全 文 ：　遗 传 学 报　Acta Genetica S inica , M ay 2005, 32 (5):487 ～ 494 ISSN 0379 -4172
收稿日期:2004 - 03 - 11;修回日期:2004 - 09 -08
基金项目:中国科学院知识创新工程项目(编号:KSCX2 -1 - 01 )和中国科学院院长青年创新基金资助课题 [ Supported by the Know ledge Inno-
vation P rogram of the Ch inese Academy of Sciences(No. KSCX2 - 1 - 01) and a President ResearchG ran t for Younger S cien tists of the Ch i-
nese Academ y of S ciences]
作者简介:施　芳(1968 -),女 ,博士 ,副研究员 ,研究方向:细胞遗传学
①　通讯作者。 E-m ail:sh ihofang@ hotm ai.l com
Inducing Rye 1R Chromosome Structural Changes in Common
Wheat cv. Chinese Spring by the Gametocidal Chromosome
2C ofAeg ilops cy lindrica
SHI Fang
1, ① , LIU Kun-Fan1 , ENDO Takashi R2 , WANG Dao-Wen1
(1.The S tateKey Labora tory o fP lant Cell and Ch rom osom e Eng ineering, Institute of Genetics and
Developm en ta lB iology, Ch inese Academ y of Sciences, Beijing　100101, China;
2.Labora tory o fP lant Genetics, Graduate S choo l of Ag ricu lture, KyotoUn iversity, Sakyoku, Kyoto　606-01, Japan)
Abstract:To generate 1R de le tion and trans loca tion lines, w e in troduced a 2C chromosom e, w h ichw as de r ived
from Aeg ilops cy lind rica and was known to have a gametocida l function when added monosom ica lly into com-
mon w hea t cv. C h inese Spring (CS) and its de riva tive, in to a w hea t-rye 1R chromosome d isom ic add ition line
(CS-1R″). W hen the ind ividua ls w ith ch romosome constitu tion 21″+1R″+2C′(2n=45) w ere se lfed, the 1R
chrom osome structu ra l changes were found to be induced w ith h igh frequency(24.1%) among the p rogen ies.
By us ing C-band ing and G ISH ana lys is, we ana lyzed 1R struc tu ra lchanges in 46 F3 ind iv idua ls, wh ich came from
23 F2 p lan ts. The rearranged 1R chromosomes cou ld be cha racter ized in abou t 85% o f the F3 ind ividua ls. Th is in-
c luded te losome 1RL (39.1%), iso-ch romosome 1RL (2. 2%), who le a rm transloca tion invo lv ing 1RL
(32. 6%), te losom e 1RS (4. 3%), iso-ch rom osome 1RS (4. 3%), and 1R de le tion mu tan tw ith b reak po int in the
long a rm (2. 2%). The mutan t 1R lines ob ta ined in th is study w ill po tentia lly be usefu l in mapp ing the ch romo-
som e loca tions o f ag ronom ica lly mi po rtan t genes loca ted in 1R. Th is study a lso demonstra ted tha tm o lecu la r
m arkersm igh t be used to identify wheat chromosom e arm invo lved in trans loca tion w ith 1R.
