Ia?iiaeueia aeaaeai?y iaoe
Oe?a?ie

?inoeooo ee?oeiii? a?ieia?? ? aaiaoe/ii? ?iaeaia???

Oa?iianueea Oaia?a Einoyioei?aia

OAeE 633.11 :
631.523 : 575

Ia?aaoaeiaa aaiiio i’yei? ioaieoe? (Triticum aestivum L)

aeey ?? aaiaoe/iiai aiae?co oa ?io?ia?an?? aai?a

Niaoe?aeuei?noue 03.00.15.-aaiaoeea

Aaoi?aoa?ao

aeena?oaoe?? ia caeiaoooy iaoeiaiai nooiaiy

aeieoi?a a?ieia?/ieo iaoe

Ee?a

1999

Aeena?oaoe??th ? ?oeiien.

?iaioo aeeiiaii a ?inoeooo? aa?iaeieia?? oa a?ioaoiieia?? OAAI.

Iaoeiaee eiinoeueoaio — Nic?iia Ieaen?e Ieaen?eiae/- aeieoi?
n?euenueeiainiiaea?nueeeo iaoe, i?ioani?, aeaaeai?e IAI Oe?a?ie oa
OAAI, caa?aeoaa/ eaai?aoi??? aeieia?/ii? aaiaoeee ?inoeoooo
aa?iaeieia?? oa a?ioaoiieia?? OAAI, aee?aeoi? ?inoeoooo aa?iaeieia?? oa
a?ioaoiieia?? OAAI.

Io?oe?ei? iiiiaioe:

aeieoi? a?ieia?/ieo iaoe, i?ioani?, aeaaeai?e OAAI Noaeueiao Aaeieueo
Oiie/, caa?aeoaa/ eaai?aoi??? niaoe?aeueii? aaiaoeee ioaieoe?
Naeaeoe?eii-aaiaoe/iiai ?inoeoooo OAAI

aeieoi? a?ieia?/ieo iaoe, i?ioani? Aiaa? Aiaoie?e Oaaei?iae/, caa?aeoaa/
eaai?aoi??? aaiaoeee ? naeaeoe?? ei?iiaeo eoeueoo? ?inoeoooo
caiea?ianoaa OAAI

aeieoi? a?ieia?/ieo iaoe, i?ioani? Oaaoeia ?ai? Aiaoie?eiae/ caa?aeoaa/
a?aeae?eo aaiaoe/ieo iniia aaoa?iceno ?inoeoooo o?c?ieia?? ?ineei ?
aaiaoeee IAI Oe?a?ie

I?ia?aeia onoaiiaa: Ee?anueeee Oi?aa?neoao ?i. O. Oaa/aiea,
eaoaae?a

caaaeueii? oa iieaeoey?ii? aaiaoeee.

Caoeno a?aeaoaeaoueny 16.09.1999?. i 13.00 aiaeei? ia can?aeaii?
niaoe?ae?ciaaii? a/aii? ?aaee Ae.26.202.01 i?e ?inoeooo? ee?oeiii?
a?ieia?? oa aaiaoe/ii? ?iaeaia??? IAI Oe?a?ie, 252143,Ee?a,
aoe.Caaieioiiai,148.

C aeena?oaoe??th iiaeia iciaeiieoenue a a?ae?ioaoe? ?inoeoooo
ee?oeiii? a?ieia?? oa aaiaoe/ii? ?iaeaia??? IAI Oe?a?ie, Ee?a, aoe.
Caaieioiiai, 148.

Aaoi?aoa?ao ?ic?neaii 12.08.1999 ?ieo

A/aiee nae?aoa?

niaoe?ae?ciaaii? a/aii? ?aaee

eaiaeeaeao a?ieia?/ieo iaoe
Oa?anaiei E.A.

CAAAEUeIA OA?AEOA?ENOEEA ?IAIOE

Aeooaeuei?noue oaie. Iioiaeaeaiiy i’yei? ioaieoe? (Triticum
aestivum L.) iia’ycaii c ia’?aeiaiiyi a iaeiiio aeiaaenaiei?aeiiio
aaiii?, AABBDD, o?ueio aaiii?a — AA (a?ae ioaieoe?-iaeiica?iyiee), BB
(aeaea?aei iaa?aeiia) ? DD (aa?eiin Oaooa). Aaaaea?oueny, ui iane?aeeii
oaei? no?oeoo?e aaiiio ? o?eie?eiaai?noue iaeiioeiieo aai?a a nenoaiao,
ui eiio?ieththoue ie?ai? iciaee. Oea oneeaaeith? ni?aa?aeiioaiiy
?ic/aieaiiy i?e aaiaoe/iiio aiae?cia? ioaieoe? ca aoaeue-yeeie iciaeaie
? iniaeeai ca iciaeaie c iaia?a?aiith i?ieea?noth —
e?euee?nieie. I?ae oei c on??? aaaaoiiai?oiino? iciae,
oniaaeeoaaiiy yeeo iio??aii ciaoe, iaeaaaeeea?oeie ? naia
e?euee?ni?, oiio ui aei oe??? eaoaai??? a?aeiinyoue a?eueo?noue
naeaeoe?eii-cia/oueo iciae.

?aeay ia?aaoaeoaaoe aaiii i’yei? ioaieoe? oaeei /eiii, uia
aaenaiei?aee a?ae??ciyeenue iaeei a?ae iaeiiai eeoa ca iaeiei c o?ueio
noaaaiii?a, oiaoi aeeiiaoe iiina?aaeeiaaio aeaiie?iei?aeecaoe?th
aaenaiei?aeii? ioaieoe?, iaeaaeaea ?.A.AEe-?iao. ?? aoei ?aae?ciaaii
aeey noaaaiiia D (AEe?ia,Oa?iianeay, 1987). Eiioeaio?aoe?y oaaae ia
noaaaiii? D iiynith?oueny aeaiia i?e/eiaie. Ii-ia?oa, c aeeth/aiiyi naia
oeueiai aaiiio o neeaae aeiaaenaiei?aea iia’ycothoue ieceo
aeanoeainoae i’yei? ioaieoe?, ui ?iaeyoue ?? iniiaiith oe?aiith
?ineeiith: aenie? oe?aiiaea?nuee? aeanoeaino? (Welsh, Hehn, 1964,
Kerber, Tipples, 1969, Dedio, Kaltsikes, 1972), ?icoe?aiiy a?aaeo
ca ?aooiie i?iieeiaiiy a i?ai?/i?o? oa iinooeea? ?aeiie (Halloran,
Boydell, 1967, Kuckuck, 1970, Ooeuiaeei,1972), a oaeiae aeaye?
iaaaoeai? iciaee: i?aeaeuaia ni?eiyoeea?noue aei a?eaieo
caoai?thaaiue (Kerber, Dyck, 1969, Ieaooiaa,1975), cieaeaiee
ai?no a?eea a ca?i? ? e?ceio a a?eeia? (Lawrence, 1958, Othoa?aa
e ae?., 1973), aaie a?a?eaeiiai iae?ico ? oei?ico (Kihara,
1951). Ii-ae?oaa, noaaaiii D aeyaeany ?aeeiei c o?ueio noaaaiii?a
ioaieoe?, A, A ? D, yeee aaeaeiny a?aeae?eeoe a?ae aeaio ?ioeo,
aeae?eeaoe oao?aiei?aeiee eiiiiiaio AAAA (Kerber, 1964, Kaltsikes et
al., 1968, Oa?iianeay, AEe?ia, 1979). E??i oeueiai, ie
ciiaee i?e?aeiaoe noaaaiii D ??cieo ni?o?a i’yei? ioaieoe?
aei iaeiiai ? oiai ae oao?aiei?aeo AAAA, io?eiaaoe aaenaiei?aei?
oi?ie i’yei? ioaieoe?, ui a?ae??ciythoueny iaeia a?ae iaeii?
noaaaiiiii D ? ye? iaeiaeia? a aaiiii?e /anoei? AA (AEe?ia,
Oa?iianeay, 1987). I?e no?auoaaii? oeeo oi?i any i?ieea?noue a
iiioeyoe?yo, ui ?ic/aieththoueny, a?aeaoaa?oueny ca ?aooiie aai?a
aaiiio D, oiaoi ? aeeiei?aeiee oei ?ic/aieaiiy. Oey ianoaaeia aea?
iiaeeea?noue aeei?enoaoe aeey aiae?co noaaaiiia D, ui ? o neeaae?
aaenaiei?aeiiai i?aai?cio, aoaeue-ye? iaoiaee eeane/iiai
aaiaoe/iiai aiae?co, a oiio /ene?, a?a?eaeieia?/iee.

Iniiaiei caaaeaiiyi oeeoiaaiaoeee i’yei? ioaieoe? ?
?ic?iaea iaoiae?a ?? iie?auaiiy ia aaie, ui eiio?ieththoue
ainiiaea?nueei-oe?ii? iciaee ye ca ?aooiie ?io?ia?an?? ie?aieo
aai?a, oae ? i?yiei eiino?othaaiiyi aaiioeio,aeei?enoiaoth/e
no/ani? iaoiaee aaiaoe/ii? ?iaeaia???. Aeey ae??oaiiy oaeiai
caaaeaiiy iio??aii aeia?a ciaoe aaiaoeeo i’yei? ioaieoe? ? iaoe
?ioi?iaoe?th i?i caeiiii??iino? oniaaeeoaaiiy iciae, ye? aaaeaii
aoei a ia?aaeaoe oe?e eoeueoo?? a?ae aeeei?ineeo ?iaee/?a oeyoii
?io?ia?an?? aai?a. Aeeiiaia aeine?aeaeaiiy i’yei? ioaieoe? ni?yiiaaii
ia aaiaoe/iee aiae?c iaei??? c? neeaaeiaeo /anoei ?? neeaaeiiai
aaiiia — noaaaiiio D ca ieceith ye?nieo ? e?euee?nieo iciae ?
ia ia?aaoaeiao oeueiai noaaaiiio oeyoii ?io?ia?aneaii? a?a?eaeecaoe??
c aeei?enoaiiyi iaoiae?a aaiiiii? ? o?iiiniiii? ?iaeaia???.

Ca’ycie aeena?oaoe?eii? ?iaioe c iaoeiaeie i?ia?aiaie, ieaiaie,
oaiaie. A aeena?oaoe?? ocaaaeueiaii ?acoeueoaoe
iaoeiai-aeine?aeieoeueei? ?iaioe, aeeiiaii? o a?aeiia?aeiino? c
oaiaoe/ieie ieaiaie i?ia?ai «?ic?iaea iaoiaeia oa oaoiieiaie o
naeaeoei?, ianiiieoeoai, ?aaeiiaiieiai? oa aa?iaeieiai? ia iniiai
aeei?enoaiiy iieaeoey?ii-aaiaoe/ieo ia?ea?ia, o?iiiniiii? oa
aaiaoe/ii? iiaeaia?i?», iiia? aea?ae?a?no?aoe?? 01930036999, «Oai?aoe/ii
iaa?oiooaaoe ? canoinoaaoe ia i?aeoeoe? caniae oa iaoiaee
aaiaoeee ? a?ioaoiieia?? aeey oe?eae ni?yiiaaiiai oi?ioaaiiy
aaiaoe/ii? eiiiiiaioe noaeeo aa?iaeinenoai», iiia?
aea?ae?a?no?aoe?? 0196U012975 a ?aieao i?iaeaie «Oai?aoe/ii
iaa?oiooaaoe ? canoinoaaoe ia i?aeoeoe? niiniae ? iaoiaee
aaiaoeee ? a?ioaoiieia?? c iaoith ni?yiiaaiiai oi?ioaaiiy aaiaoe/ii?
eiiiiiaioe noaeeo aa?iaeinenoai».

Iaoa ? caaea/? aeine?aeaeaiiy. Iaoith aeine?aeaeaiiy aoei
i?iaeaiiino?oaaoe oeyoe oa iaoiaee aeei?enoaiiy aaenaiei?aeieo oi?i
ioaieoe? c ia?aaoaeiaaiei aaiiiii aeey aaiiiiiai aiae?co i’yei?
ioaieoe?, aaiaoe/iiai aiae?co ?? noaaaiiio D oa ?io?ia?an?? aaiaoe/iiai
iaoa??aeo a?ae ?? aeeei?ineeo ?iaee/?a. Aeey aeinyaiaiiy iaoe aoei
iinoaaeaii oa ae??oaii oae? caaea/i: 1. Aeei?enoiaoth/e
aaiiiii-aeiaeai? oi?ie i’yei? ioaieoe?, ui iathoue iaeiaeiaee
oao?aiei?aeiee eiiiiiaio aaiiio AA ? ??ci? noaaaiiie D, noai?eoe
a?a?eaeiee iaoa??ae, iaiao?aeiee aeey aeeiiaiiy aaiiiiiai aiae?co
aeaio /anoei aeiaaenaiei?aeiiai aaiiio, AA oa D ? aaiaoe/iiai
aiae?co noaaaiiia D. 2. Ia noai?aiiio a?a?eaeiiio iaoa??ae?
aea/eoe aianie ie?aieo /anoei aaiiio i’yei? ioaieoe? (AA ? D) o
eiio?iethaaiiy ?icaeoeo aeayeeo e?euee?nieo iciae ? aeeiiaoe
a?iiao?e/iee aaiaoe/iee aiae?c noaaaiiio D i’yei? ioaieoe? 3.
?aeaioeo?eoaaoe aaie noaaaiiio D, ui ?o iiaeia aeei?enoaoe ye
aaie-ia?ea?e ie?aieo o?iiinii oeueiai noaaaiiio. 4. Aeei?enoiaoth/e
?aeaioeo?eiaai? aaie-ia?ea?e o?iiinii noaaaiiio D oa oaiiiai c/aieaiiy,
cae?enieoe o?iiiniiio eieae?caoe?th aieiaieo aai?a i??ieo
iciae. 5.Aeei?enooth/e aaie-ia?ea?e o?iiinii ??cieo aiiaieia?/ieo
a?oi Triticinae, eeaneo?eoaaoe io?eiai? iaie ?ai?oa aaenaiei?aei? e?i??
i’yei? ioaieoe? c i?aeaaiiiieie cai?uaiiyie o?iiinii ca iayai?noth a
ieo /oaeeiieo ?iiinii aai ?o ae?eyiie iaaii? aiiaieia?/ii? a?oie,
caeeaaoe oei naiei i?aea?oioy aeey aeei?enoaiiy e?i?e o
aaiaoe/iiio aiae?cia? i?aaenoaaiee?a Triticinae oa o naeaeoe?? ye
e?i?e-aeiii??a /oaeeiieo aai?a, ui eiio?ieththoue aaaeai? iciaee.

Iaoeiaa iiaecia. 1. Aia?oa a na?o? cae?eniaii aaiaoe/iee aiae?c
noa-aaiiio D i’yei? ioaieoe? o?ueio ??cieo ?? ni?o?a ? aeaio
??ciiaeae?a aeeei?ineiai i?aaaoueea oeueiai noaaaiiio, ui
ciaoiaeeoueny o neeaae? iiai?noth caaeainiaaieo ca e?euee?noth
o?iiinii ? a?aeiioaiith yae?i-oeeoiieacia ?ineeiieo oi?i 2. Aia?oa a
na?o? aei??yii aianie aeaio ??cieo a aaiethoe?eiiio ieai?
neeaaeiaeo /anoei aaenaiei?aeiiai aaiiio ioaieoe?: AA ? D
— a ?icaeoie e?euee?nieo iciae ioaieoe?, ia?aaaaeii naeaeoe?eii
cia/oueo, i iieacaii a?aei?iiino? a iai?yieo ae?? ??cieo /anoei
aaiiio. 3. ?aeaioeo?eiaaii aeae?eueea iiaeo aai?a i’yei?
ioaieoe?, ui eiio?ieththoue ii?oieia?/i? iciaee eieina, ? e?eueea
aieiaieo aai?a, ui aiie aa?ooue o/anoue a eiio?ie? e?euee?nieo iciae
eieina ? noaaea. 4. Aeiaaaeaii aoaeoeai?noue aeei?enoaiiy
aaiiiii-cai?uaieo aaenaiei?ae?a ioaieoe? c aaiiiiith no?oeoo?ith AAAAOO
aeey noai?aiiy /oaeeiii-cai?uaieo e?i?e i’yei? ioaieoe? c iaeeia?ieie
oa iiiaeeiieie cai?uaiiyie o?iiinii noaaaiiio D ia o?iiiniio
noaaaiiio O aaiiiii-cai?uaii? oi?ie. 5. Anoaiiaeaii ia?aaaaeio
caeaaei?noue oni?oiiai io?eiaiiy /oaeeiii-ioaie/ieo cai?uaiue a?ae
iaeaaeiino? e?eoe/ii? o?iiiniie aei iaaii? aiiaieia?/ii? a?oie o
ii??aiyii? c ?? iaeaaei?noth aeaeo-aeiii?o aaiaoe/iiai iaoa??aeo, ui
?io?ia?ano?oueny.

