani mJavebi Sedgebian luwi raodeniba naxSirbadis atomebisagan. amitom,
advili dasaSvebi iyo is, rom iseve rogorc cximovani mJavebis daSlisas, maT
sinTezSic, monawileobas Rebuloben ornaxSirbadiani acetil-CoA-s molekulebi,
magram aRmoCnda, rom es naerTi cximovani mJavebis naxSirbadovani ConCxis zrdaSi
uSualod ar monawileobs da misgan pirvelad, 3-naxSirbadiani malonil-CoA
warmoiqmneba. am ukanaskneliT iwyeba cximovani mJavebis sinTezi da mxolod,
bolo stadiebze kvlav warmoiqmneba acetil-CoA.
TviT malonil-CoA acetil-CoA-gan sinTezirdeba. Ees procesi citoplazmaSi
mimdinareobs da mas katalizs uwevs sakmod rTuli aRnagobis biotinSemcveli
regulatoruli fermenti – acetil-CoA-karboqsilaza. es ukanaskneli, 2-
naxSirbadiani acetil-CoA-s 3-naxSirbadian maloni-CoA-ad gardasaqmnelad iyenebs
CO2-s:

Aacetil-CoA + ATP + CO2 + H2O malonil-CoA + ADP + Pi + H+

COOH

CH3 CH2

C O CO 2 C O
S CoA S CoA

acetil-CoA malonil-CoA

reaqciaSi monawile naSirorJangi gamoiyeneba maloni-CoA-s karboqsilis

jgufis sinTezisaTvis. aRsaniSnavia, rom cximovani mJavebis naxSirbadovani
ConCxis zrdis procesSi es karboqsili isev Tavisufldeba CO2-is saxiT. ase, rom
cximovani MmJavebis ConCxi, sabolood mainc, acetil-CoA-gan yalibdeba.
acetil-CoA-is citoplazmuri fondi mitoqondrialuri citratidan iqmneba,
romlis Warbi raodenobac am organoebidan gamodis. citoplazmaSi igi iSleba
mJaunZmarmJavad da acetil-CoA-ad da mxolod amis Semdeg iwyeba malonil-CoA-is

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sinTezi. citrati acetil- CoA-karboqsilazis dadebiT modulators warmoadgens;
rac ufro meti citrati warmoiqmneba mitoqondriebSi krebsis ciklis
msvlelobisas, miT ufro Warbi raodenobiT gamova igi citoplazmaSi da
daiSleba acetil-CoA-ad, da Tu am SemTxevaSi acetil-CoA-karboqsilaza
gaaqtivebulia citratiT, iwyeba cximovani mJavebis intensiuri sinTezi, romlis
pirvel da malimitirebel stadias malonil- CoA-is warmoqmna warmoadgens.
cximovani mJavebis sinTezi. cximovani mJavebis sinTezs mravalfermentiani
kompleqsi – cximovani mJavebis sinTaza awarmoebs. igi 7 individualuri
fermentisagan Sedgeba. am kompleqsis centralur rgols e.w. acilebis gadamtani
cila Seadgens (agc). igi dabali molekuluri wonisaa (9000) da aqvs prostetuli
jgufi. Ees ukanaskneli, fosfopanToTenis MmJavisagan Sedgeba, romelic sakuTriv,
panToTenis mJavis (vitamini B3) garda, Seicavs sulfihidrilis jgufsac (SH).
swored am sulfihidrilis jgufis saSualebiT ikavSirebs agc cximovani MmJavebis
Sualedur produqtebs. es cila, CoA-s msgavsad cximovani mJavebis karboqsilTan
warmoqmnis kovalentur bmas. Aam sulfihidrilis jgufis garda, agc-s meore,
funqciurad mniSvnelovani SH-jgufi aqvs, romelic cilaSi Semaval amonomJava
cisteins ekuTvnis. orive jgufi monawileobas iRebs cximovani mJavebis sinTezSi.
pirvel stadiaze, agc orive sulfihidrilis jgufis saSualebiT ierTebs
acetilis da malonilis jgufebs, Sesabamisad acetil- CoA-dan da malonil CoA-
dan:

CO 2

CH 3 CO S CH3 CH2 CH 2 CO S
HS HS

COOH
CH 2 CO S CH 3 CO CH2 CO S CH 3 CH2 CH2 CO S HS

acetoacetil-acg buTiril-acg

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 am jgufebis mierTebis Semdeg, acetilis jgufi gadaitaneba malonilis
jgufze. maTi kondensaciis dros gamoiyifa CO2 da warmoiqmneba 4 naxSirbadiani
agc-Tan dakavSirebuli acetoacetili. Semdgom stadiebze agc-Tan dakavSirebuli
acetoacetili, Sidamolekuluri reaqciebis saSualebiT, gardaiqmneba buTirilis
jgufad. eleqtronebis donors am aRdgeniT reaqciebSi NADPH warmoadgens,
romelic pentozofosfaturi gziT warmoiqmneba ujredSi. bolo stadiaze,
buTirilis jgufi meore cisteinis SH-ze gadaitaneba. amis Semdeg, cximovani
mJavebis sinTaza da masSi arsebuli agc mzad arian miierTon malonilis me-2
molekula, romelic CO2 –is dakargvis Semdeg ori erTeuliT gazrdis
buTirilis jgufs. es procesi grZeldeba cximovani mJavis mTliani molekulis
warmoqmnamde.
Aamrigad, cximovani mJavebis sinTezis procesi gansxvavdeba maTi daSlis
procesisagan. pirvel gansxvavebas warmoadgens is, rom cximovani mJavebis sinTezi
mimdinareobs ara mitoqondriebSi, aramed citoplazmaSi. meore gansxvaveba
mdgomareobs imaSi, rom cximovani mJavis sinTezis dros naxSirwyalbadovani
jaWvebis gadatana xorcieldeba ara CoA-iT, aramed specialuri ciliT – agc-Ti.
garda amisa, cximovani mJavebis sinTezSi monawileobas iRebs gaaqtivebuli
acetati mxolod mas Semdeg, rac moxdeba misi gardaqmna malonilis
samnaxSirbadian molekulad; da bolos, cximovani mJavebis ZiriTad energetikul
wyaros NADPH-is aRdgeniTi potenciali warmoqmnis.
aRsaniSnavia is, rom cxovelTa organizmSi advilad sinTezirdeba mxolod
erTi ormagkavSiriani cximovani mJava. Uujer mJavebs, Semdgars ori da meti ormagi
kavSirisagan, cxovelTa ujredebi, maTSi Sesabamisi oqsigenazebis uqonlobis
gamo, ver asinTezireben. amitom, aseTi cximovani mJavebi (magaliTad linolisa da
linoleni) organizmma mza saxiT unda miiRos; maT Seucvleli cximovani mJavebi
ewodeba. Aam mJavebidan cxovelTa organizmebSi warmoiqmneba 4-ormagi kavSiris
Semcveli araqidonis mJava, romelic sawyisia mravali biologiurad aqtiuri
naerTisa da maT Soris eikozanoidebisa (tromboqsanebi, leikotrienebi,
prostaglandinebi).

triacilglicerinebis biosinTezi

triacilglicerinebis da zogierTi fosfolipidebis, kerZod,

fosfatidilqolinis da fosfatidileTanolaminis biosinTezi mzgavsad
mimdinareobs: maT ori saerTo winamorbedi – glicerinfosfati da cximovani
mJavis eTeri CoA-Tan (acil- CoA) aqvT.

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 CH 2OH CH 2OR1 CH 2OR1 CH 2OR1 R3CoA CH 2OR1
R1CoA R2CoA
CHOH OH CHOH OH CHOR2 OH CHOR2 CHOR2
CH 2 O P O CH2 O P O CH 2 O P O CH 2OH CH 2OR3
OH OH Please Register OH

glicerinfosfati monoacilglicerin- diacilglicerin- diacil- triacil-

fosfati fosfati glicerini glicerini

normaSi, zrdasrul adamianebSi da cxovelebSi, triacilglicerinebis

biosinTezi da maTi daSla-daJangva Tanadroulad mimdinareobs. myardeba
garkveuli stacionaluri wonasworoba. im SemTxvevebSi ki, rodesac organizmi
didi raodenobiT iRebs naxSirwylebs, cximebs Tu cilebs d maTi raodenoba
aRemateba moTxovnilebas, wonasworoba irRveva da zedmeti kaloriebi maragdeba
triacilgliceridebis saxiT. am SemTxvevaSi acetil-CoA-s wyaros naxSirwylebi
da aminomJavebi warmoadgenen.
triacilglicerinebi, kvebis araswori racionis garda, icvlebian rigi
hormonebis zemoqmedebiTac. magaliTad, insulini iwvevs naxSirwylebis gardaqmnas
triacilglicerinebad. Saqriani diabetis mwvave formis dros, organizmi kargavs
ara marto naxSirwylebis SeTvisebis unars, aramed cximovani mJavebisa da
triacilglicerinebis sinTezis unarsac. amitom, aseT avadmyofebs ezrdebaT
cximovani mJavebis daJangvisa da ketonuri sxeulebis warmoqmnis siswrafe, ris
gamoc adamiani xdeba. triacilglicerinebis warmoqmnaze gavlenas axdens agreTve
hipofizaruli hormonebi, glukagoni da sxva.

fosfogliceridebis biosinTezi

fosfogliceridebi, iseve rogorc neitraluri cximebi, cximovani mJavis

CoA-warmoebulsa (acil- CoA-sa) da fosfogliceratidan sinTezirdebian. am
sinTezuri reaqciis Sedegad warmoqmnili diacilglicerini an ierTebs mesame
cximovan mJavas da warmoqmnis triacilglicerins, an-fosforilirebul spirtul
komponents (mag., fosforilqolins, fosforileTanolamins da a.S.) da warmoqmnis
Sesabamis fosfoglicerids. ukanasknel SemTxevevbSi fosforilirebuli
spirtuli komponenti uerTdeba ara pirdapir, aramed igi jer aqtvirdeba
citidintrifosfatiT (CTP), xolo Semdeg uerTdeba glicerinis naSTs. Aam dros

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Sualeduri produqtis saxiT warmoiqmneba citidintrifosfatisa da spirtuli
komponentis fosfoeTeri (mag., citidindifosfoqolini):

cximovani mJava

CH 2OR1 CH 2OR1
CHOR2 CHOR2
PleaseCH 2OH
Register CH 2OR3
diacilglicerini triacilglicerini

CDP-eTanolamini CDP-qolini

CH 2OR 1 CH 2OR 1
CHOR2 O CHOR2 O
CH2 O P O CH2CH 2NH2 CH2 O P O CH 2CH2N CH3)3

OH
Please Register OH Register
Please
fosfatidileTanolamini fosfatidilqolini

analogiurad sinTezirdebian sxva fosfolipidebic: fosfatildilserini,

fosfatidilinozitoni da kardiolipini.
qolini adamiansa da cxovelebSi, ZiriTadad, eTanolaminisagan warmoqmneba
misi meTilirebis Semdeg. meTilis jgufis donors am reaqciaSi (iseve, rogorc
sxva reaqciebSi) aminomJava meTionini warmoadgens. amonomJava meTioninis
deficiti racionSi ganapirobebs fosfolipidebis sinTezis daqveiTebas, risi
nawilobrivi kompensaciac qolinis miRebis Sedegad xdeba. amitom, qolini xSirad
damatebiTi vitaminis saxiT ixmareba.
fosfolipidebis biosinTezi endoplazmuri badis fermentTa saSualebiT
warmoebs da warmoqmnili produqtebi badis membranaSive erTvian. endoplazmuri
retikolumis membranebidan fosfolipidebi gamoiyofian vezikulebis saxiT da
gadaitanebian mitoqondriebis, birTvis da sxva organoidebis membranebamde. garda
amisa, endoplazmuri membranebi goljis kompleqsis membranebis winamorbeds
warmoadgens. am ukanasknelSi, rogorc cnobilia, formirdebian “saeqsportod”

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gamzadebuli cilebisagan Semdgari vezikulebi, romlebic transportirdebian
plazmur membranasTan da egzocitozis saSualebiT aTavisufleben SigTavss.
cilisagan Tavisufali fosfolipidebis vezikula uerTdeba plazmur membranas
da mas lipidebiT amdidrebs.

steroidebis biosinTezi

centraluri roli steroidebis metabolizmSi qolesterins miekuTneba.

qolesterinisagan adamianis organizmSi sinTezirdeba naRvlis mJavebi da
sxvadasxva tipis steroiduli hormonebi. garda amisa, qolesterini monawileobas
iRebs membranebisa da sisxlis plazmis lipoproteinebis SenebaSi.
cximovani mJavebis mzgavsad qolesterinic acetil-CoA-dan sinTezirdeba.
qolesterinis sinTezis 1 etapi iwyeba Tanmimdevruli reaqciebisagan, romlis
Sedegad 3 molekula acetil-CoA-dan wamoiqmneba 6 naxSirbadiani mevalonis mJava.
Semdgom, mevalonis mJavidan miiReba 5-naxSirbadiani izopentenilpirofosfati,
romelic izoprenuli erTeulis aqtivirebul formas warmoadgens:

O O

CH2 C CH 2 CH2 O P O P OH

CH3 Please Register

OH OH
izopentenilpirofosfati

Semdgom reaqciebSi, izopentenilpirofosfatis 6 molekula, kargavs ra

pirofosfatis jgufebs, erTmaneTs uerTdeba da warmoqmnis naerTs, romelsac
skvaleni ewodeba da bolos, skvalenis sworxazovani molekula gardaiqmneba
ciklur naerTad – qolesterinad.

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 CH 3
CH3
CH CH2 CH2 CH 2 CH
CH3

OH Please Register

izopentenilpirofosfatidan, qolesterinis garda sxva naerTebic

warmoiqmnebian:

naRvlis mJavebi

steroiduli hormonebi vitamini D

qolesterini

vitamini A
vitamini E kauCuki
vitamini K izopentenilpirofosfati fitoli

karotinebi poliprenolebi

maT ricxvs miekuTvneba vitaminebi: A, E, K, kauCuki, mravali eTerzeTi da sxva

lipidTa cvlis darRvevebi

lipidTa cvlis genetikuri defeqtebi. samedicino praqtikaSi iSviaTad,

magram gvxvdeba genetikuri daavadebebi, ganpirobebuli lipidTa cvlaSi monawile
fermentTa defeqtiT. erT-erT aseT paTologias warmoadgens e.w. tea-saqsis
daavadeba, romelic gamowveulia glikolipid-gangliozidis damSleli fermentis-
acetilheqsoaminidazis uqonlobiT. amitom, gangliozidi bolomde ver iSleba da
didi raodenobiT grovdeba tvinsa da elenTaSi. es iwvevs nervul sistemaSi

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degenaraciuli procesebis ganviTarebas.
aTerosklerozi da lipoproteinebi. aTerosklerozis ganviTarebaSi did
rols TamaSobs sisxlis plazmis lipoproteinebi. isini rTul kompleqsur
warmonaqmns warmoadgenen da sakmaod Taviseburi agebuleba aqvT. lipoproteinuli
nawilaki Sedgeba cximovani wveTisagan (birTvisagan), romelic arapolarul
lipidebs Seicavs (trigliceridebs, eTerificirebul qolesterins da sxva).
birTvi garSemortymulia garsiT, romelic mdidaria polaruli lipidebiT,
Tavisufali qolesteriniT da cilebiT. am ukanasknelebs, apolipoproteinebs
eZaxian.
ganasxvaveben lipoproteinuli nawilakebis ramdenime klass: -
lipoproteinebs anu maRali simkvrivis lipoproteinebs (msl), -lipoproteinebs
anu dabali simkvrivis lipoproteinebs (dzl), pre-lipoproteinebs anu Zlier
dabali simkvrivis lipoproteinebs (Zdsl) da hilomikronebs (hm). rac ufro metia
lipoproteinul nawilakSi cilebis raodenoba, miT ufro maRali simkvrivisaa es
nawilaki da piriqiT, rac metia masSi neitraluri cximebis raodenoba, miT ufro
dabali simkvrive aqvs mas. magaliTad, msl_Si cilebis raodenoba 50% aRwevs,
xolo trigliceridebisa ki – mxolod 6-8%, maSin rodesac Zdsl-Si, piriqiT,
trigliceridebis raodenoba 40-70%-is tolia, xolo cilebisa ki – 5-15%. sisxlis
plazmaSi dabali simkvrivis lipoproteinebis mateba SesaZlebelia
aTerosklerozis erT-erTi gamomwvevi mizezTagani iyos. dsl da Zdsl
xasiaTdebian aTerogenulobiT; maT SeuZliaT SeaRwion sisxlZarRvTa kedlebSi
da Seqmnan pirveladi substrati, aTerosklerotuli wanazardebis
gasaviTareblad.

aminomJavebis da nukleotidebis biosinTezi

aminomJavebis da nukleotidebis biosinTezi mraval saerTo kanonzomierebas

eyrdnoba. es naerTebi Seicaven azotis atomebs, romelTa donorsac erTidaigive
nivTiereba warmoadgens, xolo zogierTi aminomJava nukleotidebis biosinTezSic
gamoiyeneba.

aminomJavebis sinTezi adamianis organizmSi

biologiuri saxeobebi, sxva gansxvavebebTan erTad, erTmaneTisagan

aminomJavaTa biosiTezis gzebiTac gansxvavdebian. cilovani molekulis
formirebisaTvis saWiro 20 aminomJavisagan, adamians mxolod 10-is dasinTezirebis

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unari Seswevs. danarCeni 10-is sinTezi mis organizms ar SeuZlia da amitom, man
isini unda miiRos mza mdgomareobaSi. aseT aminomJavebs Seucvleli aminomJavebi
ewodeba. am aminomJavebis sinTezi rTuli bioqimiuri gziT mimdinareobs, da
adamians, am reaqciebis Camtarebeli zogierTi fermenti ar gaaCnia. gansakuTrebiT,
es Seexeba im aminomJavebs, romelic benzolis birTvs an heterocikls Seicaven.
Seucvlel aminomJavebs warmoadgenen: izoleicini, leicini, lizini, meTionini,
fenilalanini, treonini, triftofani, valini, arginini, histidini.
DdanarCeni aminomJavebi adamianis organizmSi advilad sinTezirdebian da am
procesis centralur stadias glutaminis mJavis biosinTezi warmoadgens.
glutaminis mJava sxva aminomJavebisaTvis aminojgufebis donoria. igi, krebsis
ciklis erT-erTi metabolitis - -ketoglutaris mJavisagan miiReba
glutamatdehidrogenazuli reaqciis saSualebiT. glutaminis mJavisagan Tavis
mxriv, warmoiqmneba prolini da glutamini:

-ketoglutarati glutaminis mJava glutamini

COOH COOH COOH

C O CHNH2 CHNH2

CH2 CH2 CH2

CH 2 CH 2 CH 2
COOH COOH PleaseCONH
Register
2

COOH

CH
NH CH 2

CH
Please
2 CH 2
Register prolini

prolini ciklur aminomJavas warmoadgens, romelic ori fermentuli reaqciis

saSualebiT miiReba glutaminis mJavisagan. glutaminis mJavis gadaaminirebiT
warmoiqmneba alanini da asparaginis mJava, romelic aseve centraluri
metabolitebis, piroyurZnis mJavis da mJaunZmarmJavis warmoebulebs wamoadgenen:

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 COOH COOH
CHNH 2 CH3 C O CH 3
CH2 C O CH 2 CHNH2
CH 2 CH2
COOH COOH
COOH COOH
glutaminis piroyurZnis -ketoglutaris alanini
mJava mJava mJava

COOH COOH
COOH COOH
CHNH2 C O
C O CHNH2
CH2 CH2
CH 2 CH2
CH 2 CH 2
COOH COOH
COOH COOH
glutaminis mJaunZmarmJava -ketoglutaris asparaginis
mJava mJava mJava

aminomJava Tirozini miiReba Seucvleli aminomJava – fenilalaninisagan,

cisteini – Seucvleli aminomJava – meTioninisagan da serinisagan. Aam
ukanasknelidan, miiReba glicinic. TviT serini miiReba 3-fosfogliceratidan.

kreatinis biosinTezi

aminomJavebisagan cilebis garda miiReba sxva naerTebic. magaliTad, sami

aminomJavisagan – glicinisagan, argininisagan da meTioninisagan miiReba kreatini:

NH2 NH PO3
+
C N H2 C N H2
+

N CH3 ATP N CH3 ADP

CH 2 CH2
COOH COOH
kreatini fosfokreatini

kreatins didi mniSvneloba eniWeba kunTovani qsovilis energetikul cvlaSi,

sadac igi ATP-Tan erTad maRalenergetikuli fosfatebis samarago fonds qmnis.
kreatinis fosforilireba ATP-is saSualebiT mimdinareobs. warmoqmnili

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kreatinfosfati ujredebSi inaxeba da saWiroebis SemTxvevaSi (energiis
deficitisas) isev iSleba kreatinad. am dros gamoTavisuflebuli makroerguli
fosfati metabolur procesebSi moixmareba.

porfirinebis biosinTezi

porfirinuli birTvi hemis ZiriTadi Semadgeneli nawilia. hemproteinebSi –

hemoglobinSi da citoqromebSi es birTvi warmoqmnis koordinaciul bmas rkinis
atomTan. igi 4 monopiroluri warmoebulis - porfobilinogenisagan Sedgeba,
romelic aminomJava glicinisagan sinTezirdeba.

COOH NH N

CH 2

C CH 2CH2COOH
N HN
C CH
NH
+
CH2N H 2
porfobilinogeni hemi

hemoglobinis rkina-porfirinis birTvi, iSleba - ra elenTaSi, Tavisufldeba

rkinisagan da warmoqmnis gaxsnil tetrapirolur struqturas, romelsac
bilirubini ewodeba. bilirubini albuminis saSualebiT RviZlSi gadaitaneba,
iSleba e.w. naRvlis pigmentebad da organizmidan gamoiyofa.
bilirubini, romelic sisxlSi, elenTaSi an romelime sxva organoSi
warmoiqmneba, wyalSi sustad ixsneba, advilad adsorbirdeba Sratis cilebTan da
amitom sisxlSi misi gansazRvra, mxolod specialuri damuSavebis Sedegad xdeba.
aseT bilirubins klinikur bioqimiaSi arapirdapiri bilirubini ewodeba. sisxlSi
arsebuli bilirubimis 75% arapirdapirze modis, e.i. im bilirubinze, romelic
Sratis cilebTan aris dakavSirebuli.
bilirubinis momateba sisxlSi siyviTles iwvevs, xolo misma didma
raodenobam SesaZlebeliia mZime mowamvlis simptomebic ki gamoavlinos. xvdeba ra
RviZlSi, cilebTan dakavSirebuli bilirubini detoqsificirdeba. Mmas uerTdeba
glukuronis mJava, ris Sedegadac bilirubini SedarebiT indiferentuli da

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wyalSi xsnadi xdeba. amitom, igi advilad, yovelgvari winaswari damuSavebis
gareSe isazRvreba sisxlSi da mas pirdapiri bilirubini ewodeba. naRvelSi
mxolod es ukanaskneli identificirdeba. bilirubinis gansazRvras sisxlSi da
SardSi klinikur-diagnostikuri mniSvneloba aqvs RviZlis, elenTis, sisxlis da
sxva daavadebaTa dros.

nukleotidebis biosinTezi

purinebis biosinTezi

purinis fuZeebi (adenini, guanini) ZiriTadad aminomJavebisagan

sinTezirdebian. purinis birTvis warmoqmnaSi, monawileobas Rebulobs glicini,
asparaginis mJava da glutaminis amiduri azoti. garda amisa birTvis sinTezSi
monawileobs WianWvelamJava (formiati) da naxSirorJangi.

formiati aspartati CO2 glicini formiati

C N

N C
C

C C
N
Please Register N
glutaminis amiduri azoti

pirvel purinul naerTs, romelc am rTuli biosinTezuri reaqciiT miiReba,

inozinis mJava warmoadgens:

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 O

C N

HN C
CH

HC C O
N N
CH 2 O P OH
O
OH

Please Register
inozinmonofosforis mJava (IMP)

NH 2

C N

N C
CH

HC C O
N N
CH 2 O P OH
O
OH

Please Register
adenozinmonofosforis mJava (AMP)

C N

HN C
CH

NH2 C C O
N N
CH 2 O P OH
O
OH

guanozinmonofosforis mJava (GMP)

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 inozinis mJavis biosinTezis dros purinis birTvi Sendeba ribozo-5-fosfatis
molekulaze etapobrivad. I stadiaze ribozo-5-fosfats glutaminis amiduri
azoti uerTdeba, xolo Semdeg danarCeni atomebi. inozinmonofosforis mJavidan
warmoiqmnebian adenozin- da guanozinmonofosforis mJavebi. monofosfatebis
warmoqmnis Semdeg maT ATP-dan damatebiTi fosfatebi uerTdeba, ris Sedegadac
Sesabamisi di – da trifosfatebi warmoiqmneba:

AMP + ATP 2 ADP

GMP + ATP GDP + ADP

GDP + ATP GTP + ADP

pirimidinebis biosinTezi

pirimidinebis biosinTezi, purinebisagan gansxvavebiT, aromatuli birTvidan

iwyeba. I reaqciis dros asparaginis mJavas karbamoilfosfati ukavSurdeba N-
karbamoilaspartatis warmoqmniT. am process aspartattranskarbamoilaza
warmarTavs. igi alosterul ferments warmoadgens da inhibirdeba pirimidinebis
biosinTezis saboloo produqtiT – citidintrifosfatiT (CTP) .
pirimidinebis biosinTezis erT-erTi mTavari Sualeduri produqtia orotis
mJava:

O O NH 2

NH NH NH

O COOH O O O O O O
NH N N
CH 2 O P OH CH 2 O P O P O P OH
O O
OH OH OH OH

Please Register

orotis mJava uridinmonofosforis citidintrifosforis

mJava (UMP) mJava (CTP)

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 orotis mJavas ribozo-5-fosfatis gverdiTi jaWvi uerTdeba da sinTezirebuli
orotidilis mJavis dekarboqsilirebiT uridinmonofosforis mJava (UMP) miiReba.
UMP-is fosforilirebiT (ATP-is saSualebiT) warmoiqmneba UTP, xolo am
ukanasknelis glutaminiT amidirebiT ki -- CTP. rogorc ukve iTqva, CTP
pirimidinebis biosinTezis pirvel stadiaSi monawile fermenti –
aspartattranskarbamoilazis uaryofiT modulators warmoadgens da amitom, CTP-
is dagrovebis SemTxvevaSi jaWvi wyvets funqcionirebas. P
mesame tipis pirimidinis nukleotidi, kerZod, Timidilis mJava anu
Timidinmonofosforis mJava, mxolod dezoqsiribozuli formiT aris
warmodgenili. nukleotidebis dezoqsiribowarmoebulni Sesabamisi
ribowarmoebulebisagan miiRebian maTi aRdgenis saSualebiT. am aRdgenis reaqcias
NADP da specialuri wyalbadgadamtani cila-Tiorediqsini anxorcielebs. maTi
moqmedebis Sedegad miiReba nukleozidlifosfati, romelsac ribozis me-2
naxSirbadis hidroqsilis jgufi aRdgenili aqvs wyalbadamde. amitom,
Timidinmonofosforis mJavis warmoqmnisas, uridindifosforis mJava ( UDP) jer
gardaiqmneba dezoqsiuridindifosforis mJavad (dUDP). Semdeg mas Camoscildeba
erTi fosfati da bolos, miRebuli dezoqsiuridinmonofosforis mJavidan (dUMP),
misi meTilirebiT, warmoiqmneba dezoqsiTimidinmonofosforis mJava (dTMP).
meTilis jgufis donors ukanasknel reaqciaSi, meTilentetrahidrofolis mJava
warmoadgens:

O O

NH NH CH3
UDP dUDP dUMP

O O O O
N N
CH2 O P OH CH2 O P OH
O O
OH OH

uridinmonofosforis Timidinmonofosforis
mJava (UMP) mJava

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 purinisa da pirimidinis fuZeebis daSla

adamianis organizmSi purinebi qsantinamde iSlebian. es ukanaskneli, Semdgom

Sardis mJavad gardaiqmneba da am formiT gamoidevneba organizmidan.

OH

C
N C
C OH

OH C C
N
Please Register N
Sardis mJava
Sardis mJavis raodenoba, romelic dRe-RameSi organizmidan gamoidevneba, 0,6
gr. Seadgens. xSir SemTxvevebSi, da gansakuTrebiT xandazmul asakSi, organizmis
garkveul nawilebSi, ufro metad saxsrebSi (nekrisis qarebi) da TirkmelebSi,
Sardis mJavis marilebis (uratebis) gamoleqva xdeba. am paTologiis gamomwvevi
mizezi dReisaTvis cnobili ar aris. fiqroben, rom is SesaZlebelia gamowveuli
iyos, purinebis cvlaSi monawile fermentebis aqtivobis daqveiTebiT. aseTi
avadmyofebisaTvis mizanSewonilia dieta, romelic gamoricxavs nukleotidebiT
mdidar sakvebs (mag. RviZls, aseve yavas da Cais) radgan maT SemadgenlobaSi
Semavali kofeini da Teobromini purinebs warmoadgenen. Terapiuli efeqti
SesaZlebelia miRweul iqnas im farmakologiuri preparatebis miRebiT, romelic
purinebis damSleli fermentebis inhibitorebs warmoadgenen (mag. alopurinoliT).
pirimidinis fuZeebi iSlebian -alaninamde da -aminoerbos mJavamde.

nukleinis mJavebis da cilis biosinTezi

qromosomebis da genebis struqtura

ujreduli dnm-is 95% birTvSia moTavsebuli, romelic mosvenebul

mdgomareobaSi, interfazaSi, nukleoplazmaSia gabneuli. ormagspiraluri dnm-is
molekula eukariotul ujredebSi dakavSirebulia cilebTan da warmoqmnis
qromatinis Zafebs, qromatinis Zafebi garkveul ubnebSi mimagrebulia birTvis
matriqsze. gamravlebis dawyebis win birTvuli matriqsis bade irRveva, qromatini

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mkvrivdeba da warmoqmnis qromosomebs, romelTa raodenoba gansazRrulia yoveli
saxeobisaTvis.
qromatini Sedgeba 60% cilisagan, 35% dnm-sagan da 5% rnm-sagan. qromatinis
Zafebi qromosomebSia daxveuli, warmoqmnis mravalricxovan kvanZebs da yulfebs.
dnm-i qromatinSi mWidrodaa dakavSirebuli garkveuli tipis, dadebiTad
damuxtul cilebTan, romlebsac histonebi ewodebaT. histonebis funqcias
warmoadgens dnm-is molekulis mowesrigeba-organizacia struqturul
erTeulebSi, nukleosomebSi. arsebobs xuTi tipis sxvadasxva histoni. histonTa
oTxi tipi warmoqmnis gulas, multihistonur cilovan kompleqss, romlis
irgvlivac daxveulia dnm-is ormagspiraluri jaWvi da romelic warmoadgens
nukleosomis erT EerTuls. nukleosomebs Soris dnm-is jaWvi gaSlilia da igi
dakavSirebulia mxolod erTi tipis, H1-tipis histonTan. nukleosomaSi
warmodgenili dnm Sedgeba daaxloebiT 200 wyvili komplementaruli dnm-is
fuZeebisagan. histonebis garda qromatini Seicavs sxva tipis cilebsac.
dnm-is funqciur erTeuls geni warmoadgens. geni aris dnm-is fragmenti,
nukleotidTa Tanmimdevroba, romelSic Cawerilia informacia erTi polipeptidis
Sesaxeb. ganasxaveben genebis struqturul da regulatorul ubnebs.
struqturuli ubnebi warmoadgenen genis im nawils, romelSic Cawerilia
informacia polipeptidis Sesaxeb, xolo regulatoruli genebi ki, rTaven an
TiSaven struqturul erTeulebs. mravali struqturuli genis nukleotiodur
TanmimdevrobaSi CarTuli arian ubnebi, romlebic informaciis matarebelni ar
arian da ar translirdebian. genis aseT ubnebs intronebi ewodeba da maTi ricxvi
erT genSi SesaZlebelia 50-s aRematebodes. struqturuli genis im ubnebs,
romlebic translirdebian da romlebSic Cawerilia informacia cilis
pirveladi struqturis Sesaxeb, egzonebi ewodebaT. intronebis sigrZe
SesaZlebelia aRematebodes genis mTliani Tanmimdevrobis 80%-s. genis
transkripciis Semdeg, Sesabamisi rnm-dan intronebi amoiWrebia\n, ris Sedegadac
rCeba mxolod informaciis matarebeli genis nawili. am process splaisingi
ewodeba.
dReisaTvis mTlianad gaSifrulia adamianis genebis nukleotidTa
Tanmimdevroba.

dnm-is biosinTezi

replikacia. genetikuri informaciis gadacemis erT-erT mTavar etaps,

romlis drosac adgili aqvs dnm-is molekulis jaWvebad dayofas da TiToeul

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jaWvze komplementaruli dnm-is sinTezs, replikacia ewodeba. dnm-is replikacia
umTavresad, ujredis gamravlebis dros mimdinareobs. am process aseve adgili
aqvs rekombinaciis, (genTa gadaadgileba, erTi genis meoresTan dakavSireba da a.S.)
reparaciis (dazianebuli ubnis aRdgena da a.S.) dros.
replikaciis procesi SesaZlebelia sam stadiad gaiyos. I stadias dnm-is
ormagi spiralis daSla da mis TiToeul jaWvze komplementaruli dnm-is
sinTezis dawyeba warmoadgens. am stadias iniciaciis stadia ewodeba. II stadias
Seadgens procesi, romlis drosac dnm-is TiToeul jaWvze (matricaze)
sinTezirdeba Svileuli dnm-i. am process elongaciis stadia gansazRravs da
bolos, III terminaciis stadiaze dnm-is sinTezi mTavrdeba.
replikaciaSi monawileobs mravali cila da fermenti, romelTa funqciac
bolomde gaSifruli ar aris. replikacia iwyeba dnm-is ormagspiraluri jaWvis
gaSliT, romlis drosac komplementaruli jaWvebis erTmaneTisagan daTiSva
warmoebs. am process, specialuri fermentebi – helikazebi (helix-ingl.-spirali)
anxorcieleben. dnm-is jaWvebis urTierTdaTiSvis Semdeg, maT ukavSirdebian
specialuri dnm – damakavSirebeli cilebi. romlebic steriulad ewinaaRmdegebian
daTiSuli komplementaruli jaWvebis ukudakavSirebas. amis Semdeg dnm-is orive
jaWvi misawvdomi xdeba replikaciuri sistemisaTvis, romelic dnm-is sinTezs
iwyebs.
gaSlil jaWvebze Svileuli dnm-is sinTezi pirdapir ar iwyeba da mas win
uswrebs mcire zomis (4-10 nukleotidis) mqone komplementaruli rnm-is anu
praimeris warmoqmna, romelsac fermenti praimaza anxorcielebs. mxolod amis
Semdeg, praimeris miyolebiT, iwyeba dnm-is sinTezi, romelsac fermenti dnm-
polimeraza III atarebs. radgan, dnm-is molekula ori antiparaleluri jaWvisagan
Sedgeba, amitom dnm-polimeraza III ori mimarTulebiT aSenebs matricebze
Sesabamis komplementarul jaWvebs.
dnm-is molekula marjvniv daxveul spirals warmoqmnis da amitom misi
jaWvebis gancalkevebisaTvis saWiroa spiralis gasworeba, marcxniv Semotrialeba.
am process fermenti topoizomeraza awarmoebs da mis moqmedeba win uswrebs
elongaciis anu jaWvis dagrZelebis procesSi dnm-polimerazis moqmedebas.
dnm-is jaWvs, romelic replikaciuri sistemis moZraobis mimarTulebiT
sinTezirdeba, wamyvani jaWvi ewodeba, xolo jaWvs, romlis sinTezic
sawinaaRmdego mimarTulebiT warmoebs – CamorCenili jaWvi. wamyvani jaWvi
mTlianad sinTezirdeba, CamorCenili ki – fragmentalurad (okazakis fragmentebi).
am fragmentebis mierTeba specialuri fermentuli sistemis – dnm-ligazis --
saSualebiT xdeba.

