leqcia VII. danawevreba.

zigotis dayofa, blastulacia

dayofis tipebi. zigotis mravaljerad gayofas, romelic iweba
ganayofierebis Semdeg, dayofa ewodeba. mas mosdevs mravalujredovani
Canasaxis ganvTareba (suraTi 1).

suraTi 1. zigotis mravaljeradi dayofa.

ganayofierebisTanave zigota iyofa or ujredad, eseni Tavis mxriv

kvlav iyofian; aRmocenebuli oTxi ujredi iyofa rvad da a.S. dayofa
imdenad swrafad xdeba, rom ujredebi ver aswreben zrdas da sul ufro
da ufro wvrildebian. vinaidan dayofis dros ujredTa ricxobriv
zrdas Tan sdevs maTi zomis Semcireba, Canasaxis saerTo mouloba
TiTqmis ar icvleba.
gayofis Semdeg warmoqmnil ujredebs blastomerebi ewodeba. xolo
gadazonvras, romelic maT erTmaneTisgan TiSavs – dayofis Rarebi.
dayofis Rarebis mimarTuleba SeiZleba sxvadasxva iyos da maT
Sesabamisad aqvT saxelwodeba:
meridianulia Rarebi, rodesac isini animaluri polusidan
vegetatiurisken miimarTebian; ekvatoruli Tu zigotas ganivi
mimarTulebiT kveTen; ganedur, Tu ekvatoris paralelurni arian, da
tangenciuri, rodesac zigotas zedapiრis paralelurad miemarTebian.
dayofis Rarebis mimarTuleba ganisazRvreba dayofis TiTistaris
mdebareobiT. TiTistara Cveulebriv mdebareobs ujredis im nawilSi,
romelic TiTqmis ar Secavs CanarTebs da, amave dros, Tavisufal
plazmas meti gafeniloba aqvs. es movlena sayovelTao ar aris, magram
vinaidan sakmaod xSiria, igi dayofis wesad miiCnies. Ppirveli wesi
asaxavs ბlastomerSi TiTistaris adgil-mdebareobas, meore – mis
mimarTulebas. am or wess unda daematos mesamec dayofis Rarebis
gavlis siswrafe yovelTvis ujredSi yviTris raodenobis
ukuproporiulia.
sxvadasxva mravalujredianSi dayofa sxvadasxvanairad
mimdinareobs. es damokidebulia kvercxujredis agebulebis
Taviseburebaze, pirvelyovlisa – masSi yviTris raodenobაsa da mis
ganawilebaze. kvercxujredSi yviTri ar aris, cotaa an saSualo
raodenobiT aris, dayofas ganicdis mTeli ganayofieebuli
kvercxujredi, rac ufro metia masSi yviTri, miT metad ferxdeba
dayofis Rarebis gavleba. yviTris mniSvnelovani raodenobis dros,
zigotis mxolod is nawili iyofa, romelic yviTrisagan Tavisufalia.
imisda mixedviT, dayofas mTeli zigota ganicdis Tu mxolod misi
nawili, ganasxvaveben srul da arasrul dayofas. Amis Sesabamisad
arCeven: holoblastur kvercxebs sruli dayofiT meroblasturs –
nawilobrivi dayofiT.
sruli dayofa SeiZleba iyos Tanabari da araTanabari. esec
ganpirobebulia kvecxujredSi yviTris raodenobiT da misi ganawilebiT.
Tu yviTri cotaa da igi mTel kvercxujredis gaswvriv erTnairi
siswrfiT gaivlian da yofen mas erTnairi zomis blastomerebad. Tu
yviTri kvercxujredSi araTanabrad aris ganawilebuli, maSin is nawili,
sadac igi metia, ufro nela iyofa, vidre yviTriT Raribi nawili. Aamis
Sedegad miiReba araTanabari zomis blastomerebi. mcireni – animalur
naxevarsferoSi da msxvili – vegetatiurSi. pirvel maTgans mikromerebs
uwodeben, meores ki, - makromerebs. aseT SemTxvevaSi dayofa Tumca
srulia, magram arTanabari.
gayofis pirveli Rari animalur naxevarsferoSi Cndeba da kvercxis
ReZis gadakveTiT miemarTeba vegetatiurisaken. warmoiqmneba ori, TiTqmis
Tanabari zomis blastomeric. meore Raric meridianulia, magram
pirvelis mimarT perpendikularuli. igi or pirvel blastomers oTxad
yofs. sruli Tanabari dayofa axasiaTebs lancetas. misi kvercxi
homolecitalur tips miekuTvneba: igi Seicavs mTel ujredSi Tanabrad
ganawilebul mcire raodenobiT yviTrs.
umravlesi cxovelis kvercxujredis msgavsad lancetas
kvercxujredi polusebis mimarTulebiT odnav Sebrtyelebulia. aseTi
formis gamo mas meti ganfeniloba aqvs horizontalur sibrtyeSi da
dayofis wesis Tanaxmad, pirveli TiTistara horizontalurad Tavsdeba,
xolo dayofis Rari vertikalurad gaivlis. yovel warmoqmnil
blastomerSi citoplazmis meti ganfeniloba horizontalur sibrtyeSi
rCeba, amitom dayofis meore Rari isev meridianulad gaivlis. meore
dayofis Semdeg blastomerebSi citoplazmis meti ganfenilobis
mimarTuleba icvleba. amis Sesabamisad dayofis TiTistara vertikalurad
Tavsdeba da mesame Rari ekvatorul sibrtyeSi gaivlis. axla ukve oTxi
zeda (animaluri) blastomeric gamoeyofa oTx qveda (vegetatiur)
blastomers. meoTxe dayofa isev meridianulia. Amis Semdeg ganivi
(ekvatoruli da ganeduri) da gaswvrivi (meridianuli) dayofa erTimeores
kanonzomierad scvlis. blastomeris raodenoba ormagdeba Semdegi
TanafardobiT: prveli ori blastomeric iyofa 4-ad, Semdeg miiReba 8, 16,
32, 64, 128 da a.S. blastomeric. orive naxevarsferos blastomerebi
inarCuneben TiTqmis Tanabar zomas.
dayofis Sedegad viTardeba mravalujrediani, Rrus Semcveli,
sferos formis Canasaxi, romelsac blastula ewodeba. masSi arCeven
didi raodenobiT wvrili ujredebiT warmodgenil kedels –
blastodermas da Rrus – blastocels (sxeulis pirveli Rru).
blastulis animalur nawils bastulis saxuravs uwodeben, vegetatiurs
ki – blastulis fskers. sruli da Tanabari dayofa lancetaSi
yovelTvis mTavrdeba blastulis warmoqmniT, romlis kedeli
warmodgenilia ujredebis erTi SriT, xolo Rru centrSia moTavsebuli.
aseT blastulas celoblastula ewodeba.
sruli araTanabari dayofa axasiaTebs wyalxmeleTa cxovelebs
(suraTi 2), romelTa kvercxebi miekuTvneba telolecitarul tips,
yviTris saSualo raodenobiT.

