larisa vaSakiZe LARISA VASHAKIDZE

vazis qarTuli genotipebis mtvris marcvlis ganviTarebis biologia GEORGIAN GRAPEVINE GENOTYPES: BIOLOGY OF POLLEN DEVELOPMENT

Tbilisi Tbilisi 2010

UDC 634.8 (479.22) + 655.3.026 V. 251 soflis meurneobis mecnierebaTa doqtoris l. vaSakiZis monografiaSi ganxilulia vazis qarTuli genotipebis mtvris marcvlis Camoyalibebisa da ganviTarebis citogenetikuri da citoembriologiuri Taviseburebani. Ggaanalizebulia warmoSobis adgilis roli mtvris marcvlis kriteriumebis cvalebadobaSi. gamoyofilia jiSis identifikaciisa da Sefasebis citologiuri markerebi. mikrofoto27, cxrili-4, diagrama-2, suraTi-23. bibliografia-51. gaTvaliswinebulia praqtikos mevenaxeTa da vazis genetika-seleqciis xaziT momuSave mkvlevarTa farTo wrisaTvis, agraruli universitetis akademiuri personalisa da studentebisaTvis. wigni momzadda mebaReobis, mevenaxeobisa da meRvineobis institutSi, vazis deskriptorebis (Vitis spp.) Sesabamisad, citogenetikis laboratoriis Tematikisa da Bioversity International-is (romi, italia) proeqtis - `vazis genetikuri resursebis konservacia da mdgradi

gamoyeneba
farglebSi.

kavkasiaSi

da

Savi

zRvis

CrdiloeT

regionebSi regi onebSi

mTavari redaqtori: n. CxartiSvili, saqarTvelos soflis meurneobis mecnierebaTa akademiis akademikosi, profesori. recenzenti: a.SaTiriSvili, a.SaTiriSvili, biolog.Mmecn. doqtori, ilias universitetis emeritusi profesori.

The monograph of the Doctor of Agricultural Sciences L. Vashakidze is dedicated to cytogenetic and cyto-embryological particularities of organization and development of male generative organs in Georgian grapevine genotypes. The role of the place of origin in the variance of pollen grain criteria is analyzed. Emphasized are the cytological markers for variety identification and assessment. The book contains 27 photomicrographs, 4 tables, 2 diagrams, 23 photos, 51 bibliographical notes. The book is intended for viticulturist practitioners and broad spectrum of researchers in the sphere of genetics and selection, as well as to the academic staff and students of the Agrarian University. The monograph has been prepared within the framework of the scientific work plan of the Cytogenetic Department of the Institute of Horticulture, Viticulture and Oenology (Tbilisi, Georgia) and the project Conservation and sustainable use of grape-vine genetic resources in the Caucasus and Northern Black Sea area realized by the Plant Genetic Resources Institute Bioversity International (Rome, Italy). Editor in Chief: N.Chkhartishvili, Academician of the Georgian Agricultural Academy of Sciences, Professor. Reviewer: A.Shatirishvili, Doctor of Sciences, the Emeritus Professor of Ilia University. ISBN 978-9941-0-2553-2

rqawiTeli

winasityvaoba qveynis ekonomika, mosaxleobis ekologiurad sufTa, nedli xiliT Tu kvebis mrewvelobis produqtebiT uzrunvelyofa, mniSvnelovan wilad aris damokidebuli soflis meurneobis prioritetuli dargebis, maT Soris, mevenaxeobisa da mexileobis mdgrad ganviTarebaze. M maRali da xarisxiani mosavlis miReba moiTxovs jiSis genetikuri potencialis Rrma codnas, genetikuri resursebis monacemTa bazebSi da eleqtronul katalogebSi ganTavsebul srulyofil informacias, mis miznobriv gamoyenebas, sadac prioriteti Tanamimdevrulad: jiSis genotips, warmoSobis adgils da agrowesebis dacviT Catarebul fitoteqnikur RonisZiebebs ekuTvnis; Tumc mniSvnelovnad yuradsaRebi da gasaTvaliswinebelia is faqtic, rom jiSis sameurneo da samecniero saqmianobaSi warmatebiT gamoyeneba da mosalodneli Sedegebis winaswar prognozireba vazis mcenaris heterozigoturobis gamo, bevrad aris damokidebuli fenotipuri niSnebis vegetatiur TaobaSi damemkvidrebis xasiaTze, radgan rigi maRali sameurneo Rirebulebis niSnebi garemo faqtorTa gavleniT ganicdian ra Zlier modifikaciur cvlilebebs TaobaSi maTi damemkvidrebis xasiaTis winaswar prognozireba mxolod garegani niSnebiT gaZnelebulia. igi moiTxovs did dros da xarjebs. aRniSnuli udevs safuZvlad mcenareTa genetikuri resursebis saerTaSoriso institutis (Bioversity International), vazisa da Rvinis saerTaSoriso ofisisa (OIV) da mcenareTa axali jiSebis dacvis saerTaSoriso kavSiris (UPOV) mier SemuSavebul vazis mcenaris aRwerisa da Sefasebis deskriptorebs (Vitis spp.), romlis mixedviT jiSis Sefasebis aucilebel pirobad aris miCneuli fenotipuri niSnebis kompleqsuri _ organizmul, ujredul da molekulur doneze Seswavla; ampelografiuli,
4

citologiuri, molekuluri da bioqimiuri maxasiaTeblebis, bioturi da abioturi stresebis mimarT mgrZnobelobisa da sxvaTa mixedviT gamokvleva; Sedegad: didi mudmivobisa da mcired cvalebadi niSnebis gamovlineba, monacemTa bazis srulyofa da iseTi test-sistemis SemuSaveba, romelic safuZvlad daedeba jiSis mdgradobis Seswavlas stresoruli, anTropogenuri faqtorebis mimarT, venaxebSi gamoyenebuli pesticidebisa da biostimulatorebis skrinings, xmarebidan gamoTiSavs maRali toqsikurobisa da genetikuri aqtivobis mqone preparatebs, monitoringi dawesdeba genofondis siwmindeze, ar moxdeba misi dabinZureba da Sesabamisad, uaryofiTi klonebis ricxvis gazrda; mosaxleobas miewodeba ekologiurad sufTa produqti, vazis jansaRi nergi, mecnierulad dasabuTdeba venaxebSi Casatarebeli mTeli rigi fitoteqnikuri RonisZiebebis optimizaciis sakiTxebi; funqcionalurad mdedrobiTi sqesis jiSebisaTvis SeirCeva ukeTesi damamtverianebeli jiSebi, ganisazRvreba jiSis SejvarebebSi sawyisi masalad gamoyenebis miznobrioba da sxva. vazis qarTuli genofondi organizmul doneze sakmaod kargad aris Seswavlili, rasac ver vityviT ujredul da miT umetes, molekulur doneze Sesrulebul kvleviT samuSaoebze, romlebic Cveni mdidari genofondisaTvis Zalze mcirea. TiTqmis Seuswavlelia mcired gavrcelebuli, dakargvis safrTxis piras myofi axlad moZiebuli jiSebi da formebi. amdenad, vazis qarTuli genofondisaTvis, dRemde problematuria arsebuli Tu axlad moZiebuli TiToeuli jiSis mcenaris aRwerisa da Sefasebis deskriptorebis moTxovnaTa Sesabamisad gamokvleva, romelic srulyofs jiSis Sesaxeb arsebul informacias da realurs gaxdis misi praqtikuli da Teoriuli miznebiT realizaciis SesaZleblobas; ganTavsdeba saqarTvelos da evropis mcenareTa genetikuri
5

resursebis monacemTa bazebsa da eleqtronul katalogebSi da gamoyenebuli iqneba jiSis SemWidrovebul vadaSi identificirebis, mosalodneli Sedegebis winaswar prognozirebis, gamravlebis, miznobriv seleqciaSi mSobeli wyvilebis SerCevis, ampelografiis ganaxlebuli gamocemebisa da sxva mniSvnelovani sakiTxebis gadasawyvetad. winamdebare monografia warmoadgens vazis qarTuli genotipebis ujredul doneze Catarebul masStaburi gamokvlevebis mxolod nawils; avtoris pirvel mcdelobas, vazis qarTuli genotipebis mtvris marcvlis ganviTarebis biologia gamosaxuli yofiliyo `samuSao ekranze~ (cxrilebSi,DdiagramebSi, mikrofotoebsa da fotoebSi). Bioversity International-is, OIV-isa da UPOV-is mier SemuSavebuli, mcenaris aRwerisa da Sefasebis deskriptorebis mixedviT, instituciuri analizis formebSi. wigni cifrobrivi masalebis simravlis gamo dazRveuli ar aris teqnikuri xarvezebisagan. amdenad, yvela SesaZlo SeniSvna da winadadeba miRebuli iqneba madlierebis grZnobiT. avtori madlobas uxdis mis yofil aspirantebs, soflis meurneobis akademiur doqtorebs: d. maRraZes, v. gurasaSvilsa da i. mdinaraZes kvlevis procesSi gaweuli teqnikuri daxmarebisaTvis. L larisa vaSakiZe

saferavi

Introduction The economy of the country, providing the population with environmentally sound raw fruit and food products considerably relies on sustainable development of top-priority fields of agriculture, including viticulture and fruit growing. Obtaining heavy yield of high quality requires deep knowledge of genetic potential of a variety, availability of full information placed in the databases and electronic catalogues of genetic resources, its proper use giving continuous priority to the genotype of variety, place of origin and phyto-technical practice in accordance with agronomical normativeness, though allowing for the fact, that successful use of a variety for economic and scientific purposes or in forecasting of expected results largely depends, due to heterozygosity of grapevine plant, on the heredity of phenotypic traits in the vegetative generations, as long as series of major productive features influenced by environmental factors undergo strong modifications, and it is difficult to make forecast of their heredity in generation based on external traits. It is time-consuming and costly. The above concept underpins the grapevine (Vitis spp.) descriptors elaborated for explanation and assessment purposes by the international institutions (Bioversity International, OIV, UPOV). According to these descriptors for assessment of a variety, it is necessary to evaluate a complex of phenotypic traits on organism, cellular and molecular levels; their investigation by ampelographic, cytological, molecular and biochemical parameters; evaluation of sensibility to biotic and abiotic stress and others. This results in highly constant and hardly varying traits, perfection of database and development of such testing systems, which can be used as a basis for investigation of a varietys resistance against stress-related and anthropogenic factors, screening of pesticides and biostimulators used in vineyards; will eliminate application of highly toxic and genetically active substances; cleanliness of germplasm will be monitored, its pollution will not take place thus preventing increase in the number of negative clones; the population will be supplied with ecological products, healthy planting material; the issues of optimization of a series of phytotechnical operations in vineyards will be given scientific credence; better pollinator varieties will be selected for female
8

cultivars; target application wil be determined for initial row material in crossings of variety, etc. Georgian grapevine germplasm is quite well investigated on organism level, though the same cannot be said about the research works on cellular and molecular levels. Such works are too small in number for our rich genetic resources. Rare varieties being under the threat of extinction as well as some newly discovered varieties and forms are hardly investigated. Therefore, for the Georgian grapevine genetic resources still problematic is research into existing or newly revealed rare varieties in accordance with grapevine descriptors, a study which will make complete all the available information regarding a variety and make real the possibility for its practical or theoretical application. It will be included in the plant genetic resources databases and the electronic catalogues of Georgia and Europe and will be used for identification of varieties in a brief period, for preliminary forecast of expected results, multiplication, selection of parent pairs for breeding, for revised editions of Ampelography and for solution of other important issues. This monograph only represents part of the widescale researches into the Georgian grapevine genotypes on cellular level; the authors first attempt to show the biology of pollen development of Georgian native genotypes on the working screen (in tables, diagrams, micro-photos and photos) based on the grapevine descriptors of the international institutions (Bioversity International, OIV, UPOV) within the framework of the forms of institutional analysis which, due to abundance of figures, is not free from technical shortcomings. Therefore, the author is ready to gratefully accept all qualified remarks and suggestions and willingly acknowledge them in the following editions. The author expresses gratitude to her former post-graduate students, Doctors of Agricultural Sciences D. Maghradze, V. Gurasashvili and I. Mdinaradze for their technical assistance during the research activities. Larisa Vashakidze

sufris gorula, kloni#21

sufrisGgorula

Tavkveri
10

asureTuli Savi

I. mikrosporogenezi da qromosomaTa meiozuri meiozuri asociaciebi

farulTeslian mcenareebSi mikrosporogenezi mikrosporangiumebSi (samtvreebSi) mtvris Camoyalibebaa. erTi samtvres orive samtvre parkis arqespora warmoqmnis didi raodenobiT mtvris deda ujredebs, romelSic mimdinareobs ori meiozuri dayofa. TiToeul maTganSi warmoiqmneba 4 mikrospora da haploiduri mtvris marcvlebi. vazSi mikrosporogenezis procesi sakmaod kargad aris Seswavlili (-, 1976; , , , , 1977; Rogtehev, Terzsky, Dimova, Karageorgiev, 1999; vaSakiZe, gurasaSvili, 2000; vaSakiZe, 2006 da sxvani), magram qarTul jiSebze mniSvnelovani gamokvlevebi ZiriTadad l. xaritonaSvilis (, 1971) saxelTan aris dakavSirebuli. aRmosavleT saqarTvelos pirobebSi, aprilis bolos, maisis pirvel dekadaSi, xolo dasavleT saqarTveloSi erTi kviriT adre vazis yvavilis kokrebSi, erTnairi meristemuli ujredebisagan Semdgari borcvebis saxiT, isaxeba samtvre parkebi, romlebic mikrosporangiumis ganviTarebasTan erTad oTxnakvTian formas Rebuloben. amave periodSi uSualod epidermisis qveS formirdeba endoteciumi, mas mosdevs paracetaluri ujredebis ori Sre da tapetumi. tapetumis ujredebi samtvre parkis ganviTarebis sawyis etapebze zomiT da struqturiT arqesporiumis msgavsia, magram mas Semdeg, rac mikrosporas deda ujredebi meiozur gayofas iwyeben, maTi nawili, birTvis meiozuri gayofisa da citokinezis amovardnis Sedegad, ori an mravalbirTviani xdeba; zogjer ori an meti birTvis Serwymis Sedegad erTi didi poliploiduri birTvi viTardeba. mikrospo11