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 dnm

ZiriTadi jaWvi Svileuli jaWvi

replikacia

dnm-is sinTezi rnm-is matricaze, ukutranskripcia. zogierTi tipis

virusebs, kerZod, e.w. onkovirusebs (rausis sarkomis virusi, rauSeris virusi),
Sidsis viruss da sxva, aqvT fermenti, romelic rnm-dan dnm-is sinTezs awarmoebs.
am ferments rnm-damokidebuli dnm-polimeraza anu ukutranskriptaza anu
revertaza ewodeba, xolo aseT rnm-s Semcvel virusebs – retrovirusebi.
virusuli rnm-iT ujredis inficirebis dros, misi ”maspinZlis” citoplazmaSi
moxvedrisas, revertazis saSualebiT rnm-is molekulidan dnm-is sinTezi iwyeba.
Sedegad warmoiqmneba hibriduli molekula, romelic Sedgeba erTi jaWvi dnm-isa

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da erTi jaWvi rnm-isagan. SemdgomSi rnm-is jaWvi iSleba da darCenili dnm-is
jaWvze warmoiqmneba komplementaruli dnm-is meore jaWvi, romelic virusuli
rnm-is srul asls warmoadgens. aseTi tipis dnm-s k-dnm-i ewodeba
(komplementaruli dnm-i), radgan igi rnm-is komplementarulad warmoiqmneba.
virusuli k-dnm-i CaerTveba “maspinZlis“ dnm-Si da masTan erTad replicirdeba.
onkovirusebis moqmedebis molekuluri safuZvlebis garkveviT dadginda
avTvisebiani simsivneebis warmoqmnis zogierTi meqanizmi. aRmoCnda, rom
onkovirusebis k-dnm-i warmoadgens e.w. onkogens, romlis aqtivaciac iwvevs
ujredis araregulirebad proliferacias da xSir SemTxvevebSi mis avTvisebian
gadagvarebas. onkogenebis translaciis produqts e.w. onkocila warmoadgens.
onkocilebi, maTi warmoqmnis Semdeg moqmedeben genTa eqspresiis regulaciis
aparatze da iwveven mis darRvevas. onkocilebis umravlesobas (Tirozin)
proteinkinazuri aqtivoba aRmoaCndaT.
DdReisaTvis mravali aseTi tipis onkogenia aRmoCenili. arsebobs mosazreba
imis Sesaxeb, rom onkogenebi adamianisa Tu cxovelis genetikur aparatSi
virusebidan moxvdnen evoluciis procesSi da Taobidan TaobaSi gadadiodnen
ZiriTad genetikur masalasTan erTad. isini normalur mdgomareobaSi
represirebulni arian, magram Tu moxda maTi derepresia (aqtivacia) sxvadasxva
kanrcerogenuli nivTierebebiT SesaZlebelia, MmaT gamoiwvion
arakontrolirebadi gamravleba. zogierTi onkogeni aparatis normalur
regulaciaSia CarTuli da monawileobas iRebs diferencirebisa da
proliferaciis stadiebis monacvleobaSi. MmaT didi mniSvneloba eniWebaT
organizmis embrionalur ganviTarebaSi. Aam tipis onkogenebis mutaciis Sedegad
Sesabamisi onkocilebi kargaven genetikuri aparatis maregulirebel unars, ris
gamoc ujredebi trasformirdebian.
ukutranskriptazebis aRmoCenis Sedegad dadginda, rom informaciis gadacema
ujredSi mxolod pirdapiri mimarTlulebiT ar warmoebs da SesaZlebelia
arsebobden misi ukugadacemac:

transkripcia translacia
dnm rnm cila
ukutranskripcia

reparacia. mutacia da rekombinacia. qromosomebi inertul stabilur

struqturebs ar warmoadgenen. isini, garemos zegavleniT xSirad ganicdian
cvilebebs. magaliTad, replikaciis Sedegad, dnm-is molekulaSi xSirad

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warmoiqmnebian e.w. gawyvetebi, romlebic ori fermentuli sistemis, dnm-
polimeraza I da dnm-ligazis saSualebiT aRdgebian. dnm-is jaWvis daRrveuli
monakveTis aRdgenas reparacia ewodeba.
dnm-is struqturuli dazianeba SesaZlebelia moxdes ultraiisferi,
maionizirebuli, rentgenuli da sxva myisi gamosxivebebiT. aseTi tipis
gamosxiveba iwvevs biologiuri molekulebidan, da maT Soris dnm-dan,
eleqtronebis gamoZevebas, rasac mohyveba metad arastabiluri, maRalreaqtiuli
da maRalenergetikuli ionebisa da Tavisufali radikalebis warmoqmna. aseTi
tipis radikalebs SeuZliaT Secvalon dnm-is molekulaSi Semavali fuZvebis
struqtura. magaliTad, SesaZlebelia moxdes mezoblad mdebare ori Timinis
urTierTdakavSireba kovalenturi bmiT, ris gamoc dnm-polimeraza veRar
axerxebs replicirebas. reparaciuli sistemebis saSualebiT xdeba aseTi
dimerebisE amokveTa da maT magivrad normaluri fuZeebis Casma.
dnm-is dazianeba SesaZlebelia moxdes agreTve naerTebiT, romlebic gare
samyarodan xvdebian adamianis organizmSi. aseTi tipis narTebi mravali tipis
daavadebaTa mizezs warmoadgenen. naerTebi, romlebsac madezaminirebeli,
maalkilirebeli da sxva qimiuri zegavlena aqvT, SesaZlebelia dnm-is dazianebis
mizezi gaxdnen. xSir SemTvevebSi aseTi dazianebebi aRdgebian reparaciuli
sistemebis saSualebiT.
miuxedavad organizmSi arsebuli reparaciul sistemaTa simZlavrisa,
dazianebaTa nawili SesaZlebelia aRudgeneli darCes da gadaeces memkvidreobiT
STamomavlobas. aseTi tipis mudmiv cvlilebas mutacia ewodeba. xSirad mutacia
dnm-is mxolod erTaderT fuZes Seexeba (wertilovani mutacia) da mas cilis
molekulaze didi gavlena ara aqvs; erTi aminomJavis Secvla meoreTi cilis
funqcionaluri Tvisebebis Secvlas iSviaTad iwvevs. magram, Tu aseTi mutacia
Seexo fermentis aqtiur centrSi Semaval aminomJavas, da es fermenti erT-erT
ZiriTad metabolur gzas emsaxureba, maSin wertilovani mutacia leTaluri
xdeba. nivTierebebs, romlebic mutaciebs iwveven mutagenebi ewodebaT. mutagenebis
umravlesoba, amave dros kancerogenul nivTierebebs warmoadgenen.
genebi xSirad icvlebian erTmaneTSi an erTiandebian. aseTi Secvlili
qromosoma ar kargavs replikaciis Tu transkripciis unars. genebis aseT
gadaadgilebas rekombinacia ewodeba. calkeul genebs an genTa jgufebs,
romlebic qromosomis erTi nawilidan SesaZlebelia Caisvas specialuri
fermentebis saSualebiT qromosomis sxva nawilSi, mobiluri genebi ewodebaT.
genetikuri rekombinaciis erT-erT saintereso magaliTs warmoadgens B-
limfocitebSi antisxeulebis masinTezirebeli genebis formireba. cnobilia, rom

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antisxeuli warmoiqmneba organizmSi moxvedrili ucxo makromolekulis-antigenis
mimarT, romelTa raodenoba bunebaSi SesaZlebelia milionobiT iyos. bunebrivia,
rom B-limfocits ar gaaCnia yvela am antigenis Sesatyvisi antisxeulis
warmosaqmneli daprogramirebuli genetikuri masala. amitom, antisxeulebis
masinTezirebeli genebi ganicdian mravaljerad rekombinacias, raTa miRweul
iqnas antisxeulis is Sesabamisi struqtura, romelic komplementaruli iqneba
saTanado antigenis mimarT. antisxeulebs ori tipis ebani aqvT: variabeluri da
konstanturi. swored, variabeluri ubnis maprogramirebeli genebi ganicdian
mravaljerad rekombinacias.

rnm-is biosinTezi

m-rnm-is biosinTezi. genetikuri informaciis nakadi, romelic normalur

ujredSi mimdinareobs da romelic genTa eqspresiis saxeliT aris cnobili,
ZiriTad stadiad moicavs process, romelsac transkripcia ewodeba.
transkripciis procesi ganapirobebs genTa diferencirebul aqtivobas, ujredTa
populaciebis qsovilebad Camoyalibebas da maT normalur funqcionirebas.
transkripciis procesSi dnm-is erT-erT jaWvze komplementaruli rnm-I aigeba. igi
SesaZlebelia moicavdes erTi an ramdenime genis erTobliobas. warmoqmnili m-rnm-
idan, romelsac m-rnm (matriculi) ewodeba, SemdgomSi cila sinTezirdeba.
rogorc ukve iTqva, tipiuri geni Seicavs ubans, romelSic Cawerilia
informacia cilis pirveladi stuqturis Sesaxeb (makodirebeli ubani) da
regulatorul ubans, romelic pirvelis CarTvas iwvevs. regulatoruli ubani –
ori tipis arsebobs: promotori da enhanseri. promotori uSualod struqturuli
(makodirebeli) ubnis maxloblad mdebareobs, xolo enhanseri – SesaZlebeli
dacilebuli iyos mas.
promotoris da enhanseris gaaqtivebas da Sesabamisad struqturuli ubnis
CarTvas, awarmoeben specialuri cilebi, romlebic specifiurad ukavSirdebian
genis regulatorul elementebs. am cilebs transkripciis faqtorebi ewodebaT
da dReisaTvis maTi 200-mde warmomadgenelia aRmoCenili. maT raodenobas ujredSi
mravali piroba gansazRvravs: hormonebis moqmedeba, ujredis diferencirebis
xarisxi da a.S.
promotorebi da enhansrebi moicaven dnm-is ramdenime ubans, Semdgars
specifiuri nukleotidTa Tanmimdevrobisagan. TiToeul promotors an enhansers
aseTi specifiuri ubani SesaZlebelia 2 da ufro meti hqondes. maT motivi
ewodeba da TiToul motivs erTi transkripciis faqtori ukavSirdeba. xSirad

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struqturuli genis asamoqmedeblad aucilebelia promotorTan an enhanserTan
ramdenime transkripciuli faqtoris erTblivi dakavSireba.
promotoris gaaqtivebis Sedegad genis struqturuli ubani iwyebs
transkripcias. am process fermenti rnm-polimeraza II anxorcielebs. igi dnm-is
erT-erT jaWvze komplementaruli nukleotidebisagan Semdgar rnm-is jaWvs agebs.
sinTezi mimdinareobs 3’ 5’ mimarTulebiT. rnm-polimeraza II mTlianad mTeli
genis Sesatyvis rnm-s asinTezirebs e.i. iRebs asls, rogorc egzonebisagan, ise
intronebisagan, radgan intronebi Seicaven iseT Tanmimdevrobebs, romlebic
cilis kodirebas ar awarmoeben. isini birTvSive amoiWrebian specialuri sistemis
saSualebiT, romelsac mcire birTuli ribonukleoproteidebi ewodebaT.
intronebis amoWris Semdeg, rnm-is egzonebi erTmaneTs uerTdebian (splaisingi) da
warmoqmnian funqciur erTeuls, romlisganac cilis translacia warmoebs. dnm-is
sinTezis mzgavsad, rnm-is sinTezic Seicavs iniciaciis, elongaciis da
terminaciis stadiebs.
t-rnm-is biosinTezi. transportuli t-rnm-i ujredSi adaptorul funqcias
asrulebs. misi saSualebiT m-rnm-is translaciis SeuRleba xdeba cilis
pirvelad struqturasTan. t-rnm-i Sesabamisi dnm-dan sinTezirdeba, xSirad erTi
didi winamorbedis formiT, romelic SemdgomSi ramdenime t-rnm-ad iSleba. t-rnm-
i dnm-is jaWvze sinTezis Semdeg, modifikacias ganicdis: mas uerTdeba boloSi
cca Tanmimdevroba da aminomJavis aqceptoruli ubani.
r-rnm-is biosinTezi ribosomuli r-rnm-is sxvadasxva molekuluri formebi,
romlebic ribosomebis SenebaSi iReben Mmonawileobas erTi saerTo
maRalmolekuluri winamorbedidan warmoiqmnebian. am maRalmolekuluri rnm-is
geni qromosomaSi ramdenime asliT aris warmodgenili, rac metyvelebs ujredSi
ribosomuli rnm-is gamomuSavebis saWiroebaze.

cilis sinTezi da misi regulacia

genetikuri infomaciis gadacemis erT-erT ZiriTad process translacia

warmoadgens. translaciis Sedegad dnm-is molekulaSi Cawerili informacia
cilis molekulas gadaecema aminomJavaTa Tanmimdevrobis saxiT. igi ciklur
energodamokidebul da mravalsafexurian process warmoadgens, romlis drosac
Tavisufal aminomJavaTa polimerizacia warmoebs genetikurad determinirebul
TanmimdevrobaSi. cilis sinTezSi 200-mde sxvadasxva tipis makromolekula iRebs
monawileobas (rnm-ebi, fermentebi da sxva). es procesi pirobiTad or etapad
SeiZleba gaiyos: aminomJavebis aqtivirebisa da TviT translaciis etapi.

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 aminomJavaTa aqtivireba. cilis sinTezisaTvis aucilebel pirobas
aminomJavaTa aqtivireba warmoadgens, romelsac fermenti aminoacil-t-rnm-
sinTetaza awarmoebs. am reaqciis dros aminomJava Sesabamis transportul rnm-s
ukavSirdeba:

R-CHNH2-COOH + ATP + t-rnm R-CHNH2-CO-t-rnm + AMP + PPi

yoveli aminomJavis t-rnm-i cca TanmimdevrobiT bolovdeba. aminomJava

karboqsilis jgufiT adeninis hidroqsilis jgufTan maRalenergetikul eTerul
kovalentur bmas warmoqmnis. aseTi gaaqtivebuli formiT erTveba aminomJava
translaciis procesSi. am naerTs aminoacil-t-rnm-i ewodeba.
translaciis procesi. translaciis procesi sam stadiad iyofa: iniciacia,
elongacia da terminacia. iniciaciis dasawyebad aucilebelia: I-matricul m-rnm-s
hqondes iniciatoruli (CamrTveli) kodonebi, II-iniciatoruli (damwyebi)
aminoacil-t-rnm-i da III-iniciaciis cilovani faqtorebi.
baqteriebSi iniciatorul t-rnm-s (t-rnm-s, romelic pirveli ukavSirdeba
matricul rnm-s) formilmeTionil-t-rnm-i warmoadgens. formilmeTioninSi,
meTioninis aminojgufi formilirebulia e.i. masTan dakavSirebulia
WianWvelamJavas naSTi (HCOOH). amis gamo, pirveli aminomJavis aminis jgufi
blokirebulia da igi cilis sinTezSi monawileobas ar iRebs. pirveli
aminomJavis aminis jgufis blokireba aucilebelia imisaTvis, rom cilis sinTezi
daiwyos iniciatoruli aminomJavis karboqsilis jgufidan. cilis mTliani
molekulis formirebisa da sinTezis procesis damTavrebis Semdeg iniciatoruli

aminomJava sZvreba cilovan molekulas da momdevno, meore aminomJavis NH2-bolo

Tavisufldeba. amitom miRebulia, rom cilis sinTezi iwyeba NH2-bolodan.

eukariotul ujredebSi iniciatorul aminomJavas Cveulebrivi meTionini
warmoadgens.
cilis sinTezis dawyebis erT-erTi aucilebeli pirobaa agreTve is, rom
birTvidan gadmosul m-rnm-s iniciatoruli kodoni gaaCndes. es ukanaskneli,
iniciatoruli aminomJavis t-rnm-s unda scnobdes. pirvelad, m-rnm-i ribosomis
mcire (3O S) suberTeuls ukavSirdeba. am suberTeulSi iniciatoruli m-rnm-is
kodoni da iniciatoruli aminomJavis t-rnm-is antikodoni ukavSirdebian
erTmaneTs. mxolod amis Semdeg, ribosomis mcire suberTeuls didi (50 S)
suberTeuli uerTdeba.
ribosomebis suberTeulebis dakavSirebaSi, iseve, rogorc iniciatoruli

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kodon-antikodonis urTierTmoZebna–dakavSirebaSi wamyvan rols iniciaciis
cilovani faqtorebi asruleben (aRiniSnebian-eIF). dReisaTvis 10-mde aseTi
faqtoria aRmoCenili, romelTa Soris wamyvani mniSvneloba eniWeba eIF-2, eIF-3 da
eIF-5-s. eIF-3 monawileobas iRebs ribosomebis suberTeulebis urTierT
dakavSirebaSi, xolo eIF-2 - GTP-s hidrolizSi (es ukanaskneli ribosomebis mTavar
energetikul wyaros warmoadgens).
miuerTdeba-ra erTmaneTs iniciatoruli kodon-antikodoni, agreTve,
ribosomebis suberTeulebi, warmoiqmneba funqciurad aqtiuri 70 S ribosoma,
romelic iwyebs elongaciis process anu cilis heptiduri jaWvis warmoqmnas.
elongaciis process awarmoebs ribosomis 50 S suberTeuli, romelsac t-rnm-
is dasakavSireblad ori centri aqvs: aminoaciluri da heptiduri. Sesabamisi
aminomJava pirvelad aminoacilur centrs ukavSirdeba, xolo misi t-rnm-uli
nawili ki - antikodonis mixedviT kodons. amis Semdeg ribosomis heptiduri
centridan aminoacilurze gadmoitaneba iniciatoruli aminomJava (mag.
formilmeTionini) da peptidiltransferazis saSualebiT ukavSirdeba peptiduri
bmiT axlad Semosul aminomJavas. paralelurad ribosomidan iniciatoruli
aminomJavis (mag. formilmeTioninis) t-rnm-i gaidevneba da peptiduri centri
Tavisufldeba. am peptidur centrze aminaciluridan gadaitaneba iniciatoruli
aminomJavisa da meore aminomJavisagan Semdgari komleqsi, romelTanac
dakavSirebulia t-rnm-i (translokacia). am kompleqsis gadatanas ganapirobebs
ribisomis moZraoba matriculi rnm-is gaswvriv. es ukanaskneli, am dros erTi
kodoniT (3 nukleotidiT) gadainacvlebs. translokaciis Sedegad aminoacilur
centrSi arsebuli t-rnm-i Sesabamisi heptiduri jaWviT aRmoCndeba peptidur
centrSi. am procesebis Semdeg aminoaciluri centri mzadaa mesame aminomJaviani
t-rnm-is misaRebad.
erTi peptiduri bmis warmosaqmnelad elongaciis stadiaze ixarjeba 4
makroerguli fosfatis tolfasi energia: 2 ATP da 2 GTP; daaxlovebiT 14
kkal/molze (2 ATP ) aris saWiro erTi aminomJavis gasaaqtiveblad, 1 molekula
GTP ixarjeba aminoacil-t-rnm-is aminoacilur centrTan dasakavSireblad, xolo
erTi molekula GTP - translokaciis procesisaTvis.
cilis sinTezi mTavrdeba terminaciis stdadiiT. matricul rnm-s boloSi
terminatoruli kodoni aqvs (msg. u-a-g an u-a-a), romlis Sesatyvisi antikodonic
ujredSi ar arsebobs. amitom, mis adgils ribosomaze ikavebs terminaciis
cilovani faqtori (mag. faqtori R eukatriotebSi). amis Semdeg, peptiduri jaWvi
Camoscildeba bolo aminomJavis t-rnm-s da sinTezirebuli cila ribosomidan

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Tavisufldeba. erTi matriculi rnm-i SesaZlebelia ganicdides translacias
ramdenime ribosomaze. aseTi tipis dajgufebebs polisomebi ewodebaT.
sinTezirebuli cilebis transporti membranebSi. cilebis im nawils,
romelsac ujredi awarmoebs sxva ujredebisa Tu qsovilebisaTvis (mag. insulins-
-ujredebi an trifsinogens - kuWqveSa jirkvlis ujredebi) da romlebsac is
aTavisuflebs ujredgare sivrceSi, xSirad moepovebaT e.w. sasignalo
Tanmimdevrobani. aseTi sasignalo Tanmimdevrobani Cveulebriv, cilis
dasawyisSive sinTezirdeba (maTi raodenoba 20-30 aminomJavas ar aRemateba) da
Sedgeba umTavresad hidrofobuli aminomJavebisagan. maTi saSualebiT
sinTezirebuli cila gadis endoplazmuri retikulebis membranebs, goljis
aparatSi warmoebs misi vezikulebSi lokalizeba da igi sekretirdeba ujredgare
sivrceSi.
mitoqondrialuri cilebis sinTezi. mitoqondriebi Seicaven sakuTar dnm-s
da cilebis sinTezis mTel aparats: ribosomebs, m- da t-rnm-ebs. unda aRiniSnos,
rom mitoqondriebSi, baqteriebis mzgavsad iniciatorul aminomJavas, xSirad
formilmeTionini warmoadgens.
cilebis postsinTezuri cvlilebebi. cila ribosomebze sinTezis Semdeg
garkveul cvlilebebs ganicdis. magaliTad, fermentebi, romlebic aqtivobis
gamosamJRavneblad saWiroeben prostetul jgufs, ierTeben mas. cilis sinTezis
Semdeg cilis molekulaSi ama Tu im funqciuri jgufis CarTvas, cilis
posttranslaciuri modifikacia ewodeba. posttranslaciuri modifikacia
SesaZlebelia iyos kovalenturi da arakovalenturi. Tavis mxriv, kovalenturi
modifikacia SesaZlebelia iyos Seqcevadi da Seuqcevadi.
Seuqcevadi kovalenturi modifikaciebia: disulfiduri bmebis warmoqmna,
cilis molekulaSi prostetuli jgufebis mierTeba, misgan sasigna;lo
Tanmimdevrobebis CamoWra, cilis aminomJavebis acetilireba da sxva.
Seqcevadi kovalenturi modifikaciebi arin: fosforilireba, ADP-
ribozilireba, amidireba da a.S. Seqcevadi kovalenturi modifikaciebi did rols
TamaSoben ujredis hormonalur regulaciaSi

cilis sinTezis regulacia

cocxali organizmis arsebobis ZiriTad pirobas warmoadgens

regulaciis natifi sistemebis arseboba, romelic ganapirobebs bioqimiur
reaqciaTa warmarTvas Sesabamis drosa da sivrceSi. regulaciuri procesebis

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swori mimdinareoba aucilebelia ujredTa specializaciisTvis, qsovilTa
diferenciaciisTvis, embrionuli da postembrionuli ganviTarebisTvis.
adamians, iseve rogorc yovel biologiur saxeobas genTa kompleqti
saerTo aqvs mis yovel ujredSi. qsovilTa specializaciis mixedviT, genTa am
kompleqtidan aqtiur mdgomareobaSi imyofebian mxolod is genebi, romlebic
awarmoeben am ujredTaTvis damaxasiaTebel makromelekulaTa sinTezs.
magaliTad, kanis ujredebi, keratinocitebi, gansxvavdebian RviZlis
ujredebis – hepatocitebisagan imiT, rom masSi eqspresirebul
(gamomJRavnebul) mdgomareobaSi arian is genebi, romlebic hepatocitebSi
represirebul (daTrgunul) mdgomareobaSi imyofebian, da piriqiT. yoveli
tipis ujredis samuSao mdgomareobaSi myofi genebi gansazRvraven ujredis
struqturas, formasa da funqcias. igive keratinocitebSi, genTa
diferencirebuli muSaobis Sedegad warmoiqmneba cila keratini, xolo
hepatocitebSi - sisxlis Sratis cilebi, damJangveli mikrosomuli fermentebi,
da a.S. amrigad, ujredis damaxasiaTebeli funqcia, misi metabolizmis
Taviseburebani damokidebuli am ujredSi momuSave genTa nakrebze, romlebic
eqspresirdebian da represirdebian ujredis sicocxlis garkveul periodebSi.
genTa diferencirebuli aqtiobis sakiTxi molekuluri biologiis sakvanZo
problemas warmoadgens. garkveva im sakiTxebisa, Tu ra dros da ra
meqanizmebiT xdeba garkveuli genebis CarTva ujredSi, rogor xdeba
informaciis gadacema ujredSi, da kerZod birTvSi, ra bioqimiuri procesebi
gansazRvraven ujredebis specifikurobas da im erTaderT pasuxs, romelsac
ujredi hormonalur, antigenur Tu sxv. gaRizianebas upasuxebs- warmoadgens
Tanamedrove biologiis da medicinis erT-erT aqtualur sakiTxs.
jer kidev 60-ian wlebSi frangi mecnierebis Jakobisa da monos mier
gamoTqmuli iyo hipoTeza "operonis" Sesaxeb, romelic nawilobriv
ganmartavda genTa SerCeviTi aqtivobis sakiTxs, da romelic gansacvifreblad
winaswarmetyveluri aRmoCnda. am hipoTezis mixedviT genis aqtiuri nawili
Sedgeba ori monakveTisagan: struqturuli genisagan da gen-operatorisagan.
struqturuli geni warmoadgens genis im nawils, romelSic Cawerilia
informacia cilis pirveladi struqturis Sesaxeb, xolo regulatoruli
ubani anu gen-operatori gansazRvravs mis aqtivobas. Jakobisa da monos
mixedviT, rogorc struqturul genze, ise gen-operatorze xdeba m-rnm-is
transkripcia, romelic genis promotoridan iwyeba. gen-operatoris CarTva-
gamorTva xdeba represoris saSualebiT, romelic operatorTan
urTierTmoqmedebs. Tu represori operatorTan aris dakavSirebuli, maSin

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transkripciis procesi wydeba; da piriqiT, Tu represoris dakavSireba gen-
operatorTan SezRudulia, maSin transkripciis procesi aqtiuria da
struqturuli geni samuSao mdgomareobaSi imyofeba. represori cilas
warmoadgens, romelic specialuri genebis saSualebiT sinTezirdeba.
represorze, romelsac damuxruWebuli aqvs gen-operatori SeiZleba
imoqmedos induqtorma. induqtoris moqmedebis Sedegad represori scildeba
gen-operators da struqturul genze iwyeba aqtiuri transkripciis procesi.
induqtoris mier represoris moqmedebis neitralizacia anu derepresia
SeiZleba gagrZeldes manamde, sanam induqtoris koncentracia birTvSi maRalia.
induqtoris koncentraciis daqveiTebisas, represori uerTdeba gen-operators
da mTlianad geni gadadis represirebul mdgomareobaSi, igi wyvets muSaobas.
induqtori SesaZlebelia iyos rogorc dabalmolekuluri naerTi, aseve cila.
aseTi genebis erTobliobas, romelSic Sedian gen-operatori da
struqturuli geni an genebi, operonio ewodeba.

promotori gen-operatori struqturuli geni

represori

miuxedavad imisa, rom Jakobisa da monos Teoria principialurad swori

aRmoCnda, man ukanasknel wlebSi miRebuli mravali faqtis axsna ver SesZlo.
Tanamedrove gamokvlevebma cxadyves, rom genTa aqtivaciisTvis ar aris
sakmarisi mxolod misi derepresia, da rom induqtorebis garda arseboben
uamravi faqtorebi, romlebic genetikuri aparatis regulacias anxorcieleben.
dReisaTvis aRmoCenilia ramdenime aseuli tipis cila, romlebic e.w.
transkripciis faqtorebad an transkripciis aqtivatorebad arian
gaerTianebulni. transkripciuli faqtorebi ukavSirdebian dnm-is garkveul
Tanmimdevrobas, motivs promotorul an enhanserul ubanSi da iwveven genTa
eqspresias da transkripcias. zogierTi aseTi cilovani faqtori
individualurad moqmedebs, genebis umravlesobas ki aseTi faqtorebis
erToblioba esaWiroebaT, radgan isini ara mxolod dnm-is nukleotidur
Tanmimdevrobas ukavSirdebian, aramed erTmaneTsac. urTierTdakavSirebuli
faqtorebis dnm-is garkveul motivTan dakavSirebis Semdeg warmoiqmneba
aqtiuri transkripciuli kompleqsi.

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 mravali transkripciis faqtoris pirveladi struqtura gaSifrulia da
dagenilia maTi dnm-Tan urTierTdakavSirebis meqanizmebi. zogierTi maTganis
konformacia Seesatyviseba dnm-is ormag spirals, zogierTi amoicnobs
nukleotidTa garkveul Tanmimdevrobas, xolo zogierTi specifikuri
dakavSirebisTvis iyenebs sxvadasxva tipis dabalmolekulur naerTebs,
magaliTad TuTias. cila faqtorebi xSirad monomerebisagan Sedgebian,
romlebsac SeuZliaT warmoqmnan homodimerebi, an heterodimerebi msgavs
transkripciul faqtorTan.
transkripciuli faqtorebis moqmedeba maT koncentraciazea damokidebuli.
rac ufro Warbi raodenobiT arian isini birTvSi, miT ufro advilad rTaven
ama Tu im gens. amitom, faqtorebis moqmedeba maTi Sesabamisi genebis aqtiobiT
ganisazRvreba. Tu transkripciuli faqtoris geni intensiurad asinTezirebs
faqtors, es ukanaskneli birTvSi maRali koncentraciiT grovdeba, da iwvevs
garkveli tipis genebis CarTvas. transkripciis faqtoris genis mutaciis
Sedegad miRebuli cila, bunebrivia, veRar awarmoebs genTa eqspresiis
normalur regulacias. yovelive amis gamo, genTa eqspresiis procesi irRveva
da ujredi gadadis arakontrolirebad mdgomareobaSi, rasac xSirad ujredis
simsivnuri gadagvareba mohyveba. amitom, zogierTi transkripciis faqtoris
genebs proto-onkogenebi, anu onkologiuri transformaciis winamorbedi
genebi ewodebaT.
cilis sinTezis inhibitorebi. cilis sinTezis Zlier inhibitorebs
antibiotikebi warmoadgenen. isini moqmedeben replikaciis, transkripciis an
translaciis doneze da maTi efeqti specifikuria. magaliTad, antibiotiki
rifampicini, romelic tuberkulozis mkurnalobisas ixmareba, amuxruWebs
ferment rnm-polimerazas. igi gacilebiT ufro Zlierad moqmedebs baqteriul
fermentze, vidre adamianisaze, da amitom naklebad toqsikuria. tifozuri
infeqciebis winaaRmdeg moqmedi preparati – qloramfenikoli – baqteriebis
ribosomebSi mimdinare peptidiltransferazuli reaqciis inhibitoria.
antibaqteriuli preparatebi, streptomicini da neomicini, iwveven baqteriuli
rnm-is sinTezis procesSi “Secdomebis” warmoqmnas, ris gamoc producirdeba
anomaliuri cila, da baqteria iRupeba. tetraciklinis rigis antibiotikebi
ukavSirdebian ribosomebis aminoacilur centrs, xels uSlian masTan
transportuli rnm-is dakavSirebas, da amiT arRveven baqteriuli cilis
sinTezs. eriTromicinis msgavsi antibiotikebi ainhibireben translaciis
process translokaciis doneze.

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 cilis sinTezis inhibitorebs warmoadgenen agreTve simsivnis
sawinaaRmdego qimioTerapiuli antibiotikebi, romlebsac mkveTrad
gamoxatuli citostatikuri, antiproliferaciuli efeqti aqvT. maTi
umetesoba (mag. adriamicini, antramicini) dnm-is mcire RruSi arsebul
azotovan fuZeebs ukavSirdeba, ris gamoc fermenti topoizomeraza
inhibirdeba da adgili aqvs replikaciis gaCerebas.
cilis sinTezis erT-erT Zlier inhibitors difteriis toqsini
warmoadgens. igi iniciaciis cilovani faqtoris – eIF-2-is posttranslaciur
modifikacias, kerZod, ADP-ribozilirebas iwvevs. Sedegad am ukanasknels
ekargeba cilis sinTezSi monawileobis unari.
cilis sinTezis inhibitorebs warmoadgenen zogierTi mcenaris
toqsinebic. aseTi tipis cilebi ori suberTeulisagan Sedgebian, romelTagan
erTi plazmur membranaSi warmoqmnis arxs, xolo meore – ujredSi Sesvlis
Semdeg moqmedebs ribosomul aparatze da Trgunavs mas. MaTi tipiuri
warmomadgenelia abusalaTinis cila – ricini, fiTris cila – viskumini da
sxva.