K ey words:gametocida l ch rom osome;1R;comm on wheat;ch romosom e aberra tion
柱穗山羊草 2C染色体诱发中国春小麦
背景中黑麦 1R染色体结构变异的研究
施 　芳 1, ① , 刘坤凡 1 , 远藤隆 2 , 王道文 1
(1.中国科学院遗传与发育生物学研究所植物细胞与染色体工程国家重点实验室 , 北京　100101;
2.日本京都大学农学部植物遗传学研究室 , 京都　 606-01)
摘　要:当柱穗山羊草(Aegilops cy lindrica Hos.t )2C染色体单体添加到普通小麦品种中国春和以中国春为背景的
派生系时 , 减数分裂时 ,不含 2C染色体的配子会发生染色体结构变异。为了制备一套黑麦 1R染色体缺失系以用
于定位黑麦 1R染色体上的控制重要农艺性状的基因 ,把一条 2C染色体导入到小黑麦 1R二体附加系(21″+ 1R″)
中 , 然后让这些个体(21″+ 1R″+2C′, 2n =45)自交 , 以便产生 1R染色体结构变异体。实验共检测了 345粒 F2
种子 , 83粒种子带有结构变异的黑麦 1R染色体(24. 1 %)。通过 C分带和原位杂交检测 ,对来自于 23株 F2的 46
个 F3植株所带有的异常 1R染色体进行了归类:其中 1RL端体为 39. 1 %, 1RL等臂染色体为 2.2 %, 1RL易位系
为 32.6 %。 1RS端体为 4. 3%, 1RS等臂染色体为 4. 3%, 切点在长臂上的缺失体为 2. 2%。在 6. 5 %的植株中
同时含有 2种类型的 1R染色体结构变异。其余 8. 7 %带有异常 1R染色体的个体因为没有原位杂交结果而无法
判断是属于哪种类型。已获得的 1R结构变异株将有可能进一步发展成为一套可用于定位黑麦 1R染色体上重要
功能基因的遗传材料。另外 , 还探讨了综合应用细胞学和分子标记方法鉴定易位染色体中小麦染色体片段的尝
试 , 并对所获结果进行了讨论。
关键词:杀配子染色体;1R染色体;普通小麦;染色体结构变异
中图分类号:Q343.2　　　文献标识码:A　　　文章编号:0379-4172(2005)05-0487-08
　　G ametocida l (G c) factors in wheat and related
species are strong segrega tion d istorters tha t cause ga-
me tophy tic abo rtion in game tophy tes lacking them
[ 1]
.
They a re found in the w ild genus o f common whea t and
Aeg ilops species, and are generally carried by homeo lo-
gous g roups 2, 3 o r 4 chromosomes[ 2] . Game toc idal fac-
to rs can induce structural rearrangemen ts o f the chro-
mosomes from not on ly w heat but also those o f rye and
barley when they are added into su itable commonwheat
geno types
[ 3 ～ 5]
.
The 1R chromosome of rye is a sou rce of impo r-
tant genes fo r wheat improvemen.t It has been known
that a cluster of resistance genes ( to leaf rust, stem
rust, stripe rust, powde ry m ildew and whea t streak mo-
saic v irus) is loca ted on its sho rt arm[ 6] . In add ition , it
is a lso show n that whea t translocation lines invo lving
1RS cou ld even improve the yie ld o fwhea t
[ 7]
. By eva l-
uating the phosphorus e fficiency of a se t o f wheat-rye
disom ic addition lines (CS +1R″-7R″), Liu et al. [ 8]
deduced that there m ight be epu (enhanced phospha te
utiliza tion) gene(s) located on chromosome 1R, be-
cause the 1R addition line grew better than the remai-
ning six addition lines and thew ild type Ch inese Spring
unde r pho spha te deprived condition. Howeve r, to date,
the precise location o f the epu gene(s)w ith in the 1R
chromosome is still unknown. Formapping the potential
epu gene(s) in the long term , we have attempted to
genera te a se t o f 1R chromosome de letion lines using
gametocida l chromosome 2C orig ina lly de rived from Ae-
gilops cy lindrica. The strategy is to create firstly an ad-
dition line of Chinese Sp ring containing a pair of 1R
chromosomes and a sing le 2C chromosome (2n =
45). W hen the plan ts of this line are se lfed , the proge-
nies that do no t contain chromosome 2C w ill su ffe r
chromosome abe rrations. These structural abe rrations
may occur on either w heat o r rye 1R chromosomes o r
be tw een whea t and 1R ch romosomes. C ytogene tic anal-
ysis w ill reso lve the chromosomes w ith changed struc-
ture s and the types o f struc tura l changes. By succes-
sive ly se lecting and propagating the se lfed progenies in
wh ich thew heat chromosome sets a re intact and the 1R
chromosomes have unde rgone structural change, we
may eventua lly produce a set o f 1R chromosome dele-
tion lines tha t can be used formapping any potentially
impo rtant genes resided on 1R. In this pape r, we report
the progress achieved so fa r, together w ith discussions
on the future perspectives o f our research.
