四倍体コムギ品種エンマーおよびピラミダーレの開花特性とその遺伝的制御機構

J.CropRes.56:67-71

!tr!

Research

Article

(2011)

Flowering

traits

and

their

_genetic

basis

in

the

ancestral

tetraploid

wheat yarieties `Emmer' and `Pyramidale'

[fetsuyaNakazak{i),RyiijiMoriyamaiL HisashiKagatai),HiroyoshiINlakaharaV,Mika NaitoD,

KeisukeKatsurai),HirokiSaitoi),KeajiKato2LHidetakaNishida2),1laihachiKawahara3),

D

D

TakashiFudano

andAkiraKitajima

i)

Graduate

SchoolofAgriculture,

kyoto

Uinivensity

(Experimental

Farm,

12-1

Hatchonawate,

Talcatsuki,

Osaka

569

-

O096,

Japan)

2)

GraduateSehoog

ofIVI]tural

Sbienceand 7bchnolo{plOkcDJamaU)iiversity

(I-1-1

fsushima-naka,Kita-ku,Okayama 700

-

8530,

Japan)

s)

GraduateSbhoolqfAgricultune,1<)?otoUhiversity

(Plarit

Germ-plasmInstitute,Mozume, Muko, Kyoto617

-

OOOI,Japan)

tsi r

Summary: Thefloweringtraitsof Emmer and PyramidaEe, two ancesma1 wheat varieties tltatare new used

temake two

_types

of beerbrewedin_partnershipwith Kyoto Uniyersityscientists, were investigatedindetail. These varieties are classified as T turgidum L.ssp. dicocconand ssp. turanicum, respectively, and little inforrnatienisavailable about theiragronomic characteristics, Through evaluatjon of the internal

factors

determining

theirfloweringtime

_{(photoperiodic}

response, vernalization requirement and naiTow-sense earliness), we revealed thatthetwo varieties are spring-habit and _{photoperiod-sensitive.}Whilethenarrow-sense earliness of

'Emmer'

islessintensethanthatof `PyramidaleZ the_{photoperiodic}response of

`Emmer'

ismore intensethan thatof

`Pyramidale'.

Based on a_geneticanalysis using theF2_populatienofthe twovarieties, we concluded that

`Pyramidale'

harborsa

_{photoperiod-insensitive}

alleleintheIlpcl-Allocus,thesame allelereported

by

Wilhelm et al.

(2009),

This_{photoperiod-insensitive}alteleisexpected tobeausefu1 _geneticresource inthebreedingof

tetraploidand hexaploidwheat.

Key words: floweringtrait,tetrapleidwheat, _photoperiodinsensitivity)

Rpcl-A1

tntroduction

KyotoUniversitMINhsedaUniversityand KizakuraCo.Ltd. have

_jointly

developedthree brandsof beer:

"White

Nile"

(a

regular

beer)

in2006,"Blue

Nile"

(a

low-maLt

beer)

in

2007,

and "Ruby

Niie"

(a

regular

beer)

in

2oo8

(Fig.

1)

(http:11www.

kyoto-u.ac.jpfenlnewsLdatalh!hllnews7t2009/09e420-1.htm).

" -J.

While BlueNile is made with modern durum wheat strains

(T}'iticum

turgidum L.ssp,

durum),

theothertwoaremade with

ancestral tetraploid strains: "White Nile"with emmer wheat

CEmmer'>

and "Ruby Nite"with _pyramidaiewheat

(`Pyramidaie'),

which are classified as T turgidum L.ssp. dicocconand ssp.

turanicum, respectively, The two ancestral varieties were

cellected fromtheiroriginal areas of

distribution,

namelM _the

southern Medlterraneanbasin,and havesince been_preservedat

theLaboratoryofCrop Evolution,P]antGerm-plasm Institute, GraduateScheol of Agriculture,Kyoto University.We have produced

foundation

seeds for`Ernmer' and `Pyramidale' at

the

ExperimentalFarm of Kyoto Universitysince

2005

and

2008,

respectively, in order to distributethe seeds for

consignment cuttivation. Both varieties haye traitsthatmake them unsuitable forcu]tivation according tetraditionalmethods inthesoutheast area of Japan,such as a Jongculm, a lackof

tolerance against disease and excessive moisture, and atendency

to sprout

before

harvest.