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 adamianis bioqimia

adamiani evoluciis mwvervalze mdgom arsebas warmoadgens, da amitom misi

metabolizmi, nivTierebaTa cvla da regulacia gacilebiT rTulad aris
organizebuli cxoveluri samyaros sxva warmomadgenlebTan SedarebiT.
SesaZlebelia gamoiyos adamianisTvis damaxasiaTebeli bioqimiuri
organizaciis sami ZiriTadi aspeqti. pirvels warmoadgens misi organoebis,
qsovilebisa da ujredebis koordinirebuli muSaoba, meorea - masSi mimdinare
procesebis rTuli, ierarqiuli hormonaluri da nervuli regulacia, mesame
ki - nivTierebaTa cvlis Taviseburebani gare samyarosTan. ukanaskneli, wamyvan
da ZiriTad elementad kvebas moicavs, radgan igi adamianis qmediTunarianobas
gansazRvravs.

Uujredis regulaciis meqanizmebi

yoveli cocxali organizmis cxovelmoqmedebis ZiriTad principebs

regulaciis natifi sistemebis arseboba gansazRvravs. es sistemebi
ganapirobeben bioqimiuri reaqciebis warmarTvas Sesabamis drosa da sivrceSi.
regulaciis meqanizmebis arseboba aucilebelia ujredTa specializaciisTvis,
qsovilTa diferencirebisTvis, embrionaluri da postembrionaluri
ganviTarebisTvis.
ujredTa populaciebis Camoyalibeba da maTi koordinirebuli moqmedeba
ganpirobebulia ujredSorisi sakomunikacio sistemebis arsebobiT, romelSic
wamyvan rols ujredis mierve gamomuSavebuli, rTuli Tu martivi agebulebis
nivTierebebi asruleben. gamodian ra erTi tipis ujredebidan, isini
moqmedeben organizmis sxva nawilSi arsebul ujredebze da cvlian mis
metabolizms. yoveli maTgani efeqtorul (mimReb) ujredze specifiurad
moqmedebs da am ujredis pasuxi damaxasiaTebeli, adeqvaturia mxolod am
naerTis mimarT. mravali hormoni, mediatori, zrdis faqtori Tu sxva
romelime fiziologiurad aqtiuri nivTiereba, sisxlisa da limfis
saSualebiT vrceldeba mTel organizmSi, sadac Sesabamisi efeqtoruli
ujredebis specifikur pasuxs iwvevs. aseTi nivTierebebi ujredSoris
informaciis gadacemis ZiriTad sistemas warmoadgenen da maT pirveladi
sasignalo sistema ewodeba.
mravali mikro- da makroorganizmisTvis pirveladi sasignalo sistema
gansazRvravs qceviT reaqciebs, sqesobriv aqtivobas, garemosTan

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Segueblobas da sxva sacicoxlo funqciebs. qimiuri nivTierebebs,
romlebsac garemoSi gamohyofen erTi saxeobis organizmebi imave saxeobis
individuumebze zemoqmedebis mizniT, feromonebi ewodeba. feromonebi farTod
gavrcelebul sasignalo sistemas warmoadgens da ganapirobebs cxovelTa
samyaros mravali tipis populaciaTa qceviT aqtivobas.
TviT adamianis organizmSi, pirveladi sasignalo sistema gansazRvravs
organoTa koordinirebul moqmedebas, zrdisa da organizmis formirebis
process. am sistemebSi monawile ujredSorisi signalebi gansxvavdebian maTi
moqmedebis arealis da gavrcelebis mixedviT. umaRles cxovelebSi Sinagani
ujredSorisi sasignalo sistemebi aregulireben zrdis da qcevis procesebs,
nivTierebaTa cvlis da metabolur reaqciebs.
ganasxvaveben endokrinul, parakrinul da autokrinul sasignalo
sistemebs. endokrinuli sasignalo sistemis formirebaSi monawileobas
iReben morfologiurad gansazRvruli organoebi - endokrinuli
jirkvlebi. isini gamoimuSaveben naerTebs, romlebsac hormonebi ewodebaT da
romelTa moqmedebis samizne - efeqtoruli ujredi, sagrZnoblad
daSorebulia gammomuSavebel organosTan. endokrinul jirkvlebSi
sinTezirebuli hormoni sisxlis saSualebiT vrceldeba mTel organizmSi.
parakrinuli sasignalo sistemaSi mimRebi, efeqtoruli ujredi
sasignalo ujredis axlo imyofeba. amitom parakrinuli ujredis mier gamomu-
Savebuli naerTi moqmedebs mxolod adgilobriv, lokalur arealSi
myof ujredebze. parakrinuli sasignalo sistemebis tipiuri warmomadgenlebi
neirogadamcemebi arian, romelTa moqmedebis zRvari Semofarglulia nervuli
Tu kunTovani qsovilis erTi an ori ujrediT.
da bolos, ujredSirisi urTierTobis mesame tips warmoadgens
autokrinuli sasignalo sistema. am SemTxvevaSi, ujredis mier
gamomuSavebuli nivTiereba moqmedebs sakuTar receptorebze. autokrinul
regulacias adgili aqvs ujredis sxvadasxva fiziologiuri Tu
paTologiuri mdgomareobisas. mravali hormonismagvari naerTi, romlebic
aseT situaciebSi sinTezirdeba iwvevs autokrinuli ujredis zrdis da
ganviTarebis cvlilebebs.
ujredebi, pirveladi Suamavlebis (hormonebi, mediatorebi, zrdis
faqtorebi da sxva) saSualebiT miRebul inforamacias specialuri
receptoruli sistemebiT aRiqvaven. receptoruli sistemebi, informaciuli
molekulebis bunebidan gamomdinare, SesaZlebelia ramdenime tipisa iyvnen;
isini SesaZlebelia lokalizebuli iyvnen plazmur membranebze,

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citoplazmaSi an romelime sxva organoidze. Tu informaciuli milekulis
funqcias, efeqtorul ujredSi metabolizmis raodenobrivi cvlileba
warmoadgens, maSin igi upiratesad plazmur membranaze arsebul receptorebze
moqmedebs; xolo Tu naerTi ujredis xarisxobriv, funqciur maCveneblebs
cvlis da moqmedebis saboloo mizans ujredis genetikuri aparati
warmoadgens, maSin misi receptorebi, rogorc plazmur membranebze, aseve
citoplazmaSi SesaZlebelia iyvnen lokalizebuli.

plazmuri membranis receptorebi

plazmuri membranis receptorebi sami komponentisagan Sedgebian:

I - TviT receptori, II - SemakavSirebeli cila anu transduceri
(gardamqmneli) da III - ujredSida katalizuri komponenti.
receptori integralur membranul cilas warmoadgens. Cveulebriv igi,
hidrofoburi aminomJavebisagan Semdgari (7-8 -spiraluri hidrofobuli
domeni - monakveTi) cilaa, romelic membranas mTel sisqeze gasWolavs. misi
aqtiuri (liganduri) centri, romliTac igi hormons ukavSirdeba, ujredis
zedapirze mdebareobs. hormonis dakavSirebis Semdeg receptoruli cila
konformacias icvlis da gadadis "aqtivirebul" mdgomareobaSi.
aqtiuri konformaciis miRebis Semdeg receptori meore integralur cilas
- transducers ukavSirdeba. mas xSirad G cilasac uwodeben, radgan igi GTP-s
ikavSirebs. G-cila sami suberTeulisagan Sedgeba: , , da . receptorTan
dakavSirebis Semdeg G-cila suberTeulebad iSleba da erT-erTi maTgani,
kerZod , iwyebs moqmedebas ujredSida katalizur komponentze; paralelurad,
G-cila Slis GTP-s (GTP GDP + Pi).
G cila SesaZlebelia iyos gamaaqtivebeli (Gs) an Semakavebeli (Gi). Gs-is
katalizur komponentTan SeuRlebis Semdeg am ukanasknelis aqtivoba
Zlierdeba, xolo Gi-is SemTxvevaSi ki - qveiTdeba.
xSir SemTxvevebSi ujredSida katalizur komponents fermenti warmoadgens.
ZiriTadad, G-cilasTan urTierTmoqmedeben adenilatciklaza da
fosfolipazebi A2 da C. adenilatciklazis gaaqtivebis Sedegad miiReba
cikluri AMP:

ATP cAMP + PPi

cAMP - cikluri adenozinmonofosforis mJava

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 xolo fosfolipaza C-s gaaqtivebisas ki - diacilglicerini da
trifosfoinozitoli:

fosfatidilinozitfosfatiP diacilglicerini + trifosfoinozitoli

fosfolipaza A2-is gaaqtivebiT miiReba araqidonis mJava:

dosfatidilqolini araqidonis mJava + lizofosfatidilqolini

am reaqciebis Sedegad warmoqmnili cAMP, diacilglicerini (DAG),

trifosfoinozitoli (TPI) da araqidonis mJava (AA) ujredSi bioqimiuri
reaqciebis gaZlierebas an Sekavebas iwveven, da amitom, maT ujredSida
regulatorebi an meoradi Sumavlebi ewodebaT. aRweril SemTxvevebSi
hormonis, mediatoris Tu sxva romelime tipis pirveladi Suamavlis moqmedeba
SesaZlebelia Semdegnairad warmovidginoT:

hormoni receptori meoradi Suamavali ujredis pasuxi

radgan aseTi tipis receptorul sistemebSi ujredis metaboluri cvli-

lebebi xdeba, amis gamo maT metabotropul receptorebs uwodeben.
metabotropul receptorebSi hormonis moqmedebiT ganpirobebuli ujreduli
pasuxi drois SedarebiT xangrZlivi periodis Semdeg gamomJRavndeba
(milisekundebi da wamebi), amitom aseTi tipis receptorebs, agreTve, nel
receptorebsac uwodeben.
plazmuri membranis receptorul sistemebSi receptoruli cila G-cilis
meSveobiT an uSualod, SesaZlebelia SeuRlebuli iyos ionur arxTan. xSir
SemTxvevebSi, receptoruli cila TviTon warmoqmnis arxs. am SemTxvevaSi mas
8 -spiralis mqone hidrofoburi domeni aqvs, romelTa saSualebiTac
receptori gasWolavs membranas. aRniSnuli domenebi ise lagdebian
membranaSi, rom Sua nawilSi warmoqmnian foras. am foraSi anu arxSi, misi
gaxsnis SemTxvevaSi, gadian ionebi maTi koncentraciuli gradientis mixedviT:
Na, Ca da Cl ujredSorisi siTxidan Sedian ujredSi, xolo K - ujredidan
gamodis gareT. aseTi tipis receptorebis hormoniT gaaqtivebis Sedegad
adgili aqvs ionTa gadanawilebas ujredSoris siTxesa da ujredis
citoplazmas Soris da amitom maT ionotropuli receptorebi ewodebaT. am
receptorebis gaaqtivebas ujredis swrafi fiziologiuri pasuxi mohyveba

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(wamis meaTasedebSi) da amitom maT swraf receptorebsac uwodeben.
ionotropul receptorebs msgavsi domenuri agebuleba aqvT, isini
homologiuri aminomJavuri TanmimdevrobiT xasiaTdebian. swrafi tipis
receptorebi gavrcelebulia agznebad qsovilebSi - nervulSi da kunTovanSi.
gansxvavdebian es cilebi erTmaneTisagan mxolod ligandur (aqtiur) centrSi
arsebuli aminomJavebiT.
ionur arxebs, romlebsac receptoruli ubani ar gaaCniaT, magram
monawileobas iReben potencialis gavrcelebaSi identuri domenuri
agebuleba aqvT. xSirad, maT SemadgenlobaSi liganduri centris nacvlad
Sedian aminomJavebi, romlebic mgrZnobiareni arian potencialTa cvlilebis
mimarT. amitom, aseTi arxebi transmembranuli potencialis cvlilebis
Sedegad ixsnebian.
Tu Na+-is da Cl--is ujredSi Sesvla, an K+-is ujredidan gamosvla mxolod
membranuli potencialis cvlilebas iwveven, Ca++-is ujredSida koncentraciis
zrda, damatebiT, metabolur Zvrebsac ganapirobebs. Ca++ sxva ionebisagan
gansxvavebiT, cilebs da fermentebs ukavSirdeba, ucvlis maT aqtivobas.
amitom, mas metabotropuli efeqtic aqvs. igi meorad Suamavals warmoadgens.
rogorc metabotropuli, aseve ionotropuli receptorebi moqmedeben
mxolod im SemTxvevaSi, Tu maTi liganduri ubani dakavebulia Sesabamisi
hormonisa Tu neirogadamcemis mier. biologiurad aqtiur naerTebs, romlebic
receptors ligandur ubanSi ukavSirdebian, iwveven receptoruli sistemis
gaaqtivebas da ujredis pasuxs, am receptoruli sistemis agonistebi
ewodebaT. hormonebi, mediatorebi da neirogadamcemebi agonistebs
warmoadgenen, radgan maTi moqmedeba garkveul fiziologiur efeqtTan aris
dakavSirebuli. naerTebi, romlebic ukavSirdebian receptorul cilas,
ujredul pasuxs ar iwveven da agonistebis moqmedebas Trgunaven,
antagonistebi ewodebaT. naerTebi, romlebic receptorul, ligandur ubans ar
ukavSirdebian, magram moqmedeben agonistis Tu antagonistis fiziologiur
efeqtze, modulatorebi ewodebaT. bevri farmakologiuri preparati ama Tu im
receptoris agonists, antagonists an modulators warmoadgens.
receptorebs ganasxvaveben maTi agonistebisa da antagonistebis mimarT
mgrZnobelobis mixedviT. antagonistebs, xSir SemTxvevebSi, ufro didi
swrafva axasiaTebT receptoris mimarT, vidre agonistebs. vinaidan,
hormonebis umravlesoba agonistebs warmoadgenen da maTi moqmedeba
antagonistebiT muxruWdeba, organizmis antagonistebiT datvirTva iwvevs
hormonTa moqmedebis daTrgunvas. antagonistebi, romlebic xSir SemTxvevebSi

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xelovnurad miRebul qimiur naerTebs warmoadgenen, moqmedeben ra ufro
maRali swrafviT, ikaveben endogenuri hormonebis adgils receptorul
cilaze da amiT aneitraleben maT moqmedebas. amitom, hormonTa antagonistebi
farTod moixmarebian farmakologiur praqtikaSi.
receptori, romelic erTi da igive hormons Tu neirogadamcems
ukavSirdeba, SesaZlebelia SeuRlebuli iyos sxvadasxva tipebis
transducerebTan (Gs, Gi an sxv.) da Sesabamisad, sxvadasxva katalizur
komponentebTan. receptorTa aseTi subtipebi, xSirad erTmaneTisagan
farmakologiuri TvisebebiT gansxvavdebian da hormonTa antagonistebis
mimarT sxvadasxva Tvisobas, swrafvas amJRavneben. magaliTad, acetilqolinis
receptori erTi mxriv, SesaZlebelia warmoqmnides natriumis arxs, meore
mxriv - iyos SeuRlebuli G-cilasTan, xolo am ukanasknelis saSualebiT -
ferment fosfolipazasTan. pirveli tipis acetilqolinis receptori maRal
Tvisobas amJRavnebs nikotinTan, da amitom mas nikotinuri (n) tipis
acetilqolinis receptori ewodeba, xolo meore - muskarinTan, da mas
muskarinuli (m) acetilqolinis receptori hqvia. bunebrivia, rom
gansxvavebulia am receptorebis fiziologiuri efeqtic. Tu n-acetilqolinis
receptoris gaaqtiveba ujredSi ionTa gadanawilebas iwvevs, m-tipis
receptori, upiratesad, metabolizmis cvlilebaze moqmedebs. Tavis mxriv,
receptorTa subtipebic iyofian qvejgufebad. igive muskarinis tipis
acetilqolinis metabotropuli receptori SesaZlebelia, G-gamaaqtivirebeli
(Gs) cilis saSualebiT SeuRlebuli iyos fosfolipazasTan da gaaZlieros
misi aqtivoba, an G-mainhibirebeli (Gi) transduceris meSveobiT SeuRlebuli
iyos adenilciklazasTan da dakavSirebis SemTxvevaSi daamuxruWos es
fermenti. muskarinuli tipis acetilqolinis receptorebic ujredis
metabolizmze sxvadasxvanairad moqmedeben.
aseTive mravalferovnebiT xasiaTdebian hormonTa sxva receptorebic.
magaliTad, adrenalinis da noradrenalinis sul mcire 4 subtipi ( 1, 2, 1, 2)
arsebobs, dofaminisa - ori (D1, D2), opiaturisa - 4, serotoninisa - 3,
vazopresinisa - 2 da a.S.
receptoruli aqtivobis erT-erT damaxasiaTebel Tvisebas misi
desensitizacia da e.w. “down-regulation” (regulacia qvemoT-ingl.) warmoadgens.
desensitizaciis dros adgili aqvs receptoris mier hormonis mimarT
mgrZnobelobis dakargvas, xolo “down-regulation”-iis SemTxvevaSi plazmur
membranaze receptorTa raodenoba klebulobs. aRniSnuli fenomeni

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gamowveulia agonistis (hormonis) xangrZlivi moqmedebiT da metyvelebs am
sistemebis adaptaciur, SemgueblobiT unarze, raTa Tavidan acilebul iqnas
ujredis xangrZlivi gaRizianeba.
desensitizaciis dros warmoebs receptoruli cilis kovalenturi
posttranclaciuri modifikacia fosforilirebis saxiT, xolo “down-
regulation”-iis SemTxvevaSi - misi internalizacia (gadmotana) citoplazmaSi.
desensitizaciis fenomeniT aris gamowveuli zogierTi farmakologiuri
preparatis dozirebis gazrdis aucilebloba misi xangrZlivi miRebis Semedeg.

ujredSida regulatorebi (meoradi Suamavlebi)

meoradi Suamavlebi warmoadgenen ujredSida regulatorebs, romlebic

warmoiqmnebian ujredze hormonis, neirogadamcemis Tu sxva fiziologiurad
aqtiuri nivTierebebis moqmedebis Sedegad da iwveven ujredSi mimdinare
bioqimiuri procesebis cvlilebas. meorad Suamavlebs warmoadgenen
cAMP(cikluri adenozinmonofosforis mJava), cGMP (cikluri
guanozinmonofosforis mJava), DAG (diacilgliceridi), TPI
(trifosfoinozitoli), AA (araqidonis mJava), Ca++ (kalciumis ionebi).
cAMP da cGMP warmoiqmnebian Sesabamisad adenilatciklazuri da
guanilatciklazuri reaqciebis Sedegad ATP-sa da GTP-sagan. maTi
koncentracia ujredSi 10-7-10-8 moli/l farglebSi meryeobs, xolo receptoris
gaaqtivebis Sedegad izrdeba 10-6 moli/l-mde. cikluri nukleotidebi
ujredSida metabolizmis regulacias ferment proteinkinazebis saSualebiT
axorcieleben. koncentraciis momatebis Sedegad isini Sesabamis
proteinkinazas ukavSirdebian da am ukanasknelis gaaqtivebas iwveven:

cAMP + proteinkinaza(PK) cAMP-PK

gaaqtivebuli fermenti funqciurad mniSvnelovani cilebis

fosforilirebas iwyebs da ucvlis maT bioqimiur funqcias. Tu aseTi cila,
metabolizmSi monawile (mag. glikolizSi) ferments warmoadgens, igi icvlis
katalizur aqtivobas - stimulirdeba an inhibirdeba, xolo Tu mas
struqturuli funqcia aqvs, maSin konformacia ecvleba.
proteinkinaza

cila + ATP cila-P + ADP

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 cikluri nukleotidebis msgavsi meoradi Suamavlebi hormonaluri
signalis gaZlierebas, mis amplifikacias iwveven. amis gamo ujreduli pasuxi
hormonalur gaRizianebaze mZlavria da ujredis metabolizmis did nawils
moicavs. ase, magaliTad, hormon glukagonis an adrenalinis erTi molekula
hepatocitebSi ferment adenilatciklazas aaqtivebs, rasac daaxloebiT 100
molekula cAMP-s sinTezi mohyveba. Tavis mxriv erTi molekula cAMP aaqtivebs
ra proteinkinazas, 10-mde fermenti fosforilaza b gadahyavs a-Si. amis gamo,
damatebiT 1000-mde molekula glikogeni iSleba da Warbi raodenobiT
producirebuli glukoza glikolizSi iSleba an sisxlSi Tavisufldeba.
amrigad, 1 molekula hormonis moqmedebis Sedegad SesaZlebelia aTasobiT
molekula glukozis warmoqmna. adamianebSi, aRwerili tipis amplifikacias,
gaZlierebas adgili aqvs stresul SemTxvevebSi, rodesac Tirkmelzeda
jirkvlebidan gamoTavisuflebuli adrenalini energetikuli masalis
mobilizacias iwvevs.
cikluri nukleotidebi da Sesabamisi proteinkinazebi ujredis 200-mde
reaqciaSi iReben monawileobas. isini cvlian ujredSi naxSirwylebis,
cilebis, cximebis da nukleinis mJavebis metabolizms da moqmedeben mxolod am
ujredisTvis damaxasiaTebel nivTierebaTa cvlaze. es nukleotidebi
aZliereben an akaveben mxolod diferencirebuli ujredis metabolizms da ar
iwveven mis xarisxobriv cvlilebas.
DAG da TPI. diacilglicerini da trifosfoinozitoli warmoiqmnebian
ferment fosfolipaza C-s gaaqtivebis Sedegad fosfatidilinozitfosfatidan
(fosfolipidi). diacilgliceridi hidrofobur, cximovani bunebis mqone
naerTs warmoadgens. igi, cikluri nukleotidebis msgavsad, proteinkinazas
aaqtivebs. es ukanaskneli gansxvavdeba cAMP-damokidebuli proteinkinazisagan
rogorc molekuluri woniT, aseve substratuli specifikurobiT. mas lipid-
damokidebuli proteinkinaza an proteinkinaza C ewodeba. am tipis
proteinkinazebi moqmedeben mxolod Ca-is ionebis TandaswrebiT:

DAG + Ca++ + PKC PKC-DAG-Ca++

lipid-damokidebuli proteinkinazis substratuli cilebi, ZiriTadad,

ujredis da organoidebis membranebSi arian lokalizebuli. proteinkinaza C
afosforilirebs receptorul cilebs, transportul fermentebs, ionTa
arxebisa da sekretoruli vezikulebis cilebs da a.S.

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 proteinkinazebisTvis damaxasiaTebelia mobiluroba, isini garkveuli
fiziologiuri zemoqmedebisas advilad gadaadgildebian, translocirdebian
ujredis erTi nawilidan meoreSi. magaliTad, proteinkinaza C
fosfolipidebisa da Ca++-is Tanaobisas ujredis citoplazmidan
translocirdeba plazmur membranebSi, sadac rigi cilebis fosforilirebas
iwvevs.
fosfolipaza C-s aqtivobis Sedegad warmoqmnili meore naerTi,
trifosfoinozitoli endoplazmuri retikulumidan Ca-is ionebis
gamoTavisuflebas awarmoebs. am meoradi Suamavlis moqmedebis molekuluri
meqanizmebi jer-jerobiT ucnobia.
cikluri nukleotidebis msgavsad, DAG da TPI moqmedeben diferencirebuli
ujredis normalur metabolizmze, magram maTi efeqti gacilebiT ufro
Zlieria da, SesaZlebelia, gamoiwvios xarisxobrivi cvlilebebic.
AA. araqidonis mJava Tavisufldeba membranuli fosfolipidebidan Ca++-
damokidebuli fosfolipazebis, ZiriTadad fosfolipaza A2-is saSualebiT.
warmoqmnilma AA, SesaZlebelia, ganicados Semdegi gardaqmnebi:
1-ciklooqsigenazuri gziT warmoqmnas prostaglandinebi da tromboqsanebi
da 2- lipooqsigenazuri gziT gardaiqmnas lipoqsinebad da leokotrienebad.
araqidonis mJavis daJangvis produqtebs eikozanoidebi ewodeba da isini
mikrosomuli fermentebis saSualebiT sinTezirdebian. eikozanoidebs
fiziologiuri moqmedebis farTo speqtri axasiaTebT: isini iwveven zrdis
SeCerebas, gluvi kunTis modunebas an SekumSvas, sxvadasxva hormonebis
sekrecias da a.S. TviT araqidonis mJava moqmedebs K+-is arxebze da xsnis maT.
warmoadgenen ra lipidebis warmoebulebs, eikozanoidebi Tavisuflad
gamodian ujredidan da moqmedeben mezobeli ujredebis citoplazmaSi
arsebul receptorebze. amitom maT, SesaZlebelia, ujredSorisi meoradi
Suamavlebis funqciac hqondeT.
Ca-is ionebi. am ionebis ujredSida koncentracia 10-8- 5x10-7Mmoli/l-is tolia.
agznebad qsovilebSi (nervuli, kunTovani) kalciumis arxis gaxsnis Sedegad
maTi koncentracia SesaZlebelia 10-7Mmoli/l-mde, xolo araagznebadSi - 10-6M-
moli/l-mde gaizardos. garda amisa, kalciumis ionebis koncentraciam
citoplazmaSi, SesaZlebelia, moimatos ujreduli rezervuarebidan
(endoplazmuri bade, mitoqondriebi) gamoTavisuflebis Sedegad.
Ca-is ionebis ZiriTad aqceptors cila-kalmodulini (CaM) warmoadgens.
kalmodulinis molekuluri wona 16000-is tolia da mas 4 Ca++-

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damakavSirebeli ubani aqvs. Ca-is ionebis mierTebis Semdeg kalmodulins
konformacia ecvleba da aqtiur mdgomareobaSi gadadis. kalciumiT
gaaqtivebuli kalmodulini, Tavis mxriv, specifiur proteinkinazas
ukavSirdeba da es ukanaskneli sxvadasxva cilis fosforilirebas iwyebs:

Ca++ + CaM Ca-CaM

CaM + PK Ca-CaM-PK

CaM-damokidebuli proteinkinaza fizikur-qimiuri TvisebebiT da

substratuli specifikurobiT gansxvavdeba sxva tipis proteinkinazebisagan.
igi ZiriTadad, ujredis SekumSvadi da moZravi elementebis cilebis
fosforilirebas axorcielebs. amitom, kalmodulini da misi Sesabamisi
proteinkinaza monawileobas iReben mitozuri aparatis CamoyalibebaSi,
organoidebis ujredSi gadaadgilebaSi, SekumSvadi cilebis gaaqtivebaSi,
ujredis struqturis formirebaSi da a.S. xSir SemTxvevebSi, Ca++ da
kalmodulini cikluri AMP-s sawinaaRmdego reaqciebSi iReben monawileobas.
amrigad, plazmuri membranis metabotropuli receptoris moqmedebis
meqanizmi, SesaZlebelia, Semdegnairad gamovsaxoT:
hormoni receptori meoradi Suamavali proteinkinaza
fosforilirebuli cila ujredis pasuxi

citoplazmuri receptorebi

citoplazmuri receptorebi hormonalur signals pirdapir citoplazmaSi

aRiqvamen. citoplazmur receptorTan urTierTmoqmedi hormonebi lipofiluri
bunebis naerTebi arian, da amitom isini advilad gadian plazmur membranas.
citoplazmur receptorebSi gaerTianebuli arian steroiduli da Tiroiduli
hormonebi: estrogenebi, androgenebi, kortikosteroidebi da sxva.
hormonis (H) citoplazmur receptorTan dakavSirebis Semdeg es
ukanaskneli aqtivirdeba. xSirad, receptoris gaaqtiveba dakavSirebulia mis
fosforilirebasTan. fosforilirebiT gaaqtivebuli receptori iwyebs
gadaadgilebas-translokacias birTvisaken. birTvSi receptori ukavSirdeba
dnm-s da rTavs garkveul genTa kompleqts. steroiduli da Tiroiduli
hormonTa receptoris moqmedeba waagavs transkripciuli faqtorebis
moqmedebas, radgan orive maTgani garkveuli tipebis genTa transkripcias
uwyobs xels.

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 hormoni (H) hormon-receptoruli kompleqsi d n m ujredis pasuxi

steroiduli receptorebis moqmedeba, plazmurisagan gansxvavebiT,

mizanmimarTulia da ujredis mTlian metabolizmze ar vrceldeba, isini
meoradi Suamavlebis produqcias ar aZliereben. aseTi tipis receptorebis
gaaqtiveba specifikur genTa transkripcias da garkveuli tipis cilebis
sinTezs iwvevs.

zrdis faqtorebi da maTi receptorebi

zrdis faqtorebi ewodebaT cilovani bunebis naerTebs, romlebic ujredis

zrdas da diferencirebas iwveven. zrdis faqtorebs sxvadasxva tipis
ujredebi da qsovilebi warmoqmnian, da maTi moqmedebis areali, hormonebTan
SedarebiT ufro SezRudulia. dReisaTvis 60-mde aseTi faqtoria aRmoCenili,
da yvelaze ufro cnobils maT Soris insulini warmoadgens. isini
dabalmolekulur cilebs warmoadgenen, romelTa molekuluri wona meryeobs
6000-15000-mde. maTi receptori plazmur membranaze mdebareobs. igi
integralur membranul cilas warmoadgens da Sedgeba ori domenisagan:
ujreTgareTa, receptoruli aqtivobis mqone ubnisagan da ujredSida
nawilisagan. zrdis faqtoris receptori ar aris SeuRlebuli romelime
funqciur erTeulTan: G-cilasTan, fermentTan, Tu arxTan. mis katalizur
komponents qmnis ujredSida domeni, romelsac proteinkinazuri aqtivoba
axasiaTebs. sxva analogiuri fermentebisagan gansxvavebiT, es proteinkinaza
substratul cilebSi afosforilirebs ara aminomJava serinis naSTs, aramed
Tirozinisas. receptoruli (Tirozin)proteinkinazis endogenur ujredul
substratebs miekuTvnebian zrdis da diferencirebaSi monawile cilebi.

zrdis faqtori receptori transkripciuli aqtivoba ujredis

pasuxi

cnobilia, rom zrdis faqtorebis moqmedeba ujredze iwvevs birTvSi

transkripciuli faqtorebis gaaqtivebas, maT sinTezs. xSirad, zrdis
faqtorebis zegavleniT matulobs agreTve proto-onkogenebis produqtTa -
protoonkocilebis sinTezic, romlebic garkveul ujredebSi transkripciuli
faqtorebis funqcias asruleben. aRsaniSnavia is, rom proto-onkogenebis

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produqtebs, iseve, rogorc zrdis faqtorTa receptorebs,
(Tirozin)proteinkinazuri aqtivoba axasiaTebT.
zrdis faqtorebi da maTi receptorebi ganapirobeben ujredis
proliferaciisa da diferencirebis procesebis monacvleobas, da amdenad,
maT didi mniSvneloba eniWebaT organizmis embrionalur ganviTarebaSi.

azotis oqsidi (NO)

adamianis organizmSi azotis oqsidi (NO) ramdenime gziT warmoiqmneba, romelTa

Sorisac ZiriTads warmoadgens misi bisinTezi aminomJava L-argininidan. sisxlSi
mocirkulire azotis oqsidis 90% swored aminomJava argininidan warmoiqmneba.
arginini iJangeba ferment NO-sinTazis (NOS) saSualebiT, romlis moqmedebis
Sedegad warmoiqmneba NO da citrulini.