suraTi 2. sruli, araTanabari dayofa amfibiebSi.

yviTri mTel kvercxSia ganawilebuli, magram misi umetesi nawili

yovelTvis vegetatiur naxevarSia Tavmoyrili da yviTris marcvlebic
ufro msxvilia. polaruli diferencireba mkveTradaa gamoxatuli.
pirvel ori dayofis Rari gaivlis meridianulad. isini swrafad yofen
animalur nawils, vegetatiurs ki – ufro nela. mesame Rari gaivlis
ekvatorul sibrtyis paralelurad da blastomerebs yofs ganivad. igi
Tavsdeba animaluri polusis axlos. Ees imitom xdeba, rom dayofis
TiTistara gadainacvlebs vegetatiuri naxevarsferodan, sadac
Tavmoyrilia yviTris ZiriTadi nawili, animalurisaken. zeda
blastomerebi ufro mcire zomisani arian vidre qveda. Semdgomi dayofis
dros animaluri blastomerebi vegetatiurTan SedarebiT bevrad swrafad
iyofian, amitom maT zomaSi gansxvaveba sul ufro mkveTri xdeba.
arasruli dayofa xasiaTdeba kvercxis mxolod im nawilis
dayofiT, romelic yviTrs ar Seicavs, xolo yviTris Semcveli nawili ar
iyofa. am tipiT viTardebian telolecitaruli (Zlovani Tevzebis,
frinvelebis, qvewarmavlebis) da centrolecitaruli (mwerebis) kvercxebi.
am kvercxebis agebulebis TaviseburebaTa Sesabamisad arCeven diskoidur
da zedapirul dayofas (suraTi 3).
diskoiduri dayofa axasiaTebs Zvlovan Tevzebs, frinvelebs,
qvewarmavlebs. Aam cxovelebis kvercxebi mdidaria yviTriT, romlis didi
raodenoba ganapirobebs kvercxis did zomas, uyviTro citoplazma,
patara nawilis saxiT, mdebareobs kvercxis zemo nawilSi da ewodeba
Canasaxovani disko. vinaidan dayofas mxolod es Canasaxovani disko
ganicdis, mas uwodes diskoiduri dayofa.
Canasaxovani diskos sisqe metad umniSvneloa, igi yviTrzea
garTxmuli da misi ganlageba kvercxis zedapiris paralelur sibrtyes
emTxveva. amitom pirveli sami an oTxi dayofisas TiTistarebi
horizოntალuრad lagdeba, xolo dayofis Rarebi - vertikalurad. amis
Sedegad Cndeba ujredebis erTi Sre.
 suraTi 3. arasruli dayofa frinvelebsa da mwerebSi.