I?aeoe/ia cia/ou?noue. 1. ?ic?iaeaii iaoiae aaiiiiiai aiae?co
aeiii-e?iei?ae?a ia i?eeeaae? i’yei? ioaieoe?. A?i i?eaeaoiee aeey
aiae?co ??ciiai oeio ?ineeiiiai iaoa??aeo, eiee o ii??aithaaieo
aaiioeiao iiaeia aeae?eeoe aea? neeaaeia? /anoeie, iaeia c yeeo –
caaaeueia aeey aeae?eueeio aeine?aeaeoaaieo oi?i. Iaoiae iiaeia
aeei?enoaoe aeey ?ioeo aeiiie?iei?aeieo aeae?a ?ineei. 2. ?a-coeueoaoe
aaiiiiiai aiae?co iathoue cia/aiiy aeey iioei?caoe?? naeaeoe?eiiai
i?i-oeano oa iiaeooue aooe aeei?enoai? naeaeoe?iia?aie i?e aeai??
eiiiiiaio?a no?a-uoaaiiy. 3. ?ic?iaeaii niin?a o?iiiniiii?
eieae?caoe?? aieiaieo aai?a i??ieo iciae, yeee a?oioo?oueny ia
aeei?enoaii? ?acoeueoao?a iiia?aaeiueiai aaiaoe/iiai aiae?co iciaee,
ui aea/a?oueny, iaoiaeaie a?iiao?e/ii? aaiaoeee. Iaoiae iiaea aooe
aeei?enoaii aeey ??ciiai oeio aoiei?aeiiai ?ineeiiiai iaoa??aeo,
eiee a?a?eaee ?ic/aieththoueny ia aeeiei?aeiiio ??ai?, a ie?ai?
o?iiiniie aaiiio iathoue ?aeaioeo?eiaai? aaie-ia?ea?e. 4. Noai?aii
iiioeyoe?? F6 ?aeiia?iaioii-?ia?aaeieo e?i?e a?ae no?auoaaiiy
aaiiiii-aeiaeaieo oi?i ca ae?aeaeueiith noaiith. Aiie
a?ae??ciythoueny a?ae on?o iiioeyoe?e ??E i’yei? ioaieoe? oei, ui
o?enothoue i?ieea?noue aaenaiei?ae?a, ui ?ic/aieththoueny eeoa ca
noaaaiiiii D, oiaoi ia aeeiei?aeiiio ??ai?. Iiaeooue
aeei?enoiaoaaoenue aeey aaiaoe/iiai aiae?co iciae, ca yeeie
eiiiiiaioe no?auoaaiiy a?ae??ciythouenyiaeei a?ae iaeiiai, ? ?
oe?iiei aaiaoe/iei iaoa??aeii aeey ?aeaioeo?eaoe?? ? ea?ooaaiiy
QTL. 5. Aaenaiei?aei? e?i?? ioaieoe? c o?iiiniiaie ? o?aineieaoe?yie
a?ae Aegilops umbellulata, Ae. speltoides, Ae. sharonensis, Agropyron
glaucum oa?aeoa?ecothoueny no?ee?noth aei a?eaieo caoai?thaaiue
ioaieoe?, aeia?ith ceiino?ee?noth, i?aeaeuaiei o ii??aiyii? c i’yeith
ioaieoeath ai?noii a?eea. Aiie aac iaiaaeaiue no?auothoueny c ni?oaie
i’yei? ioaieoe? ? iiaeooue aooe aeei?enoai? ye aeiii?e aai?a, ui
eiio?ieththoue ia?a?aoiaai? iciaee, a naeaeoe?? ioaieoe?. 6. Caaaeueio
noaio aeeiiaiiy aeaiiai aeine?aeaeaiiy ? io?eiai? ?acoeueoaoe
iiaeia aeeth/eoe a eo?n aaiaoe/iiai aiae?co, ui /eoa?oueny nooaeaioai
c? niaoe?aeueiino? aaiaoeea, aeey ?ethno?aoe?? aaaeeeaino?
a?aeiia?aeiino? iaoiae?a aeine?aeaeaiiy iaoa??aeiai, aei yeiai
aiie canoiniaothoueny.

Ai?iaaoeiy ?iaioe. ?acoeueoaoe aeine?aeaeaiiy aeiiia?aeaeenue,
aai aoee i?aaenoaaeai? ? io?eiaee iiceoeaio ioe?ieo ia:
eiioa?aioe?? ?a?iiaenueeiai ia’?aeiaiiy aeine?aeaeaiue c aiaoiei?ae??
ioaieoe?, EWAC, Oai?ueia, Ia?oiiaaoa?,1981,I?aeia?iaei?e iaoeia?e
eiioa?aioe?? a/aieo e?a?i-/eai?a ?AA, Iaeana, 1981, Ananithci?e
ia?aae? c a?aeaeaeaii? a?a?eaeecaoe?? ?ineei ? oaa?ei, Iineaa,
1981, I ?in?eneiio neiiic?oi? «Iiaua iaoiaeu aei-oaoiieiaee ?anoaiee»,
Ioueii-ia-Ioe?, 18 – 20 o?aaiy, 1993, O Ana?in?enueeiio neiiic?oi?
c a?ioaoiieia?? ?ineei a Ioueii-ia-Ioe?, 22 –27 o?aaiy 1995,
I?aeia?iaei?e eiioa?aioe?? «Iieaeoey?ii-aaiaoe/i? ia?ea?e ?
naeaeoe?y ?ineei», Ee?a,10 — 13 o?aaiy 1994, I?aeia?iaeiiio
neiiic?oi? a i. Oa?eia?, 2-4 aeiaoiy. 1996 ?., I?aeia?iaei?e
eiioa?aioe?? «Aeooaeueiua i?iaeaiu a ?anoaieaaiaenoaa, aeeaioiiaiaenoaa
e aaoa?eia?ee», Iineaa, 30– 31 aeiaoiy 1996, I?aeia?iaei?e
eiioa?aioe?? «Iieaeoey?ii-aaiaoe/aneea ia?ea?u ?anoaiee», sseoa, 11 —
15 eenoiiaaea, 1996, I?aeia?iaei?e eiioa?aioe??
«Iieaeoey?ii-aaiaoe/aneea ia?ea?e ?anoaiee e aeeaioieo», Ee?a, 29-30
o?aaiy 1997.

Iniaenoee aianie aaoi?a iieyaa? o noai?aii? iniiaii?
eiioeaioe?? aeine?aeaeaiiy, ui aeeiiaii, aecia/aii? caaea/ ?
iaoiae?a aeey ?o ae??oaiiy, i?iaaaeaii? ia?aaaaeii? /anoeie
aenia?eiaioaeueii? ?iaioe, noai?aii? ii-aeaeae ? ia?iaoei io?eiaieo
aeaieo, aiae?cia? ? ocaaaeueiaii? ?acoeueoao?a.

Ioaeeeaoei? ?acoeueoaoia aeineiaeaeaiue. Ca iaoa??aeaie
aeena?oaoe?? iioa-e?eiaaii 37 ?ia?o. Na?aae ieo 28 noaoae o
oaoiaeo iaoeiaeo aeaeaiiyo, 6 c yeeo iaienaii aac ni?aaaoi??a,
2 noaooi o cai?ieeao i?aoeue oa 7 oac, ai?uaieo a iaoa??aeao
i?aeia?iaeieo eiioa?aioe?e ? neiiic?oi?a.

No?oeoo?a aeena?oaoei?. Aeena?oaoe?y i?noeoue anooi, iaeyae
e?oa?aoo?e (neeaaea?oueny c aeaio ?icae?e?a), aeeeaaeaiiy
aeanieo aenia?eiaio?a — i’youe ?icae?e?a, ui aeeth/athoue iien
iaoa??ae?a ? iaoiae?a. A ie?aiee ?icae?e aeia-naii iaaiai?aiiy
aieiaieo ?acoeueoao?a, ? caeeth/aiiy ? aeniiaee. ?iaioo aeeeaaeaii
ia 417 noi?. iaoeiiieniiai oaenoo, aeeth/ath/e 69 oaaeeoeue ? 70
iaethie?a. Ia?ae?e oeeoiaaii? e?oa?aoo?e neeaaeathoue 626 aeaea?ae. 50
oaaeeoeue ?icoaoiaaii o aeiaeaoeao.

INIIAIA *ANOEIA

?ICAe?E 1. Aaiaoe/iee aiae?c i’yei? ioaieoe? — aeinyaioo?

?acoeueoaoe ? i?iaeaie (iaeyae e?oa?aoo?e)

?icaeyiooi oae? ieoaiiy no/aniiai noaio aaiaoe/ieo
aeine?aeaeaiue ioa-ieoe?: no?oeoo?a aaiiio ioaieoe?, aoaie aea/aiiy
?? aaiaoeee c aeei?enoaiiyi aiaoiei?aeieo na??e i’yei? ioaieoe?,
noaiiaeaiiy iaoiaeo iieaeoey?ii-aaiaoe/ieo ia?ea??a a canoinoaaii?
aei aaiaoe/ieo aeine?aeaeaiue ioaieoe?. Oeae iaoiae
ioa?aeoa?eciaaii ye iaea?eueo ia?niaeoeaiee o aaiaoe/ieo
aeine?aeaeaiiyo ioaieoe?, a oiio /ene? i ca e?euee?nieie iciaeaie.
?ice?eoi iiiyooy aeaiie?iei?aeecaoe?? aeiaaenaiei?aeii?
ioaieoe?, i?yii? oa iiina?aaeeiaaii?, i?aee?aneaii ?? cia/aiiy aeey
aaiaoe/iiai aiae?co. Aeae?eaii /ioe?e aoaie aaiaoe/iiai
aeine?aeaeaiiy aeaeo T.aestivum: 1. Anoaiiaeaiiy no?oeoo?e aaiiio,
a?aeiioaiue aiiaieia?? i?ae o?iiiniiaie noaaaiii?a ? ii-oiaeaeaiiy
inoaii?o. Iniiaiee iaoiae aeine?aeaeaiue — oeeoieia?/iee, aieiaiee
i?aeoe/iee ?acoeueoao — /?oea ?icoi?iiy iecueei? ?ic??ciyth/i?
caeaoiino? eeane/iiai a?a?eaeieia?/iiai aiae?co aeey aaiaoe/iiai
aiae?co ioaieoe? (Kihara, 1937, Okomoto, 1962, Chapman, Riley, 1966,
Oeeeii/aiei, 1979). 2. Noai?aiiy iiaiiai eiiieaeoo na??e an?o iiaeeeaeo
aiaoiei?ae?a ia o?iiiniiiiio ? iea/iaiio ??ai? ia ni?o? i’yei?
ioaieoe? *aei?c Ni?eia, ?ic?iaea iaoiae?a aiaoiei?aeiiai aiae?co
ioaieoe? ? iaoiae?a oe?eani?yiiaaii? cai?ie iaaieo ioaie/ieo o?iiinii
ia aiiaieia?/i? o?iiiniie ?ioeo aeae?a ? ?iae?a i?aeo?eae
Triticinae. Iniiaiee iaoiae aeine?aeaeaiiy — oeeoieia?/iee. Iniiaiee
?acoeueoao — o?iiiniiia eieae?caoe?y «aieiaieo» aai?a, ui
a?aeiia?aeathoue ca ?icaeoie aeaeiaeo iciae ??cieo aaenaiei?aeieo
aeae?a ioaieoe?, aeniiaie i?i o/anoue an?o o?iiinii ioaieoe?
o aaiaoe/iiio eiio?iei aoaeue-yeeo ?? e?euee?nieo iciae ?
iaeiie/aiiy aaee/aciiai ianyao ?ineeiiiai iaoa??aeo, ui yaey?
niaith i?iaeoeoe ?io?ia?aneaii? a?a-?eaeecaoe?? i’yei? ioaieoe?
c ?? ?iaee/aie (Unrau, 1950, Sears, 1953, 1954, 1972, Kuspira, Unrau,
1959, Zeller, Fischbeck, 1974). 3. Aeei?enoaiiy ioe?-oao?aniiieo
e?i?e ioaieoe? aeey eieae?caoe?? aai?a, ui eiio?ieththoue neioac
a?ee?a. Iaoiae aeine?aeaeaiue — aeaeo?ioi?ac a?ee?a. Iniiaiee
i?aeoe/iee ?acoeueoao – noaiiaeaiiy iaoiaeo iieaeoey?ii-aaiaoe/ieo
ia?ea??a, «a?coae?caoe?y» an?o 42 iea/ae o?iiinii ioaieoe? ca
?aooiie eieae?caoe?? ia ieo a?eeiaeo aai?a-ia?ea??a ?
aeaiiino?aoe?y iiaeeeainoae iiaiai iaoiaeo aeey iieaaoaiiy
?ia?o, iia’ycaieo c ?io?ia?an??th /oaeeiieo aai?a a aaiii ioaieoe?
(Barber et al., 1968, McIntosh et al., 1998). 4. Aeoeaio oaaae
aeine?aeiee?a — ia iieaeoee AeIE, iie?ii?o?ci yeeo aeei?enoaii aeey
?ic?iaee aeaio oei?a iieaeoey?ieo ia?ea??a: RFLP-ia?ea?e oa
PCR-ia?ea?e ? /eneai? ?o aa??aioe. Iniiaiee iaoiae —
a?io?i?/i? iaoiaeeee aeey ?iaioe c AeIE. I?aeoe/iee ?acoeueoao
— noai?aiiy iieaeoey?ii-aaiaoe/ieo RFLP-ea?o aeey an?o
naie aiiaieia?/ieo a?oi o?iiinii ioaieoe? ? aeaeio?eo
nii??aeiaieo aeae?a oa aeaiiino?aoe?y iiaeeeaino? «aoeeaooaaiiy»
iieaeoey?iei ia?ea?ii i?aeoe/ii aoaeue-yeiai aaio (iciaee), eiee
oea iaiao?aeii aeey aaiaoe/iiai aiae?co ? ne?ei?iao
naeaeoe?eiiai iaoa??aeo; oi?ioaaiiy iiiyooy QTL (Young, Tanksley,
1989, Devos et al., 1992, 1993, Michelmore et al., 1991, Paterson et
al., 1988)). Ia aeoieo aaoi?a aeena?oaoe??, ia?o? o?e aoaie ?icaeoeo
aaiaoe/ieo aeine?aeaeaiue ioaieoe? oa?aeoa?ecoaaeenue iaaiith
aa?iii??th i?ae iaoa??aeii, ui aiae?co?oueny, ? iaoiaeaie, ui
aeei?enoiaothoueny aeey aiae?co. Ciino /aoaa?oiai aoaio —
aeaiiino?aoe?y a?ae?eao iaoiae?a aaiaoe/iiai aiae?co a?ae
iaoa??aeo, yeee iiaea aooe cai?iiiiiaaiei aeey ?aae?caoe?? iiaeeeainoae
iaoiaea. C oeei iia’ycaii caaino?aiiy oe?eaaino? aei iiooeo
aai noai?aiiy ?ineeiiiai iaoa??aeo, ia yeiio iiaeia aoei a aea/eoe
ni?eueii c iae?iiciaeaie ? iie?ii?o?ci iieaeoe AeIE.

?ICAe?E 2. Aaiiiiee aiae?c i’yei? ioaieoe?

Aeine?aeaeaiiy, ni?yiiaaia ia aeyaeaiiy aaiaoe/iiai aianeo ie?aieo
noaaaiii?a i’yei? ioaieoe? o oi?ioaaiiy oaiioei?/ieo na?aaei?o
cia/aiue ca e?euee?nieie iciaeaie, aeeiiaii ia niaoe?aeueii
noai?aieo ooo/ieo oi?iao aaenaiei?aeii? ioaieoe?, aaiiiii-aeiaeaieo
aaenaiei?aeao. Oe? oi?ie iathoue iaeiaeiao aaiiiio /anoeio AA ?
a?ae??ciythoueny iaeia a?ae iaeii? eeoa noaaaiiiii D. Aea/aiee ?ineeiiee
iaoa??ae: 1) icei? ni?oe i’yei? ioaieoe? T.aestivum L. Eaaeac (EA)
? Aeuea?aeoi 114 (AE); 2) e?i?y iceii? oaa?aei? ioaieoe?
T.durum Desf. var. muticitalicum Dorof. et Filat. (I?); 3)
i’youe aaiiiii-aeiaeaieo oi?i c iaeiei ? oei aea noaaaiiiii AA, yeee
? aaiiiii e?i?? I?, ? ??cieie noaaaiiiaie D: a?ae ni?oo
Eaeueyiniia (EN) o oi?ie Den, a?ae ni?oo EA — o Dea, a?ae ni?oo AE
— o Dae, a?ae Aegilops tauschii (Coss.) Schmal. var.
strangulata (e-28) o oi?ie Dt28 oa var. eusquarrosa (e-346) o
oi?ie Dt346; 4) F1 a?ae no?auoaaiiy ia?a?aoiaaieo oi?i ca
ae?aeaeueiith noaiith aac ?aoeei?ie?a. Aea/aii iciaee: aeaoa
eiein?iiy (e?euee?noue aei?a a?ae 1-ai o?aaiy), aeiaaeeia noaaea
(AeN, ni), aeiaaeeia aa?oiueiai i?aeaocey (AeI, ni), aeiaaeeia
eieina (AeE, ii), e?euee?noue eieine?a o aieiaiiio eiein? (EEE,
oo.), aeiaaeeia (AeE, ni) ? oe?eia (OE, ii) eenoea, ae?aiao?
noaaea i?ae eieinii (Ae, ii), e?euee?noue (ECE, oo.) ? iana
ca?ai (ICE, a) c aieiaiiai eieina, iana iaeiiai ca?ia (IIC, ia).