NO-sinTaza
Aarginini NO + citrulini

aRmoCenilia NO-sinTazis sami izofermenti, romelTagan pirveli

lokalizebulia ZiriTadad sisxlZarRvTa endoTeliumSi (eNOS), meore –
neironebSi (nNOS), xolo mesame ki - makrofagebSi da sxva imunokompetentur
ujredebSi. pirveli ori, neironuli nNOS-i da endoTeluri eNOS-i konstituciur
fermentebs warmoadgenen. endoTeluri eNOS mier sinTezirebuli azotis oqsidi
moqmedebs ra sisxlZarRvTa kedlebze ganapirobebs gluvi kunTebis relaqsacias
da iwvevs vazodilatacias. amitom eNOS-i organizmSi sisxlis wnevis ZiriTad
maregulirebel faqtors warmoadgens.
neironuli nNOS, agreTve konstituciur ferments warmoadgens da
neironebis garda aRmoCenilia, ganivzolian kunTebSi, pankreasis beta-ujredebSi,
broqniolebis, saSvilosnos da kuWis epiTelur ujredebSi. inducirebadi iNOS,
romelic ZiriTadad makrofagebSia lokalizebuli sinTezirdeba da aqtivirdeba
infeqciuri agentebis an anTebiTi procesebis zemoqmedebiT. NO, romelic
inducirebadi fermentis saSualebiT sinTezirdeba iwvevs ara marto infeqciuri
agentis ganadgurebas, aramed TviT makrofagebis apoftozsac. NO-sinTazis samive
izoforma aRmoCenilia TirkmelebSi.
NO-sinTazis konstituciuri formebi (nNOS da eNOS) aqtivirdebian
kalciumis ionebiT da kalmoduliniT. Ca-is ujredSida mateba am fermentebis

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xanmokle aqtivacias iwvevs, rac ujredSi azotis oqsidis warmoqmnas
ganapirobebs. vinaidan azotis oqsidi advilad difundirebad naerTs warmoadgens,
igi Tavisuflad gadaadgildeba ujredebSi, da misi biologiuri aqtivobis gamo
moqmedebs Sesabamis efeqtorebze rogorc neiromediatori an neiromodulatori.
konstituciuri fermentisagan gansxvavebiT inducirebadi iNOS-i mWidrod aris
dakavSirebuli Ca-kalmodulinis kompleqsTan, da amdenad ujredSida kalciumis
mateba masze ar moqmedebs.
azotis oqsidis biologiur da farmakologiur potencials am naerTis
fizikur-qimuiri Tvisebebi ganapirobebs. NO Tavisufal radikals warmoadgens,
radgan mis molekulaSi gauwyvilebeli eleqtroni Sedis. vinaidan azotis oqsidis
sicocxlis xangrZlivoba 5 wams ar aRemateba igi swrafad gadis biologiur
membranebs, moqmedebs hemis Semcvel fermentebze da iwvevs maTi aqtivobis
cvlilebebs. azotis oqsidis erT-erT ZiriTad efeqtors hem-Semcveli fermenti
guanilatciklaza warmoadgens, romlis gaaqtivebac gansazRvravs NO-s
moqmedebas sisxlZarRvTa kedlis gluv kunTebze, trombocitebze da sxva
ujredebze. azotis oqsidis narCeni nawili swrafad reagirebs wyalTan, ris
Sedegadac warmoiqmneba nitriti. es ukanaskneli cilebTan (hemoglobini,
mioglobini, rkina-Semcveli fermentebi) urTierTqmedebis gamo SesaZlebelia
gardaiqmnas nitratad da Sesatyvisi hemis met-nawarmad. magaliTad,
eriTrocitebSi azotis oqsidis difundirebis SemTxvevaSi igi urTierTqmedebs
oqsihemoglobinTan, ris Sedegadac warmoiqmneba nitriti da met-hemoglobini.
azotis oqsidi, SesaZlebelia agreTve daukavSirdes zogierTi cilis cisteinis an
Tirozinis naSTs; Dda Tu es cila fermentia gamoiwvios misi aqrivobis
cvlilebebi. cilebis nitrozilireba erT-erT mniSvnelovan post-translaciur
modifikacias warmoadgens.
rogorc ukve iTqva, makrofagebis mier dasinTezirebuli NO, rodesac is
didi raodenobiT gamoiyofa, baqteriocidul da toqsikur Tvisebebs atarebs.
mikroorganizmebis ganadgureba warmoebs mas Semdeg, rodesac makrofagebidan
. -
mikroorganizmSi Sesuli NO da superoqsidanionradikalisi (O 2 ) ujredSi jer
peroqsinitriti (ONOO-), xolo Semdeg hidroqsilis radikali (OH.) warmoiqmneba.
aRniSnuli procesi Tu normalur qsovilSi warimarTa igi mis dazianebas da
nekrozs iwvevs.
azotis oqsidis koncentracia sisxlis plazmaSi icvleba sqesis, asakis,
kvebis, organizmis sxvadasxva funqciuri mdgomareobis da paTologiebis dros.
mamakacebis sisxlis plazmaSi nitratebis koncentracia orjer aRemateba qalebis

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sisxlis plazmaSi arsebul nitratebs. garda amisa irkveva, rom mamakacebis
amonasunTqi haeri SedarebiT ufro mdidaria azotis oqsidiT. mamakacebis da
postmenopauzis Semdeg qalebis sisxlis plazmaSi azotis oqsidis raodenoba
tolia, rac imaze metyvelebs, rom NO-s metabolizmSi wamyvani mniSvneloba
estrogenebs mieniWebaT. NO-s raodenoba sisxlis plazmaSi icvleba menstrualuri
ciklis dros da igi folikulur fazaSi matulobs.
azotis oqsidis raodenoba sisxlSi da SardSi izrdeba agreTve infeqciuri
daavadebebis da anTebiTi procesebis Sedegad. nitritis maRali koncentracia
SardSi urogenitaluri infeqciebis erT-erT ZiriTad maxasiaTebels warmoadgens.
makrofaguli NO, romlis produqciac 1000-jer izrdeba anTebiTi procesebis
Sedegad, momatebulia septikuri Sokis, ciroziT gamowveuli endotoqsemiis,
sistemuri anTebiTi sindromebis, organoebis mravalmxrivi dazianebis, Sidsis
Sedegad gamowveuli oportunistuli infeqciebis dros.
NO-s mniSvneloba sisxlis mimoqcevis da arteriuli wnevis regulaciaSi
dadasturebulia mravalmxrivi gamokvlevebiT. am mxriv gansakuTrebul
yuradRebas ipyrobs endoTeluri NO-sinTaza, romlis disfunqciac
dakavSirebulia aTerogenezuli procesebis intensifikaciasTan da sagrZnob
cvlilebebs ganicdis asakis matebasTan mimarTebaSi. aRmoCenilia uaryofiTi
korelacia sisxlis plazmaSi azotis oqsidis koncentraciasa da qolesterinis
da dabali simkvrivis lipoproteinebis raodenobas Soris.
aRiniSneba kardialuri NO-s momateba gulis daavadebebis SemTxvevebSic.
ase magaliTad, nanaxia, rom kardiomiopaTiis mqone avadmyofebSi azotis oqsidis
koncentracia sisxlSi normasTan SedarebiT momatebulia. izrdeba am naerTis
Semcveloba gulis qronikuli daavadebebis drosac.
amrigad, SesaZlebelia davaskvnaT, rom azotis oqsidi koncentraciuli
meryeoba sisxlis plazmaSi mravali tipis daavadebis mniSvnelovan serologiur
korelats warmoadgens, romlis analizi saSualebas iZleva ganisazRvros
avadmyofTa mdgomareoba. inducibeluri NO-sinTazis aqtivacia da azotis oqsidis
masiuri produqcia, romelsac adgili aqvs organoTa dazianebis, infeqciis Tu
qronikuli anTebiTi procesebis dros ganapirobebs Tirkmelebis da gul-
sisxlZarRvovani sistemis destruqcias, maSin rodesac arainducibeluri
fermentiT azotis oqsidis xanmokle formireba vazodilataciur antiiSemiur
efeqtebTan aris dakavSirebuli.

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 ujredebis nekrozi da apoftozi

ujredis sikvdili biologiis da medicinis erT-erT sakvanZo problemas

warmoadgens. dReisaTvis dadgenilia, rom ujredi SesaZlebelia ori gziT
ganadgurdes, nekrozis, anu “iZulebiTi” sikvdilis, da apoftozis, anu
genetikurad programirebuli, Sinagani homeostaziT gansazRvruli sikvdilis
meSveobiT. apoftozi aucilebelia organizmSi anomaluri, dazianebuli da
daberebuli ujredebis mosacileblad. adamianis organizmSi yovel wams
milionobiT ujredi ganicdis apoftozs, da maT adgils axlad dabadebuli,
axalgazrda, janmrTeli ujredi ikavebs. Aapoftozi normalur fiziologiur
movlenas warmoadgens, xolo nekrozi mxolod paTologiuri procesebis
ganviTarebisas gvxvdeba.
apoftozi (berZn. - ganadgureba) biologebisTvis didi xania cnobili,
magram am procesma gansakuTrebuli yuradReba miipyro mas Semdeg, rodesac
gairkva, rom simsivnur ujredebs darRveuli aqvT ujredis ganviTarebis
(dabadeba gamravleba dabereba sikvdili) normaluri msvleloba, ver gadadian
apoftozSi, da amitom, ganuwyveteliv mravldebian. vinaidan yovel qsovilSi
ujredTa raodenoba limitirebulia, bunebrivia, rom gadaberebuli ujredis
adgili mxolod erTma axalgazrda ujredma SeiZleba daikavos. amitom qsovilis
normalur cxovelqmedebas, misi homeostazis SenarCunebas gansazRvravs
ujredebis gamravlebas da sikvdils Soris balansi. Tanafardoba, romelic am or
process Soris arsebobs SeiZleba dairRves, rasac aucileblad moyveba
qsovilisTvis arasasurveli procesebis msvleloba. im SemTxvevaSi, Tu apoftozi
sWarbobs viTardeba qsovilis atrofia, xolo im SemTxvevaSi, Tu gamravlebis
procesi ufro swrafad mimdinareobs adgili aqvs qsovilis simsivnur zrdas.
apoftozis aranormaluri mimdinareoba, misi Seneleba an daCqareba
safuZvlad udevs mravali daavadebis paTogenezs, romelTa Sorisac vxvdebiT
virusuli infeqciebiT gamowveul garTulebebs, autoimunur darRvevebs,
neirodegeneraciul da nerv-kunTovan paTologiebs, imunodeficitur sindromebs,
da sxva. apoftozis Seneleba an Sewyveta, magaliTad, mutaciur ujredSi,
SesaZlebelia ganapirobebdes qsovilis onkologiur zrdas, xolo imunur
ujredebSi – autoagresiuli klonebis gamravlebas, da aqedan gamomdinare,
autoimunur darRvevebs. piriqiT, apoftozis swraf mimdinareobas adgili aqvs
rogorc mwvave paTologiebis, magaliTad, toqsinis warmomqmneli
mikroorganizmebiT infecirebisas, iSemia-reperfuziuli dazianebebis, infarqtebis

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dros, ise qronikulad mimdinare paTologiebis SemTxvevebSi, magaliTad
neirodegeneraciuli paTologiebis (alcheimeris, parkinsonis, hantingtonis
daavadebebi) da Sids-is dros.
Tu apoftozi damaxasiaTebelia rogorc paTologiuri, ise normaluri
ujredisTvis, nekrozi mxolod paTologiuri dazianebebis dros viTardeba.
ujredis nekrozireba iwyeba Rrma anoqsiis, sakvebi da energetikuli nivTierebebis
miwodebis Sewyvetis (mag. hipoglikemiis), Zlieri fiziko-qimiuri zemoqmedebebis,
travmebis Sedegad. nekrozisagan gansxvavebiT apoftozSi monawileoben
regulirebadi, ujredis damSleli, kataboluri fermentebi (proteazebi da
nukleazebi). apoftozisTvis damaxasiaTebelia birTvis specifikuri
morfologiuri (qromatinis kondensacia, piknozi) da bioqimiuri cvlilebebi (dnm-
is etapobrivi fragmentacia, 200-200 fuZe-wyvilebis Semcveli monomerebis an
oligomerebis warmoqmniT). apoftozis dros ujredis garsi mTliania da iSleba
mxolod ujredis ganadgurebis saboloo etapze, maSin rodesac nekrozis dros
igi dasawyisSive irRveva. nekrozis A dros ujredis SigTavsi gareT gamodis, da
amitom am dros, apoftozisagan gansxvavebiT, anTebiTi reaqciebi iwyeba.
apoftozis Sedegad mitoqondriebis ultrastruqturuli anomaliebi iSviaTad
viTardeba, xolo nekrozul ujredSi xSiria gajirjvebuli mitoqondriebis
arseboba. apoftozis saboloo stadiaze ujredi iyofa mraval mcire nawilakebad,
romlebic axaldabadebuli ujredebis mier STainTqmeba.
mravali tipis fiziologiuri Tu paTologiuri zemoqmedeba ujredze
ganapirobebs apoftozs. ujredis programirebuli sikvdili SesaZlebelia
daiwyos zrdis hormonebis deficitiT, maionizirebeli radiaciiT, simsivnis
nekrozis gamomwvevi faqtorebiT, virusuli infeqciiT, glukokortikoidebis
zemoqmedebiT da a.S. maTi moqmedebis Sedegad ujredSi gaaqtivebas iwyebs
kataboluri reaqciebis mTeli kaskadi, romlebic ujredis daSlas, mis
danawevrebas ganapirobeben. apoftozis msvlelobaSi monawileobas iReben cilis
damSleli specialuri fermentebi – kaspazebi, da dnm-is damSleli fermentebi –
specifikuri nukleazebi, romlebic ganapirobeben makromolekulebis etapobriv
fragmentacias da gansazRvraven apoftozisTvis damaxasiaTebel fiziologiur Tu
morfologiur mdgomareobas
cnobilia, rom mitoqondriebis ZiriTadi energetikuli potencials, romlis
saSualebiTac isini ATP-is sinTezs anxorcieleben, protonTa araTanabari
ganawileba warmoadgens. vinaidan wyalbadis ionTa ufro meti raodenoba
dagrovilia mitoqondriis membranaTaSoris sivrceSi, protonTa gradienti
mimarTulia garedan SigniT, Sidamitoqondriuli matriqsisaken. protonTa

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gadaqaCva Signidan gareT sunTqviT jaWvSi eleqtronTa transportis Sedegad
gamonTavisuflebuli energiis xarjze xdeba, romlis Seferxebac
transmembranuli protonuli gradientis darRvevas, da Sesabamisad ATP-is
sinTezis SeCerebas iwvevs. mitoqondriebSi arsebuli protonuli gradienti
ganapirobebs transmembranuli potencialis arsebobas, romelic -iT aRiniSneba.
transmembranuli potenciali mitoqondriebis normaluri funqcionirebis
garantiaa da misi moSla, aucileblad jer mitoqondriebis, xolo Semdeg ki,
energiis deficitis gamo, ujredis sikvdils gamoiwvevs. Tu romelime toqsikurma
nivTierebam Sidamitoqondriul membranaSi moaxerxa naxvretis (foris) warmoqmna,
mitoqondria dakargavs “Zalovani sadguris” funqcias da veRar moamaragebs
ujreds ATP-iT.
miuxedavad amisa, mitoqondriebi Seicaven specialur forebs, romlebic
garkveul pirobebSi ixsnebian da protonTa nakads atareben garedan SigniT. sami
aseTi “naxvreti” gamoiyeneba ATP-s sinTezis procesSi da maT sunTqviTi jaWvis da
JangviTi-fosforilirebis SeuRlebis punqtebi ewodeba. isini gamudmebiT muSaoben
da maT arxebSi Sesuli wyalbadis ionebi ATP-is sinTezs ganapirobeben. rac ufro
meti wyalbadia dagrovebuli membranaTaSoris sivrceSi, miT ufro didi
raodenobiT xdeba protonebis gatareba, da Sesabamisad, miT ufro meti ATP
sinTezirdeba. da piriqiT, Tu dabalia, anu protonTa gradienti mcirea, miT
ufro mcire raodenoba ATP sinTezirdeba.
ATP-is masinTezirebeli forebis garda mitoqondriul membranaSi gvxvdeba
agreTve arxi, romelsac gardamavali ganvladobis anu megaarxi ewodeba. igi
iwyeba citoplazmis mxridan, ganWolavs orive mitoqondriul membranas, matriqss,
da mTavrdeba Sidamitoqondriul sivrceSi. am arxis gaxsna protonTa gradientis
darRvevas iwvevs, ris gamoc mcirdeba. Sedegad SeuRlebis punqtebisTvis
wyalbadis ionebi aRar arian sakmarisi, ris gamoc ATP sinTezi mcirdeba.
normalur mdgomareobaSi megaarxi, ra Tqma unda daxurulia, magram garkveul
pirobebSi igi SesaZlebelia gaixsnas. misi xangrZlivi gaxsna mitoqondriebisTvis
“dramatulad” mTavrdeba: iwyeba -is kolapsi, iTiSeba sunTqviTi jaWvi da
mitoqondriebidan gareT, citoplazmaSi, gamosvlas iwyeben sxvadasxva funqciurad
mniSvnelovani cilebi. garkveuli drois Semdeg mitoqondriebi kargaven
normaluri cxovelqmedebis unars, rasac SemdgomSi mTeli ujredis sikvdili
mohyveba. sxvadasxva toqsikuri nivTierebebi, da maT Soris zeJangebi, azotis
oqsidi, Ca-is ionebis maRali koncentraciebi ujredis nekrozs swored megaarxis
gaxsniT iwveven.

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 megaarxis TandaTanobiTi, etapobrivi gaxsna apoftozis drosac mimdinareobs,
rac SesaZlebelia gamowveuli iyos sxvadasxva toqsinebiT, sakvebi nivTierebebis
da zrdis faqtorebis deficitiT, glukokortikoidebis moqmedebiT, ujredis
garsze arsebuli specialuri apoftogenuri (Fas/Apo-1/CD95) receptorebiT da a.S.
megaarxis SemadgenlobaSi Sedian sxvadasxva tipis cilebi, romlebsac didi
mniSvneloba aqvT misi gaxsna-daxurvisTvis, da Sesabamisad ujredis sacicocxlo
programisaTvis. upirveles yovlisa aRsaniSnavia e.w. Bcl-2-is genTa ojaxis
cilebi romlebic jer kidev 80-iani wlebis dasawyisSi iyvnen aRmoCenili, da
romelTa struqturuli cvlilebebic limfoiduri ujredebis simsivnur
gadagvarebas iwvevda. dadginda, rom Bcl –Si momxdari mutaciis Sedegad ujredi
“ukvdavi” xdeba, igi veRar Sedis fiziologiur apoftozSi da ganuwyvetliv
mravldeba. ukanasknel wlebSi dadasturda, rom B Bcl mitoqondriebis megaarxis
SemadgenlobaSi Sedis da mis gaxsna-daxurvas akontrolebs. cila hidrofoburi
nawilebiT CaSenebulia megaarxis kedlebSi, da amitom arxi gamudmebiT
daxurulia. igi ixsneba mxolod im SemTxvevebSi, Tu Bcl-is moqmedeba SeizRudeba
da cila arxSi veRar CaSendeba. aseTi SemzRudveli funqcia meore cilas – Bax-s
aqvs, romelsac SeuZlia daukavSirdes Bcl-s da moacilos igi arxis kedlebidan.
Sedegad arxi ixsneba da ujredi gadadis apoftozSi. im SemTxvevaSi, Tu moxda
Bcl-is genSi mutacia, Bax mas veRar amoiRebs arxidan, ris gamoc igi gamudmebiT
iketeba. Sedegad ujredi veRar kvdeba, is iwyebs ganuwyvetel proliferacias da
viTardeba simsivne.
ujredis energetikuli potencialis daqveiTeba arxis gaxsnas iwvevs, rasac
bunebrivia mitoqondriis mier ATP-s sinTezis Semcireba mohyveba. es procesi
kidev ufro aRrmavebs ujredis energerikul mdgomareobas, ris gamoc e.w.
“mankieri wre” warmoiqmneba. aRniSnuli garemoeba metyvelebs imaze, rom apoftozi
“sul an arafris” kanonis mixedviT viTardeba, romelSic TviTgaZlierebis
meqanizmebi erTvebian, da Tu am procesma msvleloba daiwyo, mas bolo stadiebze
veRaraferi SeaCerebs.
amrigad naTelia, rom apoftozis formirebaSi wamyvani mniSvneloba
mitoqondriebs mieniWebaT. am organoidebSi arsebuli specialuri arxis gaxsna,
romelic mitoqondriebis funqciis moSlas iwvevs, SesaZlebelia warimarTos
rogorc fiziologiuri, ise paTologiuri zemoqmedebis Sedegad, da ujredis
sikvdili gamoiwvios. MpaTogenuri agentebiT misi naadrevi, araregulirebadi
gaxsna ujredis naadrev sikvdils, xolo misi Caketva – ujredis “ukvdavebas” da
simsivnur zrdas iwvevs.
megaarxis gaxsnis Sedegad mitoqondriidan citoplazmaSi gadmosvlas

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iwyeben sxvadasxva tipis cilebi da rnm-i. garda amisa, vinaidan -i vardnas
ganicdis, wydeba cilebis transporti citoplazmidan mitoqondriebSi. aRniSnul
cvlilebebs mohyveba mitoqondriis funqciis darRveva, romelic aucileblad
ujredis nekroziT mTavrdeba. ismis kiTxva, Tu rogor viTardeba apoftozi? ra
ganapirobebs ujredis etapobriv ganadgurebas? sad aris is gasayari, sadac
erTmaneTs scildebian nekrozis da apoftozis gzebi? dReisaTvis am kiTxvebze
pasuxis gacema Znelia, magram arsebobs garkveuli mosazreba, romlis mixedviTac
SesaZlebelia vivaraudoT, rom mitoqondriebidan gamosul garkveul molekulebs
sasignalo mniSvneloba aqvT. aRniSnuli molekulebi dasabams aZleven garkveuli
tipis reaqciaTa msvlelobas, romelic ganapirobebs ujredis birTvSi arsebuli
dnm-is etapobriv daSlas. erT-erT aseT molekulas, cila - holocitoqromi C
warmoadgens. igi normalur pirobebSi apocitoqromi C-s formiT
transportirdeba mitoqondriebSi, CaerTveba Sidamitoqondriul membranaSi da
monawileobas iRebs eleqtronTa gadatanis jaWvis SenebaSi. megaarxis gaxsnis
Sedegad, citoqromi C-s hemmocilebuli forma, anu holocitoqromi C gamodis
gareT citoplazmaSi, sadac is iwyebs proteolizuri, cilis damSleli
fermentebis gaaqtivebas (kerZod, kaspaza-3-is da kaspaza-6-is). proteolizuri
fermentebi ganapirobeben birTvSi e.w. dnm-is fragmentaciis faqtoris saSualebiT
specifikuri dnm-is damSleli fermentebis - nukleazebis CarTvas, romlebic
etapobrivad iwyeben 200-300 nukleotiduri wyvilisagan Semdgari fragmentebis
Camoclas dnm-is molekulidan. dasaSvebia, rom kataboluri fermentebi, romlebic
ganapirobeben dnm-is etapobriv daSlas, swrafi energetikuli “katastrofis”
pirobebSi ver aswreben gaaqtivebas, ris gamoc viTardeba nekrozi.
amrigad savaraudoa, rom apoftozi viTardeba mitoqondriebis funqciis
etapobrivi daqveiTebis Sedegad, rasac mohyveba am organoidebidan apoftogenuri
sasignalo makromolekulebis TandaTanobiTi gamonTavisufleba, da Sesabamisad,
dnm-is neli fragmentacia. masiuri energetikuli kolapsis dros, mitoqondriebis
funqcia swrafad qveiTdeba, igi ijirjveba, skdeba da misi SigTavsi swrafad
gadmodis citoplazmaSi. am dros adgili aqvs hidrolizuri fermentebis Zlier
aqtivacias da qromatinis swraf degradacias.

hormonebi

hormoni (berZn. - agzneba, gaRizianeba) qimiur naerTs warmoadgens,

romelsac mcire raodenobiT gamohyofs erTi organo, da romelic sisxlis
saSualebiT gadaitaneba sxva organoebSi da qsovilebSi. am ukanasknelebSi

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hormoni ujreduli receptorebis gaaqtivebas iwvevs, rasac ujredis
metabolizmis cvlileba mohyveba. hormonebi ZiriTad ujredSoris
sakomunikacio sistemebs warmoqmnian, da amis gamo maT pirvelad Suamavlebs
uwodeben (gansxvavebiT meoradi Suamavlebisagan, romlebic ujredSida
sakomunikacio sistemebSi iReben monawileobas). hormonebisa da msgavsi
naerTebis saSualebiT warmoebs qsovilTa da ujredTa koordinirebuli
muSaoba, maTi regulacia da kontroli.
adamianis organizmSi hormonTa gamoyofa da maTi moqmedeba
reglamentirebulia drosa da sivrceSi. yovel hormons mkacrad
gansazRvruli moqmedebis areali aqvs, romelic mxolod garkveuli tipis
samizne-efeqtorul ujredebsa da qsovilebs moicavs. hormonis gamoyofa
endokrinuli jirkvlis Tu sxva organos mier limitirebulia da mis
raodenobas akontrolebs regulaciis ierarqiuli sistema, romlis
saTaveebSic umaRlesi nervuli centrebi imyofebian:

centraluri nervuli sistema

nervuli impulsebi

hipoTalamusi

hipofizi

periferiuli jirkvlebi

efeqtoruli qsovilebi

hormonebi qimiuri agebulebis mixedviT SesaZlebelia sam jgufad daiyos:

cilovani (peptiduri) hormonebi, aminebi da steroidebi. cilovani hormonebi
ribosomebze sinTezirdebian maRalmolekuluri prohormonis formiT. isini
gamoTavisuflebisas sisxlSi hidrolizdebian specialuri proteazebiT da
warmoqmnian aqtiur hormons.
hormonebisTvis damaxasiaTebelia maTi efeqturi koncentraciis dabali
done (isini moqmedeben 10-12 - 10-8 moli/l-is koncentraciebSi; mosvenebis dros

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maTi koncentracia sisxlSi kidev ufro mcirea). hormonTa naxevardaSlis
periodi mcirea da Seadgens 1-10 wT-s.

hipoTalamusis hormonebi

hipoTalamusis hormonebi ierarqiuli regulaciis umaRles safexurze

imyofebian da maTi sekrecia Tavis tvinSi arsebuli nervuli centrebidan
imarTeba. isini mokle peptidebs warmoadgenen da Sedgebian 3-15 aminomJavuri
naSTisagan. hipoTalamusis peptidebi sisxlis mimoqcevis saerTo
cirkulaciaSi ar xvdebian, radgan pirdapir nervuli daboloebebiT
Tavisufldebian hipofizis karis venaSi da mxolod hipofizze moqmedeben.
hipoTalamusis peptidebi maTi hipofizze moqmedebis mixedviT or jgufad
iyofian: e.w. liberinebi da statinebi. liberinebi hipofizidan hormonebis
gamoTavisuflebas, xolo statinebi - maT Sekavebas iwveven. dReisaTvis
cnobilia 7 liberini da 3 statini. liberinebs warmoadgenen:
kortikoliberini -hormoni, romelic iwvevs hipofizidan
adrenokortikotropinis gamoTavisuflebas; Tireoliberini - iwvevs
Tireotropinis sekrecias; somatoliberini - hipofizidan aTavisuflebs
somatotropins; melanoliberini - aTavisuflebs melanotropins; luliberini
- aTavisuflebs maluTeinizirebel hormons; prolaqtoliberini iwvevs
prolaqtinis sekrecias; foliberini - aTavisuflebs folitropins
(folikulmastimulirebel hormons). statinebi arian: somatostatini - iwvevs
hipofizidan hormon somatotropinis gamoyofis Semcirebas; melanostatini -
akavebs melanotropinis sekrecias da prolaqtostatini - arapeptidovani
bunebis mqone naerTi (dofamini), romelic Trgunavs prolaqtinis gamoyofas.
hipoTalamusis peptidebs, maTi nervul qsovilSi sinTezirebis gamo, xSirad
neiropeptidebsac uwodeben. garda amisa maT rilizing-faqtorebic (riliz-
ingl. ganTavisufleba) hqviaT. hipoTalamusis hormonebis gamoyofa umaRlesi
nervuli centrebidan regulirdeba.

hipofizis hormonebi

hipofizi ori nawilisagan Sedgeba: wina - adenohipofizisagan da ukana -

neirohipofizisagan. maT sxvadasxva embrionuli warmoSoba aqvT. hipofizur
hormonebs maTi endokrinul jirkvlebze moqmedebis gamo tropuli hormonebi
ewodebaT. wina hipofizarul hormonebs miekuTvnebian: Tireotropini - iwvevs

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farisebri jirkvlidan Tiroiduli hormonebis gamoTavisuflebas; am hormonis
hiperproduqcia ganapirobebs hiperTireozs, xolo hipoproduqcia - meorad
hipoTireozs; kortikotropini - awarmoebs Tirkmelzeda jirkvlidan
hormonebis gamoyofas; hiperproduqcia iwvevs kuSingis sindroms, hipo -
Tirkmelzeda jirkvlis qerqovani nawilis meorad hipofunqcias;
somatotropini (zrdis hormoni) - moqmedebs TiTqmis yvela tipis qsovilebze
da aCqarebs anabolur reaqciebs (cilebis, rnm-is da sxva makromolekulaTa
sinTezs); hiperproduqcia iwvevs akromegalias, xolo hipo - zrdis SeCerebas;
prolaqtini (laqtotropini) - ganapirobebs sarZeve jirkvlebis ganviTarebas
da laqtaciis regulacias, hiperproduqcia iwvevs amenoreas, uSvilobas,
xolo hipo - laqtaciis Sesustebas. folitropini da lulitropini
(gonadotropuli hormonebi) - moqmedeben sasqeso jirkvlebze, ganapirobeben
folikulebis momwifebas, spermatogenezs, estrogenebis da progesteronis
sinTezs, testosteronis warmoqmnis gaZlierebas; hiperproduqcia iwvevs
adreul sqesobriv momwifebas, xolo hipo - sqesobrivi jirkvlebis meorad
hipofunqcias, uSvilobas; melanotropini - ganapirobebs melanocitogenezs,
kanis pigmentovani ujredebis warmoqmnas; lipotropini - moqmedebs sxvadasxva
tipis ujredebze da aCqarebs cximebis daSlas; hiperproduqcia iwvevs
dauZlurebas, xolo hipo - gasuqebas.
winahipofizaruli hormonebi, iseve rogorc cila-peptidovani hormonebis
umravlesoba, maRalmolekuluri winamorbedis formiT sinTezirdebian. am
mxriv aRsaniSnavia 260 aminomJavisagan Semdgari cila proopiokortini,
romelic hidrolizis Sedegad 6 hormons warmoqmnis: - da -melanotropinebs,
- da -lipotropinebs, kortikotropins da -endorfins (endogenuri morfini).
es ukanaskneli warmoadgens peptids, romelsac analgeziuri moqmedeba gaaCnia.
mis SemadgenlobaSi Sedis 5 aminomJaviani peptidi - enkefalini, romelsac
aseve morfinis msgavsi aqtivoba aqvs.
hipofizis ukana nawili, neirohipofizi, gamoimuSavebs or hormons:
vazopresins da oqsitocins. es peptidebi sinTezirdebian hipoTalamusSi da
sxvadasxva nervuli gzebiT gadaitanebian neirohipofizSi. orive peptidi 9-9
aminomJavisagan Sedgeba. oqsitocinis ZiriTadi funqcia orsulobis dros
saSvilosnos kunTis regulaciasa da sarZeve jirkvlebSi arsebuli kunTovani
boWkoebis stimulaciaSi mdgomareobs.
vazopresini gluvi kunTebis, maT Soris sisxlZarRvebis, SekumSvas iwvevs.
mas agreTve Zlieri antidiuretuli efeqti axasiaTebs; igi Tirkmlis
milakebidan wylis aqtiur ukuSTanTqmas ganapirobebs. neirohipofizis

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atrofiis SemTxvevaSi, sisxlSi vazopresinis simciris gamo, organizmidan
didi raodenobiT wyali gamoiyofa (uSaqro diabeti). amrigad, vazopresinis
mTavar funqcias organizmSi wylis balansis daregulireba warmoadgens.
orive peptids centraluri moqmedebac axasiaTebT. vazopresins
nootropuli efeqti aqvs, igi mexsierebis gaumjobesebas iwvevs, xolo
oqsitocini ki, piriqiT, amneziur agents warmoadgens.
vazopresinis da oqsitocinir receptorebi efeqtoruli ujredebis
plazmur membranebzea lokalizebuli. vazopresinis moqmedeba
adenilatciklazuri sistemis gaaqtivebiT aris gapirobebuli. oqsitocinis
receptoris ujredSida katalituri komponentis raoba ucnobia. varaudoben,
rom igi dakavSirebulia kalciumis ionebis cvlasTan.

farisebraxlo jirkvlis hormoni

parathormoni farisebraxlo jirkvlis peptidovan hormons warmoadgens.

parathormoni moqmedebs Zvlebze da Tirkmelebze, kontrols uwevs kalciumisa
da fosfatis balanss sisxlSi.

farisebri jirkvlis hormonebi

farisebri jirkvlis folikulebSi sinTezirdeba maRalmolekuluri cila

- iodTireoglobulini (mol.wona - 650000), romelic hormon Tiroqsinis
samarago formas warmoadgens. igi mdidaria aminomJava TiroziniT, romlis
iodirebac garkveuli fermentatiuli reaqciebis saSualebiT xdeba. Sedegad
warmoiqmneba cilis molekulasTan dakavSirebuli tetra-(Tiroqsini) da
triiodTironini, romlebic proteolizuri reaqciebis Semdeg scildebian
cilas da Tavisufldebian sisxlSi. am reaqciebis stimulators
hipofizaruli hormoni, Tireotropini warmoadgens. Tireoiduli hormonebi
qsovilebis (tvinis garda) metabolizmis gaaqtivebas iwveven. maTi receptorebi
citoplazmaSi mdebareoben da birTvSi translokaciis Semdeg genTa
eqspresiaze moqmedeben. sinTezirebuli fermentebi ki aCqareben ujredis
metabolizms.
hipoTireozi bavSvebSi iwvevs kretinizms, xolo mozrdil asakSi - apaTias,
modunebas, paTologiur simsuqnes. xSirad es ukanaskneli dakavSirebulia
miqsedemasTan - hipoTireoidul SeSupebasTan. MmTian regionSi mcxovrebi
mosaxleobisTvis, wyalSi iodis Semcvelobis simciris gamo, damaxasiaTebelia

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endemuri Ciyvis ganviTareba. HiperTireozis (bazedovis daavadeba) SemTxvevebSi
adgili aqvs nivTierebaTa cvlis gaZlierebas, da cilebis intensiuri daSlis
gamo, uaryofiT azotur balanss. damaxasiaTebelia simptomebis triada:
taqikardia, egzoftalmizmi da Ciyvi. farisebri jirkvlis meore hormons
peptidi - kalcitonini warmoadgens. is jirkvalSi arsebul C-ujredebSi
sinTezirdeba da misi moqmedeba sawinaaRmdegoa parathormonisa. kalcitonini
ZvlebSi rezorbciuli procesebis daTrgunvas iwvevs, ris gamoc
hipokalciemia da hipofosfatimea viTardeba.

kuWqveSa jirkvlis hormonebi

kuWqveSa jirkvali Sereuli sekreciis mqone jirkvals miekuTvneba. is, erTi

mxriv, saWmlis momnelebel sistemaSi gamohyofs hidrolizur fermentebs,
xolo meore mxriv - monelebis procesSi monawile hormonebs sekretirebs
sisxlSi. pankreasis A-ujredebi gamoimuSaveben glukagons, B-ujredebi -
insulins, xolo C-ujredebi - somatostatins.
insulini sinTezirdeba proinsulinis saxiT da misi sinTezis regulaciaSi
didi mniSvneloba sisxlSi glukozis koncentracias eniWeba. sisxlSi
glukozis koncentraciis momatebisas insulinis sinTezi da gamoyofa
Zlierdeba, da piriqiT, glukozis raodenobis Semcirebis dros insulinis
produqciac klebulobs. insulini moqmedebs plazmuri membranis
receptorebze da maTi saSualebiT aaqtivebs glukozis ujredSi
matransportirebel sistemebs. am ukanasknels glukoza gulisa da kunTis
ujredebSi gadaaqvs, sadac is glikogenis formiT grovdeba.
insulini tipiur zrdis faqtors warmoadgens, da amitom misi nakleboba
sisxlSi hiperglikemiis garda, mTel rig anabolur reaqciaTa darRvevas
iwvevs. misi deficiti aCqarebs cximovani mJavebis daJangvas, glikogenolizs,
amcirebs cilebis sinTezs da zrdis ketonuri sxeulebis raodenobas
sisxlSi.
glukagoni cilovan hormons warmoadgens da 29 aminomJavisagan Sedgeba. igi
proglukagonis saxiT sinTezirdeba. glukagons, insulinis sawinaaRmdego
moqmedeba aqvs da igi zrdis glukozis koncentracias sisxlSi.
glukagonis hiperglikemiuri efeqti ganpirobebulia misi moqmedebiT
glikogenolizze. es hormoni, adrenalinis msgavsad aaqtivebs hepatocitebis
adenilatciklazur sistemas da kaskaduri meqanizmebis meSveobiT iwvevs
glikogenis daSlas da glukozis gamoTavisuflebas sisxlSi.

