小麦Rht3，Gai3基因的分子标记和被导入外源染色体的部分同源群归属研究
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摘要
一、小麦矮秆基因Rht3和赤霉酸反应不敏感基因Gai3的分子标记
及其与α-淀粉酶的表达研究
小麦的Rht3是来源于中国西藏大拇指矮小麦中对赤霉酸反应不敏感的显性
矮秆基因。Rht3等位基因被认为具有对F_1代籽粒产量呈超显性，使成熟籽粒α-
淀粉酶活性水平降低等效应，因而在杂交小麦育种中受到重视。Rht3与α-淀粉
酶的表达、GA信号转导途径，以及抗穗发芽和改善面粉加工品质密切相关。因
此，研究和克隆Rht3基因具有重要理论意义和应用价值。Rht3的遗传方式已较
清楚，但长期以来有关Rht3与Gai3基因是否为两个独立的基因问题仍有争议。
本研究利用Rht3供体经与其隐性等位载体品种（扬麦3号与苏麦3号）回交22
次的近等基因系与轮回亲本杂种F_1自交产生的BC_(22)F_2较大分离群体，从经典遗
传学到分子遗传学水平，系统地研究了Rht3与Gai3的遗传关系及它们对α-淀粉
酶的表达和穗发芽的影响等问题。并筛选出与Rht3、Gai3紧密连锁的RAPD和
RFLP标记。为Rht3基因的有效利用提供了许多新的信息，也为其图位克隆奠
定了基础。
研究表明Rht3与Gai3应为极紧密连锁的两个基因，在两个近等基因系分离
群体中Rht3与Gai3间的重组值分别为0.045±0.010和0.04040±0.011。Rht3、Gai3
通过抑制α-淀粉酶基因的表达，降低α-淀粉酶活性，从而提高了抗穗发芽能力。
在重组类型中Rht3表现出对α-淀粉酶表达和抗穗发芽的更大影响。等电聚焦同
功酶分析结果显示Rht3、Gai3主要对位于第6部分同源群长臂的α-Amyl基因的
表达起作用，但群体α-Amyl同功酶谱带较为复杂，可能Rht3、Gai3调控的α-Amyl
的表达机制较复杂。RAPD分析共筛选了310个随机引物，其中UBC389、OPV-06
和S1060三个引物在高矮亲本间能稳定地扩增出多态带，但群体验证结果仅
S1060_(1900)和S1060_(2000)扩增片段与Rht3、Gai3连锁，遗传距离分别为16.7cM和
29.5cM。RFLP分析选用了53个第4部分同源群短臂探针，其中Xpsr584、XksuF8
和Xcdo38三个探针在高矮亲本间揭示多态性，接连锁分析结果，Xpsr584与Rht3、
Gai3共分离。
二、利用RFLP分于标记确走导人小壹的羹观草（k kwi）染色
休邱分同回群归回扭究
从小麦亲缘物种向普通小麦转移优良基因，并在分子水平上进行有效地检
测，对于进一步利用小麦亲缘物种中所蕴囤的丰宫基因资源具有十分重要的理论
与实际意义。yLP可以准确地确定导入外源染色体及染色体片段的大小、位置
和所涉及的外源染色体与普通小麦染色体的部分同源关系，为外源优良基因的利
用提供极有用的信息。本研究选用来自小麦7个部分同源群的26个DNA探针
对45个小麦－鹅观草衍生后代株系及鹅观草、中国春和扬麦5号进行yLP分析。
结果表明大部分探针在鹅观草基因组有3条以上的杂交带，在中国春．扬麦5
号与鹅观草之间的多态性水平较高。
16个小麦一鹅观草异附加、代换或可能的易位系中，所涉及鹅观草染色体
分别属子第1、3、5、6、7部分同源群。小麦一鹅观草异染色体系中导入的成对
鹅观草染色体能够较稳定地遗传给后代。K139、K141、K214、KZ18、KZ19、
K224 H体附加系所添加的鹅观草染色体属第 1部分同源群，但 KZI4、KZIS所
添加的鹅观草染色体与 KZIg、K224所添加的鹅观草染色体来自鹅观草不同的染
色体组。K147端体添加系涉及第！部分同源群鹅观草染色体长臂，而K139、
K141和 K147所涉及的鹅观草染色体分别来自鹅观草不同的染色体组。鹅观草 U
染色体与小麦第1部分同源群有同源关系。属第1部分同源群的鹅观草染色体尤
其是它的长臂与赤霉病抗性有关．研究中还观察到鹅观草第！部分同源群与第6
部分同源群染色体之间的可能重排。另外，K203添加的两条鹅观草染色体分别
与第！和6部分同源群同源。K166中导人的鹅观草染色体涉及第5部分同源群。
短臂。K177（Zn—41，20II－I）中，所渗入的鹅观草染色质涉及第5（SL）、6（6S）、
7（SL）部分同源群。本研究结果同时也证明了鹅观草 S、H和 Y三个染色体组
间的部分同源关系。
Molecular markers linked to Rht3 and Gai3 genes that affect the expression of
a-amylase
The wheat dwarf gene Rht3 derived from Torn Thumb, a Tibetan wheat of China,
is a dominant gene with the insensitivity to gibberellic acid. The alleles of Rht3 show
overdominance for grain yield of hybrid F1 and depress of the ~-amylase activity in
ripened grains. So it抯 possible utilization in hybrid wheat breeding is of great concern