A?aeiia?aei?noue ?iciiae?eo ?ineei ca oaiioei?/ieie eeanaie o
aea/aieo iiioeyoe?yo ca eiaeiith c iciae ii?iaeueiiio caeiiiai aoei
anoaiiaeaii. Noaiaea?oia noaoenoe/ia ia?iaea aeaieo oa ?ic?aooiee c
aeei?enoaiiyi aea?aieo e?eoa???a ? iaoiae?a aeeiiaii ca
aeiiiiiaith iaeaoo noaoenoe/ieo i?ia?ai QUATTRO PRO, aa?n?y 4.
Aeey aeyaeaiiy aaiaoe/ieo aoaeo?a noaaaiii?a D ? AA
i’yei? ioaieoe? noiniaii iecee aea/aieo iciae aeei?enoaee
eiioeaiooaeueio iniiao ? oa?i?iieia?th a?iiao?e/ii? aaiaoeee
(Mather, Jinks, 1971). Aaiii aaenaiei?aeii? ioaieoe?
?icaeyaeaee ye noeoii?noue aeaio aaiaoe/ieo iaeeieoeue, D ? AA. A
na?aaeei? eiaeii? c oeeo iaeeieoeue a?aeiioaiiy aoaeothoueny ye
i?ae aeaeyie iaeiiai aaia — aaeeoeai?noue [d] aai aeii?ioaaiiy
[h]. I?ae iaeeieoeyie iiaeooue iaoe i?noea cae/aei? i?aeaaii?
aca?iiae?? — ??ci? niieo/aiiy aoaeo?a [i] (aca?iiae?y
aiiiceaio), [j] (aca?iiae?y aiiiceaio c aaoa?iceaioaie), [l]
(aca?iiae?y aaoa?iceaio c aaoa?iceaioaie).

Aeo?aeiei aoei i?eiouaiiy i?i aaeeoeaii-aeii?iaioio ae?th
aaiii?a AB oa D. Aaeeoeaii-aeii?iaioio iiaeaeue iaiiaithaaee
?acoeueoaoaie aea/aiiy naie aaouee?anueeeo oi?i ? 21 a?a?eaea i?ae
ieie (iae. 1A) ca eiaeiith c 11 e?euee?nieo iciae. Iiaeaeue
aeeth/aea ia?aiao? m — na?aaei? cia/aiiy, [P] — aoaeo
iei?aeiino? (ine?eueee iaeia c aaouee?anueeeo oi?i aoea ia
aaena-, a oao?aiei?aeii, a io?eiai? ca ?? o/anoth a?a?eaee
— iaioaiei?aeaie), o?noue ia?aiao??a [d], aeaa’youe ia?aiao??a
[h] aeey ??cieo noaaaiii?a D oa AA. *eneia? cia/aiiy
ia?aiao??a aecia/aee, aeei?enoiaoth/e oano ia’?aeiaiiai oeaethaaiiy
Eaaaee? (oeeo. ca Mather, Jinks, 1971). Aii??e/i? oa oai?aoe/i?
na?aaei? cia/aiiy ii??aithaaee iaoiaeii c2.. Iaaaeaeaaoi?noue
aaeeoeaii-aeii?iaioii? iiaeae? aeine?aeiei aeaiei aoei iieacaii
aeey eiaeii? c aea/aieo iciae. ?icaeyae neeaaeiaeo /anoei aaee/ei c2
aeyaea, ui ia?aaeuaiiy inoaii?ie e?eoe/iiai ??aiy caaaeaee
a?aeaoaa?oueny ca ?aooiie iaeieo ? oeo aea n?iae — i?e niieo/aii?
noaaaiii?a AB ? D, eiee aaoa?iceaioiee iaeei c ieo ([j], a iaoiio
aeiaaeeo oea 8 ia?aiao??a) aai iaeaeaa (aca?iiae?y [l], 7
ia?aiao??a). Iayaia e?euee?noue n?iae ia aeaea ciiae
aeiaeaoeiai aaanoe aei iiaii? iiaeae? an? 15 ia?aiao??a
aca?iiae??. Aeey i?iaeiaaeaiiy aiae?co oanooaaee ie?aii aea? iiaeae?:
1) c 12 ia?aiao?aie oa 15 n?i’yie, ui a?aoiaothoue ?ica?aeiino?
eeoa ca noaaaiiiii D ia oe? iaeiiai ? oiai ae noaaaiiia AA (iae.
1A); 2) c 5 ia?aiao?aie oa o?noueia n?i’yie c ??cieie noaaaiiiaie
AA ia oei iaeiiai e oiai ae D (iae. 1A). Ia?oa
aaeeoeaii-aeii?iaioia iiaeaeue aeyaeeany aaeaeaaoiith aeey an?o iciae,
oiaoi aaiaoe/i? ?ica?aeiino? i?ae ??cieie noaaaiiiaie D
iiaeia iienaoe, i?eeiath/e aei oaaae eeoa aaeeoeai? ?
aeii?iaioi? aoaeoe. I?e ?ica?aeiino? noaaaiii?a AA ia oe? iaeiiai
? oiai ae noaaaiiio D aaeeoeaii-aeii?iaioia iiaeaeue aoea aaeaeaaoiith
eeoa o e?eueeio aeiaaeeao. Oea iiaea aooe iia’ycaii c iayai?noth
aca?iiae?? /anoeie AA c noaaaiiiii D aai coiiaeaii oao?aiei?aeiith
i?e?iaeith /anoeie AA. Aaiiiia no?oeoo?a eiiiiiaio?a no?auoaaiiy
(iae. 1A) aeaea ciiao aecia/eoe eeoa aca?iiae?th [j].
Aaeaeaaoi?noue iiaeae? c o?aooaaiiyi oeueiai ia?aiao?o aoei aeiaaaeaii
o a?eueoino? aeiaaee?a.

A EA
AE

.
.

Dt28 .
. I?

DAE.
. DEN

.

DEA

??ci? noaaaiiie AA oa D (iiaia noaia no?auoaaiue)

A I? . . DEN A
I?. . EA(AE)

Dt28. .DEA

.
.

DAE
DEA(AE)

A?ae??ciythoueny eeoa noaaaiiie D A?ae??ciythoueny eeoa
noaaaiiie AA

Iae. 1. Noaie aeai?o n?iae aeey iiaoaeiae ??cieo iiaeaeae:

A — 7 eiiiiiaio?a no?auoaaiiy ? 21 a?a?eae F1 ca ae?aeaeueiith noaiith
aac ?aoeei?ie?a; A — 5 eiiiiiaio?a no?auoaaiiy ? 10
a?a?eae?a F1 i?ae ieie; A — 3 eiiiiiaioe no?auoaaiiy ? 3 a?a?eaee
F1

Aaiiiiee aiae?c i’yei? ioaieoe? ca noaaaiiiaie D ? AA
aeyaea aeaye? caaaeuei? ?ene, aeanoea? eiaeiiio c ieo (iae.
2): noaaaiii D i?eai?n aei aaenaiei?aeo aaie a?eueo ?aiiueiai
aeeieiooaaiiy, ciaioaiiy e?eueeino? eieine?a o eiein? ?
e?eueeino? ca?ai o iueiio oa aaie ca?eueoaiiy aeiaaeeie ei-eina ?
ca?eueoaiiy iane iaeiiai ca?ia. Ca aaaaoaoeaieie iciaeaie aieea
noa-aaiiio D, ui iaeaaeeoue ni?oai i’yei? ioaieoe?, i?iyaey?oueny o
ca?eueoaii? na?aaei?o cia/aiue oeeo iciae, a c D a?ae aa?eiino — a
ciaioaii?. Aaeeoeaiee aoaeo noaaaiiio AA ni?yiiaaii o i?ioeeaaeiee
a?e o ii??aiyii? c ae??th D aaiiio aeey iciae aeaoa eiein?iiy,
e?euee?noue eieine?a o eiein? oa iana iae-iiai ca?ia ? an?o
aaaaoaoeaieo iciae ?ineeie.

Aaiiiiee aiae?c ii?icino?eeino? i’yei? ioaieoe? aeeiiaii i?e
ii??a-iyii? i’yoe noaaaiii?a D oa o?ueio noaaaiii?a AA. Aaie
noaaaiiio D oa?ae-oa?ecothoueny aeii?iaioiei, a aaie noaaaiiio AA —
aaeeoeaiei aoaeoii i?ae-aeaeueii? aca?iiae??. I?aeaaiiiia aca?iiae?y
aaeeoeaia. Noaaaiii AA ?iaeoue a?eueo?e aianie o ii?icino?ee?noue
aaenaiei?aeo o ii??aiyii? c noaaaiiiii D oiai ae ni?oo ioaieoe?:
Aae>ABea>Dae>Dea>Dt346>ABie>Dt28.

?ICAe?E 3. Aaiaoe/iee aiae?c noaaaiiio D i’yei?

ioaieoe? ca e?euee?nieie iciaeaie

3.1. Ae?aeaeueiee aiae?c a?a?eae?a ia?oiai iieie?iiy.
Aeine?aeaeoaaee aaiiiii-aeiaeai? oi?ie Den, Dea, Dae, Dt28 oa
Dt346 ? F1 a?ae no?auoaaiue oeeo oi?i ca ae?aeaeueiith noaiith
aac ?aoeei?ie?a (iae. 3). Aea/ai? iciaee — o? ae, ui ? a ?icae?e?
2 ca aeeeth/aiiyi aeaoe eiein?iiy ? ae?aiao?o noaaea. ?acoeueoaoe
ioe?ithaaiiy ?ineei aiae?coaaee a aeaa aoaie: 1)
aeenia?n?eiee oa ?aa?a-n?eiee aiae?c aa??ain-eiaa??ain aeey
ia?aa??ee aaeaeaaoiino? aaeeoeaii-aeii?-iaioii? iiaeae?; 2) o
aeiaaeeo aaeaeaaoiino? oe??? iiaeae? i?iaiaeeany aeen-ia?n?eiee
aiae?c aeaieo ioe?ithaaiiy c ianooiiei ?iceeaaeaiiyi
aeenia?n?? ia eiiiiiaioe. Aiae?c ae?aeaeueieo oaaeeoeue aeeiiaii
ca Oaeiaiii (Hayman, 1954), Aenaeai oa Aeaeiiniiii (Aksel,
Jonson, 1963). ?acoeueoaoe ocaaaeueiaii a oaae. 1 oa 2.

1989 oa 1990 ?iee 1993 ??e
1996 ??e

Den Dea Dea
Den Dea

Dt346 Den Dae

Dae Dt28
Dae Dt28

Dt28

Iae. 3. Noaie no?auoaaiiy aaiiiii-aeiaeaieo oi?i.

? eeoa aeaa caaaeueieo aeey an?o aea/aieo iciae aeniiaee
i?i oa?aeoa? ?o aaiaoe/iiai eiio?ieth: ia? i?noea o/anoue aai?a
ye c aaeeoeaiei, oae ? c aeii?iaioiei oeiii
i?aeaeaeueii? ae??, ? e?euee?noue aai?a, ui ca?eueoothoue
/enaeueiee ae?ac iciaee (u), i?aeoe/ii i?eiee ia aei??aith?
e?eueeino? aai?a, ui ciaioothoue oea cia/aiiy (v). Ca aanue
ia??iae aeine?aeaeaiue aoei aea/aii caaaeii 35 eiiieaen?a
«??e aeine?aeaeaiiy-iciaea». Iciaee noaaea, eenoea, a oaeiae
iciaea «aeaoa eiein?iiy» ? «aei?noe?noue eonee» oiiaii
a?aeian-ee aei aaaaoaoeaieo iciae (anueiai 18 eiiieaen?a) ?
a?aeae?eeee ?? a?ae iciae eieina ye iaea?eueo iii?oieo
eiiiiiaio?a o?iaeaeiino? — iana iaeiiai ca?-ia, aeiaaeeia
eieina, e?euee?noue eieine?a o eiein?, e?euee?noue ca?ai c
eiei-na, iana ca?ia c eieina (17 eiiieaen?a). C 18 eiiieaen?a
ca o/anoth aa-aaoaoeaieo iciae 14 iiaee aooe iienai? ca
aeiiiiiaith aaeeoeaii-aei-i?iaioii? iiaeae? aaiaoe/iiai eiio?ieth,
aea eeoa aeey 7 c 17 eiiieaen?a ca iciaeaie i?iaeoeoeaiino? aoea
aaeaeaaoiith oey ae iiaeaeue. Oiaoi aca?iiae?y i?ae aaiaie /ano?oa
ia? i?noea a eiio?ie? iciae i?iaeoeoeaiino?, i?ae iciae
aaaaoaoeaieo. Na?aae aai?a, ui eiio?ieththoue iaeaeaa oeie
iciae, a aaia-oe/iiio ioe? aea/aieo oi?i ia?aaaaeathoue aaie, ca
iacia/ieie aeeeth/aiiyie, c ?aoeaneaiei oeiii i?aeaeaeueii? aca?iiae??.
Iai?yiie ae?? aeii?iaioieo aai?a a?ae??ciy?oueny
ianoaa?euei?noth a ??ci? ?iee aeine?aeaeaiiy aeey o?ueio
iciae: aeiaaeeia noaaea, aeiaaeeia ? oe?eia eenoea. ? iaeneiaeueia
e?euee?noue aeii?iaioieo aeae?a caaaeaee aoea i?eoaiaiia oiio
eiiiiiaioo no?auoaaiiy, oaiioei yeiai a?aeiia?aeaa iai?yieo ae??
aeii?iaioieo aai?a.

3.2.Aaiaoe/iee aiae?c ca aeaieie ioe?ithaaiiy F1, F2 oa
aaee?in?a aei eiaeiiai c aaouee?a. Eiiiiiaioe no?auoaaiiy oa noaie
io?eiaiiy a?a?eae?a iaaaaeaii ia iae. 3. A 1989 ?ioe? aeey
an?o eiia?iaoe?e no?auoaaiiy aea/aii a?a?eaee ia?oiai ? ae?oaiai
iieie?iue, a on? ?io? ?iee — F1, F2 oa aaee?ine ia?oiai iieie?iiy aei
eiaeiiai c aaouee?a. Aea/ai? iciaee — o? ae, ui e o i?ae?icae?e? 3.1.
Aeey aiae?co ?acoeueoao?a ioe?ithaaiiy iciae aeei?enoaii
ia’?aeiaiee oano Eaaaee?. A iiooeao aaeaeaaoii? iiaeae?
oanooaaee aea? iiaeae?: aaeeoeaii-aeii?iaioio ? iiaeaeue c
o?aooaaiiyi aca?iiae?? aeaio aai?a. Aaeeoeaii-aeii?iaioia iiaeaeue
/anoi aeyaey?oueny i?eaeaoiith aeey iieno aaaaoaoeaieo iciae ?
aeiaaeeie eieina (iae. 4). I?e aiae?c? aaiioei?a ca aaiaie, ui
eiio?ieththoue iciaee i?iaeoeoeaiino? eieina, aeey iieno
?ica?aeiinoae iaeaea caaaeaee aeiaiaeeoueny caa?oaoeny aei
iiaeae? c o?aooaaiiyi aca?iiae?? i?ae aeaiia aaiaie. A iecoe?
aeiaaee?a i?ae?a?aoe aaeaeaaoio iiaeaeue ia aaeaeiny, ine?eueee
e?euee?noue iayaieo o iaoiio ?icii?yaeaeaii? n?iae ia aeaaaei
ciiae ?icaeyaeaoe iiaeae? a?eueo neeaaei?, i?ae c o/anoth aeaio
aca?iiae?th/eo aai?a. Iaea?eueoa e?euee?noue eiia?iaoe?e, aeey yeeo a
aeiaeai ??e aeine?aeaeaiiy ia aaeaeiny i?ae?a?aoe aaeaeaaoii?
iiaeae?, niinoa??aa?oueny aeey iciae aeiaaeeia ? oe?eia eenoea,
aeiaaeeia eieina oa aei?noe?noue eieineiai? eonee. Ia iaoo aeoieo, ?
o?e ??ci? i?e/eie oaeiai noaio ni?aa. Iciaee aeiaaeeia ? oe?eia
eenoea, ia a?aei?io a?ae ?ioeo aaaaoaoeaieo iciae, aeyaeeinue aaaeei
ioe?ithaaoe /a?ac iaeiio?ieueiaai? oa aacnenoaii? niioai?aiiy. A
eiio?ie? iciaee aei?noe?noue eonee aa?a o/anoue e?eueea aai?a ? aeaa
c ieo c/aieai? (?icae?e 4 aeena?oaoe??). Aeey noai?aiiy iiaeae?, ui
a?aoiao? c/aieaiiy aai?a, ye? aca?iiae?thoue, ia aoei iio??aii?
e?eueeino? n?iae. Iciaeo aeiaaeeia eieina ioe?ithaaoe aeoaea i?inoi, ?
yeui ia aaea?oueny iiaoaeoaaoe aaeaeaaoii? iiaeae? aeey iieno ??
eiio?ieth, oi oea iia’ycaii aai c ia?aaeuaiiyi e?eueeino? aai?a, ui
aca?iiae?thoue (a?eueoa aeaio), aai, ye aeey aei?noeino? eonee, a
eiio?ie? aa?ooue o/anoue aaie, ui c/aieai?. A aaniethoi?e a?eueoino?
aeiaaee?a a eiio?ie? an?o iciae an?o eiia?iaoe?e no?auoaaiiy cia/oueie
aeyaeythoueny iiceoeai? aaeeoeai? aoaeoe i?aeaeaeueii? aca?iiae?? ([d]).
Aeiyoe?a aoei e?eueea: ae?th aeae?a ni?yiiaaii a noi?iio ciaioaiiy
iciaee e?euee?noue ca?ai o eiein? a eiia?iaoe?? Deao Dt28 ? ia?aiao? [d]
aeyaeany iacia/ouei aeey iciae aeiaaeeia noaaea ? i?aeaocey a
eiia?iaoe?? DeaoDt346, aeiaaeeia ? oe?eia eenoea a eiia?iaoe?? Den o
Dt346, e?euee?noue ca?ai o eiein? a eiia?iaoe?? Dea o Dae,
iana ca?ai a