1　M ate rials andM ethods
1. 1　Generation o f 1R chromosome structu ral
changes
　　Common wheat (cv. CS)-rye (cv. Imperial) 1R
chromosome disom ic add ition line carrying a pair of 2C
chromosomes(2n =46, 21″+1R″+2C″)was used as
488 遗传学报　Acta Gene tica S inica　Vo .l 32　No. 5　2005
female to cross w ith CS-1R disom ic add ition line (2n
=44 , 21″+1R″). The F1 w ith chromosome composi-
tion 21″+1R″+2C′(2n =45)was selfed to gene r-
ate F2. Among F2 progenies, p lants w ith structu ra lly
changed 1R ch romosomes w ere identified by C-ban-
ding. These p lants w ith rearranged 1R ch romosomes
w ere further se lfed to obta in F3 seeds. A llF3 seedswe re
ge rm inated. The roo t-tips o f the germ ina ted seed ling s
w ere exc ised for cy to logical ana lysis. The p lants we re
grown at greenhouse under suitab le condition (F ig. 1).
F ig. 1　A scheme for screeningwheat p lants
conta iningmuta ted 1R chromosomes
The tw o starter lines hav ing Chinese Spring (CS)
backg round we re cro ssed to gene ra te F1 , which had 42
wheat chromosomes, two 1R chromosomes and one 2C
chromosome from Ae. cy lindrica. Selfing o f F1 produced
F2 p lants. Among F2 plan ts there would be th ree geno-
types containing 1R chromosomes. The 1R chromo-
somes in the F2 plant conta in ing tw o 2C ch romosomes
w ere no t expec ted to undergo struc tura l muta tions
(ow ing to the possession o f the 2C chromosomes in
both the ma le and fema le game tes whose fusion led to
the production o f the F2 plan t). The 1R ch romosomes
in the F2 p lant con taining a sing le 2C chromosome
w ou ld undergo struc tura lmu tations (ow ing to the ab-
sence o f the 2C chromosome in either the ma le o r fe-
ma le game tesw hose fusion led to the production of the
F2 plant). The 1R chromosomes in the F2 plan t con tai-
ning no 2C chromosomesw ou ld have the h ighest proba-
bility to unde rgo struc tura lmu tations(ow ing to the ab-
sence of the 2C chromosome in bo th the male and fe-
m ale gametes whose fusion led to the production o f the
F2 p lant). The F2 p lants o f the latter two genotypes
w ere allow ed to se lf to g ive rise to F3. The F3 seedlings
containing 1R, but not 2C , chromosomes w ere se lected
for investigating structurally changed chromosomes.
1. 2　Mo lecular cytogenetics analysis
C-banding and G ISH procedures we re carried ou t
fo llow ing a previous pub lication
[ 4]
. Briefly, seeds w ere
germ ina ted on the w e t filte r pape r in petri d ish a t23℃
incuba to r. The roo t tips we re gathe red and placed into
v ials w ith prev iously coo led w a te r fo r 20 ～ 24 h fo l-
low ed by fixa tive (100% ethano l∶acetic acid=3∶1)
trea tment fo r more than 2 day s. S lides fo r C-banding
and G ISH were prepa red a fte r fixative treatmen.t Fo r
C-banding, the slides w ere dehydra ted using 45% ace-
tic acid, follow ed by 5% (w /v) barium hydrate treat-
m ent for 2 m in and 2 ×SSC so lution for more than 5
m in at 48℃. The slides w ere stained in so lution (K ato
chem ica l co. Japan) prev iously prepared for a su itable
time. Good metaphases w ith good C-banding we re pho-
tog raphed by dig ital came ra (Spo t, Diagnostic instru-
m ents. Inc. ) and the slides w ere desta ined by 70%,
95%, 100% e thano l fo r 5 m in each. The slides w ere
dena tured using 70% formam ide in 2×SSC so lution a t
70℃ fo r 2m in and the probe w as applied onto i.t The
slides w ere then placed into w et box for hybrid ization
ove rnigh.t The probe w as prepared using rye genom ic
DNA labe lled e itherw ith digoxigenin-11-dUTP or bio tin-
16-dUTP w ith nick translation (Roche). The hybridization
signals were detected by anti-digox igenin conjugated Rho-
dam ine (red) or av idin conjugated Fluorescein (g reen,
Roche).