Because so

little

informationis

¢

.･･

ttt

t

tt..

_.

.

_t.･t.･..

ttt

'

' es/.

ttt

tt

x.}.･

tt/

tt.//t./tt/.//lt...

ts.t..t

.ff,l,

tt/

tt

tt

Acccepted :April 25,2011

Correspondingauthor :

'retsuyaNakazaki([email protected])

Fig.1Threebrandsof beerdevelopedbyKyotoUniversity,

Waseda Univefsityand KizakuraCo.Ltd.

"White Nile"(right),

"Blue

Nile"{left}and

"Ruby

Nile" {center}.

The Society of Crop Science and Breeding in Kinki, Japan

NII-Electronic Library Service

TheSociety of Crop Science and Breeding in Kinki, Japan

J.CropRes.

56(2011)

available about the bestcultivation methods

for

these two

varieties, such as

fertilization

with manure, or about theirbest cropping seasons, ithasbeennecessary todeterminetheoptimal proceduresthrough trialand error.

Among themany variables _thatcan

be

adjusted

in

such trials, cropping season, that is,thelengthof time from seeding to

haryest,is_particularlyimportantfor_yieldand _{grainquaiity.}Tb determinean appropriate lengthforthe cropping season, itis

essentialte clarify the

flowering

traitsof each variety. The

fiewerlngtraitsof wheat depend on three internal factors, photoperiodieresponse, vernalization requirement and narTow-sense earliness

(Yasuda

and Shimoyama 1965,Hoogendoorn 1985,Kato and Ylamagata1988).According te Kato and

Yamagata

(1988),

photoperiodlcresponse refers to the retardation or acceleration of headinginresponse to changes in daylength.Thiscan beevaluated byrneasuring thediffbrencein

the number of daysfromthefu11yvernalized stage to heading

time between _plantsraised under short-day and long-day conditions. The

_{phetoperiodic}

response inwheat is

determined

bythreehomoeologous _genes,llpcl=Ai,llpd-BJand llpd-Dl, which are orthelogeus to the_{pseudoresponse}regulatoT _gene

(Beales

et al.

2007).

Inhexaploidwheat, the_genesconferring photoperiod

insensitivity

have

been

identifiedon chromosomes

2B

and

2D

<WOrland

and Law

1986,

Mohleret al.

2004,

Beales

et al.2007).In tetraploidwheat, _two allelesofthe llptMl lecus which cause a_{photoperiod-insensitivephenetype}haverecently beenidentified

_(Wilhelm

etat. 2009).Vernalizationrequirement isevaluated by measuring theminimum durattonof chi11ing treatmentthat isnecessary forfu11vernalization

(Kato

and

Yainagata

1988).

So fhr;threekinclsof _genes

(homoeologous

genes)thatdeterminethe vernallzation response inwheat have beenisotated,YRIVI

_(Yan

et al.

2003),

VRAI2

(Yiin

et ar.

2004)

and VR?V3

(Yan

et al. 2006).Narrow-senseearliness isdefined as the earliness of

fu11y

vernalized _plantsunder optimum conditions

for

reproductlve _grewth

(lhkahashi

and Yasuda 1958).Itcan be evaluated by rneasuring thenumber of days fromthefu11yverna]ized stage to headingtime under optimum conditions forreproductive _growth,includingof 24hdaylighi

(Kato

and Yimagata 19zz).

Several

QTLs

controlling

narrow-sense earliness havebeenidentified

(Cockrarn

etal.

20e7).

Inthisstudy, thethreeintemalfactorscontrotting fiowering

.

.i

1i

tt

traitswere evaluated in Emmer and Pyramidaleusing the

rnethod deve]opedby Kato and Yamagata

(1988).

Sincethis

evaluation uncovered a iargediffereneebetween the two

varieties in_{photoperiodie}response, a segregation analysis for

thetraitwas also _performedinthisstudM using an F2_population

between`Emmer'

and `PyramidaEeZ Here,we report the flowering

traitsand the _genetic

basis

of the _{photoperiodic}response

eharacteristics ofthese twoancestralvarieties.

Materials

and

Methods

Plantmaterials

sti

,

Emmer, Pyramidale and the

hexaploid

wheat

(T

aestivum L.)variety `Norin

61'

were subjected to analysis of the three

factorscontrolling floweringtraits.A totalof 148 F2 _plants

xt

i

!

between

Emmer

and

Pyramidale

were used toanalyze the phenotypic

fiowering

response.