ras deda ujredebSi meiozis damTavrebis periodisaTvis tapetumis ujredebi daSlas iwyeben da samtvre parkSi orujrediani mtvris marcvlis CamoyalibebisaTvis degenirebulni arian. mikrosporangiumis yvelaze Sida nawili sporogenul kompleqts ukavia, romelic ganviTarebis adreul stadiaze warmodgenilia arqesporiumis pirveladi ujredebis saxiT, ganicdian ramodenime mitozur gayofas da mtvris marcvlebis deda ujredebad gardaiqmnebian, romelTa birTvebi 2 an 3 birTvaks Seicavs. mtvris marcvlis deda ujredebSi meiozi normalurad mimdinareobs. mikrosporebis tetrada simultanuri tipis mixedviT formirdeba, romelSic ujredebi tetraedrulad aris ganlagebuli. vazis orujredian mtvris marcvalSi vegetatiuri birTvi ovaluri formisaa da dnm-ze sust reqcias iZleva. generatiuli birTvi linzuri formisaa da fiolgen-dadebiTi reaqciiT xasiaTdeba. sperma _ ujredebi ovaluria, odnav wagrZelebuli da citoplazmis mniSvnelovani SemcvelobiT xasiaTdebian. Tavkveris magaliTze, funqcionalurad mdedrobiT vazis jiSebSi mikrosporogenezis procesis sawyisi stadiebi normalurad mimdinareobs. reduqciuli dayofis I metafazaSi 19 bivalenti yalibdeba. gadaxrebi aRiniSneba mtvris marcvlis ganviTarebis procesSi erTbirTvian mikrosporas formirebis Semdeg. miuxedavad imisa, rom degeneraciis procesebi erTbirTvian stadiaze iwyeba, umetesad orbirTviani mikrosporebi yalibdeba, ris Semdegac mimdinareobs birTvebis daSla. pirvel rigSi degenerirdeba generatiuli birTvi, xolo Semdeg iSleba citoplazma. zogjer vegetatiuri da generatiuli birTvebis degeneracia erTdroulad mimdinareobs. amasTan, mtvris marcvlebis Camoyalibebis dros ekzinaSi fore-

12

bi ar viTardeba da yvavilobis dasawyisisaTvis samtvre parkebi sterilur mtvers Seicaven (, 1971). vazis jiSebSi mtevnis wvrilmarcvlianobisa da parTenokarpiisadmi midrekilebis mizezebi momwifebul samtvre parkebSi xSirad morfologiurad araerTgvarovani mtvris marcvlebis Camoyalibeba, an Canasaxis parkis ganuviTareblobaa, rac xels uSlis ganayofierebisa da Sesabamisad, Teslis warmoSobis process. poliploidur mcenareebSi, maT Soris diploidurtetraploidur klonSi, citoembriologiuri kvlevis ZiriTadi etapebi diploiduri jiSebis msgavsad mimdinareobs (- 1976; , , , 1982; vaSakiZe, gurasaSvili, 2000), Tumca, mikrosporogenezis procesSi mikrosporaTa deda ujredebSi SeiniSneba zogierTi saxis darRvevebi. kerZod: I profazaSi ujredebis birTvebs Soris qromatiduli migracia-citomiqsisi; I metafazaSi bivalentebis nacvlad qromosomTa univalentebisa da trivalentebis Camoyalibeba; I anafazaSi qromosomebis polusebisaken araTanabari ganawileba, qromosomebis gasvla aqromatuli TiTistaras farglebidan da Sesabamisad, morfologiurad araerTgvarovani mtvris marcvlebis Camoyalibeba. sufris gorulas diploidur-tetraploidur klonSi, diploiduri da poliploiduri jiSebisa da formebis msgavsad, daculia meiozis procesisaTvis damaxasiaTebeli ZiriTadi etapebi (vaSakiZe, gurasaSvili, 2000; vaSakiZe, 2006) mtvris marcvlis Camoyalibeba iwyeba deda ujredebidan mikrosporocitebidan, romlebic I dayofisas gadian I profazas, xdeba qromosomebis konugacia da qiazmopaTia, mimdinareobs homologiuri qromosomebis calkeuli ubnebis gacvla. I profazis bolos qromosomebi maqsimalurad mokldebian da advilad iTvleba qromosomuli wyvilebi.
13

sur. 1. meiozuri dayofis I metafaza (2n=38)

ur. sur . 2. II metafaza (n=19)

sur. 3. metafazuri darRvevebi.

ganlagesur. 4. marTobulad ganlage buli metafazuri firfitebi

anafaza, ormagi sur. 5. I anafaza , or magi qromosomuli xidi

anafaza, sur. 6. I anafaza , sampolusiani anafaza, qromosomuli xidebi

14

I metafazaSi birTvis garsi iSleba da qromosomebi lagdebian metafazur firfitaze sur. 1. anafazaSi TiTistaras meqanizmebiT homologiuri qromosomebi scildebian erTmaneTs da polusisaken miemarTebian. anafazur aberaciaTa sixSire jiS gorulaSi 8.21.5%, xolo klonSi 12.41.4%-ia (sur. 5, 6, 7, 8), ZiriTadad warmodgenilia CamorCenili qromosomebis, qromosomuli xidebisa da polusebze araTanabrad ganawilebuli qromosomebis saxiT. klonSi aRiniSneba sampolusiani anafazebi, sadac qromosomebi samive polusis mimarT TiTqmis Tanabari raodenobiT miemarTebian.

I1 anafaza. sur. 7. I 1 anafaza . orpolusiani anafaza. CamorCenili qromosoma.

sur. 8. asinqronuli anafaza

I telofazaSi xdeba sxvadsxva polusze myofi qromosomebis birTvuli garsiT Semofargvla, citoplazma ar iyofa da miiReba orbirTviani diada. orive genotipSi normaluri diadebis gverdiT gvxvdeba mikronukleusiani diadebi da triadebi. diadebi mikronukleusiT miiReba qromosomuli CamorCenis Sedegad. ori an sami CamorCenili qromosomis garSemo yalibdeba birTvuli garsi da gadaiqceva mikronukleusad. triadebi miiReba I anafazaSi, sampolusianobis Sedegad da umetesad gvxdeba klonSi. mikronukleusiani diadebisa da triadebis wili klonSi, jiSTan SedarebiT, metia.
15

diada arsebobs mcire xnis manZilze, ris Semdegac iwyeba II ekvaciuri dayofa, romelic mimdinareobs Cveulebrivi mitozis msgavsad, mxolod profazis gareSe. meiozis meore ekvaciuri dayofisas ar xdeba qromosomebis gaormageba da mimdinareobs qromosomTa diploiduri nakrebis (2n=38) ganaxevreba, ris Sedegadac miiReba qromosomTa haploiduri nakrebi (n=19) sur. 2. II metafazaSi qromosomaTa firfitebi ganlagdeba erTmaneTis mimarT marTobuli kuTxiT. normaluri ori metafazuri firfitiani ujredebis gverdiT gvxvdeba rogorc ori normaluri da erTi patara metafazuri firfitiani ujredebi (miiReba mikronukleusiani diadebisagan), ise Tanabari zomis sami metafazuri firfitis mqone ujredebi (miiReba triadebisagan) sur. 3, 4. II anafazaSi xdeba qromosomTa polusebze ganlageba. diadebSi qromosomuli nakrebebi II metafazidan II anafazaSi araerTdroulad gadadian. xSirad gvxvdeba ujredebi, romlebSic diadis erTi qromosomuli nakrebi II anafazaSia, xolo II qromosomuli nakrebi II metafazaSia. II anafazaSi aberaciaTa sixSire jiS gorulaSi 9.52.6%-ia, xolo klonis genotipSi 20.53.2%. II anafazaSi klonSi statistikurad maRali (p<0.01) sizustiT aberaciebis sixSire orjer metia jiSis analogiur maCveneblebTan SedarebiT. darRvevebi gorulaSi umetesad warmodgenilia CamorCenili qromosomebis saxiT, xolo klonSi am ukanasknelis garda mravalpolusiani anafazebic gvxvdeba (sur. 7, 8). II telofazis Semdgom warmoiqmneba oTxi cxrametqromosomiani haploiduri birTvi, romlebic erTmaneTisgan gansxvavdebian vazisaTvis damaxasiaTebeli simultanuri tipis mixedviT, erTdrouli citokineziT, (, , , , 1977), ris Sedegadac miiReba
16

mikrosporas oTxi ujredi, romlebic tetraeduladaa ganlagebuli, Tumc klonis genotipSi aRiniSneba sakmao raodenobiT pentadebis, heqsadebis da mikronukleusiani tetradebis arseboba (sur.9).

sur. 9. mikrosporas pentadebi

dasawyisSi tetrada saerTo dediseuli garsiT aris Semofargluli, magram ganviTarebis Semdgom periodSi igi inviTarebs sakuTar celulozis garss, ris Semdegac dediseuli garsi rezorbirdeba, tapetumic iSleba da ujredebi Sordebian erTmaneTs. kargad ganviTarebul mikrosporaSi gamokveTilad Cans birTvi da protoplazma. mikrosporebis ganviTareba swrafad xdeba. misi birTvi miemarTeba garsisaken da Cveulebrivi mitozis Sedegad iyofa or nawilad, romelTagan erTi ifargleba citoplazmis Txeli garsiT da gardaiqmneba generatiul birTvad, xolo meore yalibdeba ujredis vegetatiur birTvad. amasTan, mikrosporas kedeli sqeldeba da gadaiqceva ekzinad, romelzedac yalibdeba sami fora. kedlis qveda ujredebisagan warmoiqmneba garsis meore, SigniTa Sre intina. mikrosporogenezis Sedegad miiRe-

17

ba orgarsiani da samforiani momwifebuli mtvris marcvlebi. jiS gorulaSi da mis klon #21-Si mikrosporogenezi mTavrdeba maisis bolos. meiozSi anafazur aberaciaTa sixSirisa da speqtris, Secvlili diadebisa da tetradebis gaanalizebiT ikveTeba, rom gorulas jiSsa da klonSi mtvris marcvlis momwifeba sxvadasxva xarisxiT mimdinareobs, rasac adasturebs maTi mtvris ganayofierebis xarisxi, romelic gorulas genotipisaTvis 86.41.0%, xolo klonisaTvis 70.21.1%-ia.

tetradebi, mikrosporas tetradebi , gadideba 90X10

18

usaxelouri

aleqsandrouli

wulukiZis TeTra
19

ojaleSi

II. II. mtvris marcvlis morfomorfo-fiziologiuri Tvisebebi

warmoadgenen jiSis erT-erT mTavar biologiur niSnebs, romelTa momwifebis xarisxzea damokidebuli yvavilis normaluri damtverva-ganayofierebisa da nayofwarmoqmnis procesebi, uxvi da xarisxiani mosavlis uzrunvelyofa. kulturuli vazis (Vitis vinifera ssp. sativa D.C.) mtvris marcvali jiSebis mixedviT gansxvavebulia, xasiaTdeba ampelografiulad Rirebuli xarisxobrivi da raodenobrivi niSnebiT. mniSvnelovani adgili ukavia jiSisa da formis identifikaciisa da Sefasebis saqmeSi (vaSakiZe, 2006) 1. palinomorfologia palinologia biologiis erT-erTi mniSvlenovani dargia, romlis monacemebi da meTodebi warmatebiT SeiZleba iqnes gamoyenebuli botanikis, sistematikis, filogeniis, paleobotanikasa da sxva dargebSi mravali amocanis gadasawyvetad. mkvlevarebis (yvavaZe, 2004; , , 1976; , 1966 da sxv.) da sakuTari (vaSakiZe, 2006; , 2008) eqsperimentuli gamokvlevebiT kulturuli vazis orsqesiani jiSebis normaluri mtvris marcvlebi xorblis marcvlis formis da mcire zomisaa (sur. 10, 11), romlebic sakveb areze moxvedris an dasvelebis SemTxvevaSi dasawyisSi samkuTxedis, xolo Semdgom ki sferul formas Rebuloben, xolo rac Seexeba funqcionalurad mdedrobiT vazis jiSebis mtvris marcvlebs, maT wamaxvilebuli forma aqvT da dasvelebis SemTxvevaSi erTgvarovan sferul formas Rebuloben (sur. 12).
20

sur. 10 rqawiTelis mtvris marcvali haermSral mdgomareobaSi gad. 10 20

sur. 11 goruli mwvanis mtvris marcvali haermSral mdgomareobaSi gad. 10 20

sur. 12 funqcionalurad mdedrobiTi vazis jiSis Tavkveris mtvris marcvlebi gad. 10 20

mtvris marcvlis parametrebi parametrebi - Cveni eqsperimentuli

gamokvlevebiT, vazis qarTuli genotipebis mtvris marcvlis sigrZe haermSral mgomareobaSi gansxvavebuli _ 25.00.9-38.80.3 mkm-mdea. maqsimaluri (38.80.3 mkm) sigrZiT xasiaTdeba saferavi budeSuriseburis, xolo minimaluriT (21.10.9 mkm) colikouris mtvris marcvlebi, danarCen jiSebs maT Soris Sualeduri adgili ukaviaT. mtvris marcvlis sigane 14.70.2-25.30.4 mkm-mde meryeobs. maqsimaluri siganiT xasiaTdeba jiSebis: saferavis, rqawiTelisa da xixvis, gansakuTrebulad ki, saferavis kloni #359-is (25.30.4 mkm), xolo minimaluriT (14.70.2) saferavis mtvris marcvlebi. diametris maCvenebeli 18.10.3-28.80.3 mkm-is farglebSia. maqsimaluri maCvenebe21

li (28.80.3mkm) axasiaTebs jiS bueras, xolo minimaluri (18.10.3 mkm) saferavis mtvris marcvlebs.Yyvela klons jiSTan SedarebiT gazrdili parametrebi axasiaTebs (cxr. 1). cxrilSi warmodgenili vazis qarTuli genotipebis mtvris marcvlis parametrebis maCveneblebi, fenotipuri niSnebis cvalebadobaSi, genotipis wamyvan rolze miuTiTeben.