1. 3　PCR-based wheat ch romosome identifi-
cation
　　Genom ic DNA o f whea t lines ca rry ing rearranged
489SH I Fang et a l. :Inducing Rye 1R Ch rom osom e S tructura l Changes in Comm on…
1R chromosome w ere ex tracted using CATB method.
A fte rRN ase trea tmen t and purification, the DNA s w ere
used as temp la tes for PCR.
Tw enty-one pairs o f primers fo r amplify ing chro-
mosome specific SSR loci w ere synthesized based on
the sequences pub lished by Roder et al.
[ 9]
. PCR con-
ditions w ere the same as those de term ined prev ious-
ly
[ 9]
. PCR products o f SSRsw ere run in 6% po lyacry l-
am ide ge l w ith those amplified from w ild type CS as
contro ls.
2　Results
2. 1　 F requency o f structurally changed 1R
chromosome
　　To induce structu ra l changes of 1R chromosome,
we made the crosses shown in F ig. 1. Ou t o f 345 F2
seeds exam ined, 83 seeds (24.1%) had at least one
struc tura lly changed 1R chromosome. Among these
seeds, 48 did no t bear 2C chromosome. The plan ts de-
rived from the 48 F2 seeds w ithout 2C w ere se lfed to
obtain F3 seeds for further investigations.
2. 2　 The types o f structurally changed 1R
chromosomes
　　By combining C-banding and G ISH , we identified
the types o f1R structural aberra tions in 46 F3 ind ividu-
als, which came from 23 F2 plan ts (Tab le 1). In 18
(39.1%) F3 indiv idua ls, te losom ic 1RL ch romosomes
w ere found (Fig.2, A). In tw o (4.3%) F3 ind ividu-
als, telosom ic 1RS ch romosomes w ere found (F ig. 2 ,
B). Iso-ch romosome 1RS w as found in tw o F3 p lants
(4.3%) (F ig. 2, C). Iso-ch romosome 1RL was found
in on ly one F3 p lant (2.2%) (Data not show n). In
one (2.2%) F3 p lant, a pair o f 1R chromosomes w ith
the ma jo r parts o f the ir long arms be ing deleted w as
found (F ig. 2, D). In three (6.5%) F3 p lants, the re
w ere tw o types o f mutated 1R chromosomes (Fig. 2,
E). In contrast, in 15 (32.6%) F3 ind ividua ls, there
w ere translocations involving the whole long arm of 1R
and wheat ch romosomes(or segmen ts) (F ig. 3, A and
B). The types o f abe rrant 1R chromosomes in four
(8.7%) F3 p lants w as unc lear because o f fa iled G ISH
experiments despite m any a ttempts. While some of the
F3 indiv idualsw ere a lready homozygous in both the mu-
tated 1R chromosomes and the wheat chromosomes
(Fig. 2, A), o thersw e re still he terozygous for themuta-
ted 1R ch romosomes.