Evaluation

ot

flewering

traits

Seedsof thethreevarieties were _geTrninatedon a wet filter paper at 20℃ for two days.To evaluate vernalization requirement and narrow-sense earliness, each

_germinating

seed

was _putinto a 2-crn-squarecell trayfi11ed with soil and subjected to chilling treatmentat

4

℃

for

various durations

(O,

7,

14,

21

or

28

days)under

24-h

daylightconditiens. fen _plantswere thus

treatedforeach duratlon.Afterthe chilling treatment,the cell

trayswere maintained at20℃ uncier 24-hdayliglttconditions,

We rneasured the nurnber of daysfromtheend of thechilting

treatment totheunfoldlng ofthefirstleagand thenurnber of

daysfromtheend of thechilling treatment totheunfolding of

the aag leafineach _group,From these datawe calculated the

Deg thatis,theadjusted number of daysftomthe_plantingof a

seed

in

its

cel]traytotheunfolding of

lts

flag

leaC

according to

themethed of Kato and Yamagata

(1988).

The strength of the effect of the durationof the chiiling treatment on the Dof indicatesthedegreeof a _genotype'svernalization requirement, while the Dof itselfindicatesthedegreeof that _genotype's

narrow-sense earliness

(Kato

and Yamagata1988).

Ib cstimate photoperiodic response,

30

seedtings of each

variety were _preparedas describedabove. After

42

daysofthe

chil}ing treamient

_(suMcient

forfu11vernalization),

15

seedlmgs of each variety were _grown at

20

℃ under

12-h

daylight

conditions

(SD)

while theother 15were _grown at 20℃ under

24-h

daylighr

cenditions

(LD).

The

number ef

days

until

heading,

excluding

the

time spent

in

thechilling treatment,was recorded foreach _plantas the Doh. The differenceinDoh betweenthe SD and LD seedlings isreferred tothedegreeof photeperiedicresponse inthisstudy.

For our segregation analysis ofphotoperiodic response inthe

F2_population,

148

Fzseedlings and tenparentalseedlings were preparedas describedabove. After

37

days of the chil]ing

treament, the seedlings were grown under SD conditions at

20

℃,and thenumber ofdays untilheadingwas recorded, Assay

for

lpct-A

falleles

The DNA of

'Emmei,

`Pyramidale', `Norin 61'and each

plant

intheF2

_population

was extractedfromleavesefseedlings grownforabout two weeks aftertheend ofthe chilling treatment

according to themethod of Zheng et aL

(1995).

Tbdetectthe

allele at the flpcltr41locusineach _plant,PCR analysis was performedusing three _primersdevelopedby Wilhelinet al.

68

Floweringtraitsand their_geneticbasisintheancestral tetraploidwheatvarieties

4

ftEmmer

and PyramidaIe

t

(2009):

durum-Ag5del-Fl,

durum-Ag5del-F2

and

'

Ag5delRl. PCR was conducted using BIOI:AQ DNA Polymerase

_(Bioline

LabsLtd.,London,UK) with thefo11owing PCR _profile:

95

℃

for

5

min,

foI{owed

by

35

cycres each

consistingof96 ℃ for

30

s,

55

℃ for

1

min and

72

℃

for

2min,

with a finalextension

72

℃ for

5

min. Sequencingof thePCR

ir- s

productsfrom Emmer and Pyramida]ewas ordered frem and _perfbrmedbyFASMAC Co.,Ltd.,Japan.

Results

and

Discus$ion

Flowering

traitsof `Emmer' <sub>and</sub> `Pyramidale'

The changes of Dof values inaccordance with thelengthof

the chilling treatment,observed forthreevarieties, are shown in Fig,2. In

`Emmer'

and

`Pyramidalel

forexample, the Def

resulting

from

seven

days

ofchillingtreatmentwas significarrtly

different

fromthatresulting from

14,

21

or

28

daystreatment

(P<O.05;

t-test).Theminimum durationofchi]]ing necessar}J to

inducevernalization was 14 days inbothvarieties, and the

differencesinDof between _plantssubjected to no chiiLing

treatment and thesesubjected tofuLlyvernalizing treatment were

only 1.7daysfor`Emmer' and 1.6for`Pyramidale'. Therefore we concluded

that

both

of

these

are strongly spring-habit varieties. `Norin

61',

on

the

other

hand,

did

not show aconstant

Dof

yalue

in

thisexperiment, This result indicates

that

the

minimum durationof chilling forthisvariety ismoie than28 days;this iscQnsistent with the

30-day

minimum duration repertecl ina_previousstudy

(Kato

and Yamagata 1988).