cxrili 1 vazis qarTuli jiSebisa da maTi klonebis mtvris marcvlis marcvlis parametrebi
sigrZe sigane diametri

jiSi
XSx (mkm) V (%) XSx V (%) (mkm) XSx (mkm) V (%)

10 11

1 aladasturi 2 aleqsandrouli 3 aspinZura

30.20.2 1.5 5.1

6.00.3 1.8 11.2 23.60.3 2.4 10.0

28.40.7 5.0 16.0 16.80.4 3.0 17.6 22.60.4 2.5 11.3 31.60.7 5.1 16.3 17.303 2.5 14.3 27.50.4 3.3 11.1

4 asureTuli Savi 29.00.5 3.7 12.7 22.30.4 2.8 12.5 25.40.44 3.1 12.3 5 bazaleTuri 6 budeSuri wiTeli 31.60.7 4.2 13.3 20.50.5 3.3 16.3 24.30.7 4.8 12.8 30.30.3 1.9 6.1 16.00.3 1.8 11.2 19.10.3 1.9 9.8

7 buera 8 gorula 9 gorula kloni s#21

31.51.0 7.2 20.0 20.40..3 2.1 10.4 28.80,3 1.8 6.3 30.10.4 4.4 14.6 17.10.2 1.7 9.9 26.00.5 3.3 12.6

33.90.4 4.4 14.6 7.10.2 2.5 13.2 31.00.3 2.8 9.0

22

10 11

10 goruli mwvane 11 grZelmtevana 12 Tavkveri 13 Tamaris yurZeni 14 kaxuri mcvivani 15 kaxuri mwvane 16 kraxuna 17 kraxuna #1170 kloni

30.50.5 3.9 12.7 20.40.5 3.3 16.3 24.10.5 3.6 14.7 32.50.5 3.5 10.7 19.80.3 2.1 10.7 28.40.3 2.3 8.1 7.80.5 3.3 11.9 21.80.4 3.1 14.0 21.60.3 1.8 8.7 36.60.9 6.1 16.8 21.00.3 2.2 10.6 29.30/4 2.6 8.7 31.70.3 5.7 17.9 16.50.2 1.9 11.7 25.50.3 3.4 13.2 33.00.3 1.9 5.8 18.10.3 2.0 10.9 25.90.6 4.0 15.6 30.40.4 4.1 13.6 18.70.4 2.8 14.9 24.80.3 2.5 12.9 33.20.3 1.9 5.7 7.20.3 1.8 10.4 30.50.3 2.1 7.0

18 kaWiWi 19 kumsi TeTri 20 kumsi wiTeli 21 mesxuri mwvane 22 mujureTuli 23 orbeluri ojaleSi

32.50.2 1.5 4.7 16.30.3 2.0 12.2 23.40.3 1.9 8.3 33.00.5 3.2 9.6 19.20.4 3.1 15.9 26.60.4 2.5 9.4 32.00.6 4.2 13.3 21.90.4 2.6 12.2 25.80.4 2.9 11.3 33.30.4 2.7 8.1 18.40.3 2.4 12.8 27.80.3 2.1 7.5 31.50.7 5.1 16.1 16.80.4 2.6 15.8 23.40.4 2.8 11.9 35.00.3 2.1 5.9 17.20.2 1.4 8.4 27.70.3 2.5 8.9 30.20.3 2.3 7.5 14.80.2 1.6 13.7 22.50.3 1.9 8.6

24 ojaleSi

25 ocxanuri safere 27.006 3.9 14.5 17.50.3 2.3 13.4 24.40.4 3.0 14.4 26 ocxanuri safere kloni #1200 32.00.4 3.3 8.3 19.50.4 2.4 10.0 29.3 0.7 2.2 9.3 32.20.6 4.1 12.7 17.10.3 2.2 13.0 22.50.3 2.3 10.0 35.90.8 5.6 15.5 21.40.2 1.7 7.9 28.60.4 2.5 8.9 34.50.5 3.5 10.2 20.30.3 2.2 10.6 25.70.5 3.8 14.6 30.44.2 3.0 9.8 23.00.3 2.3 10.2 23.80.3 2.1 8.7

27 rqawiTeli 28 rqawiTeli #48 29 rqawiTeli wiTeli

30 safena

23

10 11

31 saferavi 32 saferavi #359 33 saferavi budeSurisebri

30.60.3 2.2 7.1 14.70.2 1.7 11.3 18.10.3 1.9 10.6 33.70.4 3.0 8.9 25.30.4 2.7 10.5 27.70.3 2.3 8.3 38.80.3 1.8 5.3 15.40.3 2.2 13.8 23.70.3 2.2 9.3 31.90.3 2.1 6.6 15.80.3 1.8 11.1 27.90.4 2.5 9.1 31.80.3 2.2 6.8 16.80.3 1.9 11.6 24.40.3 2.2 8.8 35.90.5 3.5 9.7 20.50.3 2.3 11.1 28.10.4 2.6 9.1 31.00.5 3.3 10.6 16.50.4 3.1 18.9 21.50.4 2.6 12.2 32.00.3 1.8 5.5 16.30.3 2.0 11.8 23.00.3 2.0 8.9 26.20.6 4.1 15.4 18.10.3 2.4 13.4 22.10.3 2.5 11.1

34 sawuravi 35 usaxelouri 36 yornisTvala 37 Cinuri 38 Cxaveri 39 cicqa

40 cicqa kloni#1298 33.10.4 2.5 7.7 16.90.3 2.4 14.0 23.20.3 2.1 9.2 41 cicqa klonigrZelmtevana 33.30.2 1.7 5.2 19.403 1.8 9.0 23.30.3 2.2 9.4

42 colikouri 43 colikouri kloni #1093 cxvedianis TeTra

25.00.9 4.6 18.3 16.60.4 2.7 16.4 20.80.4 2.6 12.3 32.8 0.3 2.3 6.9 16.8 0.3 1.9 11.3 25.8 0.4 3.0 116

44

33.40..3 2.4 7.3 19.30.2 1.6 8.3 26.10.5 3.3 12.5 32.90.3 2.2 6.8 15.90.3 1.6 10.8 20.00.3 1.8 9.1

45 Wvitiluri

46 wulukiZis TeTra 32.00.4 2.7 8.5 16.80.4 2.3 13.9 22.70.3 1.9 8.4 47 xarisTvala TeTri 32.80.4 2.8 8.5 19.80.3 2.3 11.8 28,00,3 2,3 8.3 34.40.5 3.3 9.5 18.00.3 2.4 13.3 26.70.6 4.1 15.2

48 xarisTvala mesxuri 50 xixvi

49 xarisTvala Savi 34.30.6 3.6 11.4 24.90.5 3.4 13.6 27.30.3 2.4 8.7 31.30.4 2.9 9.2 16.30.3 2.1 12.8 25.70,5 3.6 13.9 24

51 xixvi #430 52 jani 53 ZaRliarWama

32.705 3.8 11.8 21.50.3 2.5 11.5 27.30.4 3.2 11.6 31.70.3 2.4 7.5 15.80.3 1.8 11.4 24.50.3 1.9 7.9 34.70.5 3.6 10.2 21.10.3 2.3 10.9 27.40.4 2,9 10.7

cxrilSi motanili eqsperimentuli monacemebis erTfaqtoriani dispersiuli analizis Sedegad (p<0.01 sizustiT), ikveTeba vazis jiSis warmoSobis adgilis gavlena mtvris marcvlis kriteriumebis cvalebadobaze, romelic sigrZis SemTxvevaSi 43.44%-s, siganis 46.07%-s, xolo diametris SemTxvevaSi 51.19%-s Seadgens. mtvris marcvlis kriteriumebis kvleviT masalebSi gansxvaveba aRiniSneba regionebis mixedviTac: mtvris marcvlis sigrZe 16.84_20.72 mkm, sigane 9,71_10.95 mkm, xolo diametri 13.40_18.11 mkm-ia (diagr. 1). cnobilia, rom kulturuli vazis jiSebis raodenobrivi niSnebi, fenotipuri cvalebadobis xarisxis mixedviT, didi mudmivobis (Cv<10%), mcired cvalebad (10%<Cv<20%) da Zlier cvalebad (Cv>20%) niSnebad iyofa (,1983).

25

vazis qarTuli genotipebis mtvris marcvlis parametrebi regionebis mixedviT

25 20 15 parametris saSualo 10 maCvenebeli 5 X 0
sigrZe sigan e di ametri

am mimarTebiT Catarebul kvleviTi Sedegebidan gamomdinare vazis qarTuli genotipebis mtvris marcvlis parametrebi raodenobrivi niSnebis konstanturobisa da cvalebadobis maCveneblebis mixedviT didi mudmivobisa (mdgrad) da mcired cvalebad niSanTa jgufs miekuTvnebian. Cvens mier gamokvleul 53 jiSidan da klonidan jiSebi: orbeluri ojaleSi, ocxanuri safere-kloni da cicqa-kloni-grZelmtevana sami niSniT, jiSebi: kaWiWi, jani, cxvedianis TeTra, wiTeli budeSuri, kraxuna-kloni, ojaleSi, safena, rqawiTeli #48, saferavikloni, saferavi budeSuriseburi, sawuravi, usaxelouri, yornisTvala, Cxaveri, cicqakloni, Wvitiluri, wulukiZis TeTra, xarisTvala TeTri _ ori niSniT da jiSebi: aladasturi, saferavi, cicqakloni, gorula, colikouri, xarisTvala TeTri, xarisTvala mesxuri, xixvi, buera, colikourikloni, kumsi TeTri, kraxuna-kloni, mesxuri mwvane, xarisTvala Savi _ erTi niSniT didi mudmivobis,
26

r ia r aW l a eC x sa um me i gr el o aW ar af a xa z eT i
regioni

i l T im er

xe

eT

ka

qa

gu

xolo danarCeni jiSebi mcired cvalebad fenotipuri niSanebis matarebel jiSebis jgufs miekuTvnebian, rac amtkicebs mkvlevarTa azrs generatiuli organoebis raodenobrivi niSnebis mcired cvalebadobis Sesaxeb da miuTiTebs aRniSnuli kriteriumebis jiSis indentifikaciisaTvis gamoyenebis SesaZleblobaze. mtvris marcvlis forianoba _ fori aris mtvris marcvlis gareTa garsze ekzinaze ganlagebuli viwro xvreli, riTac mtvris marcvali anxorcielebs kavSirs garemosTan. ekzinas struqturidan gamomdinare maTi sidide, forma da raodenoba farulTeslovan mcenareTa saxeobebSi gansxvavebulia. sqeli, ekliani ekzinis mqone saxeobebisaTvis damaxasiaTebelia mravalforianoba, romelTa ricxvi 3-dan 20-mdea. arsebuli literaturuli masalebisa (, , , , 1977; -, 1976; , 1971 da sxv.) da Cveni gamokvlevebiT (vaSakiZe, 2006; vaSakiZe, gurasaSvili, 2000; vaSakiZe, CxartiSvili, maRraZe, gurasaSvili, mdinaraZe, 2007; vaSakiZe 2008; vaSakiZe, maRraZe, 2009) vazis normaluri mtvris marcvali ZiriTadad samforiania, magram mikrosporogenezis procesSi arsebuli darRvevebis gamo gvxvdeba uforo mtvris marcvlebic, romelTa Sefardeba jiSebis mixedviT gansxvavebulia.

sur. 13 samforiani mtvris marcvali

27

cxrili 2 vazis qarTuli jiSebis mtvris marcvlis forianoba

mtvris marcvlebis saerTo raodenoba oTxforiani samforiani

jiSi

n 4

XSx 5

n XSx 6 7

n 8

XSx 9

3 180 111 126 121 108 94

1 aladasturi 2 aleqsandrouli 3 aspinZura 5 bazaleTuri budeSuri 6 wiTeli 7 buera 8 gorula gorula kloni #21 10 goruli mwvane 9 11 grZelmtevana 12 Tavkveri 13 Tamaris yurZeni 14 kaxuri mcvivani 15 kaxuri mwvane 16 kaWiWi 17 kraxuna kraxuna- kloni 18 #1170

151 83.92.7 75 67.64.4 71 5634.4 5 3 3.41.5 2.51.4

10 forebi 29 16.12.7 mkveTradaa gamoxatuli 36 32.44.4 56 44.54.4 120 96.609 118 97.51.4 25 24.34.2 12 12.73.4