eiia?iaoe?yo Deno Dae, Deno Dt28, Dea o Dae ? iana iaeiiai ca?ia a
eiia?iaoe?? Deno Dea. A oeeo aeiaaeeao cia/ouee aianie o oaiioei?/iee
ae?ac iciaee iaeaaeeoue aai aeii?iaioiiio aoaeoo aeae?a, aai
i?aeaaii?e aca?iiae??. Aeii?iaioia ae?y aeae?a aeyaey?oueny cia/ouith
oaeiae aeiaie? /anoi. Ii??aiyiiy iai?yieo ae?? aeii?iaioieo aeae?a
ciiao iiae?ey? aeine?aeaeai? iciaee ia aea? a?oie: aaaaoaoeai?, a ?o
eiio?ie? aa?ooue o/anoue aeii?iaioi? aeae?, ui ae?thoue o iai?yieo ye
ca?eueoaiiy oaiioei?/iiai ae?aco iciaee, oae ? a iai?yieo eiai
ciaioaiiy; iciaee i?iaeoeoeaiino?, aeey yeeo /ano?oa aeyaeythoueny
aeae?, ui ae?thoue a iai?yieo ciaioaiiy /eneiaiai ae?aco iciaee.
Aiaeia?/io ea?oeio niinoa??aa?ii i?e aea/aii? ia?aiao?o [i]: aeey
iciae aeiaaeeia noaaea ? i?aeaocey a?i caaaeaee iiceoeaiee ? a?eueoith
i??ith iiceoeaiee aeey iciaee iana iaeiiai ca?ia. Aeey ?ioeo iciae
i?iaeoeoeaiino? ? aei?noeino? eieineiai? eonee iaaaoeai? cia/aiiy
oeueiai ia?aiao?o cono??/aeenue /ano?oa.

Aea/aiiy aaeeei? e?eueeino? n?iae (a?ae 4 aei 10 a caeaaeiino?
a?ae ?ieo aeine?aeaeaiiy ? eiia?iaoe?? no?auoaaiiy) a ?aieao oanoo
ianooaaiiai oeaethaaiiy noiniaii caaaeueii? oa?aeoa?enoeee
aaiioei?/ieo iniaeeainoae D aaiii?a, ui aea/athoueny, aeaei, caaaeii,
oo ae ea?oeio, ui e aea/aiiy a?a?eae?a ia?oiai iieie?iiy a?ae
no?auoaaiiy aaiii?a D ca ae?aeaeueiith noaiith. Aeey iieno nenoaie
aai?a, ui aa?ooue o/anoue a eiio?ie? iciae aaaaoaoeaii? /anoeie
?ineeie, /anoi aeyaey?oueny i?eaeaoiith aaeeoeaii-aeii?iaioia
iiaeaeue, a a eiio?ie? iciae i?iaeoeoeaiino? eieina aei aai?a c
aaeeoeaii-aeii?iaioiei oeiii ae?? i?e?aeiothoueny aaie c ai?noaoe/iei
oeiii i?aeaaiii? aca?iiae??. Ii??aiyiiy ?acoeueoao?a, io?eiaieo
i?e ioe?ithaaii? i?eioeeiiai ??cieo iaai??a iiioeyoe?e ( aaouee?anuee?
iiioeyoe?? oa F1, ui ia ?ic/aieththoueny, a ae?aeaeueiiio aiae?c?
? aaouee?anuee? iiioeyoe?? oa F1, ui ia ?ic/aieththoueny ?
iiioeyoe??, ui ?ic/aieththoueny; F2, ia?oee aaee?in aei eiaeiiai c
aaouee?a, F3 ? ua o?e ??ci? iiioeyoe?? a?ae no?auoaaiiy ?ineei F2 c F1
? c eiaeiei c aaouee?a a ia’?aeiaiiio oano? Eaaaee?) aeyaeei aeoaea
ae?acii, ui caaaeueia oa?aeoa?enoeea an?o eiiiiiaio?a no?auoaaiiy ca ?o
aaiioei?/iith no?oeoo?ith a oe?eiio ia ? i?inoith noiith
oa?aeoa?enoee, yeo ie io?eio?ii a oano? Eaaaee? aeey eiaeii?
eiie?aoii? aaouee?anueei? ia?e. C?iaeaii aeniiaie, ui iniiaia
?ioi?iaoe?y, yeo iiaeia caeiaooe c ?acoeueoao?a ae?aeaeueiiai
aiae?co, noino?oueny ni?aa?aeiioaiiy ?aoeaneaieo ? aeii?iaioieo aai?a
a iiioeyoe?? ? iino?eiino? oeeo ni?aa?aeiioaiue c ?ieo a ??e. An?
?io? oa?aeoa?enoeee aea/aieo aaiioei?a iio??aii io?eiaoe c
?acoeueoao?a oanoo Eaaaee? c o??th ?noioiith ia?aaaaith ia?aae
?acoeueoaoaie, io?eiaieie c ae?aeaeueiiai aiae?co, ui aiie
noinoaaoeiooueny eiaeii? eiie?aoii? ia?e aaiii?a D, ui aea/athoueny,
a ia an??? iiioeyoe?? aea/aieo oi?i a oe?eiio.

?ICAe?E 4. Aaiaoe/iee aiae?c noaaaiiio D i’yei? ioaieoe? ca

iciaeaie ii?oieia?? ?ineeie ? aeayeeie a?eeaie

Aaiiiii-aeiaeai? oi?ie ? a?a?eaee, io?eiai? a?ae oeeee?/ieo
no?auoaaiue (iae. 3), aea/aee ca ii?oieia?/ieie
(iayai?noue/a?aenooi?noue aineiaiai iaeueioo, caaa?aeaiiy noeaeiai
eieino, ?iciiae?e caaa?aeaiiy ia eonoe? oa iiooaiiy eieineiai? eonee,
ai??aeea, inoaaeaei? a?ae?inoee, aei?noe?noue eieineiai? eonee,
aaeaaeai?noue iniiae eieineiai? eonee, caaa?aeaiiy ca?i?aee) ?
a?io?i?/ieie (aeaeo?ioi?aoe/i? niaeo?e aethoai?i?a, ae?aaeei?a,
aaoa-ai?eace) iciaeaie. Aeei?enoiaoth/e o?aaeeoe?ei? i?eeiie
a?a?eaeieia?/iiai aiae?co, aeeiiaee aaiaoe/iee aiae?c aiae?ciaaieo
noaaaiii?a D ca ia?a?aoiaaieie iciaeaie, anoaiiaeaoe e?euee?noue
aai?a, ui aa?ooue o/anoue a eiio?ie? oeeo iciae, oei i?aeaeaeueii? oa
i?aeaaiii? aca?iiae??.

Ca aaiaie, ui eiio?ieththoue iciaeo iayai?noue/a?aenooi?noue
aineiaiai iaeueioo ia ?ineei? aeine?aeaeoaai? oi?ie iathoue aaiioeie:
Den, Dea, Dae — W1(2B)W2i(2D), Dt28- W1(2B)W2I(2D), a naia iciaea iiaea
aooe ia?ea?ii o?iiiniie 2D (Tsunewaki, 1966). Iaiao?aioi?noue iiaia.
Eiio?ieue iciaee eie?? noeaei? eieineiai? eonee cae?enith?oueny aaiaie
Rg2 oa Rg3,ui c/aieai? ? ye? aca?iiae?thoue ca oeiii eiioeyoeaii?
iie?ia??? ca o/anoth ?aoeaneaiiai ?ia?a?oi?a Rg3i. Anoaii-aeaii
aaiioeie/i? oi?ioee: Den — rg2Rg3Rg3I, Dea ? Dae- rg2Rg3Rg3i, Dt28 —
Rg2rg3Rg3I. Rg2 eieae?ciaaii ia o?iiinii? 1D (Rowland, Kerber, 1974),
oiio eie?? noeaei? eonee neoao? ia?ea?ii oe??? o?iiiniie. ?iciiae?e
i?aiaioo ia ei-eineia?e eonoe? eiio?ieth?oueny aeii?iaioiei aaiii c
aeia?ith aeni?aneai?noth ? iiaiith iaiao?aioi?noth. Aai
?aeaioeo?eiaaii aia?oa. Aaiioeie: Den, Dae oa Dt28 Cd1I, Dea — Cd1i.
Iciaea iiooaiiy eieineiai? eonee eiio?ieth?oueny e?eueeiia aaiaie.
Aeaa c ieo a?aeiia?aeathoue ca aonoioo iiooaiiy (Hg oa Hgi-HgI) ? aeaa
– ca ??aiii??i?noue eiai ?iciiae?eo ii eonoe?. Aaiioeie: Den — Hg(1A)hg2
HgIhg3, Dea 1Hg(1A)hg2HgiHg3, Dae- Hg(1A)Hg2HgIhg3, Dt28- Hg(1A)
Hg2Hgihg3. sse ia?ea?ia iciaea iiaea aooe aeei?enoaia eeoa a
aeae?eueeio eiia?iaoe?yo no?auoaaiiy. Iiooaiiy iniiae /eaieea
eieiniaiai no?eaeiy (ai??aeea) ? i?ae eiio?ieai aai?a Br1, aeii?iaioiiai
? Br2, iai?aaeii?iaioiiai. Aaie ?aeaioeo?eiaaii aia?oa. Aaiioeie
aea/aieo oi?i: Den oa Dae — br1Br2, Dea — br1br2, Dt28 — Br1Br2. sse
ia?ea?iee e?aua i?aeoiaeeoue oie aai, yeee a?ae??ciy? Dt28 a?ae Den ?
1Dae. AEi?noe?noue eieineiai? eonee eiio?ieth?oueny aaiaie Tg (Kerber,
Rowland, 1974) ? Tg2 (?aeaioeoe?eiaaii aia?oa) — iai?aaeii?iaioieie ?
c/aieaieie, o?iiiniia 2D. ?aoeaneai? aeae? tg2 a?aeiia?aeathoue ca
oi?ioaaiiy aaeaaeaiino? a iniia? eieineiai? eonee. Aaiaoe/iee eiio?ieue
oe??? iciaee anoaiiaeaii a oeueiio aeine?aeaeaii? aia?oa. Aaiioeie:
Den — tgTg2en, Dea — tgtg2, Dae — tgTg2ae, Dt28 — TgTg2en. Aai R1
(/a?aiia caaa?aeaiiy ca?i?aee, oi?ie Dea ? Dt28) ia?ea?othoue o?iiiniio
3D (McIntosh et al., 1998), ?aoeaneai? aeae? eiio?ieththoue a?eee
eie?? ca?i?aee (oi?ie Den oa Dae). Ca aaiaie Glu-D1(Bietz et
al., 1975; Lawrence, Shepherd, 1980) a?ae??ciyeeny iaeei a?ae iaeiiai
an? aaiiie, e??i Dea oa Dae, ca aaiaie Gli-D1 (Mecham et al., 1978;
Niceiia n niaao., 1978) ? Gli-D2 (Shepherd, 1968) — an? aaiiie, ca
aaiaie b-Amy-D1 (Ainsworth, 1983) — aaiii Dt28 a?ae aoaeue-yei? c
o?ueio ?ioeo oi?i. Ia?a?aoiaai? eieone iiaeooue aooe aeei?enoai? ye
ia?ea?e o?iiinii 1D, 6D oa 4D, a?aeiia?aeii.

?ICAe?E 5. Aeei?enoaiiy aai?a-ia?ea??a aeey

aaiaoe/iiai aiae?co noaaaiiia D

O?iiiniiio eieae?caoe?th ?aeaioeo?eiaaieo aai?a, ui eiio?ieththoue
ii?oieia?/i? iciaee ioaieoe?, aeeiiaee ca oaeoii ?o c/aieaiiy c aaiaie,
eieae?caoe?y yeeo a?aeiia (McIntosh et al., 1998), aeei?enoiaoth/e
e?eoa??e ocaiaeaeaiino? aeey /ioe?ei?eueieo oaaeeoeue (Aaeee, 1964).
O a?aeiia?aeiino? c io?eiaieie ?acoeueoaoaie aea/ai? aaiioeie iiaeia
iienaoe iaaieie aaiioei?/ieie oi?ioeaie noiniaii an?o aea/aieo ye?nieo
iciae c aeacoaaiiyi o?iiiniiii? eieae?caoe?? aai?a (oaae. 3).

Oaaeeoey 3. Aaiioeie aaiiiii-aeiaeaieo oi?i ca aaiaie ii?oieia?/ieo
iciae,

ui aea/aeeny, oa ?o o?iiiniiia eieae?caoe?y

Iciaee Den Dea Dae Dt28

Aineiaee iae?o W2i W2i W2i W2I

Eie?? noeaeiai eieino rg2Rg3Rg3I Rg2Rg3Rg3i rg2Rg3Rg3i Rg2rg3Rg3I

?iciiae?e i?aiaioo ia eonoe? CdI Cdi CdI CdI

Iiooaiiy eonee hg2HgIhg3 hg2HgiHg3 Hg2HgIhg3 Hg2Hgihg3

Inoeno?noue BnBn+1 bnbn+1 bnbn+1 BnBn+1

Ai??aeea br1Br2 br1br2 br1Br2 Br1Br2

AEi?noe?noue eonee tgaTg2 tgtg2 tgbTg2 TgaTg2

Eie?? ca?i?aee r1 R1 r1 R1

O?iiiniiia eieae?caoe?y

1D 2D 3D 4D
6D

Rg2 W i R1
b-Amy-D1 Gli-D2

Rg3 Tg Tg3

Glu-D Tg2

Gli-D1 Cd I

Hg I Br2

Br1

I?e aea/aii? e?euee?nieo iciae c iaoith anoaiiaeaiiy o?iiiniiii?
eieae?caoe?? aieiaieo aai?a, ye? eiio?ieththoue ?icaeoie e?euee?nieo
iciae, aeei?enoaee aeaa i?aeoiaee: aiae?c aeenia?n?e ? ii??aiyiiy
na?aaei?o cia/aiue. Aeenia?n?? aiae?coaaee ca noaiith
aeaioaeoi?iiai aeenia?n?eiiai aiae?co ca oaeoi?aie aaiioei ?
iiaoi?i?noue (Eaeei, 1980). Ii??aiyiiy na?aaei?o cia/aiue aeeiioaaee ca
e?eoa???i Nouethaeaioa, aeio?eioth/enue ?ic?iaeaiiai iaie aeai?eoio: 1.
C iiioeyoe?? F2 aeae?aee aea? a?oie ?ineei, iaeia c yeeo ia?
iaoa?einueeee oaiioei ca iaaiith iciaeith-ia?ea?ii, a ae?oaa —
aaouee?anueeee ?c na?aaei?ie cia/aiiyie F2M1±sM1 oa F2M2+sM2.
sseui ??cieoey i?ae oeeie na?aaei?ie ia aoea cia/oua, an? oae? aeiaaeee
aeeeth/aee c iiaeaeueoiai aiae?co a?oiiaeo na?aaei?o, ni?yiiaaiiai ia
eieae?caoe?th aieiaieo aai?a e?euee?nieo iciae. Eiee ??cieoey aoea
cia/oua, ia?aoiaeeee aei ae?oaiai ioieoo. 2. Aii??e/ia na?aaei?
cia/aiiy a?oie, F2M1±sM1 aai F2M2±sM2, ii??aithaaee c oai?aoe/ii
i/?eoaaiei na?aaei?i cia/aiiyi, ae?aoiaaiei, aeoiaey/e c aaiioei?/ieo
oi?ioe iiii ? aeea?a?eaeiiai ?ic/aieaiue a F2, i?eeiyoeo a a?iiao?e/i?e
aaiaoeoe? (Mather, Jinks 1971). sse aaiaoe/i? aoaeoe aeei?enoiaoaaee
aii??e/i? cia/aiiy ia?aiao??a, ae?aooaai? a i?ae?icae?e? 3.2
aeena?oaoe??.

