全 文 ：收稿日期:2004-07-30
基金项目:国家自然科学基金项目(30160058)、内蒙古自然科学基金项目(2002305)、教育部留学回国人员基金项目
(2001345)资助
作者简介:于卓(1958-),男 ,内蒙古托县人 ,博士 ,教授 ,博士生导师 ,从事牧草饲用作物种质资源与遗传育种研究.
Received date:2004-07-30
Foundation item:Supported by National Natural Science Foundation of China (No.30160058), Nat ional Natural S cience
Foundation of Inner Mongolia (No.2002305)and Educat ion Ministry Foundat ion forthe Scholars Returned f rom Abroad(No.
2001345)
Biography:YU Zhuo(1958-), male , Ph.D , Professor , Supervi sor;E-mail:yuzhuo58@sina.com.
加拿大披碱草×野大麦三倍体杂种 F1
加倍植株的染色体及育性鉴定
于　卓 , 马艳红 , 李造哲
(内蒙古农业大学农学院 ,内蒙古　呼和浩特　010019)
摘要:对加拿大披碱草与野大麦 2 个四倍体亲本杂交产生的三倍体(2n=3x=21)属间杂种
F 1的自然加倍植株 、自然加倍植株 F 1代及秋水仙素诱导加倍植株的染色体数目及配对构型 、花粉
育性 、结实性进行了鉴定。结果显示:自然加倍植株 、自然加倍植株 F1 代和秋水仙素诱导加倍植
株的 RTC 染色体数目均为 42 条 , 即为六倍体(2n=6x=42);其 PMCM Ⅰ平均染色体配对构型分
别为 0.04Ⅰ +20.96Ⅱ 、0.03Ⅰ +20.98Ⅱ和 0.03Ⅰ +20.97Ⅱ , 配对行为规则;它们的花粉可育率
分别为 84.97%、94.08%和 90.21%,自然结实率分别为 77.51%、83.87%和 80.13%。证明该属
间杂种 F1 染色体已加倍 ,且染色体加倍植株的育性已得到恢复。
　　关键词:加拿大披碱草;野大麦;属间杂种 F1;染色体加倍植株;育性
　　中图分类号:Q943　　文献标识码:A　　文章编号:1000-6311(2004)05-0001-08
Identification of Chromosome and Fertility of Chromosome Doubling Plant of Triploid
Hybrid F1 between Elymus canadensis and Hordeum brevisubulatum.YU Zhuo , MA
Yan_hong LI Zao_zhe(Agronomy College of Inner Mongolia Agricul tural Universi ty ,
Hohhot 010019 , China):Grassland of China , No.5 ,2004 ,pp.1 ～ 8.
Abstract:The chromosome number and pairing configuration ,pollen fertili ty , seed set of
the natural chromosome doubling plant , chromosome_doubling F1 and colchicines induced
chromosome doubling plant of t riploid (2n=3x =21)intergeneric hybrid F1 between
E lymus canadensis and Hordeum brevisubulatum were identified.The results show ed
that the chromosome numbers in root tip cell of the natural chromosome doubling plant ,
natural chromosome-doubling F1 and cochicines induced chromosome doubling plant of
triploid hybrid F1 are 42 , which are hexaploids (2n=4x=42);their average chromo-
—1—
第 26卷　第 5期　　　　　　　　　　　　中　国　草　地　　　　　　　　　　　　2004 年 9 月
Vol.26 　No.5　 　　　　　　　　　　　Grassland of China　　　　　　　　 　　　　Sept.2004
some pairing configuration at PMCM I are 0.04 Ⅰ +20.96 Ⅱ , 0.03 Ⅰ +20.98 Ⅱ and
0.03 Ⅰ +20.97 Ⅱ respectively , the chromosome pairing behavior is regular;their
pollen fert ility are 84.97%, 94.08% and 90.21% respectively , the seed set are
77.51%,83.87%and 80.13% respect ively .It w as proved that the chromosome of in-
tergenus hybrid F1 had been doubled , and the fertility of chromosome doubling plant had
been restored.