4 asureTuli Savi 125

78 75.74.2 82 87.23.4

295 245 83.12.2 4 237 214 67.12.5 161 135 83.92.9 129 121 93.82.1 272 1 0.40.4

dafiqsirda oTxforiani 350.6 46 15.62.1 mtvris marcvali 23 32.92.5 26 16.12.9 8 6.22.1

dafiqsirda samforiani

271 99.60.4 mtvris marcvlis arseboba

110 69 62.74.6 115 93 80.93.7 110 104 92 83.64.2 86 82.73.7 175 105 98.32.6

41 37.34.6 22 19.13.7 18 16.44.2 10 0.92.6 18 17.33.7 10 8.82.7

113 103 91.22.7

28

SeniSvna

uforo

3 129 154 133 113 118 156

8 9 4

9 6.92.2 3.01.5

10

19 kumsi TeTri 20 kumsi wiTeli 21 mesxuri mwvane 22 mujureTuli orbeluri 23 ojaleSi 24 ojaleSi ocxanuri 25 safere ocxanuri safe26 re kloni #1200 27 rqawiTeli 28 rqawiTeli #48 rqawiTeli 29 wiTeli 30 rcxili 31 safena 32 saferavi 33 saferavi budeSurisebri

120 93.02.2 104 67.53,8 129 97.01.5 79 70.04.3 104 88.13.0 130 84.42.9

50 2.53.8 24 21.24.3 14 11.83.0 26 15.62.9 51 16.82.2 57 19.72.3 23 22.54.1 33 28.94.2 35 32.14.5 92 91.02.7 111 96.51.7 forebi 16 15.83.4 mkveTradaa gamoxatuli 16 14.13.2 dafiqsirda oTxforiani mtvris marcvali

298 247 83.22.3 290 102 114 109 100 115 114 113 154 119 92 151 144 130 106 125 126 118 111 233 80.32.3 79 77.54.1 81 71.14.2 74 67.94.5 8 4 8.02.7 3.41.7

96 84.23.4 97 85.83.2

34 sawuravi 35 usaxelouri 36 yornisTvala 37 Cinuri 38 Cxaveri 39 cicqa cicqa kloni 40 #31298 cicqa kloni 41 grZelmtevana 42 colikouri colikouri 43 kloni #1093 44 cxvedianis TeT

135 87.72.6 1 0.650.6 18 11.72.5 102 85.73.2 56 60.95.1 131 86.82.8 119 82.63.2 113 86.93.0 68 63.04.7 99 79.23.8 105 83.33.3 76 64.44.4 87 78.43.9 17 14.23.2 36 39.15.1 20 13.22.8

forebi 25 17.43.2 mkveTradaa gamoxatuli 17 13.13.0 38 37.04.7 26 20.83.8 21 16.73.3 42 35.04.4 24 21.63.9

29

1 2 45 Wvitiluri wulukiZis 46 TeTra xarisTvala 47 TeTri xarisTvala 48 mesxuri xarisTvala 49 Savi 50 xixvi 51 xixvi #430 52 jani 53 ZaRliarWama

3 139 124 114 110 126 115 108 118 108

4 5 6 107 77.03.6 108 87.13.0 85 74.64.1 80 72.74.2 68 54.04.4 91 70 108 74 7913.8 64.84.6 91.52.6 68.54.5

8 9 32 23.03.6 16 12.93.0 29 25.44.1 30 27.34.2 58 46.04.4 24 38 10 34 20.93.8 35.24.6 8.42.6 31.54.5

10

vazis qarTul genotipebSi samforiani mtvris marcvlebis raodenoba absoluturobisaken miiswrafvis da igi 67.64.4 _ 98.32.6%-is farglebSia. samforiani mtvris marcvlebis maqsimaluri (98.32.6%) SemcvelobiT xasiaTdeba jiSi kaWiWi da yvelaze naklebiT (54,64.4%) jiSi xarisTvala Savi, danarCen jiSebs maT Soris Sualeduri adgili ukaviaT. gamonaklisis saxiT, jiS gorulaSi aRiniSna oTxforiani mtvris marcvlebis arsebobis erTeuli SemTxvevebi. samforiani mtvris marcvlebis umniSvnelo raodenobaa (0.41.3 _ 2.51.4%) funqcionalurad mdedrobiT vazis jiSebSi: TavkverSi, safenaSi, asureTul Savsa da bazaleTurSi (cxr. 2). 1. mtvris marcvlis sic sicocxl ocxlis xlisunarianoba: isunarianoba:

fertiloba - aris mtvris ganayofierebis unari, romelic uzrunvelyofs mtvris marcvlis gaRivebis, samtvre milis zrdis, generatiuli ujredebis dayofisa da spermiebis warmoqmnis procesebis normalur msvlelobas. igi ganisazRvreba garegnulad (zoma, forma, SeRebva) da mtvris Sinagani agebulebiT, dingze da xel30

ovnur sakveb areze gaRivebiT. dakavSirebulia mtvris formirebasa da ganviTarebasTan, romelTa darRveva iwvevs mtvris marcvlis sterilurobas (ColoyaSvili,1939; vaSakiZe, 2006; , 1982; 2008; , , , 2003; , , , , 1977; -, 1976; , 1971 da sxvani). fertiluri mtveri formirdeba orsqesiani da funqcionalurad mamrobiTi yvavilis mqone mcenareebSi. aqvT wagrZelebuli xorblis forma, Sedgeba ori _ vegetatiuri da generatiuli birTvisagan, dafarulia ormagi garsiT (intina, ekzina), sami foriT, romlebic mdebareoben or paralelur napralze da uzrunvelyofen normalur ganayofierebas, xolo organizebuli steriluri mtveri, formirdeba funqcionalurad mdedrobiT yvavilebSi, aqvT wakveTili fialis forma, wawvetebuli, aweuli boloebiT. Sedgeba ujredebisagan mkvdari birTvebiTa da protoplazmiT, uforo garsisagan da napralisagan. igi ar uzrunvelyofs normalur ganayofierebas, mxolod SeuZlia mastimulirebeli gavlenis moxdena, uTeslo wvrili marcvlebis ganviTarebaze. kulturuli vazis jiSebi mtvris marcvlis fertilurobis mixedviT sam: I. dabal (<30%), saSualo (30-50%) da maRal fertilur (>50%) klasad jgufdeba (, 1983; , , , , 1977). vazis qarTuli genotipebis mtvris marcvlis Seswavlis Sedegad gamoikveTa, rom orsqesian jiSebs, rogorc wesi maRal fertiluri mtveri axasiaTebT (sur. 13,14,15, 16), romelTa raodenoba 55.22.5-dan 98.90.5%-mde cvalebadobs da absoluturobisaken miiswrafis. fertiluri mtvris maqsimaluri raodenobiT 98.90.5% xasiaTdeba jiSi saferavi, masTan SedarebiT mniSvnelovnad dabali, magram klasifikaciiT maRali 55.22.5% rcxili, xolo klonebi, yvela SemTxvevaSi, jiSTan SedarebiT ramdenadme dabali fertilobiT xasiaTdebian.
31

rac Seexeba funqcionalurad mdedrobiT jiSebs: Tavkvers, asureTul Savs, safenasa da bazaleTurs, aq fertiluri mtvris umniSvnelo (0.60.4%) raodenoba aRiniSneba, darCenili mtvris umravlesoba organizebuli steriluria (sur. 17), romelsac ar SeuZlia normaluri ganayofiereba, magram mas SeuZlia moaxdinos mastimulirebeli gavlena wvrili, uTeslo marcvlebis ganviTarebaze (cxr. 3).

sur. 14 rqawiTelis mtvris marcvlis fertiloba

sur. 15 Cinuris mtvris marcvlis fertiloba

sur. 16 goruli mwvanis mtvris marcvlis fertiloba

sur. 17 saferavis mtvris marcvlis fertiloba

sur. 18 Tavkveris mtvris marcvlis fertiloba

32

cxrili 3 vazis qarTuli jiSebis fertiloba gasinjuli fertiluri mtvris mtvris marcvlis jiSi marcvlebis n PSp % raodenoba N
2 aladasturi aleqsandrouli aspinZura asureTuli Savi bazaleTuri budeSuri wiTeli buera gorula gorula kloni #21 goruli mwvane grZelmtevana Tavkveri Tamaris yurZeni kaxuri mcvivani kaxuri mwvane kraxuna kraxuna kloni #1170 kaWiWi kumsi TeTri kumsi wiTeli mesxuri mwvane mujureTuli orbeluri ojaleSi ojaleSi ocxanuri safere 33 3 622 292 331 522 556 527 335 1546 1739 402 378 670 346 343 503 318 113 538 388 335 400 291 567 667 492 4 583 202 178 49 69 506 316 1336 1219 337 362 56 238 218 365 277 103 521 374 230 385 207 517 635 452 5 93.71.0 69.22.7 53.82.7 9.41.3 12.41.4 96.00.9 94.31.3 86.41.0 70.21.4 83.81.8 95.81.0 8.41.1 68.82.5 63.62.6 72.62.0 87.11.9 91.22.7 96.80.1 94.40.9 68.72.5 96.30.9 71.12.7 91.21.2 95.20.8 92.01.2

1 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25

1 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54

2 ocxanuri safere kloni #1200 rqawiTeli rqawiTeli #48 rqawiTeli wiTeli rcxili safena saferavi saferavi #349 saferavi budeSuris sawuravi usaxelouri yornisTvala Cinuri Cxaveri cicqa cicqa kloni #1298 cicqa kloni grZelmtevana colikouri colokouri kloni #1093 cxvedianis TeTra Wvitiluri wulukiZis TeTra xarisTvala TeTri xarisTvala mesxuri xarisTvala Savi xixvi xixvi #430 jani ZaRliarWama

3 400 609 442 343 321 457 371 518 584 570 609 377 357 653 284 275 293 340 601 305 591 402 442 397 425 676 404 569 356

4 391 539 311 219 135 3 367 491 523 532 546 262 286 568 253 241 257 322 501 272 531 379 332 254 253 547 227 533 246

5 97.00.7 87.71.3 70.42.2 83.82.6 42.12.8 0.60.4 98.90.5 94.81.0 89.61.3 93.31.0 89.71.2 69.52.4 81.02.1 87.01.3 89.11.9 87.82.0 87.60.4 94.71.2 83.41.5 89.21.8 89.81.2 94.31.2 75.12.1 64.02.4 59.5 2.4 80.91.5 55.22.5 93.71.0 69.12.4

34

mtvris marcvlis gaRiveba xelovnur sakveb areze

mtvris marcvlis unars _ gaRivdes xelovnur sakveb areze, udidesi Teoriuli da praqtikuli mniSvneloba aqvs. xelovnur sakveb areze gaRivebis procesSi SesaZlebelia imis dadgena, Tu rogor mimdinareobs mtvris marcvlis gaRiveba, samtvre milis Camoyalibeba, romelic erT-erTi umniSvnelovanesi faqtoria ganayofierebis procesis normaluri msvlelobisaTvis. kvlevebs mtvris milis xelovnur sakveb areebze gaRivebis Sesaxeb saukunovani istoria aqvs, magram man gansakuTrebuli mniSvneloba SeiZina mecnieruli seleqciis Camoyalibebis Semdeg, rogorc erT-erTma aucilebelma pirobaTaganma axali jiSebis misaRebad sawyisi masalis SerCevisa da hibridizaciis procesis miznobrivad warmarTvis saqmeSi. mtvris daTesva xelovnur sakveb areze saSualebas iZleva SemWidrovebul vadaSi iqnas gamokvleuli mtvris marcvlis cxovelunarianoba da ganayofierebis xarisxi (, , , 1982), Seswavlili iqnas spermiebis Camoyalibebisa da ganviTarebis procesi, dadgenili iqnes samtvre milis zrdis sigrZe, romelic sxvadasxva sakveb areze SeiZleba gansxvavebuli iyos. sakveb areze generatiuli ujredi iyofa Cqara, spermiebic ufro adre yalibdeba, vidre Cveulebriv bunebriv pirobebSi dingze, magram samtvre milis zrdis siCqare da sigrZe naklebia, Cveulebriv bunebriv pirobebTan SedarebiT. maT Soris arsebobs pirdapiri Sefardeba 1:5, rac SeiZleba imiT aixsnas, rom samtvre milis zrdis sigrZe butkos svetSi stimulirdeba gansakuTrebuli nivTierebebiT, romlebic imyofebian butkos svetSi da ar arian xelovnur sakveb areze. pos (Po, 1929) mixedviT mtvris marcvlis gaRivebasa da samtvre milis sigrZes Soris pirdapir proporciuli damokidebuleba
35

arsebobs. rac msxvilia mtvris marcvali, miT ufro kargad Rivdeba, Tumc iakimovis (, , , , 1977) monacemebiT saSualo zomis mtvris marcvali ufro intensiurad Rivdeba da grZel samtvre mils inviTarebs. axladaRebuli mtvris marcvali xasiaTdeba kargi gaRivebis unariT. mis gaRivebaze gavlenas axdens temperatura, tenianoba da xelovnuri sakvebi aris Sedgeniloba. xelovnur sakveb areze mtvris marcvlis gaRivebisaTvis optimaluri temperatura 20-300C da tenianoba 70%-ia. xelovnur sakveb areebze jiSebis moTxovna gansxvavebulia. rogorc Catarebulma eqsperimentulma gamokvlevebma aCvena sakvlevi jiSebis mtvris marcvali xvdeba ra sakveb areebze, maSinve Rebulobs sferul formas. cota xnis Semdeg ekzinaze mdebare erT-erT foridan xdeba citoplazmis gamonazardis gamosvla, romelic swrafad izrdeba da mtvris marcvlis zomas aRwevs. citoplazmis nawili gadadis gamonazardSi da mokle droSi warmoiqmneba mtvris mili, romelic izrdeba da ramodenime xnis Semdeg iRebs saboloo formas (sur. 18). amave periodSi mTavrdeba sasqeso ujredebis spermiebis Camoyalibeba. mtvris marcvlidan foris gavliT citoplazmis gamonazardSi gadadis generatiuli birTvi. cota xnis Semdeg amave gzas gadis vegetatiuri birTvi. isini citoplazmis gamonazardSi cota xniT yovndebian da Semdeg vegetatiuri birTvi gadadis mtvris milSi, roca igi milis Suaguls miaRwevs mtvris milSi gadadis generatiuli birTvi, ramodenime xnis Semdeg isini aRweven mtvris milis bolos (sur. 19). amis Semdeg generatiuli ujredi wyvets moZraobas, izrdeba zomaSi da mitozuri dayofiT or spermiad iyofa. sabolood mtvris milis
36