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 somatostatini, romelic wina ori hormonis msgavsad pankreasSi
sinTezirdeba, ainhibirebs, rogorc glukagonis, aseve insulinis sekrecias.

Tirkmelzeda jirkvlis hormonebi

Tirkmelzeda jirkvali morfologiurad da funqcionalurad

gansxvavebuli ori - tvinovani da qerqovani - nawilebisagan Sedgeba. tvinovani
nivTiereba miekuTvneba e.w. qromafinul sistemas, romelic eqtodermisagan
viTardeba, xolo qerqovani nivTiereba - mezenqimaluri warmoSobisaa da
epiTelialuri qsovilisagan Sedgeba. pirveli asinTezirebs aminomJavuri
warmoSobis hormonebs - kateqolaminebs, xolo meore - steroidebs.
kateqolaminebi - tvinovani nivTierebis hormonebs warmoadgenen. isini
Tirkmelzeda jirkvlis garda farTod arian gavrcelebuli centralur da
periferiul nervul sistemebSi. kateqolaminebis ZiriTadi warmomadgenlebi
arian: adrenalini, noradrenalini da dofamini. isini aminomJava
Tirozinisagan sinTezirdebian:

CH2 CHNH2 COOH CH2 CHNH2 COOH

OH
O2 CO 2

OH OH
Please Register

Tirozini dioqsifenilalanini (dofa)

CH 2 CH 2NH 2 CHOH CH2NH 2

OH O2 OH
CH 3
OH OH

dofamini noradrenalini

CHOH CH2NH CH3

OH

OH Please Register

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 adrenalini

adrenalini da noradrenalini mZlavri vazokonstriqtorebi arian, da amis

gamo, arterialuri wnevis gadidebas iwveven. sisxlSi maTi koncentraciis zrda
simpatikuri nervuli sistemis gaRizianebis analogiuria. garda amisa,
kateqolaminebi sisxlSi glukozis raodenobis matebas iwveven hepatocitebSi
maTi receptorebis adenilatciklazasTan SeuRlebisa da glikogenolizis
intensificirebis gamo.
steroiduli hormonebi. steroiduli hormonebi anu kortikosteroidebi
Tirkmelzeda jirkvlis qerqovan nivTierebaSi sinTezirdebian. isini
pirobiTad, maTi biologiuri efeqtis mixedviT SesaZlebelia, or jgufad
davyoT: glukokortikoidebad - naerTebad, romlebic cilebis, cximebis da
naxSirwylebis cvlaze moqmedeben, da mineralokortikoidebad - naerTebad,
romlebic marilebisa da wylis cvlaze moqmedeben.
kortikosteroidebi, qolesterinis, naRvlis pigmentebis, vitamini D-s,
sasqeso hormonebis msgavsad, ciklopentanperhidrofenantrenul birTvs
Seicaven da qolesterinidan sinTezirdebian. maTi warmoqmna regulirdeba
hipofizaruli hormoniT - kortikotropiniT. kortikosteroidebisaTvis, iseve
rogorc mTlianad steroiduli hormonebisTvis damaxasiaTebelia ujredze
moqmedebis citoplazmuri meqanizmi, receptorTa translokacia birTvSi da
genTa eqspresia. maTi qsovilebze zemoqmedebis Sedegad sinTezirdebian
aminomJavaTa da naxSirwylebis cvlaSi monawile fermentebi, romlebic
ujredis metabolizms aZliereben. kortikosteroidebis erT-erTi mTavari
warmomadgenelia kortizoli, romelsac rig SemTxvevebSi insulinis
sapirispiro moqmedeba axasiaTebs. glukokortikoidebs alergiisa da anTebis
sawinaaRmdego efeqti aqvT.
mineralkortikoidebis mTavar funqcias organizmSi wylis da marilebis
balansis daregulireba Seadgens. isini uzrunvelyofen qlorisa da natriumis
Sekavebas TirkmelebSi, da ganapirobeben kaliumis gamodevnas SardTan erTad.
wamyvan mineralkortikoids aldosteroni warmoadgens.
aldosteronis sekrecias Tirkmelzeda jirkvlebidan e.w. renin-
angiotenzinuri sistema aregulirebs. Tirkmlis jorjlovan aparatSi
sinTezirdeba proteolizuri fermenti - renini. es ukanaskneli, sisxlis
Sratis cilebSi Semaval 2-globulinidan CamoWris 10 aminomJavian fragments

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-angiotenzin I-s. warmoqmnil peptids SemdgomSi, specialuri fermentis
(angiotenzin-gardamqmneli fermentis) saSualebiT damatebiT ori aminomJava
Camoscildeba. miRebuli angiotenzini II aqtiur naerTs warmoadgens da
qerqovani nivTierebidan aldosteronis gamoTavisuflebas iwvevs.
aldosteroni ki, rogorc ukve iTqva, xels uwyobs Na+-is reabsorbcias
Tirkmlis milakebSi da K+-is gandevnas, ris gamoc izrdeba arteriuli wneva.
amrigad, aldosterons da mis agonistebs hipertenziuli moqmedeba
axasiaTebT, xolo antagonistebs - hipotenziuli.

sasqeso jirkvlebis hormonebi

sasqeso hormonebi mamakacis da qalis sasqeso jirkvlebSi (sakvercxeebSi)

sinTezirdebian. qalis sasqeso hormonebidan - estrogenebidan wamyvani
mniSvneloba estradiols eniWeba. mis gamoyofas sakvercxeebidan cikluri
xasiaTi aqvs da mTavar daniSnulebas qalis reproduqciuli sistemis
funqciuri uzrunvelyofa warmoadgens. estrogenebi iseve, rogorc mamakacis
sasqeso hormonebi - androgenebi da mesame Sereuli tipis hormoni -
progesteroni, qolesterinidan sinTezirdebian. qolesterinidan jer
warmoiqmneba mamakacis mTavari androgeni - testosteroni, xolo Semdeg ki
estradioli. progesteroni, iseve rogorc estrogeni, qalis yviTel sxeulSi
war- moiqmneba da xSirad, misi organizmSi gamoyofis xasiaTi estradiolis
sawinaaRmdegoa. progesteroni ganayofierebisas xels uwyobs nayofis
implantacias. orsulobis procesSi progesteronis raodenoba sisxlSi
matulobs, xolo mSobiarobis dros - qveiTdeba.
sasqeso hormonebis gamoyofa hipofizaruli hormonebiT - honadotropinebiT
regulirdeba. mamakacisa da qalis reproduqciuli sistemebis garda, sasqeso
hormonebi moqmedeben sxva qsovilebzec da organoebzec. magaliTad,
estrogenebi gavlenas axdenen tvinis garkveul ubnebze, xolo androgenebi
iwveven ConCxis kunTebis zrdas. amitom, androgenebs da mis warmoebulebs
xSirad sportsmenebi kunTovani masis gasaZliereblad iyeneben. maT anaboluri
steroidebi ewodebaT.

qsovilovani hormonebi

morfologiurad gansazRvruli organoebis - endokrinuli jirkvlebis

garda, hormonebi sxva qsovilebSic gamomuSavdebian.

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 saWmlis momnelebeli sistemis hormonebi ZiriTadad peptiduri bunebisa
arian. aRsaniSnavia is, rom am peptidebis umravlesoba gastrointestinaluri
sistemis garda aRmoCenilia centralur nervul sistemaSic. aRniSnuli
peptidebi monawileobas iReben peristaltikis regulaciaSi da sekreciis
procesSi. maTi ZiriTadi warmomadgenlebi arian: gastrini, qolecistokinini,
sekretini, somatostatini, vazoaqtiuri intestinaluri peptidi.
serotonini da histamini, kateqolaminebis msgavsad, biogenur aminebs
ganekuTvnebian. serotonini - aminomJava triftofanisgan, xolo histamini -
histidinisagan sinTezirdeba. serotonini warmoiqmneba saWmlis momnelebeli
traqtis garkveuli ujredebis (enteroqromafinuli), bronqialuri da
epifizis ujredebis mier. misi moqmedeba lokaluria da mdgomareobs
sisxlZarRvebis SeviwroebaSi, agreTve bronqebisa da gastrointestinaluri
sistemis gluvi kunTebis SekumSvaSi. histamini sinTezirdeba mastocitebSi
(poxier ujredebSi). igi lokalurad moqmedebs anTebiTi da alergiuli
reaqciebis dros, cvlis sisxlZarRvebis kedlebis gamtarunarianobas;
monawileobas iRebs SeSupebis warmoqmnaSi da qavilis SegrZnebaSi. rogorc
serotonini, aseve histamini centraluri neirogadamcemebic arian.
kininebi. kinin-kalikreinis sistema genezisisa da moqmedebis meqanizmis
mixedviT renin-angiotenzinis sistemas hgavs. am sistemis aqtiur hormons 9
aminomJaviani (nonapeptidi) bradikinini warmoadgens. igi warmoiqmneba sisxlis
plazmaSi arsebul proteolizur ferment - kalikreinis meSveobiT, lizin-
bradikininidan. es ukanaskneli, Tavis mxriv, CamoiWreba pankreasuli
kalikreinis meSveobiT plazmis 2-globulinidan. bradikinini lokalurad
monawileobs sisxlZarRvebis gafarToebaSi, anTebiTi da alergiuli
reaqciebis warmoqmnaSi, tkivilis da qavilis SegrZnebaSi.
gulis peptidebi. kardiomiocitebi gamoyofen peptidebs, romlebic diurezs
da natriurezs iwveven. amdenad, maTi moqmedeba renin-angiotenzinis sistemis
sawinaaRmdegoa. am peptidebs atriopeptinebi ewodeba. maTi receptori
ujredebis plazmur membranaze mdebareobs da SeuRlebulia
guanilatciklazasTan. isini ujredSi cGMP-is sinTezs iwveven.
prostaglandinebi araqidonis mJavisagan sinTezirdebian. prostaglandinebs
iseve, rogorc araqidonis mJavis sxva warmoebulebs - tromboqsanebs,
leikotrienebs, prostaciklinebs - Zlieri fiziologiuri aqtivoba aqvT. isini
monawileobas iReben anTebiTi procesebis ganviTarebaSi, vazokonstriqciaSi
da dilataciaSi, gluvi kunTebis spazmebSi, trombocitebis agregaciaSi da

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a.S. maTi receptorTa lokalizacia, fiziologiuri roli da moqmedebis
meqanizmi bolomde garkveuli ar aris.
zrdis faqtorebi hormonebis erT-erT saintereso da SedarebiT
Seuswavleli klasia. isini cilebs warmoadgenen da sinTezirdebian
sxvadasxva organoebSi da qsovilebSi (sanerwyve jirkvalSi, hipofizSi,
RviZlSi da a.S.). maTi gavrceleba sisxlis saSualebiT warmoebs, xolo
ujredze moqmedebis meqanizmiT hgvanan insulins. RviZlSi sinTezirdebian
faqtorebi - somatomedinebi A da B, romlebic garkveuli tipis ujredebis
zrdas da proliferacias iwveven. sanerwyve jirkvalSi warmoiqmneba cila,
romelsac nervebis zrdis faqtori ewodeba. am ukanasknels didi mniSvneloba
eniWeba simpaTikuri nervuli sistemis embrionalur ganviTarebaSi, mis zrdaSi
da diferencirebaSi. am jirkvalSi sinTezirdeba agreTve epidermisis zrdis
faqtori, romelsac kardiovaskularuli, renaluri da endokrinuli
aqtivobebi axasiaTebs. igi, garkveuli tipis ujredebis mimarT Zlier
mitogenurobas amJRavnebs da ujredTa proliferacias iwvevs. Tirkmelzeda
jirkvlebis ganviTarebasa da maT formirebaSi monawileobas iRebs e.w.
fibroblastebis zrdis faqtori, romelic aRmoCenilia tvinsa da hipofizSi.
sisxlis ujredebis zrdaSi da diferencirebaSi didi mniSvneloba eniWeba e.w.
trombocitebis agregaciis faqtors. zogierTi aseTi faqtori ujredebis
avTvisebian gadagvarebas iwvevs. zrdis faqtorebidan maT ricxvs ganekuTvnebian
TGF da TGF .
zrdis faqtorTa receptori plazmur membranazea lokalizebuli da mis
ujredSida domens (Tirozin)proteinkinaza warmoadgens. am receptoris
aqtivacia transkripciuli faqtorebis moqmedebas gansazRvravs.
imunuri sistemis hormonebi. imunuri sistemis ujredebi 50-mde cilovan
faqtors gamoyofen, romelTa saSualebiTac xdeba informaciis gadacema
imunuri reaqciis dawyebisa, formirebisa da damTavrebis Sesaxeb. imunuri
hormonebi iyofian limfikinebad - naerTebad, romlebsac limficitebi
warmoqmnian, da monokinebad - naerTebad, romlebic monocitebSi da msgavs
ujredebSi sinTezirdebian. aseTi hormonebia interferonebi, interleikinebi,
simsivnis nekrozis faqtori, ujredTa koloniebis warmomqmneli faqtorebi da
sxva.

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 organoTa bioqimia

saWmlis momnelebeli sistema

adamiansa da gare samyaros Soris arsebul nivTierebaTa cvlaSi

gaerTianebulia misi mravali organo, Tu qsovili, romlebsac esa Tu is
specialuri daniSnuleba akisriaT. am procesebSi mTavars saWmlis
momnelebeli sistema warmoadgens, radgan igi uSualo kontaqtSi imyofeba
garemosTan da mis saSualebiT axorcielebs organizmi energetikuli Tu
plastiuri masalebis cvlas.
saWmlis momnelebel sistemaSi xdeba saWmlis sami ZiriTadi komponentis -
cilebis, cximebis da naxSirwylebis daSla Semadgenel nawilebad. es procesi
aucilebelia imitom, rom ujredebi, romlebic gastro-intestinaluri
traqtis gare fenas warmoqmnian, ganvladi arian mxolod dabalmolekuluri
naerTebisTvis.
naxSirwylebis moneleba. adamianis saWmlis momnelebel sistemas SeuZlia
gadaamuSaos mcenareuli da cxoveluri warmoSobis mxolod garkveuli
polisaqaridebi - glikogeni da saxamebeli - maT ZiriTad struqturul
elementad D-glukozad. es procesi piris RruSi iwyeba, sadac nerwyvTan
erTad gamoiyofa fermenti amilaza. amilazis moqmedebis Sedegad miiReba
narevi, Semdgari glukozisa da sxva oligosaqaridebisagan.
polisaqaridebis daSla Tormetgoja nawlavSi grZeldeba aseve amilazis
saSualebiT. D-glukoza sabolood warmoiqmneba di- da sxva
oligosaqaridebidan wvrili nawlavis epiTelialur ujredebSi, sadac
fermenti saqaraza anu invertaza Slis saqarozas glukozamde da
fruqtozamde, laqtaza Slis laqtozas da a.S. garda amisa, epiTelur
ujredebSi D-fruqtozis, D-galaqtozis da D-manozis nawilobrivi gardaqmna
xdeba D-glukozad.
cilebis moneleba. cilebi saWmlis momnelebel traqtSi iSlebian
aminomJavebad. kuWSi moxvedrisas, cilebi iwveven hormoni gastrinis sekrecias,
romelic Tavis mxriv aaqtivebs Semofenili ujredebidan HCl-is da
pepsinogenis (fermenti pepsinis araaqtiuri forma) gamoyofas. dabali pH-is
gamo (pH=1,5-2,5), globularuli cilebis umravlesoba, da maT Soris
baqteruli cilebi, ganicdian denaturacias. amis Sedegad, isini advilad

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misawvdomi xdebian proteolizuri fermentebisaTvis. paralelurad,
pepsinogens Camoscildeba 42 aminomJavisagan Semdgari fragmenti, ris gamoc
pepsini aqtiur formaSi gadadis (avtokatalizi). pepsini cilebis daSlas
peptiduri bmis hidroliziT awarmoebs. misi aqtivobis mizans warmoadgens is
peptiduri bmebi, romlebic aromatuli aminomJavebisagan (Tirozini,
fenilalanini, triftofani) Sedgeba. am fermentis moqmedebis Sedegad
dabalmolekuluri peptiduri fragmentebi warmoiqmneba.
TormetgojaSi, mJave pH-is mqone sakvebis moxvedrisas iwyeba hormon
sekretinis gamoyofa sisxlSi, romelic bikarbonatis sekrecias iwvevs
kuWqveSa jirkvlidan. gamoyofili bikarbonati mJave pH-s aneitralebs pH 7 -
mde. garda amisa, Tormetgoja nawlavSi sakvebiT stimulirdeba hormoni
qolecistokininis sisxlSi gamoyofa. es ukanaskneli ki, neitralur pH-ze
moqmedi sami proteolizuri fermentis - trifsinis, qimotrifsinis da
karboqsipeptidazis gamoTavisuflebas ganapirobebs. aRniSnuli fermentebi,
pepsinis msgavsad, araaqtiuri formiT gamoiyofian da avtokatalizis
saSualebiT aqtivirdebian. trifsini Slis iseT peptidur bmas, romelic
aminomJava - lizinisa an argininisgan Sedgeba, qimotripsini Slis bmas,
Semdgars romelime aromatuli aminomJavisagan, xolo karboqsipeptidaza
awarmoebs TiToeuli aminomJavis Camocilebas cilis C-bolodan. wvril
nawlavSi damatebiT gamoiyofa fermenti - aminopeptidaza, romelsac
karboqsipeptidazis msgavsi moqmedeba axasiaTebs, magram misi aqtivoba
mimarTulia cilis sawinaaRmdego, N-bolosaken.
adamianis saWmlis momnelebel sistemaSi, zemoTaRwerili fermentebis
saSualebiT, cilis daSla aminomJavebad mimdinareobs. warmoqmnili
aminomJavebi Seiwovebian wvrili nawlavis epiTeliuri ujredebis mier da
transportirdebian sisxlis meSveobiT RviZlSi.
saWmlis momnelebeli sistemis proteolizur fermentebis disfunqciasTan
aris dakavSirebuli erT-erTi gavrcelebuli daavadeba - mwvave pankreatiti. am
paTologiis dros, kuWqveSa jirkvalSi arsebuli hidrolizuri fermentebi,
romlebic normaluri mdgomareobis dros araaqtiur formaSi imyofebian,
naadrevad aqtivirdebian da iwyeben kuWqveSa jirkvlis sakuTari ujredebis
daSlas. normaSi, es procesi rom Tavidan acilebuli iyos, kuWqveSa jirkvali
gamoimuSavebs specialur cilovan naerTs - trifsinis inhibitors. es
ukanaskneli ainhibirebs trifsinis moqmedebas, da Sesabamisad muxruWdeba
araaqtiuri fermentis aqtiur formaSi gadasvla.

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 cximebis moneleba. neitraluri cximebis - triacilglicerinebis moneleba
wvril nawlavSi warmoebs, sadac kuWqveSa jirkvlidan Tavisufldeba
araaqtiuri fermenti - prolipaza. SemdgomSi igi proteolizuri fermentebis
saSualebiT aqtiur formad - lipazad gardaiqmneba. lipaza, specialuri cilis
- kolipazis meSveobiT da naRvlis mJavebis Tanaobisas, awarmoebs
trigliceridebidan erTi an ori cximovani mJavis Camocilebas. Tavisufali
cximovani mJavebi, agreTve monogliceridi, warmoqmnis ra Na+ da K+ -is
marilebs (sapnebs) peristaltikis saSualebiT emulgirdebian patara wveTebad
(qilomikronebad) da STainTqmebian nawlavSi aresebuli wvrili limfaturi
kvanZebis mier. SemdgomSi qilomikronebi, laviwqveSa venis saSualebiT
vrceldebian cximovan qsovilebSi.
naRvlis mJavebi saWmlis momnelebeli sistemis erT-erT ZiriTad komponents
warmoadgenen, radgan maTi saSualebiT lipofiluri naerTebis da maT Soris
vitaminebis A, E da K-s emulgireba warmoebs. es ukanasknelni ki mxolod
saWmlis saSualebiT xvdebian organizmSi.
nawlavTa mikroflora. adamianis nawlavebSi, garda sakuTriv saWmlis
momnelebeli fermentebisa, aminomJavebis metabolur gardaqmnebs awarmoebs
mikroorganizmebis fermentebic. mikroorganizmebis mier aminomJavaTa
metabolizmis Sedegad warmoiqmnebian toqsikuri naerTebi, romlebic
aminomJavebTan erTad, karis venis gavliT xvdebian RviZlSi.
aseTi tipis naerTebs warmoadgens toqsikuri aminebi: fenileTilamini,
Tiramini, triptamini, putrescini, kadaverini da sxva, agreTve krezoli,
fenoli, indoli da sxva. maTi gauvnebleba RviZlSi mimdinareobs.
saWmlis momnelebeli sistemis hormonuli regulacia
gastro-intestinalur sistemaSi arsebuli organoebi endokrinul ujredebs
Seicaven, romelTa saSualebiTac sisxlSi da saWmlis momnelebel traqtSi
hormonTa sekrecia warmoebs. magaliTad, peptiduri hormonebi - gastrini,
qolecistokinini da sekretini iwveven hormoni insulinis gamoTavisuflebas
sisxlSi da fermentebis sekrecias nawlavebsa da kuWSi. garda amisa, gastrini
aaqtivebs marilmJavas gamoyofas, kuWis lorwovani garsis zrdas da mis
meqanikur muSaobas. qolecistokinini amuxruWebs kuWidan saWmlis gamosvlas
da aaqtivebs naRvlisa da proteolizuri fermentebis gamoyofas. sekretini
iwvevs bikarbonatebis sekrecias, da bolos, somatostatini - akavebs
gastrointestinaluri hormonebis moqmedebas.
gastrointestinaluri peptidebis gamoyofa nervuli kontroliT
xorcieldeba. am sistemis inervacia cTomili da Signeulobis nervebis

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saSualebiT xdeba, xolo am ukanasknelis agzneba Tu Sekaveba Tavis mxriv
hipofizuri hormonebiT regulirdeba. magaliTad, orsulobis periodSi,
organizmSi estrogenebis raodenoba matulobs. es ukanaskneli hipofizidan
peptiduri hormonis - oqsotocinis gamoyofas iwvevs. sisxlSi oqsitocini
cTomili nervis mgrZnobiare gangliebze moqmedebs, iwvevs maT gaaqtivebas, da
Sesabamisad, aZlierebs gastrointestinaluri peptidebis gamoyofas. es
peptidebi, da kerZod qolecistokinini, Tavis mxriv cTomili nervis
aferentul, centralur nervul sistemaSi mimaval gzebs aaqtivebs. amis
Sedegad, Tavis tvinSi gamoiyofa peptidebi (endorfini, oqsotocini,
qolecistokinini da sxva), romlebsac somnogenur efeqti (Zils,
moTenTilobas) aqvT.

RviZli

RviZlis roli naxSirwylovan cvlaSi.. RviZlis funqcia naxSirwylovan

cvlaSi mdgomareobs sisxlSi glukozis mudmivi koncentraciis SenarCunebaSi,
romelic glikogenis daSlisa da sinTezis regulaciiT xdeba. RviZlSi
warmoqmnili Tu transportirebuli glukoza SesaZlebelia 5 gziT
metabolizdes. centralur metabolits, romlidanac xuTive gza iwyeba,
glukozo-6-fosfati warmoadgens. RviZlSi arsebuli fermenti glukokinaza,
romelic glukozidan glukozo-6-fosfatis sinTezs awarmoebs, gansxvavdeba
sxva qsovilebSi warmodgenil analogiuri fermenti - heqsokinazisagan imiT,
rom mas jer-erTi glukozis mimarT maRali Km gaaCnia, da meorec - igi ar
inhibirdeba reaqciis produqtiT - glukozo-6-fosfatiT. RviZlSi moxvedrili
sxva naxSirwylebic glukozo-6-fosfatad gardaiqmnebian.
RviZlidan glukoza SesaZlebelia sisxlSi gadavides. glukozo-6-fosfati
ganicdis defosforilirebas da Tavisufali glukozis saxiT sisxlSi
transportirdeba. am gzas didi mniSvneloba eniWeba, radgan sxva organoebis
energetikuli momaragebisaTvis glukozis koncentracia sisxlSi mudmivi
unda iyos.
me-2 gzis saSualebiT glukozo-6-fosfati gardaiqmneba glikogenad. am
dros glukozis Warbi raodenoba glikogenis formiT Seinaxeba. glukozo-6-
fosfatis Warbi raodenoba, romelic RviZlSi rCeba misi sisxlSi
transportirebisa da glikogenad gardaqmnis Semdeg, Sedis glikolizis
reaqciebSi (me-3 gza). am procesis Sedegad warmoqmnili acetil-CoA-dan jer

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cximovani mJavebis, xolo Semdeg neitraluri da fosfolipidebis sinTezi
xdeba. cximebi, sisxlis lipoproteinebis saSualebiT gadaitanebian sxva
qsovilebSi.
glukozis moxmarebis me-4 gzas - misi saboloo daJangva warmoadgens
limonmJava ciklSi, xolo me-5 gzas ki - pentozofosfaturi gardaqmnebi,
romlis Sedegad warmoqmnili NADPH da ribozo-5-fosfati, Sesabamisad
cximovani mJavebis da dezoqsiribonukleotidebis biosinTezSi gamoiyenebian.
glukozis utilizaciisa da moxmarebis gzebis gadarTva warmoebs
regulatoruli fermentebisa da hormonebis saSualebiT.
RviZlis roli aminomJavaTa cvlaSi. RviZlSi aminomJavaTa gardaqmna aseve
ramdenime gziT mimdinareobs. pirvel gzas warmoadgens maTi CarTva cilebis
sinTezis procesSi. unda aRiniSnos, rom RviZlis ujredebSi-hepatocitebSi
sisxlis plazmis cilebis ZiriTadi nawili sinTezirdeba (albuminebis -100%,
-globulinebis - 75-90%, -globulinebis - 50%, garda amisa, RviZlSi
sinTezirdebian proTrombini, fibrinogeni da sxva).
RviZlSi aminomJavebis gardaqmnis meore gzas maTi daSla da dezaminireba
warmoadgens. am dros warmoqmnili acetil-CoA SesaZlebelia gamoyenebul iqnas
an cximovani mJavebis biosinTezSi an glukoneogenezSi. adamianis organizmSi
aminomJavebis aminuri azotidan Sardovana sinTezirdeba da misi warmoqmna
RviZlSi mimdinareobs. aminomJavebis gardaqmnis mesame gzas qolinis,
kreatinis, hemis da nukleotidebis sinTezi warmoadgens.
RviZlis daavadebaTa dros sisxlis cilebis fraqciebis raodenobriv
cvlilebebs, ara marto diagnostikuri, aramed prognostikuri mniSvnelobac
aqvs. RviZlis paTologiebis dros biosinTezuri reaqciebi am organoSi
daTrgunulia; da Tu am dros sisxlSi albuminis raodenoba Semcirda,
qveiTdeba plazmis onkozuri wneva, rasac SeSupebebis, xolo xSir SemTxvevebSi
ascitis ganviTarebac ki SeiZleba mohyves.
RviZlis roli lipidur cvlaSi. RviZlSi lipidebic 5 gziT SeiZleba
gardaiqmnas. pirveli gzis meSveobiT cximovani mJavebi, romlebic lipidTa
cvlis centralur metabolits warmoadgenen, iSlebian jer acetil-CoA-mde,
xolo Semdeg naxSirorJangamde da wylamde. am dros gamoyofili energia
biosinTezuri reaqciebisTvis moixmareba. meore gzis saSualebiT, cximovani
mJavebis daJangvis Sedegad miRebuli acetil-CoA-s Warbi raodenoba
SesaZlebelia ketonur sxeulebad - acetoacetatad, acetonad da -
oqsibuTiratad gardaiqmnas da sisxlis meSveobiT gavrceldes periferiul
organoebamde. ketonuri sxeulebi SesaZlebelia warmovidginoT rogorc

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acetilis jgufis gadamtanebi, romelTa energetikuli wvlili periferiuli
qsovilebis metabolizmSi SesaZlebelia didi iyos (magaliTad, gulis kunTis
energetikuli moTxovnilebis 1/3 am gzis gamoyenebiT kmayofildeba).
me-3, da Zalze mniSvnelovan gzas, qolesterinisa da naRvlis mJavebis
biosinTezi warmoadgens. qolesterini acetil-CoA-dan warmoiqmneba. misi erTi
nawili sxva qsovilebSi gadaitaneba steroidebisa da sxva naerTebis
biosinTezisaTvis, xolo meore - naRvlis mJavebis sinTezSi moixmareba. me-4
gziT cximovani mJavebi sisxlis lipoproteinebis gamoyenebiT
transportirdebian sxva qsovilebSi; da bolos, Tavisufali cximovani mJavebi
ukavSirdebian Sratis albumins da am cilis meSveobiT gadaitanebian
periferiul organoebamde (me-5 gza). am ukanaskneliT kunTovani qsovilebi
ZiriTad energetikul masalas iReben.
amrigad, RviZli sakveb nivTierebaTa mTavar gamanawilebels warmoadgens.
igi uzrunvelyofs sxva organoebsa da qsovilebs energetikuli masaliT,
amcirebs sakvebi nivTierebis donis meryeobas saWmlis miRebis Semdeg.

detoqsikaciuri procesebi RviZlSi. RviZlSi moxvedrili ucxo naerTebi

(qsenobiotikebi) naklebad toqsikur nivTierebebad gardaiqmnebian. ZiriTadad,
aseTi naerTebis gauvnebelyofa ori gziT xdeba: koniugirebiT (sxva naerTTan
SeerTebiT) da daJangviT. garda amisa maT SesaZlebelia ganicadon
meTilireba, acetilireba da sxva.
detoqsikaciur procesebSi wamyvani mniSvneloba daJangvas eniWeba. daJangvis
reaqciebis warmarTvisaTvis hepatocitebSi specialuri organoidebi -
peroqsisomebi arseboben. garda amisa, es reaqciebi maRali efeqturobiT
mikrosomebSic (endoplazmur badeze) mimdinareoben. daJangvis Sedegad
qsenobiotikebis umravlesoba wyalSi xsnadi xdeba da maTi daSla
hidrolizuri fermentebiT advildeba. koniugirebis reaqciebSi monawileobs
glukuronis mJava, tripeptidi glutaTioni da sxva polaruli naerTebi. maTi
mierTebis Sedegad qsenobiotiki advilad idevneba organizmidan.
RviZlSi, toqsikuri nivTierebebis garda iSlebian bunebrivi cikluri
naerTebic: steroiduli hormonebi, kateqolaminebi da sxva.
RviZlis mniSvneloba pigmentur cvlaSi. hemoglobinis daSla
retikuloendoTelur ujredebSi mimdinareobs da pirvel produqts, romelic
am dros miiReba, aris verdhemoglibini. Semdgom stadiebze verdhemoglobins
cildeba rkina, da darCenili hemi globinis molekulidan disocirdeba.
cilovani nawilidan Camocilebul hems biliverdini ewodeba. igi Sedgeba

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oTxi piroluri birTvisagan, romlebic erTmaneTTan meTinis xidakebiT arian
SeerTebulni. biliverdinis aRdgeniT miiReba bilirubini - pigmenti, romelic
SemdgomSi naRvelTan erTad gamoiyofa, da amitom mas naRvlis pigmenti
ewodeba. warmoqmnili bilirubini naklebad polarulia, wyalSi cudad
ixsneba da misi raodenobrivi analizi sisxlSi mxolod spirtiT winaswari
damuSavebis Semdeg SeiZleba. amitom aseT bilirubins arapirdapiri
bilirubini ewodeba.
RviZlSi moxvedris Semdeg, bilirubini glukuronis mJavasTan koniugirdeba
(ukavSirdeba). warmoqmnili glukokoniugati polarul naerTs warmoadgens,
wyalSi advilad ixsneba, da amis gamo sisxlSi pirdapir, winaswari
damuSavebis gareSe isazRvreba. mas pirdapiri bilirubini ewodeba. pirdapiri
bilirubini naRvelTan erTad gadadis wvril NnawlavSi. msxvil nawlavSi igi
anaerobuli mikrofloris saSualebiT gardaiqmneba sterkobilinad da
gamoiyofa organizmidan. sterkobilinis mcire nawili (mas arasworad
urobilins uwodeben) Seiwoveba nawlavebSi da Tavisufldeba SardTan erTad.
klinikaSi bilirubinisa da misi fraqciebis gansazRvras wamyvani
mniSvneloba eniWeba siyviTlis diferencirebul diagnostikaSi. hemolizuri
siyviTlis dros hiperbilirubinemia gamowveulia arapirdapiri bilirubinis
momatebiT. es ganpirobebulia imiT, rom eriTrocitebis intensiuri daSlis
gamo, RviZli ver aswrebs mis koniugirebas. arapirdapiri bilirubinis
TirkmelebSi gausvlelobis gamo, am dros SardSi misi mateba ar aRiniSneba.
RviZlis paTologiebidan ganviTarebuli siyviTle ganpirobebulia imiT,
rom hepatocitebis mier glukokoniugirebuli, pirdapiri bilirubinis
naRvlis arxebSi eqskrecia SeuZleblia, ris gamoc is sisxlis nakadSi xvdeba.
garda amisa, hepatocitebis mier glukokoniugatebis sinTezis SemcirebiT,
sisxlSi izrdeba, agreTve, arapirdapiri bilirubinic.
obturaciuli siyviTlis dros naRvlis sadinrebis gadaketvis gamo
sisxlSi mkveTrad matulobs pirdapiri bilirubini.
naRveli warmoadgens moyviTalo-moyavisfro feris Txevad sekrets, romelic
RviZlis ujredebidan gamoiyofa. NnaRvlis buStSi warmoebs naRvlis
koncentrireba eleqtrolitebisa da wylis adsorbciis Sedegad. naRveli
didi raodenobiT Seicavs bilirubins, naRvlis mJavebs, qolesterins. naRvlis
mJavebis (romlebic detergentebs warmoadgenen) ukmarisobis SemTxvevaSi
qolesterini veRar ixsneba da warmoqmnis naleqs (kenWebs).