Tab le 1　Types and frequenc ies of 1R structura l
changes in 46 F3 seedlings
*
Type of 1R m u tation
F requency (num ber
of F3 seedlings)
1RL telosom e
1RS telosom e
1RL isoch rom osom e
1RS isoch rom osom e
Delet ion 1RL (brakeage poin t in 1RL)
Who le arm tran slocation invo lving 1RL
Tw o types of 1R aberrat ion in a p lan t
39. 1% (18)
4. 30% (2)
2. 20% (1)
4. 30% (2)
2. 20% (1)
32. 6% (15)
6. 50% (3)
　N ote:*Th e types of structu ral changes in the 1R ch rom osom es w ere
not studied in detail in 4 of the 46 F3 seed lings(although they w ere i-
dentif ied by C-band ing to contain stru ctural ly changed 1R ch rom o-
som es, the G ISH experim en tsw ere fai led).
2. 3 　 Identification o f wheat chromosomes
invo lved in translocations w ith 1R
　　Sequential C-band ing and G ISH enab led us to i-
dentify most of chromosome segments or arm s involved
in transloca tions. How ever, for those chromosome seg-
m ents or a rms w ithout diagno stic C-banding character-
istics, they cou ld no t be identified using cy to log ica l
staining. To overcome this prob lem , we evaluated the
use fu lness of m arke r-assisted ch romosome identifica-
tion. F ig. 3 illustrates an examp le fo r identify ing whea t
chromosome componen t invo lved in a translocation w ith
1R using simp le sequence repeat markers (SSRs, m ic-
ro sa te llite markers). Based on C-banding (Fig. 3, A),
we identified a substitution-translocation line (2n =
42), in w hich a pair o f 1B ch romosomesw ere substitu-
490 遗传学报　Acta Gene tica S inica　Vo .l 32　No. 5　2005
F ig. 2　Structura lly changed 1R chromosomes in F3 seed lings identif ied by C-banding
A:a F3 seed ling con tain ing a pair of1RL telosom es ( ind icated by arrow s);B:a F3 seed ling contain ing a s ingle 1RS telosom e (ar-
row ed);C:a F3 seed ling con tain ing one in tact 1R ch romosom e ( ind icated by an asterisk) and one 1RS isochrom osom e (marked by
arrow head);D:a F3 seed ling con taining a pair of mu tated 1R chromosom es w i th the m ajor parts of their long arm s being deleted
( ind icated by arrow s);E:a F3 seed ling con taining one m utated 1R chrom osom e w ith the m ajor part of its short arm being deleted
( ind icated by the A rabic number “ 1”) and one 1RL telosom e ( ind icated by the Arab ic num ber “ 2”).
ted by a pair o f chromosome 1R and a translocation e-
vent also occurred invo lving one o f the tw o 1R chromo-
somes and an unknown whea t chromosome. G ISH ana l-
y sis indicated tha t the transloca tion chromosome consis-
ted of 1RL and a sho rt chromosome segment p robab ly
o riginated from an unknown whea t chromosome (F ig.
3, B). W e tried to amp lify 21 SSR markers even ly dis-
tributed in the 21 whea t chromosomes from w ild type
Ch inese Spring and the above substitu tion-translocation
line. For one SSR marker located on the v icinity o f the
491SH I Fang et a l. :Inducing Rye 1R Ch rom osom e S tructura l Changes in Comm on…
centrome re o f 3D chromosome, the expected amplifica-
tion products w ere present in the w ild type Chinese
Spring but no t the above substitu tion-translocation line
(F ig. 3, C , arrowed). How ever, by C-banding (Fig. 3 ,
A), we iden tified a pair o f 3D chromosomes present in
the w heat backg round of the substitution-transloca tion
line. Based on th is fac t, it is difficult to expla in why the
bands o f SSR on 3D were absen t if the pair of 3D chro-
mosomes are intact and to deducewhethe r the 3D chro-
mosome m igh t have been involved in the transloca tion
even.t Considering tha t the chromosome structural
changes caused by game tocidal gene on 2C chromo-
some occur random ly ove r the genome o f CS wheat
backg round and tha t if the backcross w ith CS of this
linew e re conducted incomp le tely, it is like ly tha t a fine
in terca lary de le tionm igh t have occurred on the vicin ity
of the cen tromere of 3D ch romosome. In addition, be-
cause 3D chromosome had only a few of diagnostic
C-bands and because the line illustra ted here happened
to be a complica ted one, we w ere no t able to identify
the unknownwheat chromosome segmen t is 3D chromo-
some. If the de letion and translocation happened here is
a sequen tial event invo lved the same wheat chromo-
some, we can probably deduce that the 3D chromosome
in the substitu tion-translocation line m ight have been
invo lved in the translocation even t, o therw ise, we can
not iden tify the unknown wheat chromosome. For these
reasons, whe ther the unknown transloca ted whea t chro-
mosome in this line is 3D chromosome needs further
experiments to be confirmed.