sur. 19 mtvris milSi generat gadasuli genera tiuli birTvi

sur. 20 rqawiTelis mtvris marcvlis gaRiveba xelovnur sakveb areze

sur. 21 datotvili mtvris mili

sur. 22 datotvili mtvris mili

boloSi Camoyalibdeba ori sperma ujredi da vegetatiuri birTvi. gorulas klon #21-Si dafiqsirda poliploiduri po formebisaTvis damaxasiaTebeli (-, 1976) datotvili mtvris milebis arseboba (sur. 21, 22). arsebuli literaturis (, 1983) mixedviT mtvris marcvlis gaRivebis unaris mixedviT jiSebi iyofian sam jgufad: 1) dabali (0-30%), 2) saSualo (31-70%) da 3) kargi (71-100%).
37

vazis qarTuli genotipebis mtvris marcvlis cxovelunarianobis dadgenis mizniT Catarebulma mikroskopulma gamokvlevebma aCvena, rom orsqesiani jiSebis umravlesobisaTvis (rqawiTeli, kaxuri mwvane, wiTeli budeSuri, xixvi, colikouri, cicqa, Cxaveri, aladasturi, ojaleSi, wulukiZis TeTra, sawuravi, jani, kaWiWi da sxva) optimaluria saqarozis 15%-iani agarizebuli sakvebi are, Tumc zogierTi jiSi (saferavi, goruli mwvane, Cinuri, gorula, kraxuna, ocxanuri safere, Wvitiluri, aleqsandrouli, mujureTuli, orbeluri ojaleSi, usaxelouri, saferavi budeSurisebri da sxva) maqsimalur gaRivebas saqarozis 20%-iani agarizebuli sakvebi areze aRwevs. sakvlevi jiSebidan maRali gaRivebis (76-89%) unariT xasiaTdebian: rqawiTeli, saferavi, xixvi, cicqa, kraxuna; saSualoze maRaliT (33-67%): saferavi bedeSurisebri, ocxanuri safere, sawuravi, wulukiZis TeTra, usaxelouri, jani, gorula, goruli mwvane, wiTeli budeSuri, orbeluri ojaleSi, kaWiWi da sxva, xolo saSualoze dabaliT (29,1-15.0%): Cxaveri, aladasturi, ojaleSi, aleqsandrouli, mujureTuli, Wvitiluri. danarCenebs maT Soris Sualeduri adgili ukaviaT (cxr. 4). Tavkversa da bazaleTurisaTvis optimaluria saqarozis 15%-iani, xolo safenasa da asureTuli SavisaTvis glukozis 10%-iani agarizebuli sakvebi are. yvela jiSisaTvis mtvris marcvlis gaRivebisaTvis optimaluri temperatura 28-300C da tenianoba 70-75%-ia. rac Seexeba fuqcionalurad mdedrobiT vazis jiSebs, isini arc erT jgufs ar miekuTvnebian, radgan mtvris marcvlebis didi umravlesoba steriluria da mxolod umniSvnelo nawils (0.8-1.6%) Tu axasiaTebs gaRivebis unari. gaRivebuli marcvlebis yvelaze meti raodenoba (1.6%) miiReba jiS asureTul SavSi, xolo naklebi (0.4%) jiS TavkverSi. danarCen jiSebs Sualedu38

ri adgili ukaviaT. funqcionalurad mdedrobiT vazis jiSebSi fertiluri mtvris marcvlebis arseboba dadasturda organizmis donezec. arsebuli literaturis mixedviT funqcionalurad mdedrobiT vazis jiSebSi, yvavilebis izolirebis SemTxvevaSi, mtevanze parTenokarpiuli marcvlebis gverdiT normaluri ganviTarebis Tesliani marcvlebic viTardeba. am movlenas mecnierebi gansxvavebulad xsnian. rig maTgans (ColoyaSvili, 1939; ramiSvili, 1960; Sarikhani, 2007; , 1935; , 1925 da sxvani) miaCnia, rom igi yvaviledebSi orsqesiani yvavilebis arsebobiT unda iyos ganpirobebuli, xolo meoreni (Sarikhani, 2007; , 1959; , 1925 da sxvani) ki mas Teslebis apomiqtur ganviTarebas miaweren.

sur. 23 Sewepebuli marcvlebi mtvris marcvleb i

SgaRivebuli sur. 24. klonis S gaRivebuli mtvris marcvali

Cveni eqsperimentuli masalebi (vaSakiZe, 2006; Chkhartishvili, Vashakidze, Gurasashvili, Maghradze, 2006; vaSakiZe, CxartiSvili da sxvani, 2007), arcerT versias ar uaryofs, piriqiT, maT Tanaarsebobaze miuTiTebs, radgan wlebis ganmavlobaSi funqcionalurad mdedrobiTi vazis jiSebis: Tavkveris, asureTul Savis, safenas da bazaleTuris TviTdamtveril (sur. 25) yvaviledebidan
39

mtevanze arsebuli marcvlebis 1.10,1_1.40.1% normaluri ganviTarebis wipwas Seicavs (diagrama 2). morfologiurad orsqesiani yvavilebis arseboba da marcvlis gamonaskva iqna dafiqsirebuli laboratoriul pirobebSi dafesvianebul rqebze ganviTarebul yvaviledebze (sur. 27).

cxrili 4
vazis qarTuli genotipebis mtvris marcvlis
xelovnur sakveb sakveb areze gaRiveba xelovnur sakveb areze gaRiveba(%)

sakvebi are

jiSi

SeniSvna

3 89.01.8 77.01.9 78.82.1 32.91.6 61.72.4 54.72.7 71.72.0 63.22.1 89.41.8 75.72.8

4 C-15 C-20 C-15 C-15 C-15 C-20 C-20 C-20 C-20 C-15 C-15

1 rqawiTeli 2 saferavi 3 xixvi 4 kaxuri mwvane 6 Cinuri 7 goruli mwvane 8 gorula gorula kloni 9 #21 10 cicqa cicqa kloni 11 #1298

5 grZeli milebi dafiqsirda tyupi samtvre milebi Sewepebuli mtvris erTeuli SemTxvevebi grZeli milebi grZeli milebi grZeli milebi grZeli milebi grZeli milebi

5 wiTeli budeSuri 31.52.5

grZeli milebi

40

1 2 12 colikouri colikouri 13 kloni #1093 14 ocxanuri safere 15 ocxanuri safere kloni #1200 16 Kkraxuna 17 kraxuna kloni #1170 18 Cxaveri

3 82.82.4 76.62.6 64.76.7 63.63.2 86.01.3 77.92.1 29.12.6 17.44.5 64.32.6 69.48.6 22.32.6 19.02.5 16.12.7 57.03.2 37.02.2 0.80.4 1.60.3 0.60.4 0.90.5 80.42.4 67.13.6 56.52.2

4 C-15 C-15 C-15 C-20 C-15

grZeli milebi

Mmtveri-Sewebebuli, arareducirebuli; datotvili samtvre milebi

C-15 C-15 C-15 C-15 C-20 C-20 C-20 C-15 C-20 C-20 C -15 -10 C-15 -10 C-15 C-15 C-15 deformirebuli milebi deformirebuli milebi dafiqsirda oTkuTxedis formis samtvre milis erTi SemTxveva mokle deformirebuli milebi grZeli milebi grZeli milebi grZeli milebi

19 aladasturi 20 jani 21 saferavi budeSurisebri 22 aleqsandrouli 23 mujureTuli wulukiZis 24 TeTra orbeluri 25 ojaleSi 26 usaxelouri 27 Tavkveri 28 asureTuli Savi 29 bazaleTuri 30 safena 32 citogenetika #5 33 sawuravi 34 kaWiWi

41

diagrama 2

sur. 25 Tavkveris TviTdamtveril yvaviledze normaluri marcvlebis ganviTreba

sur. 26 Tavkveris yvavikastrirebul yvavi ledze normaluri ganvimarcvlebis ganvi Tareba (apomiqsisi)

sur. 27 laboratoriaSi dafesvianebul rqebze marcvlis gamonaskva

42

aRniSnulma movlenam Tavisi asaxva hpova ujredul donezec. mtvris marcvlis sakveb areebze Tesvis dros, aRiniSneboda maTi gaRivebis erTeuli SemTxvevebi (sur. 28, 29, 30), romelic jiSebis mixedviT gansxvavdeboda da saSualod 0.4+0.1 _ 1.6+0.3%-s Seadgenda. gaRivebuli marcvlebis yvelaze meti raodenoba (1.6%) miiReboda jiS asureTul SavSi, xolo naklebi (0.4%) jiS TavkverSi. danarCen jiSebs maT Soris Sualeduri adgili ekavaT. funqcionalurad unqcionalurad mdedrobiTi sqesis sqesis vazis jiSebis mtvris marcvlis xelovnur sakvb areze gaRiveba

sur.28 asureTuli Savi

sur. 29 safena

sur. 30 Tavkveri

rac Seexeba funqcionalurad mdedrobiTi sqesis jiSebis apomiqsisisadmi midrekilebas, Cvens mier Catarebuli eqsperimentuli gamokvlevebiT 5600-ze met kastri43

rebul yvavilze srulfasovani (normaluri ganviTarebis) Tesliani marcvlebi ganiviTara jiS Tavkveris yvavilebis 0.80.2%-ma, asureTuli Savis 0.80.2%-ma, safenas 1.10.3%-ma da bazaleTuris 1.40.8%-ma (sur. 26), risi miRebis SesaZleblobis marTebulobas adasturebs embriologiuri gamokvlevebi: `TavkverSi zogierTi Canasaxis parki yvavilobidan 5-7 dRis Semdeg Seicavs kvercxujreds, romelSic ramodenime birTvakiani didi zomis birTvia moTavsebuli, xolo yvavilobidan 10-12 dRis Semdeg dafiqsirebul masalaSi rig SemTxvevaSi degenirebuli sinergidebis gverdiT Canasaxis parkis arseboba aRiniSneba, rac imaze miuTiTebs, rom vazSi Canasaxi SeiZleba ganviTardes damtvervis gareSe, parTenogenezis gziT~ (, 1971). mopovebuli eqsperimentuli masalebiT funqcionalurad mdedrobiT jiSebSi: TavkverSi, asureTul SavSi, safenasa da bazaleTuris yvavilebis izolirebis SemTxvevaSi, mtevanze, parTenokarpiuli marcvlebis gverdiT arsebuli normaluri ganviTarebis Tesliani marcvlebi SesaZlebelia miRebuli iyos, rogorc TviTdamtvervis, aseve apomiqsisis gziT.D D gamoiricxa maTi mtvris marcvlis absoluturi steriloba; dadginda sakuTari mtveriT nawilobrivi ganayofierebis SesaZlebloba, Sedegad, vazis nargaobaSi iseTi klonebis Ziebis realoba, romelTa yvaviledebi Seicaven didi raodenobiT orsqesian yvavilebs, SesaZlebeli iqneba maTi wminda nargaobebis saxiT gaSeneba da Tavidan iqneba acilebuli is sirTuleebi da xarjebi, rac xelovnur damtvervasa Tu Sereul nargaobebTan aris dakavSirebuli, Tumc aseTi klonis moZiebamde, reglamentirebuli da xarisxiani mosavlis misaRebad, aucilebel pirobad migvaCnia, citologiuri meTodebis gamoyenebiT, ukeTesi damamtverianebeli jiSis SerCeva; kvlav dadasturda funqcionalurad mdedrobiT va44

zis jiSebisaTvis regularuli apomiqsisis arseboba; daiSva normaluri ganviTarebis wipwiani marcvlebis TviTdamtvervisa da apomiqsisis gziT miRebis SesaZlebloba. vazis jiSebi mtvris marcvlis milebis sigrZis mixedviT iyofa SedarebiT mokle (114-200 mkm), saSualo (200-300 mkm) da grZeli (300-435 mkm) mtvris milis mqone jgufebad. SedarebiT mokle samtvre milebi axasiaTebT jiSebs: colikours, xixvs, Cxavers, sawuravs da sxvas; saSualo: Cinurs, cicqas, ocxanur saferes, kraxunas, aladasturs, jans, saferavi budeSurisebursa da kaWiWs, xolo grZeli: rqawiTels, saferavs, wiTel budeSurs, gorul mwvanes da sxvas.

mtvris marcvlis cxovelmyofelobis cxovelmyofelobis xangZlivoba vazis qarTuli genotipebi Cveulebriv haermSral mdgomareobaSi, laboratoriul pirobebSi, cxovelmyofelobas inarCuneben 7-10 dRis ganmavlobaSi.