ramdenime gayofis Semdeg ujredebis simaRle matulobs, ris gamoc

dayofis TiTistarebi maTSi vertikalurad lagdeba, xolo dayofis
Rarebi – kvercxis zedapiris paralelurad. amasTan dakavSirebiT
Canasaxovan diskoSi Cndeba zedapiruli ujredebi da yviTrze mdebare
ujredebi. momdevno dayofebi mimdinareobs sxvadasxva mimarTulebiT da
Canasaxovan disko ujredebis ramdenime rigisagan Semdgar firfitad
gardaiqceva. diskosa da yviTrs Soris Cndeba patara napralis magvari
Rru, romelsac blastocels uTanabreben.
zedapiruli dayofa axasiaTebs centrolecitalur kvercxebs,
romlis SuaSi didi raodenobiT yviTria. aseT kvercxebSi citoplazma
ganlagebulia ujredis periferiaze da centrSi birTvis axlos, yviTris
masaSi gadis wvrili citoplazmuri xarisxebi, romlebic akavSiebs
periferiul da birTvTan mdebare citoplazmas.
dayofa iwyeba birTvis gayofiT da citoplazmis gamocalkevebiT
axlad warmoqmnili birTvebis irgvliv. birTvebis ricxvi izrdeba.
citoplazmiT garSemortymulni isini TandaTan gadainacvleben
kvercxujredis periferiisaken. rogorc ki birTvebi moxvdeba mis
zedapirul SreSi, kvercxujredi birTvebis ricxvis Sesabamisad daiyofa
blastomerebad. Aamgvarad, dayofis Sedegad, citoplazmis mTeli
centraluri nawili gadainacvlebs zedapirisaken da Seerwymeba
periferul citoplazmas. warmoiqmneba erTiani blastoderma,
romlisganac viTardeba Canasaxi. zedapiruli dayofa, rogorc wesi
axasiaTebs mxolod fexsaxsriani cxovelebis kvercxebs.
dayofis formebis ganxilva gviCvenebs, Tu ramdenad rTuldeba es
procesi kvercxujredSi yviTris radenobis zrdasTan dakavSirebiT da
rogor TandaTanobiT icvleba umdables qordianTa sruli da Tanabari
dayofa nawilobrivi dayofiT qvewarmavlebsa da frinvelebSi.
dayofis xasiaTze did gavlenas axdens ara marto yviTris
Semcveloba da ganawileba kvercxujredSi, aramed iseTi faqtorebic,
rogoricaa citoplazmis Tvisebebi, rac gansazRvravs blastomerebis
ganlageba urTierTis mimarT, am niSnis mixedviT arCeven agreTve dayofis
ramdenime tips: radialurs, spiralurs da ormxriv simetriuls
(bilateralurs) (suraTi 4, 5).
radialuri dayofa xasiaTdeba imiT, rom yoveli zeda
blastomeric Tavsdeba zustad qveda bლastomeris zemoT, amis Sedegad
miiReba sferos radiusebis Sesabamisi rigebi. Bblastomerebis amgvari
ganlageba gamowveuliა imiT, rom dayofis TiTistarebi monacvleobiT
ikaveben xan vertikalur mimarTulebas, xan horizontalurs. Amis
Sesabamisad blastomerebi gamoiyofa xan zeviT da qveviT, xan marjvniv da
marcxniv, radialuri dayofa damaxasiaTebelia nawlavRruianebis,
ekalkanianebisa da agreTve bevr qordianebisaTvis.
 suraTi 4. radialuri da spiraluri dayofa.

suraTi 5. spiraluri dayofa.

spiraluri dayofa axasiaTebs Wiebis da moluskebis umravlesobas.