?acoeueoaoe iaoeo aeine?aeaeaiue (oaae. 3) oa a?aeiiino? i?i
oa?aeoa? oniaaeeoaaiiy ? o?iiiniiio eieae?caoe?th iaaieo aai?a
(McIntosh et al., 1998) aoei aeei?enoaii aeey o?iiiniiii?
eieae?caoe?? aieiaieo aai?a, ui aa?ooue o/anoue a eiio?ie? e?euee?nieo
iciae c /enea iaie aea/aieo. Iaea?aeai? ?acoeueoaoe ocaaaeueiaii a oaae.
4. Eiia?iaoe?yie, iaea?eueo ia?niaeoeaieie aeey iiaeaeueoi? ?iaioe c
?aeaioeo?eaoe?? ? aea/aiiy QTL, aeyaeeenue: Den o Dt28 aeey iciae
aeiaaeeia noaaea, i?aeaocey ? eenoea, Dea o Dt28 aeey iciae e?euee?noue
eieine?a o eiein?, iana ca?ai c aieiaiiai eieina, iana iaeiiai ca?ia,
Deno Dea aeey iciae aeiaaeeia eieina ? e?euee?noue eieine?a o eiein?.

?ICAe?E 6. ?io?ia?aneaia a?a?eaeecaoe?y i’yei? ioaieoe?: iaoiaee ?

ia?niaeoeae oeueiai iai?yieo aeine?aeaeaiue (iaeyae
e?oa?aoo?e)

Aiae?c no/aniiai noaio aeine?aeaeaiue a aaeoc? ?io?ia?aneaii?
a?a?eaeecaoe?? ioaieoe? aeyaea e?eueea iai?yie?a, ui a ieo
?ic?iaey?oueny i?iaeaia. Ia?o ca ana, eeoa?oueny aeooaeueiei iiooe
aeaea?ae ei?enieo aai?a. No/ani? ?iaioe oa?aeoa?ecothoueny aeaiia
iii?oieie iniaeeainoyie. Ia?oa: aei no?auoaaiue caeo/athoueny ana a?eueo
a?aeaeaeai? nii??aeiai? i’ye?e ioaieoe? aeaee ? ?iaee ceae?a ?
noai?ththoueny a?a?eaee ca o/anoth o?ueio ? /ioe?ueio aeo?aeieo
eiiiiiaio?a (Zhuang et al., 1988, Tang et al., 1997). Oea iia’ycaii c
oni?oaie aeine?aeaeaiue, ni?yiiaaieo ia iiaeieaiiy ?ieiia?oaioiino?
no?auoaaiue ca ?aooiie oaeineiiaeaiiy iaoiae?a ae?iuoaaiiy aaiaoio?o?a ?
a?a?eaeieo ca?iaee?a oa ua no/ani?oeo iaoiae?a. ? ae?oaa: oiiaith
aeinyaiaiiy iaoe a?aeaeaeaii? a?a?eaeecaoe??- ?io?ia?an?? iio??aiiai
aaia aai e?eueeio aai?a — noa? aeei?enoaiiy aai?a-

Oaaeeoey 4. O?iiiniiia eieae?caoe?y ? iai?yiie ae?? aieiaieo aai?a, ui
aa?ooue

o/anoue a aaiaoe/iiio eiio?ie? i??ieo iciae ioaieoe?

_________________________________________________________________

Iciaea1 O?iiiniia 1 O?iiiniia 2 O?ii.3 O?ii.4
O?ii.6

Glu Gli1 HgI Rg1 Br1 WI Tg Tg2 Br2 CdI1 R1
b-Amy1 Gli2

Den

AeN — 2 — + + — + +
+ — + +

AeI — + + + — + —
— + — +

AeE — + + — + + + +
+ + — +

OE + + ? — + + — —
— + — +

AeE + + + ? — +
+ + +

EEE + + + + — + + +
+ — — +

ICE — + + — + + — —
+ + + +

IIC + + + — + + + +
+ — + +

AEEE + — + + — + +
+ — — —

Dea

AeN — + + + — — + —
+ + + +

AeI — + + ? — — + —
— + + — +

OE — + ? — — + — —
— + — +

AeE + ? ? ? — — —
+ + + — ?

EEE + + + — — + + +
+ + + +

ICE ? + + ? — — + + —
+ + + +

IIC + + + ? — ? + + +
+ + + +

AEEE + + + ? — — ? + +
+ — — —

Dae

AeN — + ? + — — — —
— — + + +

AeI — + + + — + —
— — + — +

OE — + + — + + — —
— + — +

AeE + + + + + + ? —
+ — + — —

ICE + + — + ? + + —
+ + + —

IIC + + — + + ? + — —
— + + +

Dt28

AeN — ? + + ? + —
— ? — + +

AeI — + + ? + — + —
— + — ? +

AeE — + + + + ? — —
— — — — +

*CE + + + + + + — —
+ + + —

ICE + + + ? + — —
— + + +

IIC + + — + — —
+ + — —

________________________________

1 Iicia/aiiy iciae ia noi?. 5.

2 + ?ica?aeiino? i?ae na?aaei?ie F2M1±sM1 oa F2M2+sM2 cia/ou? eeoa ca
?acoeue oaoaie aeenia?n?eiiai aiae?co; — cia/oueo ?ica?aeiinoae ia
ciaeaeaii;

? QTL, ui ca?eueoo? ae?ac iciaee; ? ? QTL, ui ciaioo? ae?ac iciaee.

ia?ea??a (Gale et al., 1984, Marais, 1992, Koebner, 1995). A ia?oo
/a?ao oea noino?oueny iciae, aaciina?aaei? ioe?ithaaiiy yeeo a
a?a?eaeieo iiioeyoe?yo, ui ?ic/aieththoueny, na?eicii oneeaaeiaia
(no?ee?noue aei eaoth/i? naaeee, ooca??ico oa ?i.). Ae?oaee eiiieaen
caaaeaiue – ciaioaiiy ianyao /oaeeiiiai iaoa??aeo, ui aiineoueny a
aaiii ioaieoe?, aei oiai i?i?ioio, eiee iio??aio iciaeo ua ia
c?oeiiaaii, aea iia?aeii? iciaee aaea iaia. ?iaioe ?icai?oathoueny
iaeii/anii a e?eueeio iai?yieao. 1. Aea/aiiy oniaaeeoaaiiy iciaee, ui
ia?aaea?oueny. sseui iciaeo ia iiaeia ia?aaeaoe o ioaieoeth a oiio
aeaeyae?, a yeiio aiia aeeeeea? iaoa noaaeaiiy o nii??aeiaiiai aeaeo
(iai?eeeaae, ai?no a?eea a ca?i? ? e?ceio a a?eoe?, no?ee?noue aei
iecueeeo oaiia?aoo?), e?aua oea anoaiiaeoe caa/anii ? aecia/eoe
no?aoaa?th ? oaeoeeo ?iaioe c oaeith iciaeith aai acaaae?
a?aeiiaeoeny a?ae ?io?ia?an?? (Muramatsu et al., 1983, Porceddu et
al., 1988) . 2. ?iaeoeoe?y ?aeiia?iaoe?e ? o?aineieaoe?e. Naia
ii?aoeueiaaia aei oeueiai /ano /anoeia i?iaeaie (Sears, 1956, 1982,
Okomoto, 1957, Knott, 1961, Lapitan et al., 1984, Maan , 1988) . A
no/aniee ia??iae iniiaio oaaao iaaeathoue iaoiaeai ne?ei?iao
iiioeyoe?e, ui ?ic/aieththoueny, aeey aeyaeaiiy i?iaeoeo?a ?io?ia?an??.
Iaaoaa? cia/aiiy «aoeeaooaaiiy» iio??aieo ae?eyiie o?iiinii ia?ea?aie
??cii? i?e?iaee. Ia ia?oee ieai ooo nei?i aecia/eoueny caaaeaiiy
aeyaeaiiy ia i?inoi i?iaeoeo?a ?io?ia?an?? ??cii? yeino?, a oaeeo
o?ainoa??a, aea a?aeaoany ?aoeei?ieiee ? oaeee iacia/iee ca ianyaii
iai?i aaiaoe/iei iaoa??aeii, ui a ?acoeueoao? iaoeiaii ni?aao c
iaiaeaethth/eie aaiaie (Thomas et al., 1998). Oea — ?aeaaeueiee
?acoeueoao ?io?ia?an??.

O?ao?e eiiieaen caaaeaiue ii nai?e noo? ? i?/ei ?ioei, ye ia?ao?ae
aei oe?eani?yiiaaiiai eiino?othaaiiy aaiiio ioaieoe?, iaoa??aeueia
iniiaa aeey yeiai ae?iaey?oueny ca?ac iiceoeaieie ?acoeueoaoaie
a?aeaeaeaii? a?a-?eaeecaoe??. Ia ieoaiiy: ye? aaie iio??aii
neioacoaaoe, uia caaacia/eoe no?ee?noue, iai?eeeaae, aei ??ae? aai
aei iaaeeeoeo aethi?i?th o a?oio? — aeanoue a?aeiia?aeue eeoa aea/aiiy
oiiei? no?oeoo?e naieo aai?a. Aea ?o nii/aoeo iio??aii aoaea ciaeoe,
noai?eoe oiiae aeey ii??aiyiiy c aeueoa?iaoeaiei aeaeai, aea/eoe
iaoai?ci, yeei caaacia/o?oueny ?icaeoie iio??aii? iciaee, a aaea
iio?i ia?aoiaeeoe aei neioaco iio??aiiai aaia. Ia aeoieo iiooea/a, an?
iciaee aaeaioeaiino?, ia ye? ie oi/aii iie?aueoe ioaieoeth,
iiae?eythoueny ia aea? a?oie: oaeoi?, ui iiaaii aieeaa? ia ioaieoeth,
i?aeaea?oueny aaiaoe/ii coiiaeai?e i?ieeaino? (oai?iae, oe?aeieee);
no?aniaee oaeoi? ia i?aeaea?oueny aaiaoe/i?e i?ieeaino? ? ia aoaea
ci?ithaaoeny ?acii c aaiaie no?eeino? aei iueiai (no?ee?noue aei
ceiiaeo no?an?a, iinooe, iaiioeiaeueiiai neeaaeo a?oioo). Uia
caaacia/eoe ioaieoe? no?ee?noue aei oaeoi??a ae?oai? a?oie, ? nain
aeo?a/aoe neee ? caniae ia ia?aianaiiy aai?a no?eeino? a?ae
aeeei?ineeo ?iaee/?a aei ioaieoee oiio, ui iane?aeeaie ia?aianaiiy ie
aoaeaii ei?enooaaoeny aeoaea aeiaai. No?ee?noue ioaieoe? aei oaeoi??a
ia?oi? a?oie ie ni?iao?ii caaacia/eoe oeyoii ia?aianaiiy aai?a
no?eeino? o ioaieoeth a?ae ?iaee/?a eeoa oiio, ui i?/iai ?ioiai iiee ui
ia ai??ii. ? noai?thaaoe iaoa??aeueio iniiao c ?ineei, ia yeeo
iiaeia aeine?aeeoe oiieo no?oeoo?o aaio ? iane?aeee iaoeo ao?o/aiue
a ia?, i?ino?oa anueiai, iaaooue, aeei?enoiaoth/e ?aaeueii ?nioth/? a
i?e?iae? aaie, yeeo iaia? o i’yei? ioaieoe?, aea ye? iiaeia aianoe
aei ?? aaiiio.

?ICAe?E 7. Noai?aiiy ?io?ia?aneaieo e?i?e i’yei? ioaieoe?

c aeei?enoaiiyi aaiiiii-cai?uaieo aaenaiei?ae?a

Aaiiiii-cai?uaieie oi?iaie ioaieoe? ie iacaaee aaenaiei?aee,
io?eiai? ye cae/aei? aio?aeeiei?aee oao?aiei?aeiiai eiiiiiaioa AAAA
i’yei? ioaieoe? ni?oa Aa?i?a ? nii??aeiaieo ioaieoe? aeeiei?aeieo
aeae?a c ?iaeo Aegilops: Aa?ieaoa (AABBUU, o?ao?e noaaaiii a?ae
Ae.umbellulata), Aa?iaean (AABBSS, o?ao?e noaaaiii a?ae Ae.spelto1ides),
Aa?icen (AABBSlSl, o?ao?e noaaaiii a?ae Ae.sharonensis). Ca iaoiaeii
noai?aiiy ? caniaiia iiaeaeueoi? ?iaioe nthaee ae a?aeiineoueny oi?ia
Aa?iaa, iaiiaiee aio?aeeiei?ae oao?a-Aa?i?e ? aeaeo Agropyron glaucum
(AEe?ia, Oa?iianeay, 1984). ?acoeueoaoii aaaaoi??/ii? oe?eani?yiiaaii?
?iaioe c aaeeeith e?euee?noth e?i?e, io?eiaieo a?ae no?auoaaiiy
aaiiiii-cai?uaieo aaenaiei?ae?a i’yei? ioieoe? ? Aa?iaaa c ni?oii
Aa?i?a noaei noai?aiiy /oaeeiii-cai?uaieo e?i?e i’yei? ioaieoe?, a yeeo
a?ae iaeii? aei o?ueio ia? o?iiinii noaaaiiio D ioaieoe? cai?uaii
aiiaieia?/ieie o?iiiniiaie ia?a?aoiaaieo nii??aeiaieo ioaieoe? aeae?a.
Io?eiaii oaeiae /oaeeiii-o?aineieaoe?ei? e?i??. Oaeo iayaiino?
cai?uaiiy aai o?aineieaoe?? a e?i?yo anoaiiaethaaee oeyoii aea/aiiy
anioe?aoe?e o?iiinii a II iaeico IEI a?a?eae?a a?ae no?auoaaiiy
aiae?ciaaii? e?i?? c ni?oii ioaieoe? Aa?i?a. ?aeaioeo?eaoe?y /oaeeiieo
aeeth/aiue noiniaii ?o iaeaaeiino? aei a?oie aiiaieia?/ieo
o?iiinii aeeiiaii oeyoii ne?ei?iao anueiai iayaiiai ?icia?ooy e?i?e c
/oaeeiieie o?iiiniiaie (o?aineieaoe?yie) iaoiaeii aeaeo?ioi?aco
caianieo a?ee?a ca?ia oa iecee oa?iaioieo nenoai (a?eie-eiaeoth/?
aaie-ia?ea?e o?iiinii). C 7 aiiaieia?/ieo a?oi o?iiinii a?eeiaeie
ia?ea?aie a aeaiiio aeine?aeaeaii? aoei iii?/aii i’youe: 1, 3, 4, 6
oa 7. Ca eiaeiith c ieo a aaenaiei?aeieo e?i?yo ioaieoe? aoei aeyaeaii
cai?uaiiy ioaie/ii? o?iiiniie c noaaaiiio D ia /oaeeiio aiiaieia?/io,
ye ii iaei?e o?iiinii? (iaeeia?i? cai?uaiiy), oae ? ii ??ciei
?o ii?aeiaiiyi (iiiaeeii? cai?uaiiy). Oaeei /eiii aoei aeiaaaeaii
i?eaeaoi?noue iaoiaeo «ci?ooaaiiy» o?iiinii ??cieo aaiii?a (U+D, S+D,
Sl+D, X (ie??th) +D) a iaaeao iaeiiai aeeiei?aeiiai noaaaiiio
aaenaiei?aeiiai i?aai?cio, cai?iiiiiaaiiai ? oai?aoe/ii ?ic?iaeaiiai
AEe?iaei (AEe?ia, Oa?iianeay, 1984) aeey noai?aiiy /oaeeiii-cai?uaieo
e?i?e ioaieoe? c iaeeia?ieie ? iiiaeeiieie cai?uaiiyie. Iaie aia?oa a
na?o? io?eiaii /oaeeiii-cai?uai? e?i?? ioaieoe? c o?iiiniiaie
Ae.sharonensis, ?ioeie i?ae 4Sl (a?aeiia ye aaiaoioeeaeia o?iiiniia), ?
ye? ia i?noyoue ?? ye o?iiiniio, iaia’yceiao aeey caaacia/aiiy
ii?iaeueii? aeeoo?caeaoiino? ? ooieoe?iioaaiiy aaiao. Oea — ua iaeia
na?ae/aiiy aoaeoeaiino? iaoiaeo ci?ooaaiiy» o?iiinii a iaaeao iaeiiai
noaaaiiio aaenaiei?aea aeey noai?aiiy /oaeeiii-cai?uaieo e?i?e ?
aeayeeo eiai ia?aaaa ia?aae eeane/iei iaoiaeii, ?ic?iaeaiei O’Mara ?
Jenkins (O’Mara, 1940, Jenkins, 1968). Oii?yaeeoaaiiy ?icia?ooy iayaieo
/oaeeiii-cai?uaieo e?i?e ca aiiaieia?/iith iaeaaei?noth /oaeeiieo
o?iiinii, ui cai?noeee ioaie/i?, ca aeiiiiiaith a?eie-eiaeoth/eo
aai?a-ia?ea??a ? ianooiia aea/aiiy oeeo e?i?e ca an?ia iciaeaie
ii?oieia?? ?ineei, yeeie aiie a?ae??ciythoueny a?ae ni?oo ioaieoe?
Aa?i?a, aeaei ciiao aeyaeoe ii?oieia?/i? iciaee, iia’ycai? c
o?iiiniiaie 1, 3, 6 oa 7 aiiaieia?/ieo a?oi. Oe? ?acoeueoaoe
aeyaeythoueny aeoaea ei?enieie aeey a?coaeueiiai ioe?ithaaiiy
i?iaeoeo?a ?io?ia?aneaii? a?a?eaeecaoe?? ? aaiaoe/iiai aiae?co iciae,
iiyaa /oaeeiieo a?aaeaoe?e yeeo a i’ye?e ioaieoe? ? iaaaaeaiei
iane?aeeii ?io?ia?an?? /oaeeiiiai aaiaoe/iiai iaoa??aeo (eaie?noue
eieiniaiai no?eaeiy, aei?noe?noue eonee, /i?iee eie?? eieina).