45

Cxaveri

aladasturi

kraxuna

rqawiTeli vardisferi

46

III. III. vazis qarTuli genotipebis mtvris marcvlis Camoyalibebisa da ganviTarebis Seswavlis Sedegebis institucialuri Aanalizi vazis qarTuli genotipebis mtvris marcvlis ganviTarebis biologiis mcenareTa genetikuri resursebis saerTaSoriso institutis (Bioversity International), vazisa da Rvinis saerTaSoriso ofisisa (OIV) da mcenareTa axali jiSebis dacvis saerTaSoriso kavSiris (UPOV) vazis deskriptorebis (Vitis spp.) moTxovnaTa gaTvaliswinebiT, Catarebuli citoembriologiuri da citogenetikuri kvlevebis Sedegad dadginda, rom: vazis qarTul genotipebSi ZiriTadi embriologiuri niSnebi _ samtvre parkis agebuleba da mikrosporisa da mamrobiTi gametofitis formireba mimdinareobs V. Vinifera L.-saTvis arsebuli aRweris mixedviT. mamrobiTi gametofitebi orujrediania. vgetatiuri ujredis birTvi dnm-ze iZleva sust reaqcias. generatiuli ujredis birTvi xasiaTdeba fiolgen-dadebiTi reaqciiT. spermatogenezi mimdinareobs mtvris milSi. spermiebi ovaluri formis, erT mxares odnav wagrZelebuli; xasiaTdebian citoplazmis mniSvnelovani SemcvelobiT. funqcionalurad mdedrobiTi sqesis jiSebSi erTbirTviani mikrosporas formireba sawyis etapze normalurad mimdinareobs; mikrosporis degenerireba iwyeba maTi ganviTarebis erTbirTvian stadiaze, ris Sedegadac miiReba gansxvavebuli formis, uforo, steriluri mtvris marcvlebi. poliploidur mcenareebSi citoembriologiuri kvlevis ZiriTadi etapebi diploiduri jiSebis msgavsad mimdinareobs, Tumc mikrosporogenezis procesSi mikrosporas deda ujredebSi aRiniSneba zogierTi saxis darRvevebi: I profazaSi ujredebis birTvebs Soris qroma47

tiduli migracia-citomiqsisi; I metafazaSi bivalentebis nacvlad univalentebisa da trivalentebis Camoyalibeba; I anafazaSi qromosomebis polusebisaken araTnabari gadanawileba, qromosomebis gasvla aqromatiduli TiTistaris farglebidan da Sesabamisad, araerTgvarovani mtvris marcvlebis Camoyalibeba. I anafazaSi qromosomuli darRvevebi Ziriadad warmodgenilia CamorCenili qromosomebis, qromosomuli xidebisa da polusebze araTanabrad gadanawilebuli qromosomebis, sampolu-siani anafazebis saxiT da sxva.P poliploidur jiSebSi meiozis procesSi arsebuli aberaciaTa sixSire da speqtri, Secvlili diadebi da tetradebi ganapirobebs araerTgvarovani, momwifebis sxvadasxva xarisxisa da Sesabamisad, gansxvavebuli ganayofierebis unaris mqone mtvris marcvlis Camoyalibebas. orsqesiani jiSebi maRalfertiluria (69.22.7 _ 98.9 0.5%), xolo rac Seexeba funqcionalurad mdedrobiT jiSebs aq fiqsirdeba fertiluri mtvris marcvlis arsebobis erTeuli SemTxvevebi (0.60.4%). mtvris marcvlis gaRiveba xelovnur sakveb areze gansxvvebulia da 40-90%-is farglebSia, xolo cxovelmyofeloba, haermSral mdgomareobaSi, laboratoriul pirobebSi, 7-10 dRea. vazis qarTuli genotipebis orsqesian jiSebs axasiaTebT xorblis marcvlis formis, ZiriTadad samforiani (67.624.4-98.32.6% (danarCeni mikrosporogenezis procesSi arsebuli darRvevebis gamo uforoa), mcire zomis mtvris marcvlebi, romlebic sakveb areze moxvedrisas an dasvelebis SemTxvevaSi dasawyisSi samkuTxedis, xolo Semdeg ki sferul formas Rebuloben. maTi parametrebi jiSebis mixedviT gansxvavebulia (haermSral mdgomareobaSi): sigrZe - 21.10.9_38.80.3 mkm, sigane - 14.7
48

0.2_23.00.9 mkm da diametri (acetokarminis saRebavSi) 18.10.3_27.90.4 mkm-is farglebSia. funqcionalurad mdedrobiT sqesis vazis jiSebis mtvris marcvalebs haermSral mdgomareobaSi wamaxvilebuli forma aqvT, absolituri umravlesoba uforoa (Tumc wlebis ganmavlobaSi dafiqsirebulia samforiani mtvris marcvlebis arsebobis erTeuli (0.41.3_2.51.4%) SemTxvevebi.), dasvelebis SemTxvevaSi erTgvarovan sferul formas Rebuloben.Pparametrebi haermSral mdgomareobaSi: sigrZe - 31.60.7_27.80.5 mkm, sigane - 23.00.3_ 20.50.5 mkm, xolo diametri (acetokarminis saRebavSi) 25.404_21.603 mkm-ia. jiSis warmoSobis adgilis wili (p<0.01 sizustiT) pasuxismgebelia mtvris marcvlis sigrZis cvalebadobis 43.44%-ze, siganis _ 46.0%-ze da diametris _ 51.19%-ze. gansxvaveba fiqsirdeba regionebis mixedviTac: sigrZe 16.84_20.72-mkm, sigane 9.71_10.95-mkm, xolo diametri 13.40_ 18.11-mkm-ia. cdis saSualo cdomileba Sx dabalia da 0.010.68 mkm-s Seadgens. variaciis koeficientis mixedviT vazis qarTuli genotipebis mtvris marcvlis citologiuri maxasiaTeblebis zemoT motanili maCveneblebi didi mudmivobisa da mcired cvalebad raodenobriv niSnebs miekuTvneba, aqvT didi ampelografiuli da saseleqcio Rirebuleba da warmatebiT SeiZleba iqnen gamoyenebulni, rogorc jiSisa da formis identifikaciisa da Sefasebis citologiuri markerebi. funqcionalurad mdedrobiT jiSebSi gamoiricxa mtvris marcvlis absoluturi steriloba; dadginda sakuTari mtveriT nawilobrivi ganayofierebis SesaZlebloba, Sedegad, vazis nargaobaSi iseTi klonebis Ziebis realoba, romelTa yvaviledebi Seicaven didi raodenobiT orsqesian yvavilebs, SesaZlebeli iqneba maTi wmin49

da nargaobebis saxiT gaSeneba da Tavidan iqneba acilebuli is sirTuleebi da xarjebi, rac xelovnur damtvervasa Tu Sereul nargaobebTan aris dakavSirebuli, Tumc aseTi klonis moZiebamde, reglamentirebuli da xarisxiani mosavlis misaRebad, aucilebel pirobad migvaCnia, citologiuri meTodebis gamoyenebiT, ukeTesi damamtverianebeli jiSis SerCeva; kvlav dadasturda funqcionalurad mdedrobiT vazis jiSebisaTvis regularuli apomiqsisis arseboba; daiSva normaluri ganviTarebis wipwiani marcvlebis TviTdamtvervisa da apomiqsisis gziT miRebis SesaZlebloba.

50

colikouri

sawuravi

budeSuri wiTeli
51

xixvi

1V. wignSi gamoyenebuli terminebi

a aberacia (gadaxra) anu qromosomuli anomalia, qromosomaTa nebismieri tipis mutaciebis zogadi saxelwodebaa: delecia, translokacia, inversia, duplikacia. avtopoliploidi organizmi, romelic warmoiqmneba qromosomaTa erTidaigive anawyobis jeradi gadidebiT; individi, romelsac avtopoliploidiis safuZvelze Tavis qromosomul nakrebSi aqvs ramdenime erTnairi genomi (avtotetraploidi, avtoheqsaploidi) mas ewodeba balansuri, xolo arawyviladia (triploidi, pentaploidi), ewodeba arabalansirebuli. Tu avtoploids qromosomul nakrebSi aqvs or wyvil homologiurze meti, meiozSi irRveva homologiuri qromosomebis konugacia dacileba. bivalentebTan erTad warmoiqmneba uni, tri da tetravalentebi - dabalsicocxlisunariani gametebi.M avtopoliploidia, euploidia - erTidaigive saxeobis haploidur nakrebis orjeradze metjer gadideba. azotovani fuZe - fuZe, romelic Sedis nukleinis mJavis SemadgenlobaSi.Aarsebobs ori tipis: pirimidinuri (uracili, Timini, citozini da purinuli (adenini, guanini). aleli - genis erTi an ramdenime alternatiuli forma, romelTagan TiToeuls axasiaTebs nukleotidebis unikaluri Tanamimdevroba. aleluri genebi - homologiuri qromosomebis erTnair wertilSi moTavsebuli genebi erTi wyvili niSniT. diploidur organizmebSi erT gametaSi. ori aleli ar SeiZleba iyos.

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Aanafaza - birTvis mitozuri da meiozuri gayofis faza, romlis drosac qromosomebi an qromatidebi miemarTebian sxvadasxva polusebisaken. Aapomiqsisi - organizmis ganviTareba sasqeso ujredebis Serwymis gareSe: gaunayofierebeli kvercxujredisagan (parTenogenezisi), Canasaxis parkis vegetatiuri ujredebisagan (apogamia), an maTi qsovilis garemomcveli vegetatiuri ujredebisagan (aposporia). Aarqesporiumi - ujredebis rigi an fena, yvavilovan mcenareebSi, saidanac viTardeba mtvris marcvlis an Canasaxis parkis deda ujredebi. b bivalenti - ori konugirebuli homologiuri qromosoma. bivalentebis raodenoba qromosomebis haploiduri raodenobis tolia; yoveli bivalenti Sedgeba 4 qromatidisagan, amitom qromosomebis bivalentebs uwodeben qromatidebis tetradas.Bbivalentebi warmoiqmneba pirveli meiozuri gayofis profazaSi. birTvi - aRmoCenilia brounis mier. igi ujredis umniSvnelovanesi nawili, yvela procesisa da misi cxovelmoqmedebis marTvis centria. birTvSi Tavmoyrilia: organizmis memkvidreobis materialuri matarebeli qromosomebi, birTvakebi, birTvis wveni-karioplazma. birTvi citoplazmidan gamoyofilia birTvis garsiT. birTvi denovo ar warmoiqmneba. igi iyofa. birTvis ZiriTadi nivTiereba - qromatinia, romelic dnm-sa da cilis kompleqsuri naerTia. organoidia. sicocxlisunariania mxolod citoplazmaSi; birTvaki - ujredis momrgvalo, optikurad homogenuri organoidia, romelic lokalizebulia birTvSi. birTvaki ujredSi SeiZleba iyos ori an ramodenime. birTvaki Sedgeba cilebisa da rnm-sagan. birTvakSi yalibdeba ribosomebi.
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g gameta - momwifebuli mamrobiTi da mdedrobiTi sasqeso ujredebi, romlebic sxeulis sxvadasxva ujredebTan SedarebiT Seicaven qromosomTa haploidur (naxevar) ricxvs. gametogenezi - mtvris marcvlebSi mamrobiTi gametebis (spermiebis) da Canasaxis parkSi mdedrobiTi gametebis (kvercxujredis) formirebis procesis mimdinareoba. gametofiti - sqesobrivi Taoba yvavilovan mcenareebSi, romelic atarebs qromosomebis naxevar raodenobas. Ggametociti - ujredebi, saidanac gametogenezis dros warmoiqmneba gametebi (makrosporociti, mikrosporociti). geni klasikur genetikaSi organizmis ama Tu im niSan-Tvisebis ganmapirobebeli materialuri erTeuli; molekulur genetikaSi - qromosomis (dnm-is) ubani, romelic kodirebs romelime funqcionalurad aqtiur produqts - rnm-s an misi transilaciis produqts (polipeptidebs). genetikuri modifikacia - transgenuri organizmi, romelSic gadaitanes erTi an ramodenime geni mis mimarT aramonaTesave organizmidan. genofondi - populaciis (saxeobis) genTa erToblioba, romelic xasiaTdeba maTi garkveuli sixSiriT. genetika - mecniereba organizmis memkvidreobasa da cvalebadobaze. genotipi - mTeli genebis erToblioba, romelic gansazRvravs mcenareTa niSan-TvisebaTa ganviTarebas. genomi - qromosomTa ZiriTadi haploiduri anawyobi, Tvisobrivad gansxvavebuli qromosomTa amonakrebi, romelic Seicavs genebis srul erTmag anawyobs.