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 sisxli

organoebs Soris nivTierebaTa cvla sisxlis saSualebiT warmoebs. mas

gadaaqvs sakvebi nivTierebebi da maTi daSlis produqtebi, agreTve, Jangbadi,
naxSirorJangi, hormonebi da mediatorebi. adamianSi sisxlis moculoba 5-6 l
tolia, romlis naxevars eriTrocitebi ikaveben. SedarebiT ufro mcire
raodenobiT arian masSi leikocitebi da trombocitebi. sisxlis xsnadi
nawili, plazma, 90% wylisagan da 10% masSi gaxsnili nivTierebebisagan
Sedgeba. am nivTierebebidan daaxloebiT 70%-s e.w. plazmis cilebi
warmoadgenen.
plazmis cilebi RviZlSi da retikuloendoTelialur sistemaSi
sinTezirdebian. SesaZlebelia gamoiyos am cilebis ramdenime funqcia.
upirveles yovlisa isini monawileobas iReben koloido-osmosuri (onkozuri)
wnevis warmoqmnaSi, riTac did rols TamaSoben arteriuli da venuri
sisxlis mimarTulebaTa formirebaSi. onkozuri wnevis Semcireba iwvevs
SeSupebis ganviTarebas. meore - plazmis cilebi monawileobas Rebuloben
sisxlis SededebaSi (fibrinogeni). Mmesame - plazmis cilebi, warmoadgenen ra
amfoterul naerTebs, ganapirobeben pH-is stabilizacias. meoTxe - plazmis
cilebs satransporto funqciebi akisriaT, maT mravali naerTi gadaaqvT,
gansakuTrebiT cximovani bunebisa (steroiduli hormonebi, qolesterini,
bilirubini da a.S.). mexuTe - plazmis cilebs wamyvani mniSvneloba eniWebaT
imunitetis ganviTarebaSi.
plazmis cilebis ZiriTad masas (55-60%) albumini Seadgens. albuminis
molekuluri wona 70000-is tolia. plazmis onkozur wnevas albumini
ganapirobebs. albumini ikavSirebs da gadaaqvs bilirubini, qolesterini da
sxva naerTebi. globulinebis fraqcia Sedgeba , da - globulinebisagan,
romlebic erTmaneTisagan sxvadasxva bioqomiuri meTodebis (eleqtroforezi,
qromatografia da a.S.) saSualebiT iyofian. da -globulinebi Seicaven
lipoproteinebs, - globulinebSi ki warmodgenilia sisxlSi cirkulirebadi
antisxeulebis ZiriTadi raodenoba.
plazmis hiperproteinemia (xSirad dakavSirebulia hiperglobulinemiasTan)
ganpirobebulia wylis didi raodenobiT dakargvasTan (gulis reva, kanis didi
farTis damwvroba da sxva), xolo hipoproteinemia (xSirad dakavSirebulia
hipoalbuminemiasTan) nefrozuli sindromebis erT-erT mTavar simptoms
warmoadgens. globulinebis fraqciis cilebi ZiriTadad glikoproteinebi
arian.

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 plazmis lipoproteinebi warmoadgenen rTul kompleqsur SenaerTebs isini
Sedgebian arapolaruli lipidebis wveTisagan, romlis irgvlivac
garSemortymulia polaruli lipidebis, qolesterinis da cilis Sre. -
lipoproteini anu maRali simkvrivis lipoproteini Seicavs -globulinebs,
xolo -lipoproteini anu dabali simkvrivis lipoproteini - -globulinebs.
lipoproteinebi sinTezirdebian RviZlSi. maTi raodenoba sxvadasxva tipis
aTerosklerozul daavadebebTan korelaciaSi imyofeba. gulis iSemiuri
daavadebebis dros lipoproteinebSi arsebuli qolesterini matulobs.
sisxlis plazmis cilebidan klinikuri TvalsazrisiT garkveul interess
warmoadgenen: haptoglobini- 2 globulinis cila, romelic ukavSirdeba
hemoglobins da masTan kompleqsSi STainTqmeba makrofagebiT. am kavSiris
wyalobiT organizmidan rkina ar ikargeba; transperini - -globulinebis
fraqciis cila, romelsac rkinis atomi gadaaqvs; ceruloplazmini - spilenZis
Semcvel cilas warmoadgens, misi raodenoba hepatocerebraluri distrofiebis
dros mcirdeba; a-fetoproteini - erT-erT karcinoembrionalur cilas
warmoadgens, didi raodenobiT gvxvdeba nayofSi, qalebis sisxlSi orsulobis
dros da hepatokarcinomebiT daavadebul adamianebSi; C-reaqtiuli cila -
saxelwodeba warmoqmnilia misi unariT daleqos pnevmokokebis C-polisaqaridi,
normaSi ar aRiniSneba; misi raodenoba sisxlSi matulobs qsovilTa anTebiTi
da nekrozuli procesebis dros, ganisazRvreba 2-globulinebis fraqciaSi;
krioglobulini - -globulinebis fraqciis cila; janmrTeli adamianebis
sisxlSi ar moipoveba, magram warmoiqmneba mielomebis, nefrozebis, RviZlis
cirozis, revmatizmebis, leikozebis da sxva msgavsi daavadebebis dros.
sisxlSi isazRvreba agreTve simsivnis markeruli cilebi: prostat-
specifikuri antigeni, karcino-embrionuli antigeni, da sxva.
sisxlis plazmaSi arsebuli fermentebi SesaZlebelia or jgufad daiyos:
sekretorulad da indikatorulad. sekretoruli fermentebi RviZlSi
sinTezirdebian da maT sisxlSi garkveuli roli akisriaT. aseT fermentebs
warmoadgenen sisxlis SededebaSi monawile komponentebi, qolinesTeraza da
sxv. indikatoruli fermentebi sisxlSi sxvadasxva qsovilebidan xvdebian.
magaliTad, mwvave infarqtis dros, kardiomiocitebidan sisxlSi gadmodian
fermentebi kreatinkinaza, aspartatminotransferaza da sxv., xolo RviZlis
daavadebebis dros - alaninaminotrasferaza, glutamatdehidrogenaza da sxv.
cilovani komponentebis garda sisxlis plazmaSi warmodgenilia
dabalmolekuluri azotovani naerTebi: Sardovana, aminomJavebi, Sardis mJava,
kreatini, kreatinini, amoniaki. garkveuli paTologiebis dros aracilovani

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azotis raodenoba plazmaSi izrdeba. ganasxvaveben retenciul da produqciul
azotemias. retenciuli azotemiis dros azotovani naerTebis koncentracia
sisxlSi matulobs Tirkmlebis ukmarisobis, arteriuli wnevis Semcirebis da
sisxlis mimoqcevis daqveiTebis gamo.
produqciuli azotemiis SemTxvevaSi, azotovani naerTebis raodenobrivi
zrda ganpiribebulia qsovilebSi cilebis intensiuri daSliT, rasac adgili
aqvs vrceli anTebiTi procesebis, Wrilobebis, damwvrobis, kaxeqsiis dros.
sisxlis plazmis eleqtrolituri Semadgenloba. plazmis ZiriTad osmoaqtiur
ions Na warmoadgens. natriumis koncentracia plazmaSi tolia 132-150 mmoli/l.
hipernatriemia vlindeba parenqimatozuri nefritis, Tandayolili gulis
ukmarisobis, hiperaldosteronizmis dros. natriumis koncentracia plazmaSi
8-jer aRemateba am ionis koncentracias eriTrocitSi.
kaliumis ionTa koncentracia plazmaSi tolia 3,9-5,4 mmoli/l da igi 20-jer
naklebia eriTrocitebSi arsebuli kaliumis koncentraciaze.
hiperkaliemia aRiricxeba Tirkmlebis mwvave ukmarisobis dros da
Tirkmelzeda jirkvalis qerqovani nivTierebis hipofunqciis Sedegad. da
piriqiT, Tirkmelzedas hiperfunqciisa, da Sesabamisad aldosteronis Zlieri
gamomuSavebis dros adgili aqvs hipokaliemias.
kalciumis ionTa koncentracia plazmaSi tolia 2,2-2,8 mmoli/l da igi 1000
jer ufro metia am ionis koncentraciaze eriTrocitebSi. diagnostikuri
mniSvneloba eniWeba kalciumis gansazRvras hipokalcinemiis dros. es
mdgomareoba aRiniSneba hipoparaTireozis, agreTve raqitis, nefrozebis,
glomeluronefritebis drosac.
magniumis koncentracia plazmaSi tolia 0,8-1,5 moli/l. eriTrocitebSi ki es
maCvenebeli udris 2,4-2,8 moli/l.
fosfatebis raodenobis Semcireba plazmaSi aRiniSneba raqitis dros.
rkina. sisxlSi rkina ZiriTadad eriTrocitebSi gvxvdeba. mis mTavar
rezervuars organizmSi Zvlis tvini (adamianis mTavari eriTropoeturi
qsovili), RviZli da Tirkmelebi warmoadgenen. am organoebSi rkina
Tormetgojadan transferinis saSualebiT gadaitaneba.
sisxli mdidaria mikroelementebiT: iodiT, spilenZiT, cinkiT, kobaltiT,
seleniT da sxva.
sisxlis ujredebi23. adamianis 1 mikrolitri sisxli 5 x 106 eriTrocits
Seicavs. es ujredebi Zvlis tvinSi warmoiqmnebian. eriTrocitebi moklebuli
arian birTvs da mTlianad hemoglobiniT arian Sevsebuli. gacilebiT ufro
mcire (1000-jer) raodenobiT arian sisxlSi leikocitebi, romlebsac

167
eriTrocitebisagan gansxvavebiT, birTvi da sxva organoidebi gaaCnia.
arsebobs leikocitebis sami tipi: limfocitebi, monocitebi da
granulocitebi. granulocitebi sxvadasxva tipis saRebavebiT iRebebian,
riTic ganasxvaveben neitrofilebs, bazofilebs da eozinofilebs.
neitrofilebis mTavar funqcias fagocitozi warmoadgens; eozinofilebi
monawileobas Rebuloben parazitebisagan dacviT da alergiul reaqciebSi,
xolo bazofilebi - alergiul reaqciebSi. limfocitebis mTavar funqcias
antisxeulebis produqcia da imunuri pasuxis organizacia warmoadgens.
da bolos, sisxlSi warmodgenili arian trombocitebi, romlebic sisxlis
SededebaSi iReben monawileobas.
eriTropoezis regulacia. sisxlis ujredebi Zvlis tvinSi warmoiqmnebian
erTi saerTo winamorbedisagan - Rerovani ujredebisagan. jer kidev
gaurkveveli signalebis meSveobiT, es ujredebi iwyeben diferencirebas ori
tipis - limfoiduri da mieloiduri ujredebis warmoqmniT. mieloiduri
ujredebisagan miiRebian eriTrocitebi, neitrifilebi,
monocitebi(makrofagebi), eozinofilebi, bazofilebi da megakariociti,
romlisganac SemdgomSi trombocitebi viTardebian. limfoiduri ujredebisagan
warmoiqmnebian B da T limficitebi.
eriTropoezi (eriTrocotebis warmoqmna) sisxlSi Jangbadis raodenobiT
regulirdeba. sisxlSi Jangbadis Semcirebisas Tirkmlebi iwyeben
eriTropoetinis sekrecias. es ukanaskneli cilovan hormons warmoadgens da
misi molekuluri wona tolia 34000. sisxlSi gamoTavisuflebis Semdeg
eriTropoetini moqmedebs mieloidur ujredebze da iwyebs maTi
diferencirebis process.
msgavsi peptiduri da cilovani hormonebiT warmoebs limfocitebis
diferencirebac. am procesSi monawileobas iReben ori tipis peptiduri
hormonebi: limfokinebi da monokinebi. monokinebi monocitebSi (makrofagebSi)
sinTezirdebian da maTi produqcia imunuri reaqciis pirvelad pasuxs
warmoadgens. monokinebi SerCeviTad moqmedeben mieloidur da limfoidur
ujredebze da iwveven maTSi limfokinebis produqcias. gamomuSavebuli
limfikinebi aZliereben imunur pasuxSi monawile ujredebis proliferacias,
diferenciacias da maTgan specifiuri ujreduli klonebis Camoyalibebas.
monokinebis ricxvs ganekuTvnebian interleikini-1, simsivnis nekrozis faqtori
da sxva, xolo limfokinebisa ki - interleikini-3, klonebis warmomqmneli
faqtorebi, interferoni da sxva. am cilovani hormonebis saSualebiT
formirdeba maRalspecifiuri da swrafi imunuri pasuxi.

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 sisxlis buferuli sistema. sisxlis pH neitraluria da tolia 7,4. pH-is es
mniSvneloba ganpirobebulia sisxlis buferuli sistemebiT. sisxlis buferuli
sistema, ZiriTadad, ori tipis marilis koncentracias emyareba: karbonatebisa
da fosfatebisa. sisxlis buferuli sistemis formirebaSi didi wvlili
cilebsac SeaqvT. sisxlSi mJava-tutovani balansis darRvevis SemTxvevaSi
adgili aqvs acidozis (sisxlis mJavianobis momateba - pH-is Semcireba) an
alkalozis (sisxlis tutianobis momateba - pH-is gazrda) ganviTarebas.
ganasxvaveben sunTqviT da metabolur acidozebs ( an alkalozebs).
sunTqviTi acidozi sunTqviTi moculobis Semcirebisas warmoiqmneba (mag.,
bronqialuri astmis, filtvebis SeSupebis, emfizemis da sxva analogiur
daavadebaTa dros). am SemTxvevaSi filtvebis hipoventilirebis gamo xdeba
Pco2 (CO2-is parcialuri wneva) momateba sisxlSi. Sedegad H2CO3-is
koncentracia plazmaSi matulobs, da amitom, misi pH mcirdeba.
metaboluri acidozi dakavSirebulia nivTierebaTa cvlis darRvevebTan da
ganpirobebulia organuli mJavebis dagrovebiT. pH-is aseT cvlilebas adgili
aqvs diabetis, SimSilis, cieb-cxelebis, Sokis dros.
sisxlis Sededebis sistema. sisxlis Sededebis procesi bioqimiuri
reaqciebis rTul kaskads warmoadgens, romelSic monawileobas iRebs 15-mde
plazmis da 10-mde trombocituli faqtori. aRniSnuli faqtorebi aRiqvamen
sisxlZarRvis dazianebis keras da CarTaven reaqciaTa Tanmimdevrobas,
romlis Sedegad araaqtiuri fermenti-proTrombini gardaiqmneba aqtiur
Trombinad. Trombini awarmoebs fibrinogenidan ori peptidis (peptidi A da
peptidi B ) CamoWras. am reaqciis Semdeg miRebuli fibrin-monomeri, kalciumis
ionebis Tanaobisas, swrafad polimerizirdeba fibrin-polimerSi. es
ukanaskneli wyalSi uxsnadia.

plazmis da Trombocituli faqtorebi

proTrombini Trombini

fibrinogeni fibrin-monomeri

fibrin-polimeri
Ca2+

sisxlis
169 Sededeba
 sisxlis Sededebis sistemis erT-erT stimulators vitamini K warmoadgens.
igi iwvevs RviZlSi proTrombinisa da rigi cilovani faqtorebis sinTezs.
amitom, antikoagulantebis nawili K-s antivitaminebi arian.
farTod gavrcelebul antikoagulants heparini warmoadgens. igi xels
uSlis Trombinis moqmedebas fibrinogenze. organizmSi arsebobs fibrinuli
Trombebis damSleli mZlavri endogenuri sistema, romelsac fibrinolizis
sistema ewodeba.

Tirkmelebi da Sardi

Tirkmelebis ZiriTad funqcias warmoadgens organizmSi mudmivi da

stabiluri endogenuri pirobebis SenarCuneba. Tirkmlis qsovili or zonad
iyofa: zeda - qerqovani da qveda - tvinovani. Tirkmlis ujredSi - nefronSi
SesaZlebelia Semdegi anatomiuri warmonaqmnebis gamoyofa: eferentuli
arteriola, romelic gadadis Tirkmlis jorjalSi (glomerula), jorjlis
irgvliv arsebuli kafsula (biumenis kapsula), nefronis proqsimaluri arxi,
nefronis yulfi (henles maryuJi), nefronis distaluri arxi, Semagrovebeli
milaki. nefronSi sami ZiriTadi procesi mimdinareobs: filtracia gorglebSi,
reabsorbcia da sekrecia arxebSi.
filtracia. filtracia, romelic jorjalSi mimdinareobs pasiur process
warmoadgens. filtracias ganicdis sisxlis plazmis dabalmolekuluri
organuli naerTebi da ionebi. normalur mdgomareobaSi cilebi Tirkmlis
filtrSi ar gadian. maTi (ZiriTadad albuminebis) gasvla filtrSi
proteinuriis erT-erTi gamomwvevi mizezTagania. filtratSi gasul siTxes
pirveladi Sardi ewodeba.
reabsorbcia da sekrecia. pirveladi Sardis mxolod 1% gamodis Sardis
saxiT organizmidan. nivTierebebis danarCeni da ZiriTadi masa, gansakuTrebiT
ki wyali, nefronis arxis gavlis dros reabsorbirdeba sisxlSi. arxebSi
absorbirdeba agreTve natriumis, qloris, hidrokarbonatebis, aminomJavaTa
99%, kaliumis 93%, Sardovanas 45% da a.S. Sedegad warmoiqmneba meoradi
Sardi, romelic SardsawveTis saSualebiT gadadis Sardis buStSi.
natriumis STanTqma arxebSi aqtiur process warmoadgens e.i. mis
adsorbciaze ukan sisxlSi ixarjeba ATP-s energia. radgan sisxlSi Na+
koncentracia ufro maRalia, vidre arxebSi, am process fermenti Na,K- -
ATPaza anxorcielebs. rogorc cnobilia, am ferments natriumi gadaaqvs

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koncentraciuli gradientis sawinaaRmdegod. paralelurad mas, aseve
koncentraciuli gradientis sawinaaRmdegod,sisxlidan arxebSi Semoaqvs
kaliumi. ionTa koncentraciuli gradientis sawinaaRmdegod gadatana
energodamokidebul process warmoadgens da TiToeul molekula Na-is
gadatanaze da K-is Semotanaze ixarjeba ATP-is erTi molekula. amis gamo,
SardTan erTad idevneba mxolod K+, maSin rodesac natriumis danakargebi
Zalze mcirea. Na-is transportirebis Sedegad, endosmosuri Zalebis
moqmedebis gamo, reabsorbirdeba agreTve wyalic.
natriumis garda aqtiuri transportis meqanizmebiT, romlebSic
monawileobas iReben ATPazebi, transportirdebian sxva nivTierebebic,
magaliTad, glukoza da aminomJavebi. maTi koncentraciac, natriumis msgavsad,
arxebSi ufro dabalia, vidre sisxlSi. TirkmelebSi arseboben iseTi
sistemebic, romlebic nivTierebaTa transportisaTvis gamoiyeneben ara ATP-s
energias, aramed Na-is koncentraciuli gradientis energias. am sistemebis
meSveobiT Na SesaZlebelia kvlav Semovides arxebSi da Tavidan
transportirdes sisxlSiAATP-aziT.
organizmSi arseboben mZlavri hormonaluri sistemebi, romlebic
Sardwarmoqmnas aregulireben. aseT sistemebs warmoadgenen renin-
angiotenzinuri, gulis peptidebis natriurezuli, vazopresinis da sxva
sistemebi.
Tirkmelebs didi mniSvneloba eniWebaT sisxlis mJava-tutovani
wonasworobis SenarCunebaSi. mTavari roli am procesebSi natriumis ionebs
akisriaT, romelTa TandaTanobiTi reabsorbcia pirveladi Sardidan
gansazRvravs pH-is cvlilebebs meorad SardSi da sisxlSi. Tirkmelebi
maRali metaboluri aqtivobiT xasiaTdebian. mosvenebis mdgomareobaSi,
adamianis mier STanTqmuli Jangbadis 8-10% Tirkmlebis mier moixmareba. am
energiis udidesi nawili ATP-is warmoqmnazea gansazRvruli, da amitom
Tirkmlis ujredebSi, gansakuTrebiT ki mis qerqovan nawilSi, nivTierebaTa
cvlis aerobuli tipi mkafiod aris gamoxatuli. sinTezirebuli ATP-i,
+
ZiriTadad, Na -is, glukozis, aminomJavebis da sxva naerTTa aqtiur
transportze moixmareba.
nivTierebaTa cvlis darRvevebi TirkmelebSi SesaZlebelia ganpirobebuli
iyos infeqciuri, anTebiTi, iSemiuri da sxva mizezebiT. maT SesaZlebelia
gamoiwvion jorjalis sisxliT momaragebis blokada, filtraciisa da
reabsorbciis darRveva, Sarddenis blokada, sekreciis darRveva da a.S.

171
 Sardi. adamiani normaSi dRe-Ramis ganmavlobaSi 1-2 l Sards gamoyofs.
Sardis zrda (poliuria) paTologiur SemTxvevebSi SesaZlebelia
ganpirobebuli iyos qronikuli nefritebis, pielonefritebis, diabetis da
sxva daavadebaTa dros, xolo misi simcire - oligouria, cieb-cxelebis,
toqsikozebis, mwvave nefritebis dros. mZime mowamvlis, nervuli daZabulobis,
sadinarSi kenWebis arsebobisas SesaZlebelia Sardis gamoyofa mTlianad
Sewydes (anurea). xangrZlivi anurea iwvevs uremias.
Sardis organul nivTierebebs, normaSi, azotovani cvlis saboloo
produqtebi - Sardovana da kreatinini warmoadgenen. garda amisa, SardSi
isazRvrebian aminomJavebi, kateqolaminebi da maTi gardaqmnis produqtebi,
limonmJava ciklis metabolitebi da sxva. araorganuli nivTierebebidan
ZiriTadad warmodgenilia qloridebi da natriumi; SedarebiT mcirea
kaliumis, magniumis da kalciumis ionebi; didi raodenobiT aris amoniaki.
Sardis paTologiur komponentebs Seadgenen cila (proteinuria), hemoglobini
(hemoglobinuria), glukoza (glukozuria), ketonuri sxeulebi (ketonuria),
bilirubini (bilirubinuria). yovel maTgans sadiagnostiko mniSvneloba aqvs.

nervuli qsovili

nervuli qsovili, ZiriTadad, ori tipis ujredebs Seicavs: neironebs da

glias. neironebi Sedgebian sxeulisagan (soma), mcire zomis morCebisagan
(dendritebi) da erTi wagrZelebuli morCisagan (aqsoni), romelic boloSi
itoteba nervuli daboloebebiT. aqsoni, xSirad, garSemortymulia lipiduri
warmonaqmniT - mieliniT. es ukanaskneli gliuri ujredebidan -
oligodendrocitebidan warmoiqmneba. mielinizirebul aqsonSi moqmedebis
potenciali gacilebiT ufro swrafad vrceldeba.
neironebis mTavar funqcias infirmaciis miReba, gadamuSaveba da gadacema
warmoadgens. yoveli neironis aqsoni bolovdeba specialuri struqturiT,
romelsac sinafsuri daboloeba ewodeba. sinafsur daboloebebSi aqsoni
farTovdeba. neironis es nawili mdidaria mitoqondriebiT, neirogadamcemebis
Semcveli sekretoruli vezikulebiT da sxva specializirebuli
struqturebiT. sinafsuri daboloeba mezobel neironTan warmoqmnis e.w.
sinafsur kavSirs. sinafsuri kavSiri myardeba ori urTierTorientirebuli
neironis plazmur membranas Soris, romlebic daSorebuli arian
erTmaneTisagan 100-200 anqstremis manZiliT. am ukanasknels sinafsuri naprali
ewodeba. sinafsur napralSi Tavisuflad moZraobs ujredSorisi siTxe,

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romelsac gadaqvs energetikuli da biologiurad aqtiuri nivTierebebi.
aqsonis sinafsuri daboloebis mxares mdebare plazmur membranas
presinafsuri (sinafsamde), xolo mezobeli neironis plazmur membranas -
postsinafsuri (sinafsis Semdegi) ewodebaT.
pre- da postsinafsuri membranebi, maTi momsaxure bioqimiuri aparati da
sinafsuri naprali nervuli qsovilis mTavar funqciur erTeuls - sinafss
warmoqmnian. neironTa Soris urTierTkavSiri sinafsuri gadacemis
saSualebiT warmoebs. moqmedebis potenciali, warmoiqmneba ra neironis
somaSi Tu dendritze, vrceldeba aqsonze da aRwevs nervul daboloebas. aq
igi egzocitoziT, iwvevs vezikulebidan neirogadamcemebis gamoTavisuflebas,
romelic sinafsur napralSi difuziiT gadadis mezobeli ujredis
postsinafsur membranamde da moqmedebd mis receptorul ubanze. Tu es
ukanaskneli ionotropul receptorTa tips ganekuTvneba, aRmocenebuli
moqmedebis potenciali vrceldeba am neironzec.
moqmedebis potencialis gavrceleba ganpirobebulia neironis plazmuri
membranis transmembranuli potencialis cvlilebiT, romelic warmoiqmneba
ujredSi Na+-is ionTa SesvliT Na-seleqciuri arxebiT. am ionTa Sesvla
ujredSi membranuli potencialis vardnas iwvevs. potencialTa sxvaobis
Semcireba impulsis warmoqmnas ganapirobebs, romelic analogiuri meqanizmebiT
vrceldeba mTel ujredul membranaze. potencialis aRdgena K-is ionTa
saSualebiT xdeba, romlebic ujredidan gareT gamodian. K+-is ujredSida
koncentracia gacilebiT aRemateba am ionis ujreTgare koncentracias, da
amitom maTi gamosvla, koncentraciuli gradientis gamo, Tavisufali
difuziiT xdeba. am procesSi monawileobas iReben potencial-mgrZnobiare K+-
seleqciuri arxebi.
kaliumiT potencialis aRdgenis Semdeg, difundirebul ionTa gadaqaCva
warmoebs: natriumi gaitaneba ujredidan, xolo kaliumi Semoitaneba SigniT.
radgan aseTi gadatana ukve am ionTa koncentraciuli gradientis winaaRmdeg
warmoebs, masze ixarjeba ATP-is energia. am process fermenti Na,K-ATPaza
asrulebs. igi, ionTa gadatanas membranaze ATP-is bolokidura
fosfodieTeruli bmis hidrolizis Sedegad gamoTavisuflebuli energiis
xarjze awarmoebs. neironebSi arsebuli ATP-is 70%-mde gaixarjeba Na,K-
ATPazas muSaobaze. ATPazuri reaqciis Sedegad warmoiqmneba ADP da
araorganuli fosfati - Pi

ATP ADP + Pi

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 membranis biofizikuri TvisebebisTvis ganmsazRvreli mniSvneloba aqvs e.w.
K+-is dens, anu K+-is im raodenobas, romelic ujredidan gareT gamodis. im
SemTxvevaSi, Tu ujredidan kaliumis meti raodenoba gamovida, vidre Na+
Sevida ujredSi, damyarebuli potencialTa sxvaoba, sawyisTan SedarebiT,
ufro didi iqneba anu membrana iqneba hiperpolarizebuli. am SemTxvevaSi,
membranis Semdgomi depolarizaciisTvis da impulsis aRmosaceneblad ufro
Zlieri gaRizianeba iqneba saWiro. da piriqiT, rac ufro naklebi KK+
difundirdeba ujredidan, miT ufro gaadvildeba am neironis Semdgomi
agzneba.
nervul daboloebebSi Na-is ionTa garda, ujredSi, koncentraciuli
gradientis gamo, Sedian agreTve kalciumis ionebic. swored es ukanaskneli,
Ca-kalmodulin-damokidebuli- da Ca-fosfolipid-damokidebuli
proteinkinazebis saSualebiT awarmoeben vezikulebis transports
presinafsur membranasTan da neirogadamcemTa sekrecias sinafsur napralSi
postsinafsur membranaze. neirogadamcemebi ukavSirdebian Sesabamis
receptorebs da iwveven maT gaaqtivebas. rogorc ukve iTqva, ganasxvaveben
swrafi da neli tipis receptorebs (ix. ujreduli aqtivobis regulacia).
swrafi tipis receptorebi ionotropuli bunebis arian e.i. ukavSirdebian
ionur arxs, xolo neli tipis receptorebi ki - metabotropul funqciebTan
arian dakavSirebuli da kompleqsirdebian meorad Suamavlebis warmomqmnel
fermentebTan.
centralur da periferiul nervul sistemebSi warmodgenil ZiriTad
postsinafsur receptorebs warmoadgenen: glutamaterguli (glutaminis mJava),
gaem-erguli (gaem-gamaaminoerbos mJava), adrenerguli (noradrenalini da
adrenalini), dofaminerguli (dofamini), qolinerguli (acetilqolini),
opiaturi (enkefalinebi, endorfini). garda amisa, nervul sistemaSi
identificitdebian sxvadasxva tipis peptiderguli neironebi (somatostatinis,
qolecistokininis, substancia P-s, vazopresinis, oqsitocinis,
kortikotropinis da sxva peptidebis).
postsinafsuri neironis citoplazma mdidaria mikromilakovani da
mikrofilamenturi warmoSobis citoConCxis ZafebiT, romelsac postsinafsuri
simkvrive ewodeba. postsinafsuri filamentebi mWidrod arian dakavSirebuli
postsinafsur membranasTan da ganapirobeben mis sivrciT orientacias,
konfiguracias. garda amisa, maTi saSualebiT warmoebs membranuli

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makromolekulebis gadaadgileba, koncentrireba, internalizacia
citoplazmaSi, da degradacia.
metabotropul receptorTa aqtivaciis Sedegad warmoqmnili meoradi
Suamavali, gadis ra postsinafsur simkvrives, moqmedebs neironis
organoidebze, maT metabolizmze. meore mxriv, meoradi Suamavlebi, TviT
postsinafsuri milakebis konfiguraciasac cvlian, rasac didi mniSvneloba
eniWeba sinafsebis funcionirebaSi.
postsinafsuri simkvrivis CamoyalibebaSi da misi moqmedebis realizaciaSi
did rols, cila-tubulinisagan Semdgari mikromilakebi da aqtinisagan
Semdgari mikrofilamentebi TamaSoben. tubulini polimerizirebad cilas
warmoadgens, romelsac rTuli konfiguraciis Zafebis formireba SeuZlia.
tubulini polimerizacias da depolimerizacias aregulireben specifiuri
cilebi, romlebsac mikromilakebTan asociirebuli cilebi (mac) ewodebaT.
mravali maTgani neirospecifiur cilas warmoadgens e.i. maTi geni
eqspresirebulia da transkribirdeba mxolod nervul qsovilSi. wamyvans am
cilebs Soris warmoadgenen maRalmolekuluri (mol.wona 150000-200000)cilebi
mac-1 da mac-2.
mac-1 da mac-2 ganicdian fosforilirebas sxvadasxva tipis
proteinkinazebiT, romlebic reagireben TiTqmis yvela tipis meoreul
Suamavalze (cAMP, Ca++, fosfolipidebi). am cilebis fosforilireba scvlis
maT konformacias, da Sesabamisad, maT zemoqmedebas tubulinis
polimerizaciaze. Sedegad icvleba tubulinis milakebis forma da masTan
dakavSirebuli struqturebis (magaliTad, plazmuri membranis) konfiguracia.
proteinkinazebis garda, mac-ebis regulatorul unarze moqmedebs
proteolizuri fermenti, romelic cilis molekulidan garkveuli sigrZis
fragmentis CamoWras awarmoebs. fragmentis CamoWra ucvlis mac-s
konformacias, da Sesabamisad, tubulinis polimerizaciaze moqmedebis
efeqturobas. aseTi tipis proteolizuri fermenti, romelsac nervul
++
qsovilSi kalpaini ewodeba, Ca - iT aqtivirdeba. amdenad, es procesic,
meoradi Suamavlebis kontrolis qveS imyofeba.
aRwevs ra informacia meoradi Suamavlebis saSualebiT birTvSi, igi
cilebis sinTezis gaaqtivebas iwvevs. xSir SemTxvevebSi, jer aqtivirdebian
proto-onkogenebi Ras, Jun da Fos, da Semdeg, maT mier sinTezirebuli
transkripciuli faqtorebiT inducirdeba genTa eqspresia. nervul qsovilSi
aTasobiT qsovilspecifiuri genebi muSaoben.