?ICAe?E 8. Iaaiai?aiiy aieiaieo ?acoeueoao?a ?iaioe

Aeei?enoaiiy aeaiie?iei?aeeciaaieo aei aeeiei?aeiiai ??aiy
?ic/aieaiiy ca aaiaie noaaaiiio D aeiaaenaiei?aeieo aaiiiii-aeiaeaieo
oi?i i’yei? ioaieoe? aeaei ciiao aeeiiaoe ?o aaiaoe/iee aiae?c ca
ieceith iciae ii?oieia?? ?ineei ? aeayeeie e?euee?nieie iciaeaie.
Io?eiaaoe ieceo iiioeyoe?e a?ae no?auoaaiue 4 ? 5 aaiiiii-aeiaeaieo
oi?i ca oeeee?/iith noaiith no?auoaaiiy, aeei?enoaee eeane/i? iaoiaee
a?a?eaeieia?/iiai aiae?co aeey anoaiiaeaiiy aaiioei?a
aaiiiii-aeiaeaieo oi?i ca aeayeeie oaea a?aeiieie aaiaie, ui
eiio?ieththoue iciaee caaa?aeaiiy, iiooaiiy ? aei?noeino? eieineiai?
eonee, a?io?i?/i? iciaee ? aeey ?aeaioeo?eaoe?? aeae?eueeio iiaeo
aai?a, ?iciiae?e iiooaiiy ? caaa?aeaiiy ia eieineia?e eonoe?,
aaeaaeai?noue iniiae eieineiai? eonee, iiooaiiy iniiae /eaieea
eieiniaiai no?eaeiy. Aeey aaiaoe/iiai aiae?co ca e?euee?nieie iciaeaie
aeei?enoiaoaaee aeaa i?aeoiaee, aeia?a a?aeiieo a a?iiao?e/i?e
aaiaoeoe?: ae?aeaeueiee aiae?c ? oano noi?niiai oeaethaaiiy Eaaaee?
(Mather, Jinks, 1971). E?euee?ni? iciaee aiae?coaaee ia aeaio
??aiyo ?o aaiaoe/iiai eiio?ieth: aaiiiiiio, aeyaeyth/e aianie ie?aieo
/anoei, AA oa D, neeaaeiiai aaiiia AAD a oi?ioaaii? aea/aieo iciae oa
aaiaoe/iiio a iaaeao noaaaiiia D, aaie yeiai ?ic/aieththoueny ia
oe? iaci?iii? /anoeie AA neeaaeiiai aaiiio AAD. ?acoeueoaoe oaeiai
aiae?co aeaee ciiao aeyaeoe ?ica?aeiino? a aaiaoe/ieo nenoaiao, ui aiie
eiio?ieththoue ie?aie iciaee, o caeaaeiino? a?ae oa?aeoa?o iciaee:
ia?aaaaeaiiy aaeeoeaii-aeii?iaioii? aca?iiae?? aai?a ? aeae?a, eiee
eiio?ieththoueny iciaee aaaaoaoeaii? /anoeie ?ineeie, ? iayai?noue
ai?noaoe/ii? i?aeaaiii? aca?iiae?? (a aaniethoi?e a?eueoino?
aeiaaee?a — aeoie?eaoia aca?iiae?y), eiee eiio?ieththoueny
iciaee-eiiiiiaioe i?iaeoeoeaiino? ?ineeie. Ciaeaeaii a?aei?iiino? a
aaiaoe/ieo no?oeoo?ao noaaaiiia D o caeaaeiino? a?ae eiai iioiaeaeaiiy:
noaaaiii D, ui ? aaiiiii aeeei?ineiai aeaeo Ae.tauschii,
a?ae??ciy?oueny ca nai?th aaiaoe/iith no?oeoo?ith a?ae an?o noaaaiii?a
D, ui aa?ooue nai? iioiaeaeaiiy a?ae ni?o?a eoeueoeaiaaii? i’yei?
ioaieoe? a?eueo /anoei aeyaeaiiyi cia/oueo aaee/ei [h] o ii??aiyii? c
aaee/eiaie [d] ? a?eueoith e?euee?noth aai?a, ui aca?iiae?thoue.
Ciaeaeai? ?ica?aeiino? a aaiaoe/ieo no?oeoo?ao iio??aii ?icaeyaeaoe
ye iane?aeie o?eaaeiai ooo/iiai aeiai?o ? ?acoeueoaoe aeine?aeaeaiiy
iiaeooue aooe inieneai? oa aeei?enoai? i?e i?iaiicoaaii? ?acoeueoao?a
aieeao iai?yieo ? iaoiae?a aeiai?o ia io?eioaaio a ?acoeueoao?
aaiioei?/io no?oeoo?o iiioeyoe??.

?acoeueoaoe aaiaoe/iiai aiae?co noaaaiiio D ca ye?nieie iciaeaie
aeaee ciiao ia?ea?oaaoe aaiaie-ia?ea?aie i’youe c naie o?iiinii oeueiai
noaaaiiio ioaieoe?. Aaiaoe/iee aiae?c ca e?euee?nieie iciaeaie iaaeaa
?ioi?iaoe?th i?i oa, ye? eiia?iaoe?? no?auoaaiue ? aeey yeeo iciae
aeioe?eueii aiae?coaaoe aeae?. Aeei?enooth/e ?acoeueoaoe aaiaiae?co ca
aaiaie-ia?ea?aie ? aaiaie, ui eiio?ieththoue e?euee?ni? iciaeaie oa
oaiiiai c/aieaiiy ae?oaeo c ia?oeie, iiaeia aeacaoe o?iiiniie —
iaea?eueo a??ia?aei? iin?? QTL ca ie?aieie e?euee?niei? iciaeaie.
Iaea?aeaia a aeine?aeaeaii? ?ioi?iaoe?y i?i aaiaoe/io no?oeoo?o aea/aieo
oi?i, c iaeiiai aieo, caaaa/o? iao? ciaiiy i?i aaiaoe/io aoaeiao
i’yei? ioaieoe?, c ae?oaiai — aea? iiooiao i?iaeiaaeaiith
aeine?aeaeaiue, ni?yiiaaieo ia ea?ooaaiiy QTL, ui aaea ciaeoee naith
o?iiiniiio eieae?caoe?th. Aeyaeaii iioeiaeuei? eiia?iaoe?? no?auoaaiue
c /enea aea/aieo, c yeeie ? nain i?aoethaaoe aeae?, caeo/ath/e
iieaeoey?ii-aaiaoe/i? ia?ea?e.

Iao? aeine?aeaeaiiy, iia’ycai? c a?aeaeaeaiith a?a?eaeecaoe??th,
iiaeia oa?aeoa?ecoaaoe ye aeei?enoaiiy i?iaeoeo?a iai?yii?
aeaiie?iei?aeecaoe?? ioaieoe? aeey iioei?caoe?? i?ioeano ?io?ia?an?? ia
o?iiiniiiiio ??ai? ia?aaea/? /oaeeiiiai aaiaoe/iiai iaoa??aeo. An?
?ica?aeiino? i?ae /oaeeiii-cai?uaieie aaenaiei?aeieie e?i?yie
iia’ycai? c aaiioei?/iith ??cieoeath eeoa ca iaeiei c o?ueio
noaaaiii?a: /anoeia AAAA o ieo iaeiaeiaa ? ? iaeiieiaiiei
oao?a-eiiiiiaioii ni?oo i’yei? ioaieoe? Aa?i?a, a a aeeiei?aeiiio
noaaaiii? DD a?aeaoeeny ? aoee iaie ?aeaioeo?eiaai? ci?ie ia
o?iiiniiiiio (cai?uaiiy oe?ei? o?iiiniie) aai noao?iiiniiiiio
(o?aineieaoe??) ??aiyo. Io?eiaiee a ?acoeueoao? ?ineeiiee iaoa??ae ?
oe?eaaei a aeaio a?aeiioaiiyo. Ii-ia?oa, eiai caaaa/aii aeeth/aiiyie
/oaeeiiiai aaiaoe/iiai iaoa??aeo a ??cieo ianyaao ? a ??cieo niieo/aiiyo
? iiaea aooe aeei?enoaii ye aaciina?aaei? aeaea?aei aai?a, ui
eiio?ieththoue ei?eni? iciaee ? ye? aaea ia?aianaii ia aaiaoe/ia oei
i’yei? ioaieoe?. E?i?? a?aeiinii noaa?euei?, naiioa?oeeuei? ?
no?auothoueny aac iaiaaeaiue c aoaeue-yeeie ci?oaie i’yei? ioaieoe?.
Ii-ae?oaa, na?aae oeeo aaenaiei?aeieo e?i?e, ye? a?ae??ciythoueny
iaeia a?ae iaeii? ia aeeiei?aeiiio ??ai?, iiaea aeae-?aoeny
?ineeiiee iaoa??ae, i?eaeaoiee aeey ne?auoaaiue c iaoith aaiaoe/iiai
aiae?co ioaieoe? ca ie?aieie iciaeaie. Iaeiae aeei?enoiaoaaoe oe? e?i??
a aia-e?oe/ieo aeine?aeaeaiiyo iio??aii c o?aooaaiiyi o??? ianoaaeie,
ui a eiio?ie? iciaee a?aoeiooue o/anoue aaie, o?aineieiaai? ia o?iiiniie
ioaieoe? o neeaae? ??ciiai ianyao /oaeeiiiai o?iiaoeio. ? oea ia iiaeia
caaoaaoe i?e ?ic?iaoe? iaoiaeeee aenia?eiaioo ? ?ioa?i?aoaoe??
?acoeueoao?a, ui niinoa??aathoueny.

Caeeth/aiiy

?acoeueoaoe aaiaoe/iiai aiae?co noaaaiiia D, aeeiiaiiai a aeai?e
?iaio? aeai-ia cian?i ??cieie niiniaaie: cae/aeiee a?a?eaeieia?/iee
aiae?c ca o/anoth oi?i, ye? eiio?anoi? ca iciaeith, ui aea/a?oueny, ?
aiae?c ca oaiioei?/iei aoaeoii a?aenooiino? iaaii? D-o?iiiniie — i? a
/iio ia anooieee o i?ioe??//y iaeei c iaeiei. Iiaeia c?iaeoe
aeniiaie, ui iai?yia aeaiie?iei?aeecaoe?y aeiaaenaiei?aeii? i’yei?
ioaieoe? c iaoith caaaeaiiy ??aiy ?ic/aieaiiy aei aeeiei?aeiiai oi/a a
aeey iaeiiai aaiiio ? aoaeoeaiei niiniaii noai?aiiy ?ineeiiiai
iaoa??eo, iaeneiaeueii i?enoiniaaiiai aeey eiai aaiaoe/iiai aiae?co
an?ia iiaeeeaeie ? aeinooiieie iai iaoiaeaie no/anii? aaiaoeee.
Aeaiie?iei?aeecaoe?y aeiaaenaiei?aeo, ui i?ecaiaeeoue aei
aeeiei?aeiiai ??aiy ?ica?aeiinoae ca iaei??th c e?eueeio /anoei
neeaaeiiai aaiiia, ?ic?iaea ? ai?iaaoe?y noai aeei?enoaiiy oaeiai
iaoa??aeo a aaiaoe/ieo aeine?aeaeaiiyo ? aianeii a oai??th ? i?aeoeeo
aaiaoe/iiai aiae?co ? iiaea neoaoaaoe iaoiaeii aiae?co ia eeoa i’yei?
ioaieoe?, aea e ?ioeo aeiiie?iei?aeieo aeae?a-naiicaieethaa/?a.

AENIIAEE

1. I’yea ioaieoey Triticum aestivum L. cia/ii iinooia?oueny ?ioei
i?ia?aeiei n?eueneiainiiaea?nueeei eoeueoo?ai ca nooiaiai aaiaoe/ii?
aea/aiino? /a?ac neeaaei?noue no?oeoo?e naiai aaiiio; ia aeoieo
oao?aoe?a, no/aniee aaiioiiae i’yei? ioaieoe? ca?aei?eee ia aaie oa ?o
niieo/aiiy, ui caaacia/othoue no?ee?noue i’yei? ioaieoe? aei aa?ioe/ieo
oa a?ioe/ieo iani?eyoeeaeo oaeoi??a. Iaea?eueo aaaeeeaeie
caaaeaiiyie oeeoiaaiaoeee ioaieoe? ? noai?aiiy ?ineeiiiai iaoa??aeo,
yeee aeicaiey? aeeiioaaoe aaiaoe/iee aiae?c i’yei? ioaieoe?
iacaaaeath/e ia aaenaiei?aeio i?e?iaeo ?? aaiiio, oa io?eiaiiy
?io?ia?aneaieo oi?i i’yei? ioaieoe?, ui aiie iathoue o nai?io aaiii?
??ci? ianyae /oaeeiiiai aaiaoe/iiai iaoa??aeo.

2. Cai?iiiiiaaii oa ai?iaiaaii ia a?aeiia?aeiiio ?ineeiiiio
iaoa??ae? iaoiae aaiaoe/iiai aiae?co aeiiie?iei?aeiiai ?ineeiiiai
i?aai?cio, yeee aea-iie?iei?aeeciaaii iai?yiei niinia?i: aiae?co
i?aeeyaathoue aaenaiei?aei? oi?ie ioaieoe? (AAAADD), aaiii yeeo
neeaaea?oueny c iaeiaeiai? aeey ieo on?o oao?aiei?aeii? neeaaeiai?
(noaaaiiio) AAAA oa aeeiei?aeii? neeaaeiai? (noa-aaiiio) DD, ui ca iath
an? aaenaiei?aee a?ae??ciythoueny iaeei a?ae iaeiiai. ??cieoey i?ae
oi?iaie ioaieoe?, ui aiae?cothoueny, iaiaaeaia aeeiei?aeiei noaaaiiiii
D.

3. Aaiaoe/iee aianie noaaaiii?a AA oa D i’yei? ioaieoe?
o oaiioei aaenaiei?aeo AAAADD ?ic??ciy?oueny ca oeiii ae??
iaeiieiaiieo noaaaiii?a (aaeeoeaii-aeii?iaioia aeey D oa
aaeeoeaii-ai?noaoe/ia aeey AA) oa iai?yieo ?o ae?? (ca?eueoaiiy aai
ciaioaiiy /eneiaiai ae?aaeaiiy oaiioei?/ii? ioe?iee iciaee) o
caeaaeiino? a?ae iciaee (aaaaoaoeaia /anoeia ?ineeie aai aeaiaioe
i?iaeoeoeaiino? eieino) oa a?ae iioiaeaeaiiy noaaaiiio D. ?acoeueoao
aeine?aeaeaiiy ei?enii a?aoe aei oaaae, aeae?ath/e aeaea?aei aaiaoe/ii?
i?ieeaino? aeey no?auoaaiue, ye? ni?yiiaaii ia caaaa/aiiy aaiioiiaeo
i’yei? ioaieoe?.

4. Aaiiiiee aiae?c i’yei? ioaieoe? ca iciaeith ii?icino?ee?noue
aeyaea i?i-a?aeio ?ieue noaaaiiio AA o ii??aiyii? c noaaaiiiii D o
caaacia/aii? aeniei? ii?icino?eeino?. Aeoiaey/e c iioiaeaeaiiy i’yei?
ioaieoe? oa no/anieo ciaiue i?i ii?icino?ee?noue ?? aeeiei?aeieo
i?aaaouee?a, ye ia?niaeoeaiiai eiiiiiaioo no?auoaaiiy a ?io?ia?aneai?e
a?a?eaeecaoe?? ioaieoe? iiaeia ?aeiiaiaeoaaoe aio?aeeiei?ae AADD
(ioaieoey-iaeiica?iyiea o aa?eiin Oaooa).