It抯 found that Rht3 is also closely related with the x-amylase activity, GA
transduction, resistance to preharvest sprouting and processing quality of wheat. The
irnpoitance of studies on molecular mapping and isolation of Rht3 is obvious The
hereditary mode of Rht3 has been clarified, but whether Rht3 and Gai3 are two
independent genes still remains as a problem to be answered In this dissertation
systematic studies on the genetic relationship between Rht3 and Gai3 and how they
affect the ci-amylase activity were made using large BC22F2 populations derived from
two set of dwarf isogenic lines including Yangmai3 and Suirnai3 as well as their
prototypes The dissertatior~ also present RAPD and RFLP markers linked to Rht3 and
Gai3 which could be of use for the applications and map-based cloning of Rht3
The results of this paper revealed that Rht3 and Gai3 are tightly linked to each other
with the recombination rate of 0 045眥) 010 and 0 140? 011 in the two segregating
populations, respectively Rht3 and Gai3 could depress the gene expression of
i-arnylase, reduce its activity and hence enhance the resistance to pre-harvest
sprouting Higher effects of Rht3 compare to Gai3 on ni-amylase activity and
resistance to pre-harvest sprouting were discovered from data of the recombinants.
The results of isoelectric focusing isoenzyme (IEF) of a-amylase verified that Rht3
and Gai3 mainly inhibit the expression of a-Amy 1 gene located on the long arms of
hornoeologous group 6 of wheat. The results also expressed that a-Amy 1 isoenzyme
electrophesis patterns within the populations are very complicated which probably
show the cornplex expression mechanism of a-Amy 1 regulated by Rht3 and Gai3. In
the RAPD analysis, out of 310 random primers (lObp) screened, only 3 primers e.g.
UBC3 89, OPV-0 6 and 51060 revealed polymorphisms in NIL, and fragments
SIO6OI9a6 and S10602000 amplified by primer S1060 were shown as to be linked to Rht3
and Gai3 with a genetic distance 16.7cM and 29.5cM. respectively. In RFLP analysis,
53 probes specific for short arms of homoeologous group 4 were screened, and
Xpsr584, XksuF8 and Xcdo3S showed polymorphism between the bilLs. The linkage
analysis showed that Xpsr5S4 co-segregated with Rht3 and Gai3 genes.