Theaverage Dofvalues of fulLyvernalized _plantsgrownunder

24-h

daylight

conditions correspond tonarrow-sense earliness. In

this

study,

the

narrow-sense earliness va]ues of `Emmer' and

`Pyrarnidale'

were estimated at

26.5

and

30.3,

respectively; the

differencebetweenthem was significant

(t=8,69,

_p<O.OOI).This

4 t result suggests thatthenarrow-sense earliness of Pyramidale

islessstrong thanthatof `Emmer',

Doh values under theSD and LD conditions were significantly

different

inatlthethreevarieties: thediffl]rencesin

`Emmer',

37 35 33go-31o 29 27 25 Flg.2 O 7 t4 21 2B

Durationof eh"Hng treatment (days) Changes inDof according toduration ofchHlhg

treatment observed forthreewheat varieties.

soo Fig,3 80i

_d

l

cC 6o

I

4ol

b

. ba I i i20

I.

I I i

'

O Iww.

...

.umww

.-

-.-...

....

ma.

Pylamidare Emmer Norln61 Doh values of thethreevarieties under SD _tolack)

anct LD

(white)

eonditions.

Dch values were compared using Tukey's tesL

Lettersoverthebars _indicatesignificantdMerence

each other atthe5% level.

`Pyramidale'

and 'Norin

61'

were

40,1,

20,O

and

6,8,

respec-tiyely

_(Fig.

3).

AlthoughDoh,unlike Dog iscalculatedwithout

an adjustment _procedure,thedifferenceinDoh

_between

theSD and LD _groups ismainly caused

by

the _intensityof their

photoperiodic

response. Therefore, _the strengths of their

lt

photoperiodie responses can be ranked as fo11ows:Emmer >

`Pyramidale'

> 'Norin

61'.

Thisfurther suggests that XEmmer'

-

t

and Pyramidaleare _{photoperiod-sensitive}varieties and elearly shows thatthe degreeof _photoperiodsensitivity is_greaterin

iJ

.;

J

Emmer thanm Pyramidale

,

Thephotoperiod-insensitivity of

tNorin

61'

cou}d becaused bya mutation intheIlpdiDllecus

on chromosome

2D

_(Tanio

et aL

2005);

the_geneticbasesof the

photoperiodicresponses in

`Emmer'

and

`Pyramidale'

remain unclear, howeyer.

Alleles

of thel

_PdLA

Ilocus of `Emmer'

and `Pyramidale'

Intetraploidwheat varieties, twe alleles ofllpd-Al thaicenfer

phetoperiodinsensitivjtyhave _beenreported; thesehave

1,027-bp

or

1,117-bp

deletions,respectively, upstream of the coding

region

(Wilhelm

et al.

2009),

To clarify the_genotypesof

`Emmer'

and

'Pyramidale'

atllpc-A

1,

a

PCR

assaywas conducted

according to

the

method

by

Wilhelm

et al.

(2oo9),

Two

types

of

PCR _product were obtained foreach of thethreevarieties

(Fig.

4).

WhilethePCR _productfrom `Pyramidale'

islikelytobethe

same as thatfromthe1,117-bpdeletion-typealleLe, thesize of which is

290

bp,the PCR _productsfrom 'Emmer'

and `Norin

61'

arelikelyto

_be

_thesame as that

from

theno-deletien-type

aLlele,

the

size ofwhich

is

452

bp.

Sequencinganalysis ef thePCR _productsfrom `Emmer' and

i

t

Pyramidalerevealed thatthesequences of theirPCR _products

are identiealto thoseof the ne-deletion-type allete

(acc.

EUI17148) and the

1,117-bp

deletion-typeallele

(ace.