Key words:Elymus canadensis;Hordeum brev isubulatum ;Intergeneric hybrid F 1;
Chromosome doubling plant;Fertility
1　Introduction
Hordeum briv isubulatum Link (2n=4x =28 , genomic st ructure H1H1H2H 2)is native to
Inner Mongolia grassland , which is a w ild perennial g rass w ith the characteristics of short rhi-
zome , more tillers , early maturing , st rong salt tolerance and drought resistance ,but its defect is
that the seeds is easy to fall of f the plant af ter maturing , therefore i t is not easy to harvest seeds
and to cultiv ate artificially;Elymus canadensis L.(2n=4x =28 , genomic st ructure SSHcHc)was
int roduced from North America , which is a cultivated perennial grass wi th the characterist ics of
more leaves , high yield , st rong winter resistance and high nutrit ive value , but it s defect is that it
is poo r in drought resistance and salt tolerance[ 1] .In order to integrate the advantages of the tw o
g rasses and to create (or breed)new g rass germplasm (or new cultivar)of good quality , high
y ield , strong resistance and wide adaptability , the artificial wide cross w as conducted and inter-
generic hybrid F 1 was abtained successfully
[ 2] .The research showed that the form of hybrid F1 is
in the middle of the parents and hybrid F1 has sho rt rhizome and st rong ability of reg row th from
paternal Hordeum brev isubulatum , the g row th rate tends to maternal , heterosis is strong , but
the anther of hybrid F 1 is small and thin , and does no t fracture , the pollen fertility rate is only
1.19%, does not seed under open pollination[ 3] .It w as found by cy tological observation that vari-
ation of chromosome number of hybrid F1 occurred , seven chromosomes were lost , it is a t riploid
(2n=3x =21), this phenomenon is seldom seen in studies on wide hybridization of perennial
g rasses both in China and overseas
[ 4] .In order to find out the chromosome constituents in hybrid
F 1 , the genomic DNA of hybrid F1 was hybridized in situ w ith probing DNA of Hordeum
brevisubulatum labled with biotin using genomic DNA of Elymus canadensis as blocking DNA ,
the results show ed that there were 14 chromosomes belonging to genome H1H2 f rom paternal
Hordeum brevisubulatum and 7 chromosomes belonging to genome S from Elymus canadensis in
21 chromosomes of t riploid hybrid F1 , 7 chromosomes belonging to genome HC of maternal
E lymus canadensis were lost
[ 5] .Furthermore , in order to restore fertili ty to hybrid F1 , doubling
chromosome of the t riploid hybrid F 1 was induced w ith 0.2% colchicines solut ion in 2002 , mean-
while some tall plants of great value w ere found in hybrid garden for propagat ing F1 by agamogen-
esis , they were confirmed to be natural chromosome doubling plants through mo rphological and
isoenzymic identificat ion , some seeds harvested in the same year w ere sow ed in Spring and natural
—2—
中国草地　2004 年　第 26卷　第 5期
chromosome doubling plants w ere got[ 6] .The chromosome , pollen fertility and seed set of natural
chromosome doubling plant , natural chromosome doubling F1 , colchicines induced chromosome
doubling plant were identif ied compared with hybrid F 1 and parents to find out the cy togenet ic
characters and status of fertility resto ration and to provide scientific basis for selection and utiliza-
tion of excellent lines of cross descendents.
2　Materials and Methods
2.1　Tested materials
The tested materials w ere natural chromosome doubling plant , natural chromosome doubling
F 1 and colchicines induced chromosome doubling plant of Elymus canadensis × Hordeum
brevisubulatumF1 , Elymus canadensis , Hordeum brev isubulatum .and their hybrid F1 were
used as check.All the materials w ere sow ed in the Experimental Crop Farm of Inner Mongolia A-
g ricultural University , this farm is located in eastern suburb of Hohhot , annual rainfall is about
400mm , f rostless period is 145days , it is sandy loam chestnut soil , pH is 7.8 ～ 8.2 , soil fertility
is middle with i rrigation conditions.