Homoeologous grouping of R. kanwji chromosomes introduced in wheat via
RFLP analysis
Twenty six DNA probes located on 7 homoeolgous groups of wheat were
screened to reveal the RFLP between 45 wheat-R.kamoji derivatives, R.kamoji,
Chinese Spring and Yairnai5. The results showed that most probes could produce
more than 3 hybridized bands showing distinct polymorphisms between Chinese
Spring, Yangmai 5 and 1? .kamoji. The introduced R.kamoji chromosomes in the 16
wheat-R.kamo,i chromosome lines including addtions, substitutions or possible
translocations were grouped into homoeologous group 1, 3, 5, 6 and 7. Alien
chromosome pairs could be readily transmitted into the descendants. Of the 16 alien
chromosome lines, the added chromosomes in K139, K141, K214, K218, K219 and
K224 disomic ad
及其与α-淀粉酶的表达研究
小麦的Rht3是来源于中国西藏大拇指矮小麦中对赤霉酸反应不敏感的显性
矮秆基因。Rht3等位基因被认为具有对F_1代籽粒产量呈超显性，使成熟籽粒α-
淀粉酶活性水平降低等效应，因而在杂交小麦育种中受到重视。Rht3与α-淀粉
酶的表达、GA信号转导途径，以及抗穗发芽和改善面粉加工品质密切相关。因
此，研究和克隆Rht3基因具有重要理论意义和应用价值。Rht3的遗传方式已较
清楚，但长期以来有关Rht3与Gai3基因是否为两个独立的基因问题仍有争议。
本研究利用Rht3供体经与其隐性等位载体品种（扬麦3号与苏麦3号）回交22
次的近等基因系与轮回亲本杂种F_1自交产生的BC_(22)F_2较大分离群体，从经典遗
传学到分子遗传学水平，系统地研究了Rht3与Gai3的遗传关系及它们对α-淀粉
酶的表达和穗发芽的影响等问题。并筛选出与Rht3、Gai3紧密连锁的RAPD和
RFLP标记。为Rht3基因的有效利用提供了许多新的信息，也为其图位克隆奠
定了基础。
研究表明Rht3与Gai3应为极紧密连锁的两个基因，在两个近等基因系分离
群体中Rht3与Gai3间的重组值分别为0.045±0.010和0.04040±0.011。Rht3、Gai3
通过抑制α-淀粉酶基因的表达，降低α-淀粉酶活性，从而提高了抗穗发芽能力。
在重组类型中Rht3表现出对α-淀粉酶表达和抗穗发芽的更大影响。等电聚焦同
功酶分析结果显示Rht3、Gai3主要对位于第6部分同源群长臂的α-Amyl基因的
表达起作用，但群体α-Amyl同功酶谱带较为复杂，可能Rht3、Gai3调控的α-Amyl
的表达机制较复杂。RAPD分析共筛选了310个随机引物，其中UBC389、OPV-06
和S1060三个引物在高矮亲本间能稳定地扩增出多态带，但群体验证结果仅
S1060_(1900)和S1060_(2000)扩增片段与Rht3、Gai3连锁，遗传距离分别为16.7cM和
29.5cM。RFLP分析选用了53个第4部分同源群短臂探针，其中Xpsr584、XksuF8
和Xcdo38三个探针在高矮亲本间揭示多态性，接连锁分析结果，Xpsr584与Rht3、
Gai3共分离。
二、利用RFLP分于标记确走导人小壹的羹观草（k kwi）染色
休邱分同回群归回扭究
从小麦亲缘物种向普通小麦转移优良基因，并在分子水平上进行有效地检
测，对于进一步利用小麦亲缘物种中所蕴囤的丰宫基因资源具有十分重要的理论
与实际意义。yLP可以准确地确定导入外源染色体及染色体片段的大小、位置
和所涉及的外源染色体与普通小麦染色体的部分同源关系，为外源优良基因的利
用提供极有用的信息。本研究选用来自小麦7个部分同源群的26个DNA探针
对45个小麦－鹅观草衍生后代株系及鹅观草、中国春和扬麦5号进行yLP分析。
结果表明大部分探针在鹅观草基因组有3条以上的杂交带，在中国春．扬麦5
号与鹅观草之间的多态性水平较高。
16个小麦一鹅观草异附加、代换或可能的易位系中，所涉及鹅观草染色体
分别属子第1、3、5、6、7部分同源群。小麦一鹅观草异染色体系中导入的成对
鹅观草染色体能够较稳定地遗传给后代。K139、K141、K214、KZ18、KZ19、
K224 H体附加系所添加的鹅观草染色体属第 1部分同源群，但 KZI4、KZIS所
添加的鹅观草染色体与 KZIg、K224所添加的鹅观草染色体来自鹅观草不同的染
色体组。K147端体添加系涉及第！部分同源群鹅观草染色体长臂，而K139、
K141和 K147所涉及的鹅观草染色体分别来自鹅观草不同的染色体组。鹅观草 U
染色体与小麦第1部分同源群有同源关系。属第1部分同源群的鹅观草染色体尤
其是它的长臂与赤霉病抗性有关．研究中还观察到鹅观草第！部分同源群与第6
部分同源群染色体之间的可能重排。另外，K203添加的两条鹅观草染色体分别
与第！和6部分同源群同源。K166中导人的鹅观草染色体涉及第5部分同源群。
短臂。K177（Zn—41，20II－I）中，所渗入的鹅观草染色质涉及第5（SL）、6（6S）、
7（SL）部分同源群。本研究结果同时也证明了鹅观草 S、H和 Y三个染色体组
间的部分同源关系。
Molecular markers linked to Rht3 and Gai3 genes that affect the expression of