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genotipi - (berZ. Genos- dabadeba, typos-gamosaxuleba (anabeWdi) aris organizmis yvela genis erToblioba, misi memkvidreobis materialuri safuZveli. d degeneracia - niSnavs gauaresebas ujredis, ujredis qsovilis organos struqturis Sedgenilobis cvlilebebs, ris gamoc sustdeba cxovelmyofeloba da funqcia. delecia - aberacia, qromosomuli mutacia, romlis drosac ikargeba qromosomebis fragmenti. diferenciacia - embrionaluri ganviTarebis procesi, romlis ujredebi Rebuloben specifikur formas da iwyeben garkveuli funqciebis Sesrulebas. diploidi - individi, romelsac somatur ujredebSi qromosomaTa ori homologiuri anawyobi aqvs (2n). zigotaSi erTi Setanilia mdedrobiTi, xolo meore mamrobiTi formisagan. dnm - dezoqsiribonukleinis mJava, memkvidreobis ZiriTadi materialuri matarebeli, romlis molekula Sedgeba spiralurad daxveuli ori polinukleotiduri jaWvisagan. dnm-is calkeuli nukleotidis SemadgenlobaSi. Sedis azotovani fuZeebi, Saqari - dezoqsiriboza da fosforis mJavis narCenebi. e endomitozi - mitozis saxesxvaoba, qromosomebis gaormageba birTvis gayofis gareSe, rac iwvevs poliploiduri ujredis warmoSobas. ekvaciuri dayofa - meore meiozuri dayofa, romelic mimdinareobs mitozis msgavsad. ekzina - mikrosporis gareTa garsi. ekgzinaze ganviTarebulia forebi anu kvlebi, romliTac xorcieldeba mtvris marcvlis kavSiri mis irgvliv myof garemosTan.
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v variacia - individebs Soris modifikaciuri an genomuri gansxvaveba. z zigota - ganayofierebuli kvercxujredi. warmoiqmneba ori gansxvavebuli sqesis SerwymiT, romelic sawyiss aZlevs axali organizmis ganviTarebas. aqvs qromosomaTa ormagi diploiduri ricxvi. zigonema - pirveli meiozuri gayofis profazis mesame stadia. iwyeba wyvili homologiuri qromosomis konugaciiT. konugacia xasiaTdeba gansakuTrebuli sizustiTa da specifikurobiT: homologiuri qromosomebidan erT-erTis TiToeuli wertili uTavsdeba meore qromosomis Sesabamis wertils, TiToeuli qromomera ki Sesabamis qromomeras. T TiTistara - eukariotul ujredebSi cilovani Zafebisagan Seqmnili struqtura, romelic axorcielebs qromosomaTa moZraobas mitozsa da meiozSi. i inversia - qromosomuli mutacia, romelic warmoiqmneba qromosomis ormagi gaglejiT da am amovardnili nawilis 1800-iT SemobrunebiT, romlis drosac genebis Tanamimdevroba Semdegnairad icvleba abcd-acbd. interfaza - eukariotuli ujreduli ciklis faza, romlis drosac ar xdeba mitozi, e.i. Sualedi or mitozs Soris. interfaza moicavs sam stadias: G1-presinTezuri, S- sinTezuri da G2- postsinTezuri. intina - mikrosporas Sida garsi

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k kariotipi - somaturi ujredis qromosomTa erToblioba, maTTvis damaxasiaTebeli ricxviT, formiT da sididiT. kloni - mevenaxeobaSi warmoadgens kvirtuli mutaciis vegetatiur Taobas, romelic ZiriTadi jiSisagan gansxvavdeba erTi an ramodenime niSniT, memkvidrulia da ucvlelad gadaecema Taobebs. vegetatiuri Taoba, romelic aRniSnul pirobas ar akmayofilebs, klonad ar CaiTvleba. konugacia konugacia qromosomebisa - homologiuri qromosomebis qromatidebis dakavSireba mTel sigrZeze pirveli meizuri gayofis profazaSi. krosingoveri - homologiuri qromosomebis araSvileuli qromatidebs Soris ubnebis gacvla pirveli meiozuri gayofis profazaSi. m meiozi - ujreduli dayofis gansakuTrebuli tipi, romelic mimdinareobs sasqeso ujredebis an sporebis ganviTarebis dros qromosomaTa ricxvis SemcirebiT (reduqcia). meiozis procesSi xdeba birTvis ori Tanamimdevruli dayofa, xolo qromosomebi mxolod erTxel ormagdeba. meiozSi homologiuri qromosomebi konugirdeba. meiozis profaza - meiozis pirveli dayofis pirveli faza, romlis drosac mimdinareobs homologiuri qromosomebis konugacia da maT Soris ubnebis gacvla (krosingoveri). metafaza - mitozis meore stadia, romelSic spiralizebuli qromosomebi lagdebian ekvatorul sibrtyeSi.

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mikrospora - meiozis ori Tanamimdevruli dayofis Sedegad warmoqmnili haploiduri ujredi, romelic Semdeg mtvris marcvlad gadaiqceva. mikrosporogenezi - farulTeslian mcenareTa samtvre parkebSi mtvris marcvlebis ganviTareba. haploiduri ujredebi (mikrosporebi, romlebic warmoiqmneba ori meiozuri dayofiT) viTardebian mtvris marcvlebSi. mikrosporociti - mikrosporas deda ujredi mitozi - somaturi ujredebis gayofa, romlis drosac diploiduri ujredidan miiReba ori Svileuli diploiduri ujredi. modifikacia - cvalebadi garemo pirobebis gavleniT romelime gansxvavebuli niSan-Tvisebis gamovlena. . mutacia - axali memkvidruli cvlileba, romelic warmoiqmneba Sejvarebisagan damoukideblad da dakavSirebulia qromosomis dnm-is SecvlasTan. . organizmze moqmedebisa da gavlenis mixedviT igi SeiZleba iyos: morfologiuri, fiziologiuri da bioqimiuri mutaciuri cvalebadoba - genebisa da qromosomebis struqturuli cvlilebebi, romlebic iwvevs organizmis axali niSnebisa da Tvisebebis gamovlenas. . n nukleinis mJavebi mJavebi - biologiuri polimerebi maRalmolekuluri naerTi, romelic uzrunvelyofs memkvidrulobis informaciis Senaxvasa da gadacemas. igi organizmSi warmodgenilia ori ZiriTadi saxiT: ribonukleinis (rnm) da dezoqsiribonukleinis (dnm) mJavis saxiT. nukleotidi - rTuli organuli nivTiereba, romelic Sedgeba azotovani fuZeebis, ribozis an dezoqsiribozisa da fosforis mJaveebisagan. igi Sedis dnm-sa da rnm-is SemadgenlobaSi.

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nucelusi - Teslkvirtis centraluri nawili, Sedgeba Txelgarsiani meristemuli ujredebisagan. p parTenokarpia - qalwulebrivi gamravleba e.i. uTeslo nayofebis warmoqmna: 1. kvercxis ganviTarebis xelovnuri stimulaciiT (stimulaciuri parTenokarpia, roca nayofis warmoqmnisaTvis saWiroa dingis gaRizianeba ucxo mtveriT, an daumtveravad ucxo mtveris gareSe, 2. ganayofierebis procesis gamovardna SeuTavseblobasTan dakavSirebiT, 3. embrionis ganviTarebis amovardna zigoturi sterilobis Sedegad. paqinema - pirveli meiozuri gayofis profazis mesame stadia, romlis drosac xdeba bivalentebis, qiazmebisa da krosingoveris warmoSoba. populacia - erTi saxeobis individTa erToblioba, romlebic dasaxlebulni arian garkveul teritoriaze: erTmaneTs Tavisuflad amtverianeben da ama Tu im xarisxiT izolirebulni arian sxva jgufisagan. seleqciaSi populacia esmiT, rogorc individTa jgufi, romelsac gansxvavebuli memkvidreoba aqvs. polisomatia (miqsoploidia) - erT qsovilSi an erT individSi diploiduri da poliploiduri ujredebis erTdrouli arseboba. r reduqcia - qromosomTa somaturi (diploiduri) raodenobis orjer Semcireba meiozSi. s saxeoba - reproduqtulad izolirebuli, urTierTSejvarebis unariani populaciebis erToblioba. simultanuri - mikrosporis tetradebis (erTdrouli) warmoSobis tipi.
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sinapsisi - qromosomebis konugacia meiozSi. sinaptonemaluri kompleqsis struqturebi. warmoiqmneba meiozSi, romelic uzrunvelyofs mWidro kontaqts homologiur qromosomebs Soris. sinergidebi - ori haploiduri Svileuli ujredebia, romelic moTavsebulia kvercxujredis orive mxares mikropilur nawilSi. sinergidebis birTvebi kvercxujredis birTvebisagan gansxvavebiT moTavsebulia im adgilas, sadac koncentrirebulia citoplazmis ZiriTadi masa. sinergidis qveda nawilSi viTardeba vakuoli. sinergidebi xels uwyobs mtvris milis moZraobas Canasaxis parkisaken da maTi garsebis gaxsnas, radganac Seicavs fermentebs: citazas da peqtazas, mtvris milis wveris Sexebisas sinergidebTan, isini ixsneba da maTi Semcveloba iRvreba Canasaxis parkSi. spermia - mtvris mamrobiTi sasqeso ujredi (gameta). spermia warmoiqmneba mtvris marcvalSi an samtvre milSi. sxvadasxva saxeobaSi, spermiis zoma farTo farglebSi varirebs.Migi umravles SemTxvevaSi damokidebulia spermiis citoplazmis masaze. . steriloba - sqesobrivi gziT STamomavlobis mocemis unaris Semcireba an daTrgunva.Aarsebobs: 1. gameturi steriloba ganpirobebulia gametebis warmoqmnis funqcionaluri uunarobiT, 2. genuri steriloba ganpirobebulia sterilobis genis moqmedebiT; recesiuli sterilobis genis moqmedebas uwodeben pirdapirs; sterilobis genis moqmedeba, romelic ganpirobebulia fizioologiuri darRveviT da vlindeba nawilobrivi an sruli sterilobiT uwodeben arapirdapir sterilurobas. steriloba qromoso qromosomuli - steriluri gametebis warmoSoba SeimCneva Soreuli hibridizaciis, trisomiisa da qromosomebis rigi darRvevebis dros, romelic meiozis normalur msvlelobas arRvevs.

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t tapetumi tapetumi - ujredebi, romelic samtvre parkSi gars akravs mtvris marcvlis dediseul ujreds da maTTvis sakvebi nivTierebis gadacemas emsaxureba. telofaza - mitozis an meiozis meoTxe faza, romlis drosac xdeba axali Svileuli birTvebis warmoqmna. tetrada - ujredebis jgufi, romelic mcenareebSi warmoiSveba meiozis Sedegad sporis dediseul ujredebSi. yvavilovan mcenareebSi mamrobiTi xazis tetradebi Sedgeba 4 mikrosporisagan, romelic mogvianebiT erTmaneTs scildeba da mtvris marcvlebad gadaiqcevian. tetraploidi - ujredi an organizmi, romelic Seicavs qromosomebis oTxmag raodenobas da orjer aRemateba qromosomTa diploidur nakrebs. Tu tetraploidis genotipSi gaormagebulia erTidaimave saxeobis qromosomebis diploiduri nakrebi, aseT organizmebs avtopoliploidi (4n=AAAA), xolo. Tu tetraploidi warmoiqmneba sxvadasxva saxeobis genomebis gaerTianebisa da gaormagebis Sedegad, alotetraploidi an amfidiploidi (4n=AABB) ewodeba. triploidi - ujredi an organizmi qromosomaTa haploiduri nakrebis sammagi raodenobiT. u univalenti - erTeuli qromosomebi, romelic meiozis pirvel dayofaSi konugacias ar ganicdian da ujredis polusebze nawildebian SemTxveviT. . f fenotipi - organizmis mTel niSan-TvisebaTa erToblioba, romelic formirdeba, genotipis safuZvelze garemo pirobebTan urTierTzemoqmedebiT.
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fragmentebi - qromosomis centruli da acentruli ubnebi, romlebic warmoiqmnebian qromosomebis dawyvetis Sedegad. acentruli fragmentebi SeiZleba sruliad moscildes qromosomas (Tavisufali fragmenti) an darCes wvrili ZafiT (fiqsirebuli fragmenti). fiqsirebuli fragmentebi anafazaSi miemarTeba erT-erTi polusisaken. Tavisufali ubani rCeba ekvatorze da eliminirdeba. q qimera - mcenare, romelic Sedgeba sxvadasxva genotipis qsovilebisagan. miiReba somaturi mutaciis Sedegad, agreTve mynobis dros, rodesac Sexorcebis Sezrdis adgilas yalibdeba kvirti, romelSic qsovilTa nawili ekuTvnis saZires da nawili ki sanamyenes. . qiazma - bivalentebis qromatidebis gadajvaredineba, pirveli meiozuri gayofis profazaSi, maTi konugaciis dros. qloroplasti - mcenaris eukariotuli ujredis organeli, romelSic mimdinareobs fotosinTezi. qromatida - gaormagebuli qromosomebis ori Svileuli Zafi, romlebic centromerebiT aris dakavSirebuli. misi struqturuli erTeulia nukleoproteinis Zafi - qromonema. isini Sedgebian mikrofibrilebisagan, romelic Cans eleqtronul mikroskopSi. mitozis anafazaSi Svileuli qromatidebi erTmaneTisagan dacilebis Semdeg gadaiqcevian damoukidebel qromosomebad. . qromonema - nukleotiduri Sedgenilobis ujreduli birTvis ZiriTadi Sedgeniloba, kargad iRebeba fuZe saRebavebiT. Sedgeba dnm-sa da histonebisagan, umniSvnelo raodenobiT Seicavs rnm-is cilebs. ansxvaveben or qromatins: 1. heteroqromatins, romelic intensiurad iRebeba saRebaviT, ZiriTadad lokalizebulia centromerasTan axlos da genetikurad pasiuria, ar Seicavs genebs,
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romlebic kontrols uweven organizmis niSnebis ganviTarebas da mxolod gavlenas axdenen maT ricxobriv gamovlinebaze 2. euqromatini, romelic sustad iRebeba, masSi ganTavsebulia niSan-Tvisebebis matarebeli genebi da genetikurad aqtiuria. qromosoma - ujredis birTvis elementi, romelic iRebeba fuZe saRebaviT. Sedgeba dnm-sa da cilebisagan. igi aris organizmis memkvidreobis informaciis ZiriTadi matarebeli. sami tipisaa: 1. Tanabarmxriani metacentruli, 2. araTanabarmxriani - submetacentruli, 3. araTanabarmxriani - akrocentruli. qromosomebis lokusi - qromosomebis nawili, romelSic genia moTavsebuli. qromosomuli aberaciebi - qromosomebis sxvadasxva struqturuli cvalebadoba (ukmarisoba, translokacia, inversia, dublikacia). qromosomuli kompleqti - mocemuli saxeobebisaTvis damaxasiaTebeli qrmosomaTa anawyobi. qromosomuli anawyobi - qromosomaTa erToblioba, romelic damaxasiaTebelia mocemuli organizmis ujredisaTvis, cnobilia ori tipis anawyobi: 1. haploiduri (n) - momwifebuli sasqeso ujredebi da 2. diploiduri (2n) - somaturi ujredebi. c cvalebadoba cvalebadoba - individebs Soris sxvadasxva niSnis mixedviT sxeulis anda misi calkeuli oraganoebisa (zoma, forma, Seferva, qimiuri Sedgeniloba) da maTi funqciebis gansxvavebis warmoqmnis procesi. igi SeiZleba iyos memkvidruli da modifikaciuric. citogenetika - mecniereba, romelic Seiswavlis organizmis memkvidreobisa da cvalebadobis movlenebs ujreduli struqturis gansakuTrebiT, qromosomebis qcevebTan dakavSirebiT.
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citokinezi - ujredis gayofa. citologia - mecniereba ujredis Sesaxeb, romelic Seiswavlis mis struqturasa (Senebas) da funqciebs (cxovelmoqmedebas). citoplazma - ujredis mTeli masa birTvis gareSe. Seicavs organoidebs, romlebic asruleben sxvadasxva funqciebs (endoplazmuri bade, mitoqondriebi, ribosomebi, plastidebi da sxva). h haploidi - organizmi, romlis ujredi Seicavs orjer nakleb qromosomas (n), vidre maTi sawyisi forma. hibridi - organizmi, romelic miRebulia genetikurad gansxvavebuli niSan-Tvisebebis mqone mSoblebis SejvarebiT heterozigotuli organizmi - individi, romlis sxeulis ujredebi Seicavs, mocemuli aleluri wyvilis sxvadasxva genebs. homologiuri qromosomebi - ganayofierebis Sedegad miRebuli msgavsi wyvili qromosomebi, romlebic meiozSi normalur konugacias ganicdian. B B Bioversity international - mcenareTa genetikuri resursebis saerTaSoriso instituti (romi, italia). O OIV vazisa da Rvinis saerTaSoriso ofisi (parizi, safrangeTi). U UPOV - mcenareTa axali jiSebis dacvis saerTaSoriso kavSiri (Jeneva, Sveicaria).
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profesor d. tabiZisa da mecnierebaTa mecnierebaTa doqtoris l. vaSakiZis seleqciiT miRebuli hibriduli hibriduli formebi