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 mexsierebis bioqimiis zogierTi sakiTxi. mexsiereba rTul, ramdenime
etapisagan Semdgar, dinamiur process warmoadgens. Tavis tvinSi informaciis
miReba, Cawera da reproduqcia warmoebs ara erT romelime makromolekulaSi,
an neironSi, aramed nervul ujredTa erTobliobaSi. wamyvani mniSvneloba am
procesebSi sinafsur kavSirebs eniWebaT.
ganasxvaveben xanmokle da xangrZliv mexsierebas. mexsierebis es ori stadia
erTi procesis - mexsierebis Caweris, konsolidaciis Semadgenel nawils
warmoadgens. dReisaTvis gamokvleulia mexsierebis procesSi monawile
bioqimiur procesTa mxolod umniSvnelo nawili. fiqroben, rom mexsierebis
procesi warimarTeba saganTa xatismieri aRqmiT, anu iseTi procesiT, romlis
drosac saganze Tu movlenaze informaciis Cawera nawilebad xdeba. aseTi
nawilobrivi aRqmis Semdeg, isini tvinSi asocirdebian, erTiandebian erT
xatad (movlenad, sagnad). xatis warmosaxva aseve elementebis saSualebiT
warmoebs da misma erTma nawilma SesaZlebelia mTeli xati gagvaxsenos.
warmosaxvis aseTi forma metyvelebs imaze, rom mexsiereba asociaciuria e.i.
igi Cawerilia sxvadasxva elementis saxiT da Cawerili kvalis aRdgena, misi
warmosaxva am elementebis gaerTianebiT, maTi asociaciiT xdeba.
mexsierebis CaweraSi gansakuTrebuli mniSvneloba dendritebs eniWebaT.
neironebSi arsebuli dendritebi Zlier datotvil struqturas warmoqmnian,
romelzec SesaZlebelia 100000-200000-mde sinafsi mTavrdebodes tvinis
sxvadasxva nawilebidan. yoveli konkretuli sensoruli gaRizianeba mxolod
garkveuli sinafsebis amoqmedebas iwvevs, da amitom, erT neironSi Cawerili
informacia SesaZlebelia Zalzed didi (praqtikulad usasrulo) iyos.
dReisaTvis aRmoCenilia erTaderTi fiziologiuri fenomeni, romelic
savaraudebelia safuZvlad edos mexsierebis Caweris garkveul etaps. am
fenomens posttetanuri potenciacia ewodeba da mdgomareobs imaSi, rom
zogierTi neironis zRurblovani, ganmeorebadi gaRizianeba iwvevs sinafsuri
gadacemis efeqturobis gaZlierebas. aseTi gaZliereba rCeba neironis
gaRizianebis Semdegac da igi grZeldeba saaTebisa da dReebis ganmavlobaSi;
neirons "axsovs" Zlieri gaRizianeba.
am fenomenis formirebaSi glutaminis mJavis receptorebi iReben
monawileobas. glutaminis mJava centraluri nervuli sistemis ZiriTad
amgznebel neirogadamcems warmoadgens da mas receptorTa ramdenime subtipi
gaaCnia. posttetanur potenciaciaSi monawile glutamatis receptori
warmodgenilia ori subtipiT: NMDA da AMPA subtipebiT, romlebic
erTmaneTisagan gansxvavdebian, rogorc farmakologiur nivTierebebTan

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urTierTobiT, ise ujredze moqmedebis meqanizmebiT. orive tipis receptori
SeuRlebulia ionur arxTan, romlebic erTmaneTisgan gansxvavdebian
seleqtiurobiT gasatarebeli ionis mimarT: NMDA-tipis arxi Tavisuflad
atarebs Ca-s da sustad - Na-s, xolo AMPA-tipisa ki - piriqiT, SedarebiT
ukeTesad atarebs Na-s. NMDA-tipis arxi mosvenebis mdgomareobaSi da
normaluri transmembranuli potencialis (-60 milivolti) arsebobisas
blokirebulia Mg-is ionebiT, xolo AMPA-tipisa - ara. Mg++ scildeba NMDA-
arxs mxolod membranis depolarizaciis Semdeg.

postsinafsuri
membrana

NMDA-receptori

dabali sixSiris gaRizianebis dros, rodesac sinafsur napralSi

gamoiyofa glutaminis mJavis mcire raodenoba, aqtivirdeba mxolod AMPA-
receptori, radgan neirogadamcemis koncentracia ar aris sakmarisi, rom
moxdes NMDA-s gaaqtiveba. amitom, ujredSi Sedis mxolod Na+ da impulsis
gavrceleba erTi mimarTulebiT xdeba. maRali sixSiriT gaRizianebis
SemTxvevaSi, AMPA-s mier gamowveuli depolarizaciis talRa, gacilebiT
Zlieria da vrceldeba NMDA-receptoramde, romelic mis maxloblad
mdebareobs. membranis depolarizacia iwvevs Mg-is ionebis mocilebas NMDA-s
arxidan, ris Sedegad postsinafsur membranaze aqtivirdeba agreTve es
receptoric. am ukanasknelis gaaqtiveba iwvevs arxis gaxsnas, saidanac neironSi
Sedis didi raodenobiT Ca-is ionebi. Ca++-is koncentraciis zrda postsinafsur
neironSi iwvevs bioqimiur gardaqmnaTa jaWvs, romlebSic ZiriTad rols ori
Ca-damokidebuli proteinkinazebi TamaSoben.
Ca++-is koncentraciis zrda postsinafsur simkvriveSi da misi dakavSireba
proteinkinazebTan iwvevs am fermentebis gadaadgilebas (translokacias)
citoplazmidan plazmur membranisaken. membranasTan dakavSirebis Semdeg,
magaliTad proteinkinaza C iwyebs K-is arxis erT-erTi cilis
fosforilirebas, ris gamoc K+-is gasvla neironidan iTrguneba. Sedegad,

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membranis am ubanSi mcirdeba membranuli potenciali da igi ufro
mgrZnobiare xdeba Semdgomi gaRizianebisa da impulsis formirebisaTvis.
meore mxriv, kalciumis koncentraciis mateba, iwvevs agreTve Ca++-
kalmodulin-damokidebuli proteinkinazis gaaqtivebas, romelic
mikromilakebis cilebis, da kerZod mac-2-is fosforilirebas iwyebs. mac-is
fosforilirebiT icvleba mikromilakebis polimerizaciis unari da masTan
dakavSirebuli dendritTa plazmuri membranis konfiguracia.
amrigad, membranis biofizikuri maxasiaTeblebis, misi morfologiuri
organizaciis Secvla, ganapirobebs im rTul funqciur cvlilebebs, romlebic
informaciis Caweras emsaxurebian. miuxedavad amisa, NMDA-receptoris
xangrZlivi gaxsna paTologiuri procesebis formirebas ganapirobebs.
sxvadsxva tipis nevrologiuri darRvevebis, tvinis iSemiuri paTologiebis
dros NMDA-receptori hiperglutamaterguli zegavlenis gamo, xangrZlivi
droiT ixsneba, rac Warbi kalciumis Sesvlas da neironebis degeneracias
iwvevs. Tu ra didi mniSvneloba aqvs aRweril sistemas ara marto mexsierebis
formirebaSi, aramed nevrologiuri darRvevebis etiopaTogenezSic,
metyvelebs is faqti, rom Tanamedrove antiepilepsiuri, antiparkinsonuli da
antiiSemiuri preparatebi NMDA-tipis glutamatis receptoris damTrgunvel
naerTebs warmoadgenen. aucilebelia aRiniSnos, rom zogierT maTgans
halucinagenuri efeqtic gaaCnia, rac miuTiTebs am sistemis urTierTkavSirze
tvinis sxva umaRles funqciebTanac.
aRwerili magaliTebidan gamomdinareobs agreTve erTi arsebiTi daskvna,
romelic adamianis yvela tipis ujredis regulaciur sistemebs Seexeba:
kalciumis ujredSida koncentraciis etapobrivi, Seqcevadi cvlilebebi
aucilebelia ujredis normaluri metabolizmisaTvis, misi zrdisa da
diferencirebisaTvis, magram am ionebis Warbi, arakontrolirebadi mateba
ujredis gadagvarebas da paTologias iwvevs.

kunTovani qsovili

kunTovani qsovili sxeulis 40-42% Seadgens. mas dinamiuri funqcia aqvs da

igi, SekumSvisa da modunebis saSualebiT, ganapirobebs organizmisa da misi
elementebis Zvradobas. SekumSvis dros kunTis ujredis qimiuri energia
meqanikurSi gadadis.

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 ganasxvaveben kunTovani qsovilis sam tips: ConCxis, gulis da gluv
kunTebs. garda amisa kunTovani qsovili SeiZleba daiyos ganivzolian da
gluv kunTebad.
ganivzoliani kunTis morfologiuri organizacia. ganivzoliani kunTi
Sedgeba wagrZelebuli, boWkos formis ujredebisagan, romlebsac giganturi
zoma, maRalspecializirebuli elementebi da ramdenime birTvi moepoveba. igi
Semofarglulia plazmuri membraniT-sarkolemiT, romelzec nervuli
boWkoebi mTavrdebian. nerv-kunTovani gadacema sinafsuris msgavsia da
xorcieldeba nervul daboloebebSi arsebuli mediatorebis saSualebiT.
sarkolemis postsinafsuri ubani Seicavs receptorebs, romelTa agznebis
Semdeg xdeba impulsis gadacema mTel ujredze.
kunTis ujredis citoplazmaSi - sarkoplazmaSi ganlagebulia Zafisebri
warmonaqmnebi, romlebsac miofibrilebi ewodebaT. ganivzoliani kunTebis
miofibrilebi Sedgebian ganmeorebadi elementebiT-sarkomerebiT. sarkomerebi
Seicaven msxvil Zafebs, Semdgars maRalmolekuluri cila-miozinisagan, da
wvril Zafebs, romlebic cila-aqtinisagan Sedgeba. SekumSvis procesSi
wvrili da msxvili Zafebi (filamentebi) iwyeben urTierTsawinaaRmdego
mimarTulebiT moZraobas, riTac kunTis ujredis konformaciul cvlilebas
(SekumSvas) iwveven.
kunTis cilebi da aracilovani azotovani naerTebi, energetikuli cvla.
kunTis ujredSi didi raodenobiT arian warmodgenilni SekumSvadi cilebi.
maT miofibrilaruli cilebi ewodebaT. miozini- Seadgens miofibrilebis
masis 50-55%. mas wagrZelebuli forma aqvs da Sedgeba ramdenime
suberTeulisagan: ori "mZime" polipeptiduri jaWvisagan, romelTa
molekuluri wonaa 210000 da mokle "msubuqi" jaWvebisagan - molekuluri
woniT 20000. "mZime" jaWvebi warmoqmnian grZel daxveul -spirals, romelic
boloSi "mokle" jaWvebs uerTdeba. miozinis molekulis am adgilze
formirdeba globula, romlis saSualebiTac SekumSvis procesSi aqtinis
molekulasTan urTierTmoqmedebs. miozinis molekulas fermentuli aqtioba
(ATPazuri) aqvs, rac imis maniSnebelia, rom ATP-s hidrolizis Sedegad
gamoyofili energia moixmareba miozinis konformaciul cvlilebebze da
aqtinTan urTierTqmedebaze e.i.meqanikur muSaobaze. aqtini-miofibrilebis masis
20% Seadgens. igi warmodgenilia ori formiT: globularuli (G-aqtini) da
fibrilaruli (F-aqtini). F-aqtini G-aqtinis polimerizirebul formas
warmoadgens. miozinTan urTiermoqmedebs mxolod F-aqtini. troponini da
tropomiozini - miofibrilebis regulatorul cilebs warmoadgenen.

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troponini ikavSirebs kalciums da am ionis dakavSirebis Semdeg cvlis
wvrili filamentebis struqturas.
aracilovani azotovani naerTebi - kunTis ujredebSi didi raodenobiT
moipoveba ATP da misi daSlis produqtebi ADP da AMP. garda am
nukleotidebisa, kunTi Seicavs kreatins da kreatinfosfats. es ukanasknelni,
kunTis aracilovani azoturi naerTebis 60% Seadgenen. kreatini RviZlSi
sinTezirdeba da sisxlis saSualebiT gadaitaneba kunTebSi. aq igi
fosforilirdeba da warmoqmnis kreatinfosfats. kreatinfosfati
maRalenergetikul naerTs warmoadgens da makroerguli fosfatebis samarago
fonds qmnis. kunTis intensiuri muSaobisas ATP-is resinTezi ZiriTadad am
fondis xarjze xdeba:
kreatinkinaza.

kreatinfosfati + ADP kreatini + ATP

mosvenebis mdgomareobaSi, kunTis energetikul substratebs, ZiriTadad,

RviZlSi sinTezirebuli cximovani mJavebi da ketonuri sxeulebi Seadgenen;
zomieri datvirTvisas sisxlis glukozac iSleba, xolo Zlieri muSaobis
dros, ATP-s gamoyeneba iseTi intensivobiT mimdinareobs, rom sisxli veRar
aswrebs energetikuli masalis miwodebas. amitom, kunTis ujredebi iwyeben
sakuTari glikogenis daSlas, romelic am dros, Jangbadis ukmarisobis gamo,
anaerobuli gziT katabolizirdeba. kunTebSi warmoqmnili rZis mJava sisxlis
saSualebiT RviZlSi gadaitaneba da glukoneogenezis saSualebiT misgan jer
glukoza, xolo Semdeg glikogeni warmoiqmneba. kritikul situaciebSi, ATP-s
warmoqmna kreatinfosfatidan warmoebs.
kunTis SekumSvis meqanizmi. ConCxis kunTebis SekumSva-moduneba
citoplazmuri Ca++-is koncentraciiT regulirdeba. mosvenebis mdgomareobaSi
ujredSi Ca-is koncentracia dabalia (10-7-10-8 moli/l). kunTis boWkos
nervuli gaRizianebisas sarkoplazmuri retikulumidan (e.w.T-sistemidan)
Tavisufldeba Ca-is ionTa didi raodenoba da misi koncentracia
citoplazmaSi 10-6-10-5 moli/l-mde izrdeba. gamoTavisuflebuli Ca++ cila-
troponins (troponini kalmodulinis homologiur cilas warmoadgens)
ukavSirdeba, romelic, konformaciuli cvlilebis Semdeg, moqmedebs miozinis
globularul nawilze. am ukanasknels, troponinTan dakavSirebis Semdeg Ca-
ATP-azuri aqtivoba ezrdeba da igi iwyebs aqtinis mimarT gadanacvlebas anu
SekumSvis process. kunTis moduneba iwyeba nervuli gaRizianebis Sewyvetisas,
da Sesabamisad, ujredSi Ca++-is ionebis daklebis Semdeg. kalciumis

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koncentraciis Semcireba citoplazmaSi ganpirobebulia maTi
transportirebiT T-sistemebSi, romlebsac Ca-is saqaCavi anu transportuli
Ca++-ATPaza anxorcielebs.
amrigad, sistema aqtini-miozini-troponini unikalur qimiur Zravas
warmoadgens, romelic ATP-s qimiur energias pirdapir meqanikurad gardaqmnis
mudmivi wnevisa da temperaturis pirobebSi.
gulis kunTis iSemia. gulis kunTis iSemiuri mdgomareoba gamowveulia masSi
mimdinare energetikuli procesebis daqveiTebiT. miokardiumis ujredTa
dazianeba safexurebrivi procesia, romlis drosac adgili aqvs rogorc
struqturul, aseve funqciur cvlilebebs. ganasxvaveben iSemiis adreul
Seqcevad, gardamaval da Seuqcevad cvlilebebs.
koronaruli sisxlis mimoqcevisa da mikrocirkulaciis daqveiTebis dros
mcirdeba gulis kunTis momarageba JangbadiT da energetikuli masaliT.
Jangbadis ukmarisoba mitoqondriebSi sunTqviTi jaWvis, fermentebis
Senelebasa da ATP-is sinTezis Semcirebas iwvevs. iSemiis sawyis, Seqcevad
stadiaze energetikuli deficiti anaerobuli procesebis, kerZod
glikolizis gaaqtivebiT kompensirdeba. am procesis xangrZlivoba
damokidebulia miokardiumSi glikogenis maragze, radgan sisxlmomaragebis
Semcirebis gamo gulis kunTs endogenuri glukoza dabali intensiobiT
miewodeba. garda amisa, anaerobuli reaqciebi mTlianad ver uzrunvelyofen
miokardiums energetikuli masaliT, kerZod ATP-iT. amitom, gulis kunTSi
iwyeba Seuqcevadi procesebis ganviTareba.
gulis, iseve rogorc ConCxis kunTebSi, ATP-is fondi, ZiriTadad or
procesze ixarjeba: miozinis SekumSvasa da meqanikur muSaobaze, agreTve
modunebis dros Ca++-is gandevnaze citoplazmidan, mis aqtiur transportze
sarkoplazmur retikulumSi (Ca-saqaCavi anu Ca-ATPaza). amitom, ATP-s deficiti,
upirveles yovlisa, am procesebis intensivobaze moqmedebs.
Ca++-is ujredSi dagroveba, misi koncentraciis zrda rigi arasasurveli
fermentuli reaqciebis stimulirebas iwvevs, romelTa Soris wamyvani
mniSvneloba fosfolipazebs eniWebaT. fosfolipazebis umravlesoba
kalciumis maRali koncentraciiT aqtivdeba da Sedegad membanebis
fosfolipidebis daSlas iwyebs. amitom, mravali organoidis, da maT Soris
mitoqondriebis membranuli struqtura irRveva, rac Seuqcevadi procesebis
ganviTarebas aRrmavebs. garda amisa, fosfolipazebis mier
gamoTavisuflebuli lizolipidebi da cximovani mJavebi, ganicdian ra
peroqsidacias (zeJangur daJangvas), damatebiT azianeben kardiomiocitebis

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ZiriTad struqturebs (zeJanguri procesebis aqtivacia gamowveulia Jangbadis
deficitiT, romelic mitoqondriebSi wyalbadis arasrul aRdgenas iwvevs).
Ca-is ionebiT gaaqtivebuli fosfolipaza A2, damatebiT aTavisuflebs
araqidonis mJavas, romelic moqmedebs K+-is arxebze, iwvevs maT gaxsnas da
gulis kunTis ujredebidan am ionTa gandevnas. amis Sedegad, icvleba
sarkolemis membranuli potenciali da misi funqcia qveiTdeba. am Rrma
Seuqcevadi procesebis fonze viTardeba gulis kunTis ukmarisoba. amrigad,
centraluri roli, gulis kunTis iSemiuri paTologiebiis ganviTarebaSi,
kalciumis ionebs eniWebaT. isini aRrmaveben daavadebas da iwveven
kardiomiocitebSi Seuqcevadi procesebis warmarTvas.
Ca++-is ionTa mniSvnelobaze gulis kunTis metabolizmSi metyvelebs
agreTve e.w. "kalciumi paradoqsi". es fenomeni mdgomareobs SemdegSi: Tu
anoqsiuri gulis perfuzia, romelic warmoebs ukalciumo fiziologiuri
xsnariT Seicvala normaluriT (kalciumiani), adgili aqvs gulis kunTis
ujredebis swraf degeneracias. anoqsiur gulSi kalciumis ionTa Sesvla misi
ujredebis sikvdils iwvevs. amitom iTvleba, rom Ca++-is didi raodenobiT
dagroveba ujredSi, mis saboloo metabolur gzas warmoadgens. aqve unda
iTqvas, rom kalciumis arakontrolirebadi zrda, aramarto gulis kunTis
ujredebze moqmedebs. neironebSi, es ionebi iwveven epilepsiur ganmuxtvebs,
tvinis iSemiis Sedegad neirodegeneraciuli procesebis ganviTarebas, mravali
topis qsovilTa avTvisebian gadagvarebas, anTebiTi procesebis gaRrmavebas da
a.S. amitom, farmakologiuri preparatebi, romelTa moqmedeba ganpirobebulia
Ca++-is arxebis blokirebiT - dihidropiridinebi da fenilalkilaminebi, an
kalmodulinis inhibirebiT - ftorfenazinebi, gamoiyenebian ara marto kardio-
da neirodegeneraciuli paTologiebis sawinaaRmdegod, aramed farTod
moixmarebian klinikur praqtikaSi.

SemaerTebeli qsovili

SemaerTebeli qsovili farTodaa gavrcelebuli adamianis organizmSi da

misi mTliani masis 50%-s Seadgens. miuxedavad maTi morfologiuri
gansxvavebisa (myesebi, Zvali, kani, da a.S.), isini erTi saerTo principiT arian
Camoyalibebuli: sxva qsovilebTan SedarebiT, SemaerTebeli qsovili faSarad
ganlagebuli ujredebisagan Sedgeba, romlebic erTmaneTTan fibrilaruli
struqturebiT - kolagenuri, elastikuri da retikuluri boWkoebiT arian

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dakavSirebuli. SemaerTebeli qsovili mdidaria ujredSorisi nivTierebiT. mas
rTuli qimiuri aRnagoba aqvs da mZlavri organuli matriqsi axasiaTebs.
kolageni. SemaerTebeli qsovilis erT-erT damaxasiaTebel elements
ujredSorisi kolagenuri boWkoebi warmoadgenen. kolagenuri boWkoebi,
adamianis organizmSi sakmaod gavrcelebul (mTliani masis -6%) fibrilarul
cila-kolagenisagan Sedgebian. am cilisTvis, ZiriTadad, sami tipis
aminomJavaa damaxasiaTebeli: glicini, prolini da oqsiprolini (oqsiprolini
mxolod cilovani molekulis aSenebis Semdeg warmoiqmneba). kolagenis
TiToeuli polipeptiduri jaWvi 1000-mde aminamJavis naSTisagan Sedgeba, igi
tipiur fibrilarul cilas warmoadgens.
elastini. elastini elastikuri boWkoebis ZiriTadi komponentia. masSi,
kolagenis msgavsad, prolinisa da glicinis Warbi raodenoba aRiniSneba. am
aminomJavebis garda, elastinis molekula didi raodenobiT valinsa da
alanins Seicavs. elastinisTvis damaxasiaTebelia unikaluri qimiuri
struqturis - desmozinis arseboba. igi 4 lizinisa da pirimi- dinis
birTvisagan Sedgeba:

R R

+
N

R (R-lizinis naSTebi)

desmozini

desmozini lizinis radikalebis saSualebiT 4 polipeptidur jaWvs

erTmaneTTan akavSirebs. amis gamo, elastinis molekula, sxva fibrilaruli
cilebisagan gansxvavebiT ori mimarTulebiT iWimeba.

Proteoglikanebi. SemaerTebeli qsovilis ujredSorisi matriqsi

maRalorganizebuli struqturiT - proteoglikaniT aris warmodgenili.
proteoglikanis ZiriTad rgols polisaqariduli glikozaminoglikanuri
jaWvi warmoadgens (umetesoba SemTxvevaSi jaWvi hialuronis mJavis
monomerebisagan Sedgeba). glikozaminoglikanuri jaWvis mTavar RerZs

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mravalricxvovani ganStoebebi aqvs cilebisa da SedarebiT mokle
polisaqariduli jaWvebis saxiT. glikozaminoglikanuri RerZi mokle
polisaqaridul ganStoebebs cilis (damakavSirebeli cila) saSualebiT
ukavSirdeba. Tavis mxriv, damakavSirebel cilas, e.w. "centraluri cila"
uerTdeba, romelzec mokle polisaqaridebia mimagrebuli:
proteinglikanebSi cilis wili SedarebiT mcirea da igi tolia mTliani
masis mxolod 1-2%-isa. proteoglikani labisebr matriqss warmoqmnis,
romelzec SemaerTebeli qsovilis ujredebia ganlagebuli.
glikozaminoglikanebi. rogorc ukve iTqva, SemaerTebeli qsovilis
glikozaminoglikanebi sworxazovan, disaqariduli erTeulebisagan Semdgar
polimers warmoadgenen. maT SemadgenlobaSi monomerebis saxiT glukozamini
an galaqtozamini Sedis. erT-erTi gavrcelebuli glikozaminoglikania -
hialuronis mJava, romlis ganmeorebadi disaqariduli erTeuli glukuronis
mJavisa da N-acetilglukozaminisagan Sedgeba.
glukozaminoglikanebis warmomadgenli arian agreTve
qondroitinsulfatebi. maT SemadgenlobaSi disaqaridul erTeulad Sedian
glukuronis mJava da sulfatirebuli (me-4 an me-6 mdgomareobaSi)
acetilgalaqtozamini. kanis qsovilisTvis damaxasiaTebelia
dermatansulfati, romelic iduronis mJavisa da sulfatirebuli
acetilgalaqtozaminis polimers warmoadgens. es glukozaminoglikani
mdgradia hialuronis mJavis damSleli fermentis - hialuronidazis mimarT.
Tvalis rqovanaSi, xrtilSi da sxva qsovilebSi warmodgenilia
glikozaminoglikani - keratansulfati. adamianis organizmSi gavrcelebulia
agreTve SemaerTebelqsovilovani warmoSobis (poxieri ujredebi)
glikozaminoglikani - heparini. mas antikoa- gulanturi moqmedeba aqvs.
proteoglikanebis funqcia damokidebulia maT SenebaSi Semaval
glikozaminoglikanze. is glikozaminoglikanebi, romlebic sulfatirebuli
arian naxSirwylis ne-4 am me-6 naxSirbadze, uaryofiTad damuxtul polimers
(polianionur polimers) warmoqmnian. amitom, maT mTavar funqcias
ujredSoris siTxeSi arsebuli kaTionebis (aseve wylis) dakavSireba da
gadanawileba warmoadgens. magaliTad, Zvlis formirebisas, osifikaciur
kerebSi koncentrirdebian qondroitinsulfatebi, romlebic Tavis mxriv, Ca++-
is intensiur dagrovebas ganapirobeben.
SemaerTebeli qsovilis metabolizmi hormonebiT da sxva biologiurad
aqtiuri nivTierebebiT regulirdeba. magaliTad, glukokortikoidi
kortizoli fibroblastebSi kolagenisa da glikozaminoglikanebis sinTezs

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amuxruWebs. mineralkortikoidi aldosteroni ki - fibroblastebis
proliferacias da biosinTezuri reaqciebis gaaqtivebas iwvevs. xandazmul
asakSi, glikozaminoglikanebis, da upirveles yovlisa hialuronis mJavis
raodenoba mcirdeba, xolo kolagenisa - izrdeba. garda amisa, kolagens
ecvleba fiziko-qimiuri Tvisebebi, igi naklebad elastiuri da
struqturulad stabiluri xdeba.
SemaerTebeli qsovilebis paTologiebs Soris gansakuTrebuli adgili
kolagenozebs ukaviaT. miuxedavad imisa, rom am paTologiebisTvis
damaxasiaTebelia simptomTa erTianoba, ganpirobebuli qsovilis boWkoebis,
ujredebis da ujredSorisi nivTierebebis daSliT, maTi etiologia da
paTogenezi SesaZlebelia gansxvavebuli iyos. kolagenozebSi aerTianeben
revmatizms, revmatoidul arTrits, sistemur wiTel mgluras, sistemur
sklerodermias, dermatomiozits da a.S. zogierTi maTgani SesaZlebelia
infeqciur-alergiuli da autoimunuri warmoSobisa iyos.

Zvlovani qsovili

Zvlovani qsovili SemaerTebeli qsovilis gansakuTrebuli tipia. mis

ZiriTad ujredebs, osteoblastebi, osteocitebi da osteoklastebi
warmoadgenen. osteoblastebSi, Zvlovani qsovilis ujredSorisi nivTierebis,
kolagenis, proteoglikanebisa da glikozaminoglikanebis biosinTezi
mimdinareobs. Zvlis formirebis procesSi osteoblastidan osteociti
warmoiqmneba; es ukanaskneli ZvalSia Catanebuli. mesame tipis Zvlovani
ujredebi - osteoklastebi, gigantur, mravalbirTvian ujreds qmnian, romlis
mTavar funqciasac Zvlis ganviTarebisa da zrdis procesSi gamokiruli
xrtilis rezorbcia warmoadgens.
kompaqturi Zvlis ujredSorisi organuli matriqsi Zvlis mxolod 20%-s
Seadgens. Zvlis masis udidesi nawili (70%) araorganul marilebs uWiravs.
Rrublovan ZvalSi organuli nivTierebebis wili metia (50%).
Zvlis araorganuli nawilis ZiriTadi Semadgeneli nawilia
hidroqsilapatiti - Ca10(PO4)6(OH)2, mis marilebs Cxirebisa da firfitebis
forma aqvT. mineraluri fazis meore Semadgenelia - SedarebiT amorfuli
kalciumis orTofosfati Ca3(PO4)2. orTofosfatis raodenoba mzard
organizmebSi Warbobs hidroqsilapatitisas, xolo zrdasrul asakSi - piriqiT,
hidroqsilapatitis wili Zvlis masaSi aRemateba kalciumis fosfatisas.
hidroqsilapatitis mier warmoqmnil kristalur meserSi gvxvdeba sxva erT-

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Tu orvalentiani ionebic. Zvlovani qsovilis mTliani ganaxleba yovel 10
weliwadSi erTxel warmoebs.
organuli matriqsis ZiriTadi komponentia kolageni. matriqsi Seicavs aseve
glikozaminoglikan - qondroitinsulfats. kolagens didi mniSvneloba
eniWeba osifikaciis procesebSi; misi boWkoebi kalcifikaciis (kalciumis
marilebis gamoleqvis) mimarTulebas gansazRvraven, radgan am cilebis
urTierTdakavSirebis adgilze warmoiqmneba fosfatebis gamoleqvis
pirveladi kerebi. aseT kerebSi, proteoglikanebi, hidrolizuri fermentebis
saSualebiT iSlebian da gamoidevnebian marilebis mier. maT adgils
hidroqsilapatitis meseri ikavebs.
Zvlovani qsovilis katabolizmic, sakmaod rTul procesebTan aris
dakavSirebuli. rogorc fiziologiuri, aseve paTologiuri procesebis dros
Zvlovani qsovilis rezorbcias win uswrebs misi organuli da araorganuli
struqturebis "gawova". osteolizis (osteolizi warmoadgens process,
romlis drosac mimdinareobs organuli matriqsis destruqciacia sxva
qsovilebiT Canacvlebis gareSe) Sedegad warmoqmnili organuli mJavebi
kalciumis marilebis disociacias iwveven, riTac xels uwyoben Zvlis
demineralizacias.
Zvlis mineralebi sisxlis plazmis kalciumTan da fosfatTan mWidro
qimiur wonasworobaSi imyofebian. kalciumis fosfatis marilebis Zvlovani
qsovilidan gamoTavisufleba, maTi ZvlebSi deponireba an gamoZeveba
regulirdeba parathormonisa (farisebraxlojirkvlis hormoni) da
kalcitoninis ( farisebri jirkvlis hormoni) saSualebiT. sisxlis plazmaSi
kalciumis koncentraciis daqveiTeba parathormonis sekrecias aZlierebs; es
ukanaskneli osteoklastebis metabolizms aaqtivebs da iwvevs Zvlis
rezorbcias. rezorbciis Sedegad mineralebi xsnad mdgomareobaSi gadadian
da plazmaSi Ca++-isa da fosfatis raodenoba matulobs. garda amisa,
parathormoni TirkmelebSi kalciumis ukuSTanTqmas aZlierebs pirveladi
Sardidan. sisxlis plazmis kalciumis koncentraciis zrda, piriqiT, hormon
kalcitoninis sekrecias aCqarebs. kalcitonini xels uwyofs am ionTa
mineralizacias, ris Sedegadac misi raodenoba sisxlSi mcirdeba.
kalciumis cvlis regulaciaSi didi mniSvneloba eniWeba vitamin D-s.
vitamini D genetikuri aparatis saSualebiT Ca-damakavSirebeli cilebis (maT
Soris kalmodulinisac) sinTezs zrdis. am cilebis mateba ganapirobebs
kalciumis ionebis intensiur Sewovas nawlavebSi, maT reabsorbcias Tirkmlis
milakebSi da Zvlebidan mobilizacias. amitom, vitamini D-s xangrZlivi miReba

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Zvlebis demineralizacias ganapirobebs Zvlebis normaluri ganviTarebisTvis
saWiroa agreTve vitaminebi A da C.
tramvuli daavadebebis garda gamohyofen Zvlis anTebiT (osteomieliti,
ostiti da sxva), distrofiul da displastikur paTologiebs.

kvebis bioqimia

bioqimiis erT-erT mTavar miRwevad kvebis Tanamedrove Sexedulebebis

formireba SeiZleba CaiTvalos. Tanamedrove adamianis racioni Semdeg ZiriTad
produqtebs unda Seicavdes: energiis wyaros (naxSirwylebi, cilebi, cximebi),
Seucvlel nivTierebebs (aminomJavebs, cximovan mJavebs, vitaminebs,
araorganul naerTebs) da wyals.
energetikuli nivTerebebi.. srulfasovan kvebis racionSi energetikul
nivTierebebad iTvlebian cilebi, cximebi da naxSirwylebi, romelTa daJangvis
procesSic energia gamoiyofa. sakvebi produqtebis am energias kaloriebSi an
sakveb kaloriebSi gamoxataven. dadgenilia, rom 1 g naxSirwylebi Seicaven 4,2
kkalorias, cximebi - 9,5 kkalorias, xolo cilebi - 4,3 kkalorias.
naxSirwylebi. naxSirwylebi TavisTavad, kvebis Seucvlel produqtebs ar
warmoadgenen, magram radgan isini SedarebiT xelmisawvdomi arian, farTod
moixmarebian adamianebis mier. naxSirwylebiT mdidaria mcenareuli sakvebi da
igi mosaxleobis zogierTi populaciis ZiriTad produqts warmoadgens.
ganviTarebul qveynebSi, sadac ZiriTad sakvebs rZisa da xorcis produqtebi
Seadgenen, naxSirwylebis wili mxolod 45% tolia.
cximebi. sakvebi lipidebis 98% neitralur cximebs anu acilglicerinebs
ganekuTvneba. cxoveluri warmoSobis triacilglicerinebSi bevria najeri
cximovani mJavebi da maT myari konzistencia aqvT, xolo mcenareulSi ki -
upiratesad warmodgenilia ujeri cximovani mJavebi da isini Txevad
mdgomareobaSi imyofebian. triacilglicerinebis daJangvisas gamoiyofa
orjer ufro meti siTbo, vidre naxSirwylebis dawvis Sedegad.
cxovelTa organizmi linolisa da linolenis mJavebs ver asinTezirebs, da
amitom isini mza mdgomareobaSi unda iqnan miRebuli. es ukanaskneli sakmao
raodenobiT arian mcenareul zeTSi. linolis mJavidan warmoiqmneba
araqidonis mJava, romelic cxovelur organizmebSi ujredSida regulators
warmoadgens, da romlisganac sinTezirdebian fiziologiurad aqtiuri
nivTierebani: prostaglandinebi, tromboqsanebi da leikotrienebi.