maTi kvercxebis animaluri nawilis blastomerebis citoplazma yoveli
dayofis win gadainacvlebs ganze, ris gamoc dayofis TiTistaras
mimarTuleba vertikaluris nacvlad gadaixreba daaxloebiT 450 kuTxiT.
amis Sedegad, zeda blastomerebi qveda blastomerebis Tavze ki ar
ganlagdeba, rogorc radialuri dayofis dros xdeba, aramed ganlagdeba
maT Soris. amave dros yvela animaluri blastomeris citoplazma erTი
gayofiს dros gadainacvlebs marjvniv, momdevno gayofisas ki marcxniv.
Tu dayofis TiTistarebis mimarTulebis xazs gagrZelebulad
warmovidgenT, maSin igi spiraluri aRmoCndeba.
bilateraluri dayofa xasiaTdeba imiT, rom zigotis dayofisas
masze SeiZleba mxolod erTi sibrtyis gavleba, romlis orive mxareze
lagdeba erTmaneTis Sesabamisi blastomerebi. am tipis dayofa axasiaTebs
mrgvali Wiebisa da ascidიebis kvercxebs.

blastula da morula. rogorc zemoT iyo naTqvami, dayofis procesi

mTavrdeba blastulis CamoyalibebiT (suraTi 6). zogierT cxovelSi
dayofis Sedegad waroiqmneba morula, romelic blastulisagan
gansxvavebiT warmoadgens mTlian sferos, romelsac SigniT Rru ar
gaaCnia (suraTi 7).

suraTi 6. blastula.
 suraTi 7. morula.

blastulis ama Tu im formis warmoqmna damokidebulia

citoplazmis Tvisebebze. Tu igi saკmaod blantia, maSin blastomerebi
inarCuneben mrgval formas da mxolod erTmaneTis Sexebis adgilas
odnav brtyeldebian. amis Sedegad, ukve 4 da 8 blastomeris stadiaze maT
Soris Cndeba naprali, romelic dayofis procesSi izrdeba, ivseba
siTxiT da blastocelad iqceva. citoplazmis mcire siblantis
SemTxvevaSi blastomerebi ar mravldebian da ise mWidrod lagdebian
erTmaneTTan, rom maT Soris naprali ar rCeba da Rruc ar warmoiqmneba.
mraval saxelmZRvaneloSi morulas iseT stadias uwodeben,
romelic aucileblad win uswrebs blastulas. Pivanovis SexedulebiT ki
morula da blastula stadiebia, rolebic sxvadasxva cxovelebSi
warmoiqmneba blastomerebis citoplazmis Tvisobrivi gansxvavebiს gamo.
blastulebi Tavisi agebulebiT metad gansxvavebulni arian, rac
damokidebulia dayofis tipze, romlis Sedegadac isini warmoiqmneba.
ganasxvaveben celoblastulas, amfiblasტulas, steroblastulas,
diskoblastulas da periblastulas.
celoblastula – blastulaa didi blastoceliT da Tanabari
sisqis, erTSriani kedliT. igi warmoiqmneba sruli, Tanabari dayofis
dros (magaliTad, lancetaSi) (suraTi 8).
 suraTi 8. celoblastula.

amfiblastula (suraTi 9) celoblastulisagan imiT gansxvavdeba,

rom misi kedeli ujredebis ramodenime Srisgan Sedgeba da animalur
nawilSi ufro Txelia, vidre vegetatiurSi, misi blastoceli
gadanacvlebulia animaluri polusisaken. amfiblastula warmoiqmneba
sruli, magram arTanabari dayofis Sedegad (magaliTad amfibiebSi).

suraTi 9. amfiblastula.

steroblastula Sedgeba didi zomis blastomerebisagan, romlebic

mis kedelSi erT rigad arian ganlagebulni. blastulis RruSi
blastomerebis Rrmad SeWris gamo, Rru mniSvnelovnad mcirdeba da
zogjer sruliad qreba. steroblastula axasiaTbs zogierT
fexsaxsrianebs.
diskoblastula (suraTi 10) warmoiqmneba diskoiduri dayofisas.
dayofis Rru viwro napralis saxiT imyofeba Canasaxovan diskosa da
yviTrs Soris. Diskoბlastula aqvT Zvlovan Tevzebs, qvewarmavlebsa da
frinvelebs.

suraTi 10. diskoblastula.