a-amylase
The wheat dwarf gene Rht3 derived from Torn Thumb, a Tibetan wheat of China,
is a dominant gene with the insensitivity to gibberellic acid. The alleles of Rht3 show
overdominance for grain yield of hybrid F1 and depress of the ~-amylase activity in
ripened grains. So it抯 possible utilization in hybrid wheat breeding is of great concern
It抯 found that Rht3 is also closely related with the x-amylase activity, GA
transduction, resistance to preharvest sprouting and processing quality of wheat. The
irnpoitance of studies on molecular mapping and isolation of Rht3 is obvious The
hereditary mode of Rht3 has been clarified, but whether Rht3 and Gai3 are two
independent genes still remains as a problem to be answered In this dissertation
systematic studies on the genetic relationship between Rht3 and Gai3 and how they
affect the ci-amylase activity were made using large BC22F2 populations derived from
two set of dwarf isogenic lines including Yangmai3 and Suirnai3 as well as their
prototypes The dissertatior~ also present RAPD and RFLP markers linked to Rht3 and
Gai3 which could be of use for the applications and map-based cloning of Rht3
The results of this paper revealed that Rht3 and Gai3 are tightly linked to each other
with the recombination rate of 0 045眥) 010 and 0 140? 011 in the two segregating
populations, respectively Rht3 and Gai3 could depress the gene expression of
i-arnylase, reduce its activity and hence enhance the resistance to pre-harvest
sprouting Higher effects of Rht3 compare to Gai3 on ni-amylase activity and
resistance to pre-harvest sprouting were discovered from data of the recombinants.
The results of isoelectric focusing isoenzyme (IEF) of a-amylase verified that Rht3
and Gai3 mainly inhibit the expression of a-Amy 1 gene located on the long arms of
hornoeologous group 6 of wheat. The results also expressed that a-Amy 1 isoenzyme
electrophesis patterns within the populations are very complicated which probably
show the cornplex expression mechanism of a-Amy 1 regulated by Rht3 and Gai3. In
the RAPD analysis, out of 310 random primers (lObp) screened, only 3 primers e.g.
UBC3 89, OPV-0 6 and 51060 revealed polymorphisms in NIL, and fragments
SIO6OI9a6 and S10602000 amplified by primer S1060 were shown as to be linked to Rht3
and Gai3 with a genetic distance 16.7cM and 29.5cM. respectively. In RFLP analysis,
53 probes specific for short arms of homoeologous group 4 were screened, and
Xpsr584, XksuF8 and Xcdo3S showed polymorphism between the bilLs. The linkage
analysis showed that Xpsr5S4 co-segregated with Rht3 and Gai3 genes.