2.2　Methods
2.2.1　The method for colchicines induced chromosome doubling
At the early stage of tillering in Spring , a clump of seedlings of Elymus canadensis ×
Hordeum brev isubulatumF1 was excavated from hybrid garden in the field , the soil on the roo ts
w as w ashed aw ay , separated into severall small clumps wi th 3 ～ 4 tiller buds , yellow leaves and
old roots were cut , the small clump w as inversely put into 10ml test tube in diameter of 1.5cm ,
0.2% cochicines solution w as added , induced at 20℃ for 6 ～ 8h , took it out , the plants w ere
w ashed w ith w ater , then t ransplanted to f lowerpot filled w ith sandy soil , then put it into g row th
chamber to cultivate.The day and night temperature in grow th chamber was 25±2℃/20±2℃,
relative humidity w as 65±7%, light intensity w as 11000 ～ 13000 lx.The seedlings w ere trans-
planted to field , when the seedlings survived and had five leaves.
2.2.2　Observation of chromosome in root tip cell
The roo t tips of parents , natural chromosome doubling plant , natural chromosome doubling
F 1 were got f rom germinated seeds , the t ips of hybrid F1 and colchicines induced chromosome
doubling plant were go t f rom new ly generated adventitious roots.The roo t tips were put into fin-
ger shaped tube , added some w ater , lidded and put into mix ture of ice and w ater for 20 ～ 24h ,
transited into Cano s solution to f ix fo r 18 ～ 24h.Then put into 1N HCl at 60℃ for 10 ～ 15min ,
washed w ith distilled w ater , stained with improved alkaline fuchsine and slice was made.Exam-
ined with microscope , and the slice on which the chromosomes w ere w ell scattered was selected to
take picture wi th microphotog raph meter Olymupus-BH2.
2.2.3　Observation of chromosome in pollen mother cell meiosis at metaphase I
The young ears of all the tested materials w ere taken in the field betw een 9:00 ～ 11:00 in
the morning in sunny day , put into bot tle filled w ith Cano s solution quickly , brought to room
—3—
于　卓　马艳红　李造哲　加拿大披碱草×野大麦三倍体杂种 F1加倍植株的染色体及育性鉴定
and put into f reezer to fix for more than 24h.Anther were put on the slid , anther wall w as taken
of f , stained wi th improved alkaline fuchsine , covered w ith lid , knocked light ly , heated slightly
over alcohol lamp and pressed , the chromosome number in cell was counted up , and the slice on
which the chromosomes in PMCM Ⅰ were w ell scat tered w as selected to take picture with mi-
crophotog raph meter.
2.2.4　Pollen fertil ity and seed set
2.2.4.1　Pollen fertility
30 ears of each tested materials were taken randomly in flowering stage , brought to room and
examined w ith microscope.The anther was taken out and put on slid , added a lit tle 1% acet ic
acid carmine , anther wall w as taken off , covered with lid , the fertile and infertile pollens w ere
counted up , Over 100 visual fields of each tested materials w ere observed.The criterion for judg-
ing fertile o r infertile pollen w ere that the pollens which are large in size , full and stained color is
dark are fertile , the pollens which are small and thin , stained color is light o r not stained are in-
fertile.Pollen fertili ty rate =(To tal fertile pollens/Total pollens observed)×100%.
2.2.4.2　Seed set
50 ～ 80 ears of each population of natural chromosome doubling plant , natural chromosome
doubling F 1 and colchicines induced chromosome doubling plant were taken randomly in seed ma-
turing stage , seeds and florets were counted up , seed set =(To tal seeds/Total f lorets observed)
×100%.
3　Results and analysis
3.1　Chromosome characters in cell
3.1.1　Chromosomes in root tip cell(RTC)
Table 1 and Fig.1_a ～ f and i ～ k showed that , the RTC chromosome number of Elymus
canadensis(♀)and Hordeum brivisubulatum (♂)used as check are all 28 , i.e the parents are
tetraploid(2n=4x=28);the RTC chromosome number of intergeneric hybrid F 1 is 21 , which is
triploid(2n=3x =21);the RTC chromosome number of 87.62% cells in the observed natural
chromosome doubling plant of t riploid hybrid F1 are 42 (generally considering , the chromosome
number w hich mo re than 85% of observed cells have is the chromosome number of the
species[ 7～ 9]), while the percentages of tetraploid cell and triploid cell are 12.21% and 0.17%
rrespectively;the RTC chromosome number of 98.26% cells in the natural chromosome_doubling
plant F 1 cells are 42 , while the percentage of t riploid cell is only 1.74%;the RTC chromosome
number of 94.69% cells in the colchicines induced chromosome doubling plant cells are 42 , while
the percentages of tet raploid cell and pentaploid cell are 1.18% and 4.13% respectively.