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 ganviTarebuli mosaxleobis kvebis racioni mdidaria cxoveluri
warmoSobis cximebiT. dadgenilia korelacia saWmlis cximianobasa da
zogierT kardiovaskularul daavadebaTa Soris. najeri cximovani mJavebi da
qolesterini iwveven aTerosklerozuli wanazardebis warmoqmnas
sisxlZarRvTa kedlebze, riTac amcirebs sisxlZarRvTa diametrs, da
Sesabamisad, organoebis sisxliT momaragebas. amis Sedegad, SesaZlebelia
ganviTardes hipoqsiuri movlenebi, Trombozebi da garkveul organoTa iSemia.
am dros sisxlSi matulobs dabali simkvrivis lipoproteinebis raodenoba.
cilebi. adamianis organizmSi 10 aminomJavis sinTezi ver warmoebs da amitom
maT Seucvleli aminomJavebi ewodebaT. esenia: arginini, histidini, izoleicini,
leicini, lizini, meTionini, treonini, triftofani da valini. argininis
miwodeba mxolod axalSobilebisTvis aris aucilebeli, radgan mozrdil
asakSi adamianis organizmi asinTezirebs am aminomJavas Sardovanas ciklis
saSualebiT.
cilebis kvebiTi Rirebuleba, misi xarisxi damokidebulia or faqtorze: 1.
mis aminomJavur Semadgenlobaze da 2. maT SeTvisebadobaze. amitom, sakveb
cilas qimiuri da biologiuri Rirebuleba gaaCnia. cilebis qimiuri
Rirebuleba damokiidebulia masSi Semaval aminomJavaTa kompleqtze, xolo
biologiuri - mis unarze SeinarCunos organizmis azoturi balansi
(magaliTad, TeTr purSi Semavali cilebis qimiuri Rirebuleba tolia 47,
xolo biologiuri - 30; saqonlis xorcisTvis es maCveneblebi tolia,
Sesabamisad 98 da 93). mcenareul cilebi, miuxedavad imisa, rom bevr maTgans
sakmaod maRali qimiuri Rirebuleba aqvs, organizmis mier Znelad iTvisebian,
radgan daculi arian celuloziT da sxva arahidrolizebadi
polisaqariduli garsiT.
sakveb produqtebSi Seucvleli aminomJavebis deficiti organizmis
(gansakuTrebiT mzardi) ganviTarebas aqveiTebs; iwvevs kunTebis atrofias,
sisustes da anemias. gansakuTrebiT, maTi defici sazianoa nervuli sistemis
formirebisa da misi Camoyalibebis periodSi, rodesac adgili aqvs sinafsuri
kavSirebis da neironuli jaWvebis warmoqmnas. am dros, SesaZlebelia tvinSi
ganviTardes Seuqcevadi cvlilebebi, romlis koreqciac SemdgomSi rTul
problemebTan aris dakavSirebuli.
vitaminebi. dReisaTvis aRmoCenili da identificirebulia yvela vitamini.
vitaminebi, organizmis mier maTi moTxovnilebis mixedviT, SesaZlebelia or
klasad daiyos: vitaminebi, romelTa ukmarisobac sagrZnoblad moqmedebs
adamianis janmrTelobaze, da vitaminebi, romelTa naklebobac iSviaTad

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aRiniSneba. kvebis normaluri racionis pirobebSi I jgufs miekuTvnebian:
Tiamini, nikotinamidi, riboflavini, folisa da askorbinis mJavebi, xolo
meores - piridoqsini, biotini, vitamini B12 da vitaminebi A, D, E da K.
organizmis moTxovnileba vitaminebze damokidebulia sakvebis tipze, saWmlis
momnelebeli traqtis mikroorganizmebze, genetikur faqtorebze. ase
magaliTad, Tu nikotinamidze moTxovnileba damokidebulia sakvebis xarisxze,
mis cilebSi aminomJava triftofanis arsebobaze, biotinis, panToTenis mJavis
da B12 ukmarisobas organizmi iSviaTad ganicdis, radgan isini mikrofloris
saSualebiT sinTezirdebian da miewodebian organizms sakmarisi raodenobiT.
wyalSi xsnadi vitaminebis umravlesoba regularulad unda miewodos
adamians, radgan isini SedarebiT swrafad gamodian organizmidan an ixarjebian
fermentul reaqciebSi. amitom maTi Warbi raodenoba organizmisTvis saSiSi ar
aris, gansxvavebiT cximSi xsnadi vitaminebisagan (gansakuTrebiT A da D),
romelTa zedmeti doza SesaZlebelia toqsikuric ki aRmoCndes.
Tiamini (B1). Tiaminis ukmarisobisas viTardeba nevrologiuri daavadeba beri-
beri. gavrcelebulia aziis qveynebSi, sadac ZiriTad sakveb produqts brinji
Seadgens. am vitaminis ukmarisoba gvxvdeba ganviTarebul qveynebSic da
mJRavndeba encefalopatiis (vernikes sindromi) an gul-sisxlZarRvovani
sistemis upiratesi dazianebiT (veisis sindromi). specifikuri simptomebi
dakavSirebulia gul-sisxlZarRvovani da nervuli, aseve saWmlis momnelebeli
sistemis darRvevebTan. B1-avitaminozis adreul simptomebad SeiZleba
CaiTvalos: umadoba, nawlavTa peristaltikis Seneleba (atonia), agreTve
fsiqikis Secvla, mexsierebis daqveiTeba, gul-sisxlZarRvovani sistemis
darRvevebi. avitaminozis Semdgomi ganviTarebisas mJRavndeba periferiuli
nervuli sistemis dazianebis, gulis ukmarisobis simptomebi. B1-is
biologiuri roli ganisazRvreba misi monawileobiT fermentul reaqciebSi,
kerZod, iseT mniSvnelovan kompleqsebSi, rogorebic arian
piruvatdehidrogenaza da -ketoglutaratdehidrigenaza.
am vitaminiT mdidaria Savi puri, lobio, mcenareuli zeTi, RviZli da
Tevzeuloba. vitamini B1 mJave areSi uZlebs kulinarul damuSavebas, xolo
tute da neitralur pH-ze ki - hidrolizdeba da kargavs aqtivobas,
gansakuTrebiT maRal temperaturaze. rekomendirebul yoveldRiur normas -
1,2-2,2 mg Seadgens.
riboflavini (B2). wyalSi xsnad vitamins warmoadgens. mdgradia mJave aris
mimarT; advilad iSleba tute da neitralur pH-ze. riboflavini Sedis

189
flavinadeninmononukleotidis (FMN) da flavinadenindinukleotidis (FAD)
SemadgenlobaSi. igi monawileobas Rebulobs Jangva-aRdgeniT re- aqciebSi.
ukmarisobis simptomebs warmoadgens - enis lorwovani garsis (glostiti),
aseve kanis epiTeliumis (gansakuTrebiT piris areSi) anTebiTi procesebi,
Tvalis rqovanas anTeba (keratiti), kataraqta. avitamonozisTvis
damaxasiaTebelia gulis kunTis da saerTo sisuste. riboflavini farTod
aris gavrcelebuli bunebaSi. gvxvdeba TiTqmis yvela saxis produqtSi.
mdidaria am vitaminiT Savi da Talxi puri, kvercxi, xorci, rZe, bostneuli da
a.S.
yoveldRiur rekomendirebul normas - 1,7 mg warmoadgens.
nikotinamidi (niacinamidi, PP) wyalSi cudad ixsneba, Sedis NAD-is da NADP-
is SemadgenlobaSi, monawileobas iRebs kofermentis formiT dehidrogenazul
reaqciebSi. vitamin PP-s ukmarisoba iwvevs pelagras, romelsac sami ZiriTadi
simptomi axasiaTebs: kanis anTebiTi procesi (dermatiti), diarea da umaRlesi
nervuli moqmedebis daqveiTeba (demencia). gavrcelebulia iseT regionebSi,
sadac upiratesad simindis produqtebiT ikvebebian. es vitamini didi
raodenobiTaa warmodgenili brinjSi, purSi, kartofilSi, lobioSi, xorcSi;
mcirea - simindSi.
yoveldRiuri normaa - 19 mg.
askorbinis mJava, vitamini C. wyalSi kargad ixsneba. warmoadgens 6-
naxSirbadian naxSirwyals me-2 da me-3 naxSirbads Soris ormagi bmiT. Zlieri
mJavaa. ormagi bma SesaZlebelia SeqceviTad daiJangos da aRdges. es
ukanaskneli ganapirobebs askorbinis mJavis antioqsidantur Tvisebebs.
daJangvis Sedegad miRebuli dihidroaskorbinis mJava ar kargavs biologiur
Tvisebebs, magram tute da neitralur areSi advilad iSleba. vitamini C-s
biologiuri funqcia jer-jerobiT gaurkvevelia, radgan ar aris nanaxi is
fermentuli sistema, romlis kofermentsac es vitamini warmoadgens. fiqroben,
rom igi monawileobs Jangva-aRdgeniT reaqciebSi.
am vitaminis ukmarisoba adamianebSi iwvevs suravands (skorbuts, cingas).
paTologia xasiaTdeba sisxlZarRvTa kedlebis da sayrdeni aparatis
dazianebiT, rac ganpirobebulia ujredSorisi adheziuri nivTierebebis
(upiratesad cilebis) biosinTezis daqveiTebiT. am dros mcirdeba kolagenis,
elastinis da sxva struqturuli cilebis sinTezic. amis Sedegad,
sisxlZarRvTa kedlebi araelastikuri da ganvladi xdebian, rac ganapirobebs
kanSi da lorwovan garsebSi sisxlCaqcevebs.
gavrcelebulia xil-bostneulSi, yoveldRiur normas - 60 mg warmoadgens.

190
 folis mJava. wyalSi sustad ixsneba. mis SemadgenlobaSi Sedis pteridinis
naSTi, romlis aRdgenili forma monawileobs bioqimiur reaqciebSi
pteridinis garda vitaminis SenebaSi monawileobas iRebs paraaminobenzois mJava
da glutaminis mJava. folis mJava warmoadgens koferments fermentul
reaqciebSi, romlebic erTatomiani naxSirbadovani naSTebis gadatanas
emsaxurebian. radgan es reaqciebi umTavresad nukleotidebis biosinTezSi
iReben monawileobas, folis mJavis ukmarisoba Trgunavs nukleinis mJavebis
sinTezs. amitom, avitaminozi, upirveles yovlisa, iwvevs eriTropoezis
inhibirebas da anemias.
folis mJavis zogierTi antagonisti (qimiuri analogi) gamoiyeneba
onkologiaSi, rogorc dnm-is sinTezis damrTgunveli nivTiereba (mag.
aminopterini - leikozis saSinaaRmdegod). garda amisa, antibaqterialuri
preparatebi - sulfanilamidebi, warmoadgenen am vitaminSi Semavali
paraaminobenzois mJavis qimiur analogs. fiqroben, rom sulfanilamidebi
konkurentulad ukavSirdebian ferments, romelic mikroorganizmebSi folis
mJavas asinTezirebs. Sedegad, es ukanaskneli ver warmoiqmneba da
baqterialuri ujredis gamravleba muxruWdeba.
folis mJava gavrcelebulia bostneulSi (foTlebSi), xorcSi. yoveldRiur
normas - 0,4 mg Seadgens.
piridoqsini(B6). tuteebSi da mJavebSi mdgradia, wyalSi kargad ixsneba,
mgrZnobiarea sinaTlis mimarT. B6-is ukmarisoba iSviaTad gvxvdeba. igi iwvevs
dermatits, romelic PP-s saSualebiT ar inkurneba. am vitaminis deficitiT
gamowveuli avitaminozis forma aRwerilia axalSobil bavSvebSi. is
upiratesad nervuli sistemis dazianebaSi (epileptoformuli Seteva)
gamoixateba. organizmSi piridoqsinidan warmoiqmneba piridoqsalfosfati,
romelic aminotransferazebis da dekarboqsilazebis prostetul jgufs
warmoadgens.
B6 farTodaa gavrcelebuli bunebaSi, gvxvdeba bostneulSi, xorcSi da a.S.
mis yoveldRiur normas - 2,2 mg Seadgens.
biotini, (vitamini H). biotini Sardovanas ciklur warmoebuls warmoadgens.
misi avitaminozis klinika kargad ar aris Seswavlili, radgan igi adamianis
mikrofliraSi arsebuli biotinis masinTezirebeli sistemebis gamo iSviaTad
gvxvdeba. biotini, prostetuli jgufis saxiT monawileobs im fermentul
reaqciebSi, romelTa saSualebiT CO2-is jgufis gadatana warmoebs (mag. pi-
ruvatdekarboqsilaza). frinvelTa kvercxis gulSi warmodgenilia cila

191
avidini, romelic specifikurad ikavSirebs biotins da aqveiTebs vitaminis
moqmedebas.
biotinis yoveldRiuri moTxovnileba Seadgens 0,15 mg.
pantotenis mJava (B3). wyalSi kargad ixsneba; advilad hidrolizdeba sust
mJavebSi da tuteebSi. B3 farTod gavrcelebulia bunebaSi, da amitom, misi
ukmarisoba iSviaTad gvxvdeba. deficiti iwvevs dermatitebs, umadobas, gulis
da Tirkmlebis paTologiebs. aseTi farTo suraTi ganpirobebulia panToTenis
mJavis biologiuri roliT; igi monawileobs erT-erTi mniSvnelovani
kofermentis, CoA-s SenebaSi.
yoveldRiuri moTxovnilebis norma tolia - 5-10 mg.
kobalamini (B12). vitamin B12-s rTuli agebuleba aqvs. igi Sedgeba oTxi
piroluri birTvisagan, romlebic koordinaciuli bmiT arian kobaltis
atomTan dakavSirebuli. kobalamini erTaderTi vitaminia, romlis
SemadgenlobaSic metalis atomi Sedis. B12 sinTezirdeba mxolod garkveuli
tipis baqteriebiT, romlebic adamianis saWmlis momnelebel traqtSi
imyofebian. adamianis da mcenareebis ujredebs am vitaminis sinTezi ar
SeuZliaT.
adamianebSi, B12-is ukmarisoba iwvevs avTvisebiani makrocitaluri
(megaloblasturi) anemiis ganviTarebas. sisxlmbadi funqciis Semcirebis
garda avitaminozisTvis damaxasiaTebelia nervuli moqmedebis daqveiTeba da
kuWis wvenis mJavianobis mkveTri dacema. ukmarisoba viTardeba ara vitaminis
simciris gamo, aramed misi matransportirebeli cilis - transkorinis
(kaslis Sinagani faqtori, gastromukoproteini) dabali Semcvelobis mizeziT.
transkorini B12-Tan warmoqmnis kompleqss, romlis saSualebiTac es vitamini
nawlavebSi Seiwoveba. amitom, perniciozul (avTvisebian) anemias mkurnaloben
vitamini B12-is peroraluri SeyvaniT transkorinTan da kuWis ganeitralebul
wvenTan erTad. aRiniSneba kavSiri avTvisebian anemiasa da kuWis lorwovani
garsis ujredebis mier cilebis (da maT Soris transkorinis) sinTezis
Semcirebas Soris.
vitamini B12 RviZlSi maragdeba, da amitom, misi avitaminozi iSviaT
daavadebas warmoadgens. kobalamini monawileobas iRebs transmeTilirebis da
izomerizaciebis reaqciebSi. am vitaminis yoveldRiuri moTxovnileba - 0,003
mg.
vitamini A (retinoli). cximSi xsnad vitamins warmoadgens. adamianis
organizmSi warmoqmnis aldehidur dormas - retinals. es ukanaskneli Sedis

192
mxedvelobiTi pigmentis - rodofsinis SemadgenlobaSi da monawileobas iRebs
sinaTlis aRqmaSi. vitamini A izoprenoiduli erTeulebisagan Sedgeba.
vitamin A-s ukmarisoba iwvevs mxedvelobis daqveiTebas, kanisa da
lorwovani garsis specifikur dazianebebs (garqovanebebs). damaxasiaTebelia
Tvalis rqovanas simSrale (qseroftalmia), romelic gamowveulia
cremlsadinrebis epiTeliumis garqovanebiT da mis daxSobiT. garda amisa,
SesaZlebelia ganviTardes koniuqtiviti, SeSupeba da Tvalis sxva dazianebebi.
avitaminozis adreuli stadiebisTvis damaxasiaTebelia qaTmis anu Ramis
sibrmave - mxedvelobis daqveiTeba bindSi. avitaminozebis dros qveiTdeba
agreTve winaaRmdegoba infeqciebis mimarT. vitamin A-s hipovitaminozebis
garda gvxvdeba misi hipervitaminozebic. A-s hipervitaminozis specifikur
simptomebs warmoadgenen Tvalis anTeba, hiperkeratozi, Tmis dacvivna.
vitamini A-s biologiuri funqcia mdgomareobs, jer-erTi, mis
monawileobaSi sinaTlis aRqmis procesebSi, da meore, ujredis genetikuri
aparatis regulaciaSi. retinolis damakavSirebel cilas transkripciuli
aqtivoba aqvs, igi ujredebis diferencirebaSi iRebs monawileobas. amitom,
vitamini A xSirad gamoiyeneba, rogorc antiproliferaciuli agenti garda
amisa, vitamini A moqmedebs Jangva-aRdgeniT procesebzec.
am vitaminis winamorbediT-karotiniT mdidaria e.w. yviTeli mcenareebi:
stafilo, gogra, mwvane Cai da sxva. yoveldRiuri normas 2,7 mg Seadgens.
vitamini A maragdeba RviZlSi.
vitamini D (kalciferoli). cximSi xsnad vitamins warmoadgens. igi
ciklopentanperhidrofenantrenuli birTvisgan Sedgeba; adamianis organizmSi
ultraiisferi sxivebis zegavleniT, qolkalciferolisagan warmoiqmneba.
vitamin D-s ukmarisoba bavSvebSi raqits iwvevs. am dros organizmSi
kalciumisa da fosforis balansis darRveva aRiniSneba, ris gamoc ZvlebSi
Ca-fosfatis gamoleqva ver warmoebs da osteogenezis procesi qveiTdeba.
Sedegad, viTardeba Zvlebis garbileba - osteomalacia. mozrdil asakSi am
vitaminis ukmarisoba iwvevs osteoporozs - marilebis gamorecxvas (ris gamoc
Zvlebi myife xdebian).
ukanasknel wlebSi gairkva, rom vitamin D-s funqcia ufro humoraluria,
vidre kofermentuli. aqtiur substanciebs, am vitaminis metabolitebi 1,25- da
24,25-dioqsiqolkalciferolebi warmoadgenen. isini aregulireben Ca-s da
fosfatis ionebis Sewovas nawlavebSi, maT rezorbcias Zvlovan qsovilSi da
reabsorbcias Tirkmlis arxebSi. sxvadasxva tipis ujredebSi aRmoCenilia

193
qolkalciferolebis damakavSirebeli cila. fiqroben, rom es cila moqmedebs
ujredebis genetikur aparatze da garkveuli cilebis induqtiur sinTezs
awarmoebs. arsebobs monacemebi vitamin D-s antiproliferaciul aqtivobaze.
vitamini D-Ti mdidaria karaqi, kvercxis guli, RviZli da cximovani
qsovilebi. yoveldRiur rekomendirebul dozas - 0,01 mg warmoadgens.
vitamini K. cximSi xsnadi naerTia. Sedgeba izoprenoiduli blokebisagan da
cikluri naftoqinonuri struqtura aqvs.
vitamini K antihemoragiul faqtors warmoadgens, romlis biologiuri
funqcia sisxlis SededebasTan aris dakavSirebuli. amitom, misi avitaminozis
SemTxvevaSi adgili aqvs parenqimatozur da kapilarul sisxldenebs.
adamianebSi am vitaminis ukmarisoba iSviaTad gvxvdeba misi mikrofloraSi
sinTezis gamo. vitamini K proTrombinisa da sisxlis SededebaSi monawile
sxva cilovani faqtorebis sinTezis regulacias awarmoebs RviZlis
ujredebSi. K-s antivitamini, dikumaroli, gamoiyeneba Trombozebis,
Tromboflebitebis da sxva paTologiebis SemTxvevebSi, romlebic viTardebian
sisxlis Sededebis zrdis fonze.
vitamini K farTod aris gavrcelebuli mcenareebSi. yoveldRiuri
sadReRamiso norma 1 mg-s Seadgens.
vitamini E (tokoferoli). cximSi xsnad vitamins warmoadgens, Sedgeba
benzoluri birTvisa da izoprenoiduli gverdiTi jaWvisagan. igi farTodaa
gavrcelebuli mcenareul samyaroSi da amitom misi avitaminozi iSviaTad
gvxvdeba. vitamini E moqmedebs ujredebis genetikur aparatze, xels uwyobs
qsovilebis diferencirebas da normalur funqcionirebas. eqsperimentul
cxovelebSi misi ukmarisoba embriogenezis darRvevas, reproduqciuli
organoebis degeneracias iwvevs.
vitamini E Zlieri antioqsidantia, da amitom, icavs membranis cximovan
mJavebs daJangvisagan. varaudoben, rom es vitamini monawileobas iRebs
mitoqondrialur eleqtronTa gadatanis jaWvSi.
yoveldRiuri rekomendirebuli dozaa - 10 mg.

makro- da mikroelementebi. ZiriTadi eqvsi elementis - naxSirbadis,

wyalbadis, azotis, Jangbadis, fosforis da gogirdis garda, adamianis
organizms sxva qimiuri elementebic esaWiroeba. isini SesaZlebelia makro- da
mikroelementebad daiyos. maT Soris sakvebi Rirebuleba SesaZlebelia
mxolod im qimiur elements hqondes, romelic produqtebSi, marilebisa Tu
sxva xsnadi naerTebis formiT imyofebian.

194
 fostori da kalciumi. zrdasruli adamiani, kbilebsa da ZvlebSi,
daaxloebiT 1 kg-mde kalciums Seicavs. aq es ionebi uxsnad minerals-
kalciumis hidroqsilapatits warmoqmnis. kalciumis ionebi ujredis
metabolizmSi iReben monawileobas. isini meorad Suamavlebs warmoadgenen da
akontroleben rig procesebs: hormonTa sekrecias, ujredTa proliferacias,
meqanikur muSaobas da sxva.
kalciumiT mdidaria rZe, yveli, nigozi, bostneuli da sxv. Zvlovani
sistemis formirebasTan dakavSirebiT, moTxovnileba kalciumSi didia
bavSvebSi da orsul qalebSi. wa-is Sewova nawlavebis mier vitamin D-s
saSualebiT regulirdeba.
Ca-is sadReRamiso moTxovnileba - 800 mg-s Seadgens.
kalciumis cvla organizmSi mWidroT aris dakavSirebuli fosfatebis
cvlasTan. es ukanaskneli, cocxali sistemebis erT-erT ZiriTad anions
warmoadgens. maTi bunebaSi farTod gavrcelebis gamo fosfatebiT gamowveul
ukmarisobas iSviaTad vxvdebiT.
magniumi. magniumis raodenoba adamianis organizmSi 25 g ar aRemateba da misi
ZiriTadi raodenoba ZvlebSia Tavmoyrili. ujredSi Mg-is koncentracia
maRalia da Seadgens 3-7 mM-s. igi monawileobas iRebs mraval fermentul
reaqciebSi, gansakuTrebiT iseTebSi, romlebic ATP-s Tanaobisas
mimdinareoben.
Mg-is yoveldRiuri moTxovnilebis normaa - 350 mg.
natriumi da kalciumi. Na ZiriTad ujreTgare, xolo K- ujredSida kaTions
warmoadgens. maT mTavar funqcias eleqtrolituri da buferuli balansis
regulacia Seadgens. organizmSi am ionTa raodenoba hormonebiT
(mineralkortikoidebiT) regulirdeba. NaCl-is Warbi raodenobiT miReba iwvevs
hipertoniul daavadebebs. K+-is gamodevna organizmidan warmoebs diabetis,
kuWis aSlilobis, diuretuli preparatebis miRebis Sedegad.
kaliumi bevria pamidorSi da citrusebSi.
rkina da spilenZi. rkina, romelic mxolod Fe++-is formiT Seiwoveba
nawlavebis mier, sakmaod dabali efeqturobiT moixmareba.am kaTionis mxolod
umniSvnelo nawili xvdeba adamianis organizmSi. rkinis utilizaciis xarisxi
damokidebulia sakvebi produqtis raobaze da im formaze, romliTac es
kaTioni biologiur obieqtSi imyofeba.
rkinis ioni aucilebelia rkina-porfirinuli cilebis funqcionirebisTvis
(hemoglobini, mioglobini, citoqromebi). sisxlSi igi cila-transferinis
saSualebiT gadaitaneba, xolo ujredebSi ki grovdeba cila-feritinTan

195
erTad. feritini didi raodenobiT sisxlmbad organoebSi - Zvlis tvinSi,
RviZlSi da elenTaSi grovdeba. rkinis ionebi SardTan erTad ar
gamoidevnebian, maTi eqskrecia naRvelTan kompleqsSi warmoebs. rkinis
ukmarisoba rkinadeficitur anemias iwvevs. am dros, eriTrocitebis
normaluri koncentraciis miuxedavad, sisxlSi hemoglobinis raodenoba
Semcirebulia.
spilenZi, adamianis organizmisTvis erT-erT aucilebel ions warmoadgens,
radgan igi mravali metaloproteinis SemadgenlobaSi Sedis. misi
Tanapovniereba aucilebelia SemaerTebeli qsovilebisa da sisxlZarRvTa
ganviTarebisaTvis.
yoveldRiurad adamians 2,5-5,0 mg spilenZi esaWiroeba.
iodi. am anionis ukmarisobiT gamowveul paTologias Ciyvi warmoadgens. es
paTologia gavrcelebulia im regionebSi, sadac iodis raodenoba niadagSi
(da Sesabamisad wyalSi) mcirea. iodis ZiriTad momxmarebel organos,
farisebri jirkvali warmoadgens. am ukanasknelSi iodi gamoiyeneba
Tireoiduli hormonebis sinTezisTvis.
ftori. ftoris ionebi monawileoben Zvlis da kbilebis Camoyalibeba-
formirebaSi. isini uerTdebian hidroqsilapatits da warmoqmnian
ftorapatits:

[Ca3(PO4)2]3.Ca(OH)2 + 2NaF ____ [Ca3(PO4)2]3.CaF2 + 2NaOH

hidroqsilapatiti ftorapatiti

miuxedavad ftorapatitis simcirisa, isini Zvlebsa da kbilebs

mJavegamZleobas da simtkices aniWeben. naturaluri da rafinirebuli
produqtebi ftors mcire raodenobiT Seicaven, da amitom, ganviTarebul
qveynebSi wylis ftorirebas awarmoeben. ftoris ukmarisobis erT-erT
gamomJRavnebaa kbilebis kariesi.
mikroelementebi. adamianisTvis aucilebel ZiriTad mikroelements TuTia
warmoadgens. igi monawileobas iRebs ujredTa genur aqtiobaSi, Sedis 80-mde
fermentis prostetuli jgufis SemadgenlobaSi. misi yoveldRiuri
moTxovnileba 15 mg-iT ganisazRvreba.
adamianis organizms esaWiroeba agreTve kala, nikeli, vanadiumi, qromi da
siliciumi; kobalti Sedis vitamin B12 SemadgenlobaSi, seleni - erT-erTi
wamyvani fermentis- glutaTionperoqsidazis SenebaSi iRebs monawileobas,
xolo molibdeni - oqsidazebis prostetuli jgufebis Semadgeneli nawilia.

196
Sesavali. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
.

S i n a a r s i

Sesavali..........................................................1
z o g a d i b i o q i m i a
ujredis molekuluri komponentebi..............................5
wyali........................................................6
cilebi da maTi biologiuri funqcia.................................8
cilebis agebuleba............................................8
aminomJavebi-cilebis struqturuli erTeulebi...................9
cilebis pirveladi struqtura.................................11
cilebis meoreuli struqtura..................................12
cilebis mesameuli struqtura.................................13
cilebis meoTxeuli struqtura.................................14
cilebis funqcia (zogadi mimoxilva)..........................15
antisxeulebi da imunuri reaqciebi (zogadi mimoxilva)........16
cilebis posttranslaciuri modifikaciebi......................17
fermentebi, maTi funqcia da daxasiaTeba..........................18
fermentTa klasifikacia......................................19
fermentTa kofaqtorebi.......................................19

197
fermentuli katalizi.........................................20
fermentebis moqmedebis meqanizmi da maTi regulacia..........23
lipidebi.........................................................25
cximovani mJavebi...........................................26
neitraluri cximebi..........................................26
fosfogliceridebi............................................27
sfingolipidebi da glikolipidebi.............................28
steroidebi..................................................29
terpenebi...................................................29
prostaglandinebi............................................30
lipiduri micelebi, lipoproteidebi da elementaruli membrana..30
Saqrebi..........................................................32
monosaqaridebi..............................................33
disaqaridebi................................................35
polisaqaridebi..............................................35
Saqrebi da ujredis garsi....................................36
nukleotidebi da polinukleotidebi.................................37
nukleotidebi................................................37
polinukleotidebi............................................40
dnm.........................................................41
rnm.........................................................42
virusebi....................................................44
m e t a b o l i z m i
adenozintrifosfati da ujredis bioenergetika.................45
katabolizmi......................................................49
glikolizi........................................................49
glikogenolizi....................................................53
limonmJavas cikli................................................57
acetil-woA-s warmoqmna......................................57
limonmJavas cikli...........................................58
glukozis katabolizmis meoradi gzebi-pentozofosfaturi gza.........62
eleqtronTa gadatana da JangviTi fosforilireba....................64
sunTqviTi JaWvi.............................................64
ATP-is sinTezi..............................................68
mikrosomuli daJangva.............................................70
cximovani mJavebis daJangva......................................71

198
 cximovani mJavebis transporti mitoqondriebSi................71
cximovani mJavebis daJangva.................................72
ketonuri sxeulebi da maTi metabolizmi.......................74
aminomJavebis daJangva da Sardovanas cikli.......................75
transaminazebi..............................................75
glutamatdehidrogenaza.......................................77
Sardovanas cikli............................................79
aminomJavebis dekarboqsilireba..............................80
aminomJavebis naxSirbadovani ConCxis katabolizmi............82
zogierTi memkvidreobiTi daavadebebi, dakavSirebuli aminomJavebis
katabolizmTan......................................82
anabolizmi.......................................................84
naxSirwylebis biosinTezi cxovelTa organizmSi.....................84
glukoneogenezi..............................................84
glikogenis sinTezi..........................................86
naxSirwylebis cvlis darRvevebi..............................87
lipidebis biosinTezi.............................................89
cximovan mJavaTa sinTezi....................................89
triacilglicerinebis sinTezi.................................92
fosfogliceridebis biosinTezi................................93
steroidebis biosinTezi......................................95
lipidTa cvlis darRvevebi....................................96
aminomJavebis da nukleotidebis biosinTezi........................97
aminomJavebis sinTezi adamianis organizmSi..................97
kreatinis biosinTezi........................................98
porfirinebis biosinTezi.....................................98
nukleotidebis biosinTezi....................................99
purinebis biosinTezi........................................99
pirimidinebis biosinTezi...................................101
purinisa da pirimidinis fuZeebis daSla..........................102
nukleinis mJavebis da cilis biosinTezi..........................103
qromosomebis da genebis struqtura..........................103
dnm-is biosinTezi..........................................104
rnm-is biosinTezi..........................................108
cilis sinTezi da misi regulacia............................110
a d a m i a n i s b i o q i m i a

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ujreduli aqtivobis regulacia....................................118
plazmuri membranis receptorebi.............................119
ujredSida regulatorebi (meoradi Suamavlebi)................123
citoplazmuri receptorebi...................................127
zrdis faqtorebi da maTi receptorebi........................127
hormonebi.......................................................129
hipoTalamusis hormonebi....................................130
hipofizis hormonebi........................................130
farisebraxlo jirkvlis hormoni..............................132
farisebri jirkvlis hormonebi...............................132
kuWqveSa jirkvlis hormonebi................................133
Tirkmelzeda jirkvlis hormonebi.............................134
sasqeso jirkvlebis hormonebi...............................136
qsovilovani hormonebi......................................136
organoTa bioqimia...............................................139
saWmlis momnelebeli sistema................................139
RviZli.....................................................142
sisxli.....................................................146
Tirkmelebi da Sardi........................................151
nervuli qsovili............................................153
kunTovani qsovili..........................................159
SemaerTebeli qsovili.......................................164
Zvlovani qsovili...........................................167
kvebis bioqimia.................................................169
energetikuli nivTierebebi..................................169
vitaminebi.................................................170
makro- da mikroelementebi..................................177
sakveb produqtTa bioqimiuri toqsikologia........................180
bioqimiuri toqsikologiis ZiriTadi principebi...............180
qsenobiotikTa metabolizmi da maTi eliminacia...............181
garemos minarevebi sakveb produqtebSi......................184

leqciaTa kursSi gamoyenebulia masalebi wignebidan:

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 leqciaTa kursi ibeWdeba eqimTa daxelovnebis institutis samecniero
sabWos rekomendaciiT

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