Homoeologous grouping of R. kanwji chromosomes introduced in wheat via
RFLP analysis
Twenty six DNA probes located on 7 homoeolgous groups of wheat were
screened to reveal the RFLP between 45 wheat-R.kamoji derivatives, R.kamoji,
Chinese Spring and Yairnai5. The results showed that most probes could produce
more than 3 hybridized bands showing distinct polymorphisms between Chinese
Spring, Yangmai 5 and 1? .kamoji. The introduced R.kamoji chromosomes in the 16
wheat-R.kamo,i chromosome lines including addtions, substitutions or possible
translocations were grouped into homoeologous group 1, 3, 5, 6 and 7. Alien
chromosome pairs could be readily transmitted into the descendants. Of the 16 alien
chromosome lines, the added chromosomes in K139, K141, K214, K218, K219 and
K224 disomic ad
引文
1．蔡士宾 曹阳 1997 小麦品种抗穗发芽性及其主要农艺性状的配合力分析 江苏农业科学 (1)：22-25
2．陈爱大 朱传统 1997 小麦穗发芽敏感性测定方法初探 江苏农业科学 (3)：14-16
3．陈佩度 1995 将大赖草种质转移给普通小麦的研究 Ⅲ．抗赤霉病异附加系选育。遗传学报 22(3)：206-210
4 陈兆夏 蒋国梁 1997 小麦白粒品种抗穗发芽性遗传的初步研究 南京农业大学学报 20(3)：1-6
5 耿以礼 1963 国产鹅观草属 Roegneria CKoch 之订证 南京大学学报 (1)：1-92
6．郭保宏 1996 我国 46 个小麦品种的矮秆基因分析 国外农学-麦类作物 NO．5：4-6
7．贾继增 1997 小麦分子标记研究进展 生物技术通报 2：1-5
8 贾继增，丁寿康，李月华等 1992 中国小麦的主要矮杆基因及矮源的研究 中国农业科学 25(1)：1-5
9 贾继增，丁寿康，李月华等 1994 小麦新品系宛原 50-2 矮基因的染色体定位 作物学报 20(3)：297-301
10．金善保等 1996 中国小麦学 中国农业出版社
11．李继耕 1980 植物同工酶及其在作物遗传研究中的应用 作物学报 6(4)：245-252
12．李竟雄，宋同明 1993 植物细胞遗传 北京：科学出版社 P102-111
13．李振声 1985 小麦远缘杂交，北京科学出版社
14．李杏普等 1998 不同矮秆基因对冬小麦农艺性状的影响 作物学报 24(4)：475-479
15 刘秉华，杨丽 1994 矮败小麦及其在矮化育种中的应用 中国农业科学 27(5)：17-21
16．刘秉华等 1999 小麦核不育的遗传研究和雄性不育遗传模式的建立 中国农业科学 32(2)：18-23
17 刘大钧 1994 向小麦转移外源抗病性的回顾与展望 南京农业大学学报 17(3) 1-7
18 刘大钧，1994 外源基因在小麦育种中的利用 作物杂志6：1-7
19．刘大钧等 1989 从鹅观草和巨大冰麦草向普通小麦转移赤霉病抗性的研究。江苏农业科学 增刊(1)：97-100
20．刘大钧等 1996 导入小麦的外源染色体片段的准确鉴定及外源抗性基因的稳定性分析 遗传学报 23(1)：18-23
21．刘文轩，陈佩度，刘大钧 1998 将大赖草种质转移给普通小麦的研究 Ⅶ．一个普通小麦-大赖草等壁染色体异附加系的选育与鉴定 遗传学报25(4)：351-355
22．刘文轩，陈佩度，刘大钧 1999 利用减数分裂成株电离辐射选育小麦-大赖草易位系的研究 植物学报 41(5)：463-467
23．陆瑞菊，陈佩度，刘大钧 1995 将大赖草种质转移给普通小麦的研究 Ⅳ 通过花药选育双重二体异附加系和代换-附加系 南京农业大学学报 18(4)：1-6
24．任晓琴，孙文献，陈佩度，刘大钧 1996 普通小麦-大赖草 6N 二体异附加系的选育与鉴定 南京农业大学学报 19(3)：1-5
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