EUI17149), respectively. Therefore we concluded that

`Pyramidale'

hasa _{phetoperiod-insensitive}aElele at theJlpd-Ai

Lt

The Society of Crop Science and Breeding in Kinki, Japan

NII-Electronic Library Service

TheSociety ofCrop Science and Breeding in Kinki, Japan

J.CropRes.56(2011)

soe

bp

rleebp

Pyramidale ErnrneF NeTln6a

M

Fig.4 PCR assay tordetecting a[leles atthe PPdLA1

Iocusof`Pyramidale'. `Emmer:

and "Norin 61

1

Arrows indlcatethetargetPCR products. M:1OO-bpDNA laddermarkei,

thesequenees of theremaining regions still need tobeanalyzed in

a

future

study.

Genetic

analysis of

tlowering

varlation

in

the

F2

population

We co/nducted a_geneticanalysis of Deh using theF2_population between

`Emmer'

and `Pyramidale'. Atotalof148 F2_plantswere clearly dividedintoearly- and ]ate-fiowering

types

(Fig.

5).The segregation of earty-type

(111

_plants)and late-type

(37

_plants) showed a _good fitto

3:1

ratio

(x2

==

O.32,

P==

O.57),

_suggesting

thata single locusisresponsible forDof variation. InthePCR assay

for

_genotypes at Ili]clzdlin

the

F2 _population,the

segregation ofplants havinghomozygous

deletion-type

alleles

(34

plants),heterozygousalleles

(77

plants),

and homozygous no-deretion-type alleles

(37

_plants)fitthe

1:2:I

ratioexpected for one-locus segregation

(

_xZ=:

O.36,

P=:

O,83).

The PCR _assay also

showed that the _presence of the deletion-typeallele was

completely correlatedwith early flowering

(Fig.

5).Thus,itcan

beconcluded

that

theIow

level

of photoperiod sensitivity

in

ut-cNa-eLooE=z5o302e10 D

d

e

n

:/l

70 Fig. se 6o 7o so Deh

5 Frequencydistributionot Doh in

the

F2 _plants

and the_parents

FEmmer'

and

`Pyramidale'.

Parent plants are shown on the upper row;

`Pyramidate'

are shown inbtack and

tErnmer'

ln

nthite.In theF2popalation,plantshomozygoustor

the deletion-typeallele are sbown inbLackand

plantshomozygous forthene-deietion-type allele are

shown invvhite on thelowerrow, Heterozygotes are

shown ingey inthisrow.

': _the

category intowhich more than 86 of Doh

values feli.

*

`Pyramidale'

ismainly due tothe effectofthe deletion-type

alleleofllpd-Al.

The F2 plantshomozygeus for the deletion-typeallele underwent headingearlier thanthe

`Pyramidale'

_plantsdid,and

the F2 _plantshomozygous for the no-deletion-type a)lele

it

underwent heading later than the Eminer plants did.This

discrepancybetweentheFz_p]antsand the_parentsmay be_partly

dueto theirmixed _geneticbackgroundcensjsting of

tEmmer'

and

`Pyramidale'

_genes,

but

itwas most likelycaused

by

an

unfavorable condition inthegrowth chamber

that

affected the

growh of

the

F2plants.Nevertheless,theeffect ofthe

deletion-type

allele on earLiness, includingthe mode of incomplete dominance

_(Fig.

5),iscompatible with theresults reported by Wilhelm et al.

(2009).

Thissuggests that `Pyramidale'

has a

photoperiod-insensitive allele at theIlr)d-Allocus,which might

be

the

1,117-bp

deletion

typeallele.

Conc[usion

Inthisstudy, the fioweringtraitsof the ancestral yarieties

4t i t

Emmer and Pyramidalewere reyealed and basicinformation

on a suitable cultivation _periodwas obtained. Both varieties are

spring-habit wheat strains. The narrow-sense earliness of

`Pyramidale'

is_greaterbyaboct fivedaysthan thatof

`Emmer'

,

whose narrow-sense earliness issimilar to thatof `Norin

61'.

Although both of the ancestral varieties show _photoperiod

sensitivity,tPyvarnidale'hasamore intensereaction than `Emmer' dose.This diillerenceiscaused by the effects of the I,117-bp

.t

)

deletion-typeallele in Pyramidaleand the no-deletien-type allele in

`Emmer'

attheIlpd=tl1locus,Inthisstudy vve identified

ti

r

t

a _photoperiodinsensitivityallele m Pyramidale thatwilE be usefu1 as a genetic resource

in

the

breeding

of earlyrnaturing varieties of tetraploidand hexaploidwheat.