Above ment ioned results show ed that the chromosome of the natural chromosome doubling
plant , natural chromosome_doubling F1 and colchicines induced chromosome doubling plant w ere
hexaploid(2n=6x =42), which proved that the chromosome of triploid intergeneric hybrid F 1
between Elymus canadensis and Hordeum brevisubulatum had been doubled in cy togenetics.
—4—
中国草地　2004 年　第 26卷　第 5期
Table 1　RTC chromosome number of tested plants
Grasses name
Observed
cell No.
Tetraploid
cell No.
2n=4x=28
Pentaploid
cell No.
2n=5x=35
Triploid
cell No.
2n=3x=21
Hexaploid
cell No.
2n=6x=42
Proportion of
di ff erent ploidy
cell(%)
Natural doubling plan t 606 74 0 1 531 6x　87.62
Natural doubling F1 516 0 0 9 507 6x　98.26
Induced doubling plant 678 8 28 0 642 6x　94.69
E.canadensis(♀) 122 122 0 0 0 4x　100
H .brevisubulatum (♂) 146 146 0 0 0 4x　100
In tergeneric hybrid F1 210 0 0 210 0 3x　100
Fig.1 Chromosomes of tested plants
a.RTC chromosome number of E.canadensis(2n=4x=28);b.RTC chromosome number of H.brevisubulatum(2n=4x=28);
c.RTC chromosome number of intergeneric hybrid F1(2n=3x=21);d.RTC chromosome number of natural doubling plant(2n=6x=42);
e , f.RTC chromosome number of natural doubling F1(2n=6x=42);g , h.Chromosome pairing configuration of natural doubling plant and
natural doubling F1 at PMCMⅠ(2n=6x=42=21Ⅱ);i , j , k.RTC chromosome number of colchicines induced doubling plant(2n=6x=42);
l.Chromosome pairing conf igurat ion of induced doubling plant at PMCMⅠ(2n=6x=42=21Ⅱ).
—5—
于　卓　马艳红　李造哲　加拿大披碱草×野大麦三倍体杂种 F1加倍植株的染色体及育性鉴定
3.1.2　Chromosome pairing behavior of pollen mother cell at metaphaseⅠ(PMCMⅠ)
The chromosome pairing configuration at PMCM Ⅰ of the wide hybrid is relative to the plant
fertilei ty , as the homologous bivalent(Ⅱ)increased , pollen fertilei ty and seed set increased obvi-
ously in general[ 8 、9] 。The results in this study showed that:comparing w ith tet raploid parents ,
the f requency of medial bivalent in each cell of the natural chromosome doubling plant , chromo-
some_doubling F 1 and colchicines induced chromosome doubling plant of t riploid hybrid F 1 between
E lymus canadensis and Hordeum brevisubulatum were very high , which were 20.96%,20.98%
and 20.97% respect ively , and ring bivalent w as dominant(92.22%, 93.28% and 94.56% of
total bivalents respectively), the f requency of univ alent is very low , which w as 0.03 ～ 0.04
(Table 2 , Fig.1-g , h , l), and multivalent and lag ging chromosome were not observed.It was
proved that the chromosome pairing behavio r at PMCM Ⅰ of the natural chromosome doubling
plant , natural chromosome_doubling F1 and colchicines induced chromosome doubling plant is reg-
ular , and this is the important cytogenetic basis of thei r fertility restoration.
Table 2　Chromosome pairing configuration at PMCMⅠ of tested plants
Grasses name
Observed
cell No.
Chromosome
No.