Tavkveri X pino Savi

asureTuli Savi X kaberne sovinioni

mxargZeli X rislingi
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safena X xixvi

gamoyenebuli literatura da eleqtronuli informaciuli masalebi

aTarbegova a., ustinova e., (1974), mcenareTa citologia. Tbilisi, 279 gv. guliaevi g.m., (1989), genetika, 457 gv. goginaSvili q., Saria S., (2009), zogadi da samedicino genetikis leqsikoni. Tbilisi, 130Ggv. vazis deskriptorebi (Vitis spp.), (2005), Tbilisi, 47 gv. vaSakiZe l., (2006), mtvris marcvlis Taviseburebani //sadoqtoro disertacia `vazis qarTuli genotipebis identifikaciisa da zogierTi fitoteqnikuri RonisZiebis optimizaciis mecnieruli safuZvlebi~, Tbilisi, gv. 102-112. http://www.nplg.gov.ge/dlibrary/coll/0002/000109/ vaSakiZe l., (1964), `saRvine vazis jiSebis axali hibridebis gamovlineba qarTlis pirobebisaTvis~ // sakand. disertacia biologiur mecnierebaTa samecniero xarisxis mosapoveblad. Tbilisi. vaSakiZe l., gurasaSvili v., (2000), vazis jiS sufris gorulas da misi klonis mikrosporogenezi // `vazi da Rvino~, 3 (27), Tbilisi, gv. 74-80. vaSakiZe l., CxartiSvili n., maRraZe d., gurasaSvili v., mdinaraZe i., (2007), funqcionalurad mdedrobiTi vazis jiSebis TviTdamtverva da apomiqsisi // saq. mecn.Akademiis moambe, Tbilisi, gv. 174-177. vaSakiZe l., (2008), saqarTvelos vazisa da xexilis genofondis citoembriologia da citogenetika // `vazisa da xexilis genofondi saqarTveloSi~ (mmmi, samecn. SromaTa krebuli), Tbilisi, gv. 34-48. vaSakiZe l., maRraZe d., (2009), kaxeTis vazis jiSebis palinomorfologia // saq. soflis meurneobis mecnierebaTa akademiis `moambe~, tomi #23, Tbilisi, gv. 149-153. leJava T., (2004), ujredis genetika. Tbilisi.
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ramiSvili m., (1960), Tavkveri // saqarTvelos ampelografia. Tbilisi, gv. 157. yvavaZe e., (2004), vales arqeologiuri masalis palinomorfologiuri Seswavlis Sedegebi // arqeologiuri kvlevis centris arqivi. saqarTvelos mecnierebaTa akademia, Tbilisi. yvavaZe e., (2004), ai-ilias arqeologiuri gaTxrebis palinologiuri Seswavlis pirveladi Sedegebi // arqeologiuri kvlevis centris arqivi. saqarTvelos mecnierebaTa akademia, Tbilisi, ColoyaSvili s., (1939).Mmevenaxeobis saxelmZRvanelo. wigni II. ampelografia. Tbilisi, saxelgami, sas.-sam. literaturis seqtori. 477 gv. joxaZe d. (1992), molekuluri genetikis Sesavali. Tbilisi. 299 gv. . ., (1958), . // , , 1, .25-28. .., (1982), // . . . . , c. 11. .., (2008), // 3, , .46-48. .., ., (1982), // - . . . . , c. 71-73. .., .., .., (2003), // . , . 1516.

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 ., , (1967), . . .., (1966), // . , .71. .., . ., . ., (2008), // : " "; , . 55-58. . ., . ., . ., . ., (1977), . . .., (1971), // . . . , . Akira Wakana et al., (2007). Charakteristics of Seedless Berries of Triploid Grapes( Vitis vinifera Complex) Derived from Eighteen Grosses .J.Fac. Agr., 29. Kyushu.Univ.52 (2), pp.337-344. Hirayanagi, H., (1929) Chromosome arrangement. 111 The pollen mother cells of the vine-Mem. Coll sci. ser.B. 4,3. Hiramatsu, M. A. and et.al, (2003) Production of triploid plants from crosses betveen diploid and tetraploid grapes ( Vitis complex) through immature seed culture and subsequent embrio culture. J Fac. Agr.,Univer.,48:51-57. Vashakidze, L. K. (2007), Improvement of Grapevine and Fruit Varieties in Georgian //International Center for Agricultural Research in Dryland Areas (ICARDA). Report of the Multi-Stakeholder Workshop. Elements of a National Integrated Strategy for Plant Genetic Resources Managment and USE Summary of the workshop prezentations. Tbilisi, p.10. Chkhartishvili, N., Vashakidze, L., Gurasashvili, V.,. Maghradze, D., (2006), Type of pollination indices of fruit sets of some Georgian grape-vine varieties // VITIS, 45, issue, (4 ), p . 153-156. lbanez, M,D.,Velez, M.T. de Andres J.Borrego // Bulletin de l, O.I.V. Vol 80 (No 914-916)

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Nitsh, J., (1970), Hormonal factors in growth and development. In the Biochemistry of fruits and their product, Vol. 11. ed. By A. C.Hulme. Academic press, London, pp.427-472. Rogtehev, V., Terzsky, B., Dimova, P., Karageorgiev, S., (1999), Scanning electron microscopy study of pollen morphology in seedless grape, Vitis vinifera L. cultivars. Vitis, 33, Pp. 105-108. O,Neill, S.D. and J. A. Nadeau, (1997), Postpollination flower development. Horticultural Reviews, 19: 1-58. Rombough, L. (2002), The Grape Grower. Vermont. USA. Kimura, P.H;. Okomoto, C; Hirano, K., (1998), The Mode of Pollination and Stigma Receptivity in Vitis coignetiae Pulliat. American Journal of Enology and Viticulture. USA. 49 (1) pp 1-5. Sarikhani, H., (2007), Studi on the seedles in Vitiss with references to parthenocarpy and fertilization-independent seed formation. Ph.D. disser-tation, Kyushu.University, pp.121. Sartorius, O., (1926), Zur Entwicklung und Phisiologie der Rebblute angew Bot 8. www.vivc.bafz.de http://basis.zadi.de www.jki.bund.de www.foodprom.ru www.vitis-vea.zadi.de www.oiv.int http://nplg.gov.ge www.bofz.de www.newcrops.uq.edu.ou www.boom.ge www.fao.org. www.icarda.org. www.vir.nw.ru www.ukntb.kz www.acnet.ge

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Larisa Vashakidze Georgian Grapevine Genotypes: Biology of Pollen Development SUMMARY In the monograph of the Doctor of Agricultural Sciences L. Vashakidze Georgian Grapevine Genotypes: Biology of Pollen Development discussed are cytological and cyto-embryological particularities of organization and development for male generative organs of Georgian grapevine genotypes; meiotic associations of chromosomes during the process of microsporogenesis. Based on the grapevine (Vitis ssp.) descriptors of international institutions (Bioversity International, OIV, UPOV) 53 Georgian grapevine varieties and their clones have been analyzed according to pollen palynology and their morphological-physiological parameters; importance of a varietys place of origin for its phenotypic variation. High constant and low variable traits were detected having ampelographic and breeding value, and cytological markers for identification and assessment of varieties. The main embryological characters in the Georgian grapevine genotypes construction of pollen bag and formation of microspores and male gametophytes occur as reported for V. vinifera L. in general. The mail gametophyte is dual-core. The nucleus of vegetative cell has weak reaction on the test of the DNA. The nucleus of generative cell has Fiolgenpositive reaction. Spermatogenesis proceeds in the tube of pollen. The sperms have oval shape, prolonged on one side, with enough amount of cytoplasm. Forming of mononuclear microspore proceeds in normal way in the female varieties, degeneration of microspore takes place on the mononuclear stage of its development, and as a results forms pollen grains with diverse shape, poreless and sterile. In the polyploid plants the main stage of cyto-embriological development proceeds similarly to diploid varieties, but in the mother cells of microspore some defect occurs during microsporogenesis, in particular: chromatin migration cytomixes between nucleuses of cells in the I Prophase; origin of univalents and trivalents in spite of bivalents in the I Metaphase; uneven distribution of chromosomes to poles; going of chromosomes out of achromatic spindles and therefore, organization of dissimilar pollen grains. In the I Anaphase of chromosomal abnormalities are mainly presented by lagging chromosomes, chromoso-

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me bridges and three-pole anaphases with not equally disseminated chromosomes on the poles and others. The frequency and spectrum of aberrations, changed diads and tetrads provide origin of dissimilar pollen grains with various levels of maturation and correspondently, germination ability in the polyploid varieties. The pollen of hermaphrodite varieties is highly fertile. Among pollen of female varieties, single number of fertile pollen grains (0.60.4%) was discovered. Pollen germination of artificial nutrient is within the range of 40-90%; their vitality is 7-10 days in the air-dry conditions of laboratory. Hermaphrodite varieties of Georgian genotypes have small size, grainshape, mainly three-pore (from 67.64.54% to 98.32.6%) pollen grains others are pore-less due to abnormalities during the process of microsporogenesis. The normal pollen grains on the artificial nutrient or in water became triangular at the beginning, and then sphere-shaped. Their parameters in air-dry conditions are as follows: length from 21.20.9m to 38.80.3 m, width from 14.70.2 m to 23.00.9 m, and diameter of Carmine-coloured grains from 18.10.03 m to 27.90.9 m. The air-dry pollen grains of female varieties are sharp-shape, mostly poreless, but not very frequently, three-pore pollen grains have been also discovered (from 0.41.3% to 2.51.4%) among them in various years. The pollen, in case of soakage, becomes homogenous and sphere-shaped. Their parameters in the air-dry conditions are as follows: length is from 31.60.7 m to 27.80.5 m, width is from 23.00.3 m to 20.50.5 m, and diameter of Caremine-coloured is from 25.40.4 m to 21.60.3 m. The place of origin of the varieties is responsible for 43,4% of variability of pollen length (p< 0.01), 46% of width and on 51.2% of diameter. The difference is discovered by region as well: length (16.84-20.72 m), width (9.71-10.95 m) and diameter 13.40-18.11 m). The average error of the test Sx is low in range of 0.01-0.68 m. According to the coefficient of variation the cytological parameters of Georgian grapevine genotypes are high-constant and slightly variable quantitative traits. They have ampelographic value and can be used successfully as cytological markers for identification and evaluation of varieties and forms.

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sarCevi
winasityvaoba . . . . . . . . . . . . . . . . . . . . . . . . . . . . mikrosporogenezi da qromosomaTa meiozuri .asociaciebi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . II. mtvris marcvlis morfo-fiziologiuri Tvisebebi . 1. palinomorfologia . . . . . . . . . . . . . . . . . . . . . . . . mtvris marcvlis parametrebi . . . . . . . . . . . . . . . . mtvris marcvlis forianoba . . . . . . . . . . . . . . . . . 2. mtvris marcvlis sicocxlisunarianoba . . . . . . . . . fertiloba . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . xelovnur skveb areze gaRiveba . . . . . . . . . . . . . . . mtvris marcvlis cxovelmyofelobis xangZlivoba . III. vazis qarTuli genotipebis mtvris marcvlis Camoyalibebisa da ganviTarebis institucionaluri analizi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . IV. wignSi gamoyenebuli terminebi . . . . . . . . . . . . . . . . gamoyenebuli literaturuli da eleqtronuli informaciis masalebi . . . . . . . . . . . . . . . . . . . . . . reziume . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. 4 11 20 20 21 26 30 30 35 45

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