Reterences

Beales,

J.,

A.

Turner,

S.

Griyths,

J,

W.

Snape

and D.A. Laurie

(2007)

A pseudo response regulator ismisexpressed in the

phetoperiodinsensitiveJlpd-Dlamutant of wheat

(7)"iticum

aestivum L.).Theor.Appl. Genet.115:

721

-

733.

Cockram,J.,H.Jones,F.J.Leigh,D.O'Sullivan,SMPowell,D.

A. Laurie and A. J.Greenland

(2007)

Controjof flowering

timeintemperate cereals: _genes,domestication,and sustainable

productivity.J.Exp.Bot.58 :

1231

-

1244.

Hoogendoorn, 1.

(1984)

A comparison of differentvernalizatien

techniques inwheat

(friticum

eastivum L,)

.

J.PlantPhysiol.

116 :11

-

20.

Kato,K. and H. Yarnagata

(1988)

Method forevaluation of chilling requirement and narrow-sense earliness of wheat cultivars. Jap.J.Breed.

38

:172

-

186.

Flowering　traits　and 由 eir　genetic　basis　in　the日ncestral　tetraploid　wheat 　varieties

画

Emmer

’

and

’

Pyramida亘e

’

Mohler

_，

　V

．

，

　R

．

　Lukman

，

　S

．

　Ortiz

−

lslas

，

　M

．

　William

，

　A

，

亅

．

　Worland

，

　J．

van 　Beem　and　

G ．

　Wenrel （

2004

）Genetic　and　

physical

　mapping 　of 　photoperiod　insensitive　gene　Ppd

．

別 in　common 　wheat

．

　Euphytica　

138

：

33− 40．

Takahashi

，

　R

．

　and 　S

．

　Yasuda （

1958

）Genetic　studies 　on　heading

　

time　in　bar且ey （in　Japanese）

，

　ln

」

’

Studies　on　bulk　me 止od　of

　plant　breeding

”

Sakai

，

　K

．

，

　R

，

　Takahashi　and　H

，

　Akemine （eds

．

）

，

　Ybkendo

_，

　Tokyo

．

　

PP．44− 64．

Tanio

，

　M

．

，

　K

．

　Kato

，

　N

．

　Ishikawa

，

　Y　Tamura_，　M

．

　Sato_，｝H

．

　

Takagi

　and　M

．

　Matsuoka （

2005

）Genetic　analysis 　of　photoporiod

　response 　in　wheat 　and　its　relation 　with 　the　earliness 　ofheading 　in　the　southwestern 　part　of　Japan

．

　Breed

．

　Sci

．

55

：

327− 334，

Wilhelm

_，

　E

．

　P

．

，

　A

．

　S

．

　Turner　and 　D

．

　A

．

　Laurie （

2009

）Photo

・

　

period

正nsensitive 　P_ρd

．

Ala　mutations 　in　tetraploid　wheat

　

（Triticum　durum　Desf

．

_）

．

　TheoL　AppL　Genet

．

118：285

−

294

，

Wbrland

，

　A

．

　J

．

　and　C

．

　N

．

　Law （

1986

）

Genetic

−

analysis　of

　chromosome 　

2D

　of　wheat

．

1

．

　the　location　of　genes　affecting

　height

，

　day

−

length　

insensitivity

，

　

hybrid

　

dwartlsm

　and　yellow

−

　rust　resistance

，

　Z

．

Pflanzenzuchtung　

96

：

331 − 345．

Yan

_，

　L

．

，

　D

．

　Fu

，

　C

．

　Li

，

　A

．

　Blechl

，

　G

．

　Tranquilli

，

　M

．

　Bonafede

，

　A

，

　

Sanchez

，

　M

．

　Valarik

，

　S

．

　Yasuda　_and　J

．

　Dubcovsky （

2006

_）The 　wheat 　and 　barley　vernalization 　gene _配

RIV3

　is　an　orthologue　of

　FT

，

　Proじ

．

　Natl

．

Acad

．

　Sci

．

　USA 　

103

：

19581−

19586

．

Yan

_，

　L

．

，

　A

．

　Loukoianov

，

　

G ．

　Tranquilli

，

　M

．

　Helguera

，

　