periblastulas faqtiurad Rru ar gaaCnia, radgan igi mTlianad

yviTriT aris amovsebuli. misi blastoderma Sedgeba ujredebis erTi
Srisagan, romelic gars ertymis yviTrs. Pperiblastula gxvdeba zogierT
mwerebSi.
garemos gavlena dayofaze. TiToeuli organizmis ganviTareba
mWidrოd aris dakavSirebuli garemo pirobebTan, romelSic igi
yalibdeboda xangrZivi filogenezis procesSi, organizmi garemos
umniSvnelo cvlilebebs egueba, magram xdeბa tipuri ganvიTarebisagan
gadaxra, xolo garemos mkveTri cvlilebebi ki mis daRupvas iwvevs.
ganviTarebis ZiriTad pirobebs ganekuTvnea: 1. M siTxis Tvisebebi,
romelSic viTardeba (qimiuri Sedgeniloba, osmosuri wneva), 2. garemos
temperatura da 3. garemoSi Jangbadis Semcveloba.
 marilebis xsnarTa moqmedeba ganviTarebis adreul stadiebSi
Cveuleriv iwvevs dayofis formis Secvlas, magaliTad, hipertonul
xsnarSi citoplazma kargavs wyals, rac mis siblantes zrdis. Aam
SemTxvevaSi dayofis Rarebs imdenad didi winaaRmdegoba xვdeba, rom
blastomerebis dayofa ferxdeba. kvercxis sruli dayofa aმgvar areSi
Tavisi formiT, arasruls uaxlovdeba. didi mniSvneloba aqvs wyalSi
gaxsnil marilTa Sedgenilobas: kalciumis ionebis SemTxvevaSi
blastomerebi scildeba erTmaneTs.
temperaturis awevisas aRiniSneba dayofis aCqareba, misi daweva ki
iwvevs dayofisa da mTeli ganviTarebis Senelebas. magram mainc arsebobs
garkveuli sazRvrebi, romlis farglebSic dayofa xorcieldeba:
civsisxliani cxovelebisaTvis sazRvrebi gacilebiT farToa, vidre
TbilsisxlianebisaTvis. temperaturis gavlena mJRavndeba adgilobriv
zemoqmedebaSic: Canasaxis is nawilebi, romlebic ufro maRali
temperaturis pirobebSi xvdeba, ufro swrafad viTardeba.
gayofis stadiaSi myofi kvercxebi erTob mgrZnobiareni arian
garemos Jangbadis mimarT. uJangbadod dayofa ar xdeba. marTalia,
amfibiebisa da kalmaxis kvercxebs anaerobul pirobebSic SeuZliaT
dayofa, magram isini mxolod blastulis stadiamde aRweven. aseT
garemoSi Semdgomi ganviTareba SeuZlebeli xdeba. Aam dros Cveulebriv
aRiniSneba simaxinjeebi, romelic sabolood Canasaxis daRupvas iwvevs,
normaluri ganviTarebis aseTi darRveva xSiria wyalxmeleTa cxovelebis
kvercxebis xelovnur pirobebSi dayofis dros (magaliTad, akvariumSi
kvercxebis didi raodenobis Tavmoyris SemTxvevaSi) sunTqvisTvis
Jangbadis ukmarisobis dros (magaliTad, rodesac wyalsatevi iyineba
Tevzebis mTeli qviriTi iRupeba).
gadaxrebi, romelic SeiniSneba Canasaxis uCveulo pirobebSi,
ganviTarebis dros damokidebulia am pirobebis xasiaTze, TviT Canasaxis
ganviTarebasa da bolos im stadiaze, romelSic xdeba zemoqmedeba.
 dayofis dros Canasaxi nakleb mgrZnobiarea garemos cvlilebebis
mimarT. imave faqtorebis zemoqmedeba bevrad ufro mkveT darRvevas iwvevs
ganviTarebis gvian stadiebSi, rodesac masalis morfologiuri
diferencireba iwyeba. ganviTarebis ufro gvian stadiebSi mgrZnobelobis
mateba Jangbadis Semcvelobisa da Sxamiani airebis mimarT, rogorc Cans,
gamowveulia ganviTarebis procesTan dakavSirebul sunTqvis
TandaTanobiTi gaZlierebiT.
dayofis umniSvnelo aCqareba an Seneleba iwvevs Canasaxis
ganviTarebaSi rame darRvevas, amitom faqtorebi, romlebic iwvevs am
procesis tempis Secvlas, SeiZleba gamoyenebul iqnas adamianis
sasargeblod.