3　D iscussion
In the wo rk　 repo rted　 in th is pape r, we　 at-
temp ted to induce structu ra l changes in 1R chromo-
somes that have been added 　 to 　 the common
F ig. 3　 Ident ifica tion of the wheat chromosomal
component in a 1R trans loca tion chromosome u-
s ing C-band ing, G ISH and amplifica tion ofwheat
chromosome spec ificm icrosa te llite markers
A:C-band ing analysis id ent ified a F3 seed ling in w h ich 1B ch ro-
m osom esw ere sub stitu ted by a pair of translocation ch rom osom e 1R
( ind icated by arrow and arrow head). The trans location even t oc-
curred involving one (m ark ed by arrow head) of th e tw o 1R ch ro-
m osom es and an unknow n wheat chrom osome ( the identi ties of al l
chrom osom esw ere labeled to facilitate the ana lysis);B:G ISH a-
na lysis indicated that the tran slocation ch rom osom e con sisted of
1RL and a short chromosom e segment originated from an unknow n
w heat ch rom osom e;C:The p roduct of a ch rom osom e 3D specific
m icrosatellite m arker w as am p lified from the w ild type Ch inese
Sp ring p lan t( left lane) bu tnot from the F3 seed ling con taining th e
tran slocation ch rom osom e ( righ t lane, the arrow indicates the pos i-
tion where th e am plif ication p roduct ism issing).
wheat va riety Chinese Sp ring using the gameto-
492 遗传学报　Acta Gene tica S inica　Vo .l 32　No. 5　2005
cidal chromosome 2C from Ae. cy lindrica. The proce-
dures used formak ing the gene tic crosses and fo r cy to-
gene tic analy sis o f the chromosomal abe rrations we re
genera lly sim ilar to those employed by prev ious stud-
ies. The resu lts obta ined in this w ork share bo th sim i-
larities and diffe rencesw ith those repo rted previously.
In common w ith previous reports show ing that the
2C chromosome is effective in inducing structu ra l chan-
ges to a lien chromosomes added to hexaplo id whea t, we
found that this chromosome caused a high frequency
and a w ide range o f structu ra l abe rrations in the 1R
chromosome p resent in common whea t varie ty Chinese
Spring. A s shown in the section of results, we found
that 83 seeds (24.1%) had a t least one structu ra lly
changed 1R ch rom some out of 345 F2 seeds. 48 F2
plants w ithout 2C chromosome but carry ing structural
changed 1R chromosomew e re used fo r the nex t experi-
men ts. By random ly identify ing some of these 48 lines
by G ISH , we knew tha tmost o f rearranged 1R chromo-
somes are e ither de le ted or translocated one. Based on
these pre lim inary results, we have expected that su ffi-
cien t lines w ith dele ted 1R chromosome w ou ld be ob-
tained and these de le tion lines would enable us tomap
the resistance genes o r epu gene(s), which located on
1R o r 1RS. Unfortunate ly, the v iab ility of mo re than
ha lf of thesemuta tion lines(25 out o f 48 F2)was ve ry
bad and died during the differen tdeve lopmenta lphase.