T．

　Fahima

　

and 亅

．

　Dubcovsky （

2003

）Positional　cloning 　of　the　wheat

　vemalization 　gene　VRIVI

．

　Proc

．

　Nat［

．

　Acad

．

　Sci

，

　USA 　lOO：

　

6263− 6268，

Yan，　 L

．

，

　 A

．

　 Loukoianov

，

　 A

．

　 Blech1

，

　 G

．

　 Tranquilli

，

　 W

．

　

Ramakrishna

，

　

P．

　

SanMiguel

，

　

J．

　L

，

　Bennetzen

，

　V

．

　Echenique　and

　

J

．

Dubcovsky （

20

（  The　wheat 　ifRIV2　_gene　is　a 月owering 　repressor 　down

−

regu 且ated　by　ve  lization

．

Science　

303

：1640

−

　1644

，

Yasuda

，

　S

．

　and　H

．

　Shimoyama （1965）Analysis　of　internal

　 factors　influencing　thc　heading　time　of　wheat　varieties

．

　Ber

．

　Ohara　Inst

．

　Landw

，

　Biol

．

　Okayama　Univ

．

13

：

23 − 38，

Zheng，

　

K ．

，

　P

．

　K

．

　Subudhi

，

」

．

　Domingo

，

　G

．

　Magpantay　and　N

，

　

Huang （

1995

）Rapid

　

DNA 　isoIation　fbr　marker 　assisted

　selection 　in　rice　breeding

．

　Rice　Genet

．

NewsL　

12

：

255 −257，

四

倍 体

コ

ム

ギ 品種

エ

ン

マ

ー

お よ

び

ピ ラ

ミ

ダ

ー

レ の

開 花 特 性 と そ

の

遺

伝 的

制 御

機 構

中崎 鉄 也D

・

森 山 竜二 L）　

・

加 賀田 恒D

・

若原浩

義

1）

・

内藤 実 加1｝

・

桂 圭 佑1｝

・

齊藤 大 樹 D

・

加 藤 鎌 司2〕

・

西田英 隆2）

・

河 原太八3）

・

札埜 高志 1）

・

北 島 宣1〕 1） 京都 大学大 学 院 農 学 研 究科 （〒569　

一

　

0096

高 槻 市八町 畷 町

12 − L

_附属農_場） 2〕 岡山大 学 大 学 院 自然 科 学 研 究 科 （〒

700− 8530

岡 山市 北 区 津島中

3−1−1

） 3〕 京都大学 大 学 院 農 学 研 究 科 （〒

617 − 0001

向日市物集女町中条 1

，

植物生殖 質 研 究 施 設 ） 要 旨：_京都大学の ブ ラン ドビ

ー

ルの原料であ る占代コ ムギ 品種 エ ンマ

ー

および ピラ ミ ダ

ー

レ の開花特 性 につ い て詳細 な解析を行っ た

，

これ らの品種は そ れ ぞ れ Tturgidum 　L

，

　ssp

．

　dicocconお よ びssp

．

　turanicum に分類 さ れ

，

これ までに そ

の栽培に必要な農 業特性に 関 す る 情 報 は ほ とんど得られ てい ない

．

その開 花 期 を 決 定 する要 因 （日 長 反 応性

，

春 化要求 性

．

純 粋早 晩性 ）の評価を行っ た結果

，

これ ら2品種はいず れ も春 播き 型の感光性 品種であ り

，

純 粋 早 晩 性 程 度 は

，

エ ン マ

ー

が ピラ ミダ

ー

レ よ りの小さいの に対し て

，

感 光性程 度 はエ ンマ

ー

の方が大きい ことが明ら か になっ た

．

さ らに

，

2品種 問の F2を 用い た 解 析 か ら

．

ピ ラ ミ ダ

ー

レ はPpd

−

A1”

A

にコムギ 育 種に有 用 な 不 感 光 性 遺伝子 を もつ こ と が判 明 した

．

キ

ー

ワ

ー

ド ：_{開 花 特 性}

，

_{四 倍 体コ ムギ}

，

_{不 感 光 性}

，

Ppd

−

_Ai 　 　 　 　作 物研 究　　56：67

−

71_（2011_） 連絡 責 任 者 ：_{中 崎 鉄}也 （  kazaki＠kais

．

kyote

・

u

．

ac

．

jp

）