5. O aaiaoe/ieo nenoaiao noaaaiiia D i’yei? ioaieoe? aeii?iothoue
aeae? «u», ui ca?eueoothoue na?aaei? cia/aiiy iciae aaaaoaoeaii? /anoeie
?ineeie, oa aeae? «v», ye? ciaioothoue na?aaei? cia/aiiy iciae-aeaiaio?a
i?iaeoeoeaiino?. Iciaee ia?oi? a?oie eiio?ieththoueny o a?eueoino?
aeiaaee?a aaiaie c aaeeoeaii-aeii?iaioiith i?aeaaiiith oa
i?aeaeaeueiith aca?iiae??th; o eiio?ie? iciae i?iaeoeoeaiino? eieino
aa?ooue o/anoue aaie c aeoie?eaoiei oeiii aca?iiae??. Iaeiie/aiiy
niieo/aiue aeoie?eaoieo aai?a o aaiioiiae? aeaeo, yeee eoeueoeao?oueny,
iiynith?oueny ?ioaineaiith naeaeoe??th ia iciaee, ui aiie ciaoiaeyoueny
i?ae eiio?ieai aeii?iaioieo aeae?a «v». ?acoeueoaoe aaiaoe/iiai aiae?co
aeioe?eueii a?aoiaoaaoe i?e aecia/aii? ianyao aeai?ie iiioeyoe?e, ui
?ic/aieththoueny, ? ye? i?ecia/ai? aeey aeiai?o o?aina?aneaieo
aaiioei?a.

6. ?aeaioeo?eiaaii aaie, ui eiio?ieththoue a?io?i?/i? (neioac
ae?aaeei?a, aeth-oai?i?a oa aaoa-ai?eace) oa ii?oieia?/i? iciaee, ye?
iiaeia aeei?enoaoe ye aaie-ia?ea?e iaaieo o?iiinii noaaaiiio D: o?noue
aai?a aeey o?iiiniie 1D, i’youe — aeey 2D, aeaa aaie aeey 3D oa ii
iaeiiio aeey o?iiinii 4D ? 6D. Na?aae ieo – oia?oa iienai?: Rg3 (eie??
noeaei? eieineiai? eonee), HgI (?ia?a?oi? iiooaiiy eieineiai? eonee),
Br1 i Br2 (iiooaiiy /eaieea eieiniaiai no?eaeiy), Tg2 ? Tg3
(aei?noe?noue eieineiai? eonee), CdI (?iciiae?e i?aiaioo ii eieineia?e
eonoe?).

7. Cai?iiiiiaaii oa ai?iaiaaii niin?a o?iiiniiii? eieae?caoe??
aieiaieo aa-i?a e?euee?nieo iciae, ui a?oioo?oueny ia ?acoeueoaoao
iiia?aaeiueiai aaiaoe/iiai aiae?co iciaee, yea aea/a?oueny, aeey
aecia/aiiy aaiaoe/ieo aoaeo?a [d], [h], [i], [j], [l]: c/aieaiiy aaio,
o?iiiniiia eieae?caoe?y yeiai aea/a?oueny, c aaiii-ia?ea?ii niaoeeo?/ii?
o?iiiniie oanoo?oueny oeyoii ii??aiyiiy ia?e na?aaei?o cia/aiue. Ia?oee
/eai ia?e ? aii??e/ia na?aaei? cia/aiiy a?oie ?ineei F2 c ia?ea?iei
oaiioeiii, a ae?oaee – oai?aoe/ia, ui ?ic?aoiao?oueny c
o?aoo-aaiiyi aaiaoe/ieo aoaeo?a oa caeiiii??iinoae oniaaeeoaaiiy
aaio-ia?ea?a.

8. Eiia?iaoe?? no?auoaaiue, ui ?o i?ecia/aii aeey eieae?caoe??
aieiaieo aai?a e?euee?nieo iciae, iiaeii? a?aeiia?aeaoe oaeei aeiiaai:
eiio?anoi? oaiioeie aaouee?a ca iciaeith, ui aeine?aeaeo?oueny, oa
iino?ei?noue ni?aa?aeiioaiiy na?aaei?o cia/aiue iciaee c ?ieo a ??e;
noaoenoe/ii cia/ou? aaiaoe/i? aoaeoe iiaeae?, ui aaeaeaaoii iieno?
aaiaoe/i? ?ica?aeiino? i?ae aaoueeaie ca e?euee?niith iciaeith,

yea aea/a?oueny.

9.O a?aeiia?aeiino? c? no?oeoo?ith aaiioei?a ca ia?ea?ieie aaiai ?
aeiiaaie, ui aenoaathoueny aei eiiiiiaio?a no?auoaaiiy noiniaii
e?euee?nii? iciaee, yea aea/a?oueny, ia?niaeoeaieie aeey iiaeaeueoi?
?iaioe c ?aeaioeo?eaoe?? ? aea/aiiy QTL ? eiia?iaoe??: Den o Dt28aeey
iciaee aeiaaeeia noaaea, i?aeaocey ? eenoea, Dea o Dt28 aeey iciae
e?euee?noue eieine?a o eiein?, iana ca?ai c aieiaiiai eieina, iana
iaeiiai ca?ia, Deno Dea aeey iciae aeiaaeeia eieina ? e?euee?noue
eieine?a o eiein?.

10.Io?eiaii iiioeyoe?? F6 ?aeiia?iaioii-?ia?aaeieo e?i?e a?ae
no?auoaaiiy on?o aaiiiii-aeiaeaieo oi?i ca ae?aeaeueiith noaiith. Aiie
o?enothoue i?ieea?noue aaenaiei?ae?a, ui ?ic/aieththoueny ia
aeeiei?aeiiio ??ai? ca aaiaie noaaaiiio D, ? aoaeooue
aeei?enoiaoaaoeny aeey ?aeaioeo?eaoe?? ? ea?ooaaiiy aieiaieo aai?a
e?euee?nieo iciae.

11.Noi?iiaaii iaai?e aaenaiei?aeieo e?i?e ioaieoe? c cai?uaiiyi
o?iiinii ?? noaaaiiio D ia aiiaieia?/i? o?iiiniie (ae?eyiee o?iiinii)
o?ueio aeae?a aa?eiino oa iaeiiai aeaeo ie??th; ie?ai? e?i??
aeeth/athoue iaeia, aeaa aai o?e i?aeaaiiiieo cai?uaiiy o?iiinii,
?aeaioeo?eiaaieo ca iaeaaeiinoth aei iaaii? aiiaieia?/ii? a?oie. Oeae
?ineeiiee iaoa??ae i?eaeaoiee aeey eiai aaiaoe/iiai aiae?co ca
iciaeaie, ui eiio?ieththoueny e?eoe/ieie o?iiiniiaie, oa iiaea
aeei?enoiaoaaoeny o naeaeoe?eiiio i?ioean? ye aeaea?aei aai?a, ye?
eiio?ieththoue no?ee?noue aei eenoiai? ??ae?, ai?ioienoi? ?ine,
i?aeaeuaio ii?icino?ee?noue oa ai?no a?eea o ca?i?.

NIENIE ?IA?O, IIOAE?EIAAIEO CA OAIITH AeENA?OAOe??

1. Oa?iianeay O.E. Aaiii D iyaeie ioaieoeu. Aeeaeeaeueiue aiaeec
aaaaoaoeaiuo i?eciaeia ?anoaiee ioaieoeu // Oeeoieiaey e aaiaoeea. —
1994 — 28, N 1. — N. 33-41.

2. Oa?iianeay O.E. Aaiii D iyaeie ioaieoeu. Aeeaeeaeueiue aiaeec
ia?iuo i?eciaeia eieina ?anoaiee ioaieoeu // Oeeoieiaey e aaiaoeea. —
1994. — 28, N 2. — N. 36-42.

3. Oa?iianeay O.E. Caiauaiea o?iiinii aaiiia D iyaeie ioaieoeu
o?iiiniiaie aaiiiia U eee Sl yaeeiinia // Oeeoieiaey e aaiaoeea. — 1994.
— 28, N 5. — N. 50-55.

4. Oa?iianeay O.E. Aaiii D iyaeie ioaieoeu. Ianeaaeiaaiea
iaeioi?uo i?eciaeia ii?oieiaee eieina // Oeeoieiaey e aaiaoeea. 1997. —
31, N 4. — N. 9-6.

5. Oa?iianeay O.E.Aaiaoe/aneee aiaeec aaiiia D ioaieoeu n
eniieueciaaieai aeeaeeaeueiuo ne?aueaaiee // Oeeoieiaey e aaiaoeea.
1999. – 33, N 2. — N. 3-10.

6. Oa?iianeay O.E. N?aaieoaeueiue aiaeec noaaaiiiia D e AB iyaeie
ioaieoeu a ioiioaiee aaiaoe/aneiai eiio?iey iaeioi?uo eiee/anoaaiiuo
i?eciaeia // Aeiiiaiaei IAI Oe?a?ie. – 1999, N 4. — N. 187-192.

7. AEe?ia A.A., Oa?iianeay O.E. Aaiiiiay eiaeaia?ey o ioaieoeu //
Aanoiee n.-o. iaoee. — 1984. — N 10. — N. 58-66.

8. AEe?ia A.A., Oa?iianeay O.E. Ia?aiin aaiiia D io iyaeie
ioaieoeu a oaa?aeoth // Aeiee. AI NNN?. – 1987. – 296. — N. 1252-1254.

9. Zhirov E.G., Ternovskaya T.K., Bessarab K.S. Investigation on
wheat cyto-genetics at Krasnodar Lukyanenko Research Institute of
Agriculture // EWAC Newsletter. – Plant Breeding Inst., Cambridge,
1986. — P. 48-52.

10. Zhirov E.G., Ternovskaya T.K. Substitution of D-genome in wheat
// European Wheat Aneuploid Co-operative Newsletter. — Agricultural
Res.Inst. of the Hungarian Acad Sci., Martonvashar and Inst. of Plant
Sci. Res.Cambridge Lab., 1987. — P. 60-65.

11. Zhirov E.G., Ternovskaya T.K. Transfer of D genome from common
wheat to durum wheat // Wheat Inform. Serv., 1988. — N 65.- P. 4-6.

12. AEe?ia A.A., Oa?iianeay O.E..Iia ee auoue eiieo?aio o aaiiia D
ioaieoeu // Oeeoieiaey e aaiaoeea. — 1989. — 24., N 3. — N. 45-48

13. AEe?ia A.A., Oa?iianeay O.E. Aanna?aa E.N. Niaoeaaiea aaiiiia a
o?eaa Triticeae. Aaiii D iyaeie ioaieoeu e aaiii iu?ay // Oeeoieiaey
e aaiaoeea. — 1990. — 24, N 4. — N. 15-19.

14. AEe?ia A.A., Oa?iianeay O.E. Aiaeec eiiuthaaoeee o?iiinii o
aea?eaeia ioaieoeu a nayce n i?ienoiaeaeaieai aa aaiiiia. Aeeieieaeiua
aea?eaeu // Aaiaoeea – 1993. — 29, N 1. N. 125-134.

15. AEe?ia A.A., Oa?iianeay O.E. Aiaeec eiiuthaaoeee o?iiinii o
aea?eaeia ioaieoeu a nayce n i?ienoiaeaeaieai aa aaiiiia. O?eieieaeiua
aea?eaeu // Aaiaoeea. – 1993. – 29, N 1. — N. 135-143.

16. AEe?ia A.A., Oa?iianeay O.E. Aiaeec eiiuthaaoeee o?iiinii o
aea?eaeia ioaieoeu a nayce n i?ienoiaeaeaieai aa aaiiiia.
Oao?aieieaeiua aea?eaeu // Aaiaoeea. – 1993. — 29, N 1. N. 144-153.

17. AEe?ia A.A., Oa?iianeay O.E. Ia?aaea/a ioaieoea Triticum
aestivum L. o?iiiniiu Aegilops sharonensis Eig., i?eaeathuae ae
onoie/eainoue e io/ienoie ?ina // Aaiaoeea. — 1993. — 29, N 4. — N.
639-645.

18.Oa?iianeay O.E., AEe?ia A.A. Aaiiiiue aiaeec ii?icionoie/eainoe
ioaieoeu // Aeieeaaeu ?innaeueoicaeaaeaiee. — 1993. — N 6. — N. 3-4.

19.Oa?iianeay O.E., AEe?ia A.A. Aaiii D iyaeie ioaieoeu. Aaiaoe/aneee
eiio?ieue i?eciaeia aineiaie iaeao, iiooaiea eieineiaie /aooe e ie?anea

c?aeiai eieina // Oeeoieiaey e aaiaoeea. — 1993. — 27, N 3. — N. 15-20.

20. Oa?iianeay O.E., AEe?ia A.A. Aaiii D iyaeie ioaieoeu.
Aaiaoe/aneee eiio?ieue i?eciaeia iyaeinoue eieineiaie /aooe e
aaeaaeaiiinoue aa iniiaaiey // Oeeoieiaey e aaiaoeea. — 1993. — 27, N 5.
N. 78-83.

21. Aioiithe I.C., Oa?iianeay O.E., Niceiia A.A. Eaeaioeoeeaoeey
aeieia yeaeo?ioi?aoe/aneeo eiiiiiaioia caianiuo aaeeia, eiaee?oaiuo
aaiaie o?ao aeaeia yaeeiina. — Oeceieiaey e aeioeiey eoeueoo?iuo
?anoaiee. — 1994. — 26, N 5. — N. 474-481.

22. Aioiithe I.C., Oa?iianeay O.E. Ecioa?iaiou aaoa- e
aeueoa-aieeacu aeey eaeaioeoeeaoeee aaiaoe/aneiai iaoa?eaea o?ao aeaeia
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aaiaoeea. — 1995. — 29, N 2. — N. 3-9.

23. Oa?iianeay O.E., Aioiithe I.C., Niceiia A.A. Caiauaiiua eeiee
iyaeie ioaieoeu n iaeiie-o?aiy ia?aie o?iiinii Aegilops sharonensis
// Aeiiiaiaei IAI Oe?a?ie. – 1995, N 12. — N. 118-120.

24. Oa?iianeay O.E., Aioiithe I.C. Aaiu aeioeie/aneeo i?eciaeia eae
ia?ea?u /oaea?iaeiiai aaiaoe/aneiai iaoa?eaea a aaiiia ioaieoeu //
Oeeoieiaey e aaiaoeea. — 1996. — 30, N 3. — N. 71-85.

25. Aioiithe I.C., Oa?iianeay O.E., Ceaoeeay A.A. Eaeaioeoeeaoeey
aaiia ca?iiaie ia?ieneaeacu o o?ao aeaeia yaeeiina // Aa?iaeieiaiy oa
aiioaoiieiaiy. Ca. iaoe.i?aoeue ei-oa aa?iaeieiai? oaaiioaoiieiai?.
Ee?a, Aa?a?ia iaoea, 1996. — N. 239-245.

26. Oa?iianeay O.E., Aioiithe I.C., Niceiia A.A.Eaeaioeoeeaoeey
/oaea-?iaeiiai aaiaoe/aneiai iaoa?eaea a eeieyo T.aestivum
/Ae.sharonensis, nicaeaiiuo ni/aoaieai o?iiinii Sl e D a ?aieao iaeiiai
aaiiia // Aa?i-aeieiaiy oa aiioaoiieiaiy. Ca. iaoe.i?aoeue ?i-oo
aa?iaeieiai? oa aiioaoiieiai?. Ee?a, Aa?a?ia iaoea, 1996. — N. 216-224.

27. Aioiithe I.C., Oa?iianeay O.E. I?eciaee ii?oieiaee ?anoaiee eae
ia?ea?u aiiaieiae/aneeo a?oii o?iiinii Triticeae // Oeeoieiaey e
aaiaoeea. — 1997. — 31, N 4. — N. 105-112.

28. Aioiithe I.C., Oa?iianeay O.E., Niceiia A.A. Eaeaioeoeeaoeey
o?iiinii iu?ay a /oaea?iaeii-caiauaiiuo eeieyo ioaieoeu n
eniieueciaaieai i?eciaeia ii?oieiaee ?anoaiee e aeioeie/aneeo ia?ea?ia
//Aeieeaaeu ?inneeneie aeaaeaiee naeueneioicyenoaaiiuo iaoe. — 1997, N
1. — N. 4-6.

29. Oa?iianeay O.E. Eco/aiea eiiaeiaoeeiiiie niiniaiinoe
oao?aieieaeiuo eiiiiiaioia AAAA iyaeie ioaieoeu ii iaeioi?ui
oicyenoaaiii-oeaiiui i?eciaeai // Naeaeoeey e aaiaoeea ioaieoeu . —
E?aniiaea?, 1982. — N. 184-191.

30. AEe?ia A.A., Oa?iianeay A.E., Eaaiia A.E. Aaiiiii-caiauaiiua
oi?iu ioaieoeu // Naeaeoeey e aaiaoeea ioaieoeu. — E?aniiaea?, 1985. —
(Na. iao/.o?./ EIEENO). — N. 84-102.

31. AEe?ia A.A., Oa?iianeay O.E. Caiauaiea aaiiia D iyaeie
ioaieoeu

T.aestivum L. aaiiiii Ae. speltoides Tausch. // Ananithciia niaau. ii
ioaea-

eaiiie aea?eaeecaoeee ?anoaiee e aeeaioiuo: Oac.aeiee.-I., 1981.- N.
126-127.

32. AEe?ia A.A., Aanna?aa E.N., Oa?iianeay O.E., Eaaiia A.E.
Nicaeaiea iiaiai enoiaeiiai iaoa?eaea aeey naeaeoeee ioaieoeu e
o?eoeeaea n iiiiuueth iaoiaeia aaiiiiie e o?iiiniiiie eiaeaia?ee//
Oai?. e i?eee. aniaeou naeaeoeee e naiaiiaiae. ioaieoeu, ?aee e
o?eoeeaea: Oac. aeiee. Iaaeaeoia?. iao/. eiio. no?ai-/eaiia NYA. —
Iaeanna, 1981. — N. 29.