This re sultw as unusua l based on the previous study by
autho rs in barley. W e consider that the influence of
both outer facto r (such as the env ironmenta l condition
in g reen house) and inner fac to r (chromosome mu ta-
tion is too severe to g row)m igh t result in this phenom-
enon. The re fore, avoiding the factors tha t affec t the v ia-
b ility o fmutation lines, to screen mo re lines w ith de le-
ted 1R chromosome by 2C chromosome fo rmapping is
our continuous e ffo r.t The abe rration types of 1R chro-
mosome shown in this paper a re from F3 progeny, which
derived from the rema in ing 23 F2. Interesting ly, a large
proportion of F3 plan ts conta ined 1RL te losomes, indi-
ca ting that 1RL te losomesmay bemore stable than oth-
er types of structurally changed 1R chromosomes. A l-
though the overa ll frequency of the rea rranged 1R ch ro-
mosomes ob tained in th is study w as higher than those
repo rted by Endo et a l.
[ 4]
and Friebe et al.
[ 5] , we did
no t find de letions of 1R w ith break po in ts in the short
a rms, which w ere found in low propo rtions by the tw o
prev ious studies
[ 4, 5]
. How ever, we found one de letion
o f 1R chromosome w ith breakpoint in the short arm e-
ven though there exists a te losome 1RL in the same
line. It seem s that the de le tion of 1R w as induced by
2C chromosome, the te losome 1RL was just the result of
m isdiv ision. In stead, we found only one F3 plan t pos-
sessing a de letionmutant of 1RL w ith the de letion po in t
in the long arm. In o rde r to identify more 1R de letions
w ith break po ints in eithe r the long or sho rt arm s, we
w ill have to analyze mo re F3 plan ts in fu ture experi-
m ents.
Ano ther interesting finding in this wo rk is tha tw e
de tected the presence of 1RL or 1RS isochromosomes
in severa l F3 p lants. Togethe rw ith 1RL and 1RS telo-
somes, these lines w ould constitute an idea l set of ge-
ne ticma terials fo r locating genes( for examp le, the epu
genes) in either the long or short arm s of 1R and fo r
testing the dosage effect of se lected genes located in the
long or short a rm s of 1R. How ever, we need to check
the stab ility of the 1RL and 1RS isoch romosomes in
more successive gene ra tions in order to be confiden t
tha t the mate ria ls can indeed be used fo r future genetic
investigations.
M arke r-assisted iden tification o f translocation
chromosomes has been reported by prev ious pa-
pe rs
[ 10, 11]
. In these papers, th is technique w as used to
identify alien ch romosome segmen ts invo lved in the
transloca tion using a lien ch romosom e specificmo lecu lar
markers. W e found that this strategy m ight also be em-
ployed for iden tify ing the w heat chromosome involved
in the translocation. The chromosome composition of
the particular line used to illustra te marker-assisted i-
dentification o f the wheat chromosome invo lved in
transloca tion in th is w ork happened to be fairly com-
493SH I Fang et a l. :Inducing Rye 1R Ch rom osom e S tructura l Changes in Comm on…
plex, so more experimen ts are needed to verify the re-
su lt ob tained so far.
In summary, we have a variety of structural chan-
ges o f the 1R chromosome added into the common
wheat varie ty Chinese Spring using the game toc idal
chromosome 2C from Ae. cy lindrica. Several of these
lines may be furthe r deve loped into suitab le gene tic
stocks that can poten tially be used fo rmapp ing the lo-
ca tions of agronom ica lly impo rtant genes in the 1R
chromosomes. Looking into the future, the availability of
a se t of genetically stable 1R de letion mu tants may a lso
be ve ry usefu l for func tiona l genom ics resea rch in
w heat and re lated species.
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494 遗传学报　Acta Gene tica S inica　Vo .l 32　No. 5　2005