Ⅰ
Univalent
Ⅱ Bivalent
Ring Rod Total
Natural doubling plant
　
211
　
42
　
0.04
(0～ 4)
19.33
(11～ 21)
1.63
(0～ 10)
20.96
　
Natural doubling F1
　
311
　
42
　
0.03
(0～ 4)
19.57
(13～ 21)
1.41
(0～ 8)
20.98
　
Induced doubling plant
　
483
　
42
　
0.03
(0～ 4)
19.83
(14～ 21)
1.14
(0～ 7)
20.97
　
E.Canadensis(♀)
　
122
　
28
　
0.04
(0～ 4)
12.72
(10～ 14)
1.26
(0～ 3)
13.98
　
H.brevisubu latum(♂)
　
146
　
28
　
0.05
(0～ 4)
12.21
(6～ 14)
1.60
(0～ 4)
13.81
　
Intergeneric hybrid F1
　
210
　
21
　
7.45
(7～ 9)
2.73
(2～ 4)
3.85
(3～ 5)
6.58
　
　　Note:In order to comparing each other , quoted the result s of the chromosome pairing configurat ion of parents and their F1 at
PMCM Ⅰ in the table[ 4] .
3.2　Pollen fertility and seed set
It s known that the rate of pollen fertili ty and seed set of triploid intergeneric hybrid F 1 be-
tween Elymus canadensis and Hordeum brevisubulatum are 1.19% and 0 respectively , and is
highly sterile[ 3] ;the rate of pollen fert ility of the natural chromosome doubling plant , natural
chromosome -doubling F1 and colchicines induced chromosome doubling plant increased to
84.97%, 94.08% and 90.21% respectively , and the rat of seed set increased to 77.51%,
—6—
中国草地　2004 年　第 26卷　第 5期
83.87% and 80.13%respectively.Hence it w as proved that the fertility of chromosome doubling
plant had been resto red , this is relative to the pairing rate of the cognate chromosome(Table 3).
Table 3　Pollen fertility and seed set of tested plants
Grasses name
Pollen fert ili ty
Ferti le
No.
Infert ile
No.
Total
　
Fertilit y
rate(%)
Seed set
Seed No. Total
f lorets
Seed set
(%)
Natural doubling plant 4301 761 5062 84.97 3257 4202 77.51
Natural doubling F1 9232 581 9813 94.08 8064 9615 83.87
Induced doubling plant 8062 875 8937 90.21 6781 8463 80.13
4　Discussion and conclusions
4.1　The results in this study show ed that the natural chromosome doubling plant , natural chro-
mosome-doubling F1 and colchicines induced chromosome doubling plant w ere hexaploid(2n=
6x=42), which proved that the chromosome of t riploid intergeneric hybrid F1 betw een Elymus
canadensis and Hordeum brevisubulatum had been doubled in cy togenetics.
4.2　The main cause of sterili ty of w ide hybrid is that there are large differences in chromosomes
between the parents , and the homologous chromosome pairing f requency of pollen mother cell at
metaphase Ⅰis lower[ 8～ 10] .Through analysis on the chromosome at PMCM Ⅰof the F1 chromo-
some doubling plant , the average chromosome pairing conf iguration at PMCM Ⅰ of the natural
chromosome doubling plant , natural chromosome_doubling F1 and colchicines induced chromosome
doubling plant are 0.04 Ⅰ +20.96 Ⅱ , 0.03 Ⅰ +20.98 Ⅱ and 0.03 Ⅰ +20.97 Ⅱ respectively ,
and thei r bivalent frequency are all high , the f requency of univalent is very low , and multivalent
and lagging chromosomes w ere not observed.That is the important cy togenet ic basis for their fer-
tility restorat ion and evaluation.
4.3　The rate of pollen fertility of the natural chromosome doubling plant , chromosome_doubling
F 1 and colchicines induced chromosome doubling plant are 84.97%, 94.08% and 90.21% re-
spectively , and the rate of seed set are 77.51%, 83.87% and 80.13% respectively .It was
proved that the fertility of chromosome doubling plant had been restored , and this provide the
condition and the basis fo r breeding and utilization of excellent st rains o r varieties.
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—7—
于　卓　马艳红　李造哲　加拿大披碱草×野大麦三倍体杂种 F1加倍植株的染色体及育性鉴定
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界草学精华 ,设有草地资源与利用 、牧草研究 、草地畜牧业 、草地农业 、草地保护 、草坪园艺 、草业论坛 、业界信
息 、草业琐谈等栏目。立足全国 , 面向世界 ,努力为从事草地科研 、生产及管理的专业人员或大专院校的师生
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中国草地　2004 年　第 26卷　第 5期