33. Aioiithe I.C., Oa?iianeay O.E. Eaeaioeoeeaoeey aeieia
yeaeo?ioi?a-oe/aneeo eiiiiiaioia aeeaaeeiia e aunieiiieaeoey?iuo
aethoaieiia, eiio-?iee?oaiuo eieonaie /oaea?iaeiiai aaiiia a
aaiiiii-caiauaiiuo oi?iao ioaieoeu. — Iiaua iaoiaeu aeioaoiieiaee
?anoaiee // Oac.aeiee. I ?inneeneee neii. 18-20 iay, 1993, Ioueii. —
Ioueii, 1993. — N. 70.

34. Aioiithe I.C., Oa?iianeay O.E. Eaeaioeoeeaoeey eieonia a-Amy
e b-Amy a aaiiiao aaenaieieaeiuo aioeaeeieieaeia, iauaaeeiythueo
aaiiiu AAAA iyaeie ioaieoeu e aaiii iaeiiai ec o?ao aeeieieaeiuo aeaeia
yaeeiina // Oac. aeiee. ia niaau. «Iieaeoey?ii-aaiaoe/aneea ia?ea?u e
naeaeoeey ?anoaiee». Eeaa, 10-13 iay 1994. – N.9-10.

35. Ceaoeeay A.A., Aioiithe I.C., Oa?iianeay O.E. Aaiiiiue aiaeec
i?eciaea niaea?aeaiea aaeea a ca?ia ioaieoeu n eniieueciaaieai aa
aeaiieeieieaeece?iaaiiuo oi?i // Oac. aeiee. iaaeaeoia?. eiio.
«Iieaeoey?ii-aaiaoe/aneea ia?ea?u ?anoaiee». sseoa, 11-15 iiya?y,
1996. — Eeaa, Aa?a?iua iaoee: 1996. — N. 22.

36. Aioiithe I.C., Oa?iianeay O.E., Niceiia A.A. Eaeaioeoeeaoeey
o?iiinii iu?ay a /oaea?iaeii-caiauaiiuo eeieyo ioaieoeu n
eniieue-ciaaieai i?eciaeia ii?oieiaee ?anoaiee //Oac. aeiee.
iaaeaeoia?. eiio. «Iieaeoey?ii-aaiaoe/aneea ia?ea?u ?anoaiee». sseoa,
11-15 iiya?y, 1996. — Eeaa, Aa?a?iua iaoee: 1996. — N. 11-12.

37. Oa?iianeay O.E. Aaiaoe/aneee aiaeec ioaieoeu ii eiee/anoaaiiui
i?eciaeai, eiio?iee?oaiui aaiaie noaaaiiia D // Oac.aeiee. iaaeaeoia?.
eiio. «Iieaeoey?ii-aaiaoe/aneea ia?ea?u ?anoaiee». sseoa, 11-15
iiya?y, 1996. — Eeaa, Aa?a?ia iaoea: 1996. — N. 37.

Oa?iianueea O.E. Ia?aaoaeiaa aaiiio i’yei? ioaieoe? (Triticum aestivum
L.)aeey ?? aaiaoe/iiai aiae?co oa ?io?ia?an?? aai?a. — ?oeiien.
Aeena?oaoe?y ia caeiaoooy ia-oeiaiai nooiaiy aeieoi?a a?ieia?/ieo iaoe
c? niaoe?aeueiino? 03.00.15 — aaiaoeea. ?inoeooo ee?oeiii? a?ieia?? ?
aaiaoe/ii? ?iaeaia??? IAI Oe?a?ie, Ee?a, 1999.

Aeena?oaoe?th i?enay/aii ?ic?iaoe? oeyo?a oa iaoiae?a aeei?enoaiiy
aaena-iei?aeieo oi?i ioaieoe? c ia?aaoaeiaaie aaiiio aeey aaiaoe/iiai
aiae?co i’yei? ioaieoe? ? caaaa/aiiy ?? aeaeiaiai aaiaoe/iiai ioeo ia
aaie a?ae aeeei?ineeo ?iaee/?a, ui eiio?ieththoue, aa?iiii?/ii aaaeeea?
iciaee. Anoaiiaeaii aianie

noaaaiii?a AA oa D i’yei? ioaieoe? a eiio?ieue aeayeeo e?euee?nieo iciae
eieino oa noaaea. Aeyaeaii caeaaei?noue aaiaoe/ii? no?oeoo?e iiioeyoe??
aaouee?anueeeo oi?i ca aaiaie, ui eiio?ieththoue ??ciiiai?oi? iciaee,
a?ae oeio iciaee (iciaee-eiiiiiaioe i?iaeoeoeaiino? — iciaee
aaaaoaoeaii? /anoeie ?ineeie). Iaea?aeaii ?aeiia?iaioii-?ia?aaei? e?i??
F6 a?ae no?auoaaiiy aaenaiei?ae?a c ia?aaoaeiaaiei aaiiiii ca
ae?aeaeueiith noaiith. Noai?aii iaai?e /oaeeiii-cai?uaieo e?i?e
ioaieoe?, aaiii yeeo aeeth/a? a?ae iaei??? aei o?ueio o?iiinii aeae?a
aa?eiino aai ie??th cai?noue aiiaieia?/ieo o?iiinii noaaaiiio D
ioaieoe?. E?i?? oa?aeoa?ecothoueny aeayeeie aa?iiii?/ii-aaaeeeaeie
iciaeaie ? iiaeooue aeei?enoiaoaaoeny o naeaeoe?? i’yei? ioaieoe? ye
aeiii?e ei?enieo aai?a.

Eeth/ia? neiaa: i’yea ioaieoey, aaie-ia?ea?e, aaiaoe/iee aiae?c,
ii?oieia?/i? iciaee, e?euee?ni? iciaee, ?io?ia?an?y aai?a.

Oa?iianeay O.E. Ia?ano?ieea aaiiia iyaeie ioaieoeu (Triticum aestivum
L.) aeey aa aaiaoe/aneiai aiaeeca e eio?ia?annee aaiia. — ?oeiienue.
Aeenna?oaoeey ia nieneaiea o/aiie noaiaie aeieoi?a aeieiae/aneeo iaoe ii
niaoeeaeueiinoe 03.00.15 — aaiaoeea. Einoeooo eeaoi/iie aeieiaee e
aaiaoe/aneie eiaeaia?ee IAI Oe?aeiu, Eeaa, 1999.

Aeeiaaenaieieaeiay i?e?iaea aaiiia iyaeie ioaieoeu iaoneiaeeaaao
o?eieeoee?iaaiiinoue aaiaoe/aneeo nenoai, eiio?iee?othueo ioaeaeueiua
i?eciaee. N yoei naycaia neiaeiinoue aa aaiaoe/aneiai aiaeeca e
iio?aaiinoue a niaoeeaeueii nicaeaaaaiii aaiaoe/aneii iaoa?eaea —
aiaoieieaeiuo eeieyo iyaeie ioaieoeu, aaiii eioi?uo ia?ano?iai ii
iaeiie-aeaoi o?iiiniiai. Aeenna?oaoeey iinayuaia eco/aieth aiciiaeiinoe
eniieueciaaoue aeey aaiaoe/aneiai aiaeeca iyaeie ioaieoeu
aaenaieieaeiua oi?iu, ia?ano?ieea aaiiia eioi?uo einioeanue ia
ioaeaeueiuo o?iiinii, a oeaeiai noaaaiiia — D, iaeiiai ec o?ao
noaaaiiiia, ninoaaeythueo neiaeiue aaiii ioaieoeu (A, B e D). Yoi —
aaiiiii-aeiaaaeaiiua oi?iu n iaeiei e oai aea eiiiiiaioii AA e ?aciuie
noaaaiiiaie D, e aaiiiii-caiauaiiua oi?iu n iaeiei e oai aea eiiiiiaioii
AA e o?aoueei noaaaiiiii, iieo/aiiui io iaeiiai ec aeeei?anooueo
ni?iaee/ae ioaieoeu Ii?aaeaeaiu iooe eniieueciaaiey aaenaieieaeia n
ia?ano?iaiiui aaiiiii aeey aaiaoe/aneiai aiaeeca ioaieoeu. Noaiaea?oiue
aea?eaeieiae/aneee aiaeec a naiai eeanne/aneii aeaea i?eaieai aeey yoiai
iaoa?eaea, iineieueeo a ?anuaieythueony iiioeyoeeyo io ne?aueaaiey
aaiiiii-aeiaaaeaiiuo oi?i ana ?aciiia?acea i?ienoiaeeo ca n/ao
?anuaieaiey oieueei ii aaiai noaaaiiia D — aeeieieaeiue oei ?anuaieaiey.
A ne?aueaaiea aiaeaeathony ?anoeoaeueiua oi?iu, naaeaine?iaaiiua ii
/eneo o?iiinii (a ioee/ea io iiii- e ioeeeniieeia) e acaeiiioiioaieth
yae?i-oeeoiieacia (aaenaieieaeiue o?iaaiue aaiiia ?anoaiee a oeaeii i?e
aeeieieaeiii oeia ?anuaieaiey). Aeey aiaeeca eiee/anoaaiiuo i?eciaeia
i?eiaieiu ethaua iaoiaeu, ?ann/eoaiiua ia aeeieieaeiue i?aaieci. A
?aaioa eniieueciaaiu aeeaeeaeueiua ne?aueaaiey e iauaaeeiaiiue oano
Eaaaeee aeey aaiaoe/aneiai aiaeeca ?anoeoaeueiuo oi?i ii
eiee/anoaaiiui i?eciaeai. Onoaiiaeaia caaeneiinoue aaiaoe/aneie
no?oeoo?u aaiiia ii nenoaiai aaiia, eiio?iee?othueo ioaeaeueiua
i?eciaee, io oa?aeoa?a i?eciaea. A nenoaiao aaiia, eiio?iee?othueo
i?eciaee aaaaoaoeaiie /anoe ?anoaiey, i?aiaeaaeatho aaiu n
aaeaeeoeaii-aeiieiaioiui oeiii iaaeaaiiiai e iaaeaeeaeueiiai
acaeiiaeaenoaey e /anoiou aeiieiaioiuo aeeaeae, oaaee/eaathueo eee
oiaiueoathueo cia/aiea i?eciaea, ?aaiicia/iu. Aeey aaiaoe/aneeo nenoai,
eiio?iee?othueo i?eciaee i?iaeoeoeaiinoe eieina, oa?aeoa?ii o/anoea
aaiia n aeoieeeaoiui yienoacii e i?aiaeaaeaiea aeiieiaioiuo aeeaeae,
oiaiueoathueo /eneiaia au?aaeaiea i?eciaea. Eaeaioeoeoee?iaai ?yae
iiauo aaiia, eiio?iee?othueo i?eciaee ii?oieiaee eieina ioaieoeu.
Onoaiiaeaii, eaeea aaiu iiaeii eniieueciaaoue eae ia?ea?u
ii?aaeaeaiiuo o?iiinii noaaaiiia D. Yoi aaiu, eiio?iee?othuea iaeioi?ua
ii?oieiae/aneea i?eciaee, e aeioeie/aneea aaiu. Iieacaia aiciiaeiinoue
eniieueciaaiey aaiia-ia?ea?ia ioaeaeueiuo o?iiinii a ni/aoaiee n
?acoeueoaoaie aaiaoe/aneiai aiaeeca ?anoaiee ii eiee/anoaaiiui i?eciaeai
aeey o?iiiniiiie eieaeecaoeee aeaaiuo aaiia, o/anoaothueo a eiio?iea
?yaea ia?iuo i?eciaeia. Ii?aaeaeaiu iioeiaeueiua eiiaeiaoeee ne?aueaaiee
aeey ea?oe?iaaiey QTL. Iieo/aiu iiioeyoeee ?aeiiaeiaioii-eia?aaeiuo
eeiee F6 io ne?aueaaiey aaiiiii-aeiaaaeaiiuo aaenaieieaeia ii
aeeaeeaeueiie noaia. Eeiee i?aaenoaaeytho niaie oeaiiue iaoa?eae aeey
aaiaoe/aneiai aiaeeca noaaaiiia D iyaeie ioaieoeu.

Aeey nicaeaiey /oaea?iaeii-caiauaiiuo eeiee iyaeie ioaieoeu
eniieueciaaii ne?aueaaiea iyaeie ioaieoeu e aaiiiii-caiauaiiuo oi?i n
aaiiiaie AABBUU (UU — aaiii Aegilops umbellulata), AABBSS (SS- aaiii
Ae.speltoides), AABBSlSl (SlSl — aaiii Ae.sharonensis), AABBXX (XX —
iaeei ec ianeieueeeo noaaaiiiia Agropyron glaucum). Aeieacaia
yooaeoeaiinoue iaoiaea «niaoeaaiey» o?iiinii ?aciuo aaiiiia a i?aaeaeao
iaeiiai noaaaiiia iieeieieaeiiai aaiiia aeey nicaeaiey
/oaea?iaeii-caiauaiiuo eeiee. Eiaaony i?aeiouanoai ia?aae o?aaeeoeeiiiui
iaoiaeii: e?iia eeiee n iaeiie caiauaiiie o?iiiniiie iieo/aiu eeiee n
caiauaieai ianeieueeeo ioaie/iuo o?iiinii ia /oaea?iaeiua aiiaieiae/iua
o?iiiniiu. Oaeo caiauaiey eiinoaoe?iaaee ii iaee/eth o ?anoaiee
/oaea?iaeiuo i?eciaeia a ni/aoaiee n ?acoeueoaoaie eco/aiey iaeica
aea?eaeia /oaea?iaeii-caiauaiiuo eeiee n iyaeie ioaieoeae. Aeey
eaeaioeoeeaoeee /oaea?iaeiuo o?iiinii ii eo aiiaieiae/aneie
i?eiaaeeaaeiinoe eniieueciaaee aeioeie/aneea aaiu-ia?ea?u ?aciuo
aiiaieiae/aneeo a?oii Triticinae. Nicaeaiu /oaea?iaeii-caiauaiiua
eeiee, aeeth/athuea a naie aaiii io iaeiie aei o?ao ia? /oaea?iaeiuo
o?iiinii ethauo aiciiaeiuo ni/aoaieyo. Iieacaii, /oi /anoioa ia?aaea/e
/oaea?iaeiuo o?iiinii 1, 2 , 3, 4, 6 e 7 aiiaieiae/aneeo a?oii
eae iiiaeeii/ea, oae e a o?iiinii a iyaeoth ioaieoeo caaeneo io
aiiaieiae/aneie a?oiiu e ia caaeneo io aeaea-aeiii?a o?iiiniiu. N?aaee
nicaeaiiuo eeiee anoue iaeaaeaoaee iieiie onoie/eainoe e io/ienoie ?ina,
ao?ie ?aeaa/eia, eeiee n aunieei niaea?aeaieai aaeea a ca?ia e
ceiinoieeinoueth, i?aauoathuae ceiinoieeinoue iyaeie ioaieoeu. Eeiee
oeeoieiae/anee noaaeeueiu, naiioa?oeeueiu, ne?aueaathony n iyaeie
ioaieoeae, aeaaay oa?oeeueiia iioiinoai, e iiaoo iaiin?aaenoaaiii
eniieueciaaoueny a naeaeoeee iyaeie ioaieoeu eae aeiii?u /oaea?iaeiuo
aaiia, eiio?iee?othueo oicyenoaaiii-oeaiiua i?eciaee ioaieoeu.

Eeth/aaua neiaa: iyaeay ioaieoea, aaiu-ia?ea?u, aaiaoe/aneee aiaeec,
ii?oieiae/aneea i?eciaee, eiee/anoaaiiua i?eciaee, eio?ia?anney aaiia.

Ternovskaya T.K. Reconstruction of common wheat (Triticum aestivum L.)
genome for wheat genetical analysis and genes introgression. —
Manuscript. Thesis on a scientific title of the Doctor of Biological
Science, specialization 03.00.15 — Genetics. Institute of Cell Biology
and Genetical Engineering , National Academy of Sciences of Ukraine,
Kyiv, 1999.

Dissertation is devoted to the elaboration of the lines and
methods of genetical analysis of common wheat and enrichment of its
genetical pool through useful genes from wild related species using
wheat hexaploids sharing the genome reconstructed.

The genetical contribution from subgenomes AB and D to control of some
quantitative traits of wheat ear and stem was determined. The
dependence of genetical structure of parents population for genes that
control different traits from type of traits studied (traits that are
the components of productivity or vegetative traits) has been revealed.
The families of recombinant-inbred lines F6 from diallel crosses of
hexaploids with reconstructed genomes has been derived. The sets of
wheat alien substitution lines that share in their genomes from one to
three chromosomes of Aegilops species or Agropyron glaucum instead
homoeological chromosomes of wheat D genome were developed. These lines
are characterized with agriculture useful properties and can be used in
common wheat breeding as the donors of useful genes.

Key words: common wheat, marker genes, genetical analysis, morphological
characters, quantitative traits, genes introgression.

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