白菜类蔬菜遗传多样性的AFLP分子标记和大白菜发育的差异显示研究
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摘要
本论文工作是我室大白菜分子育种研究的重要组成部分，主要包括了两个部分的研究内容：第一部
分是利用AFLP分子标记技术研究白菜类蔬菜的遗传多样性，第二部分是利用mRNA差异显示技术研究大
白菜结球过程中的基因表达。
白菜类蔬菜包括大白菜和小白菜，原产于我国，是我国在世界各国蔬菜品种和蔬菜栽培中独具风格
的种类。白菜类蔬菜是我国最早的栽培植物之一，它们分布广泛，品种资源非常丰富。对其进行遗传多
样性分析，明确白菜类蔬菜之间的分类关系，有助于人们进一步开发利用其种质资源和更加有效的选择
育种材料。
大白菜的叶球是典型的变态器官。根据叶片发育的形态特征，大白菜的营养生长可分为幼苗期、莲
座期、包心期和结球期四个时期。从莲座期至包心期，是大白菜叶片形态特征和生理功能发生转变的关
键时期，它标志着叶球发生的开始。对大白菜结球过程的研究不仅可以揭示其独特的发育机制和代谢途
径，而且能够为白菜和其它芸薹属蔬菜的分子育种打下基础。
本实验采用AFLP的荧光标记检测技术和银染检测技术，对135份大白菜、小白菜、芜菁材料进行
了遗传多样性和分类研究。在此基础上 进一步以大白菜“北京桔红心”为材料，对其莲座期和包心期
的叶片进行了DD-PCR分析，为揭示由莲座期至包心期发育过程中大白菜叶片基因表达的变化作了一些初
步的尝试。现将主要结果总结如下：
第一部分，利用AFLP分子标记技术研究白菜类蔬菜的遗传多样性
1采用银染检测方法，比较了E+3/M+2、E+3/M+3引物组合对白菜类蔬菜DNA的扩增条带数，发现前者
的扩增条带过于密集，而后者扩增条带数目适中，电泳图谱清晰。
2采用银染检测方法，比较了64对E+3/M+3引物组合对白菜类蔬菜DNA的扩增总带数和多态性检出率，
发现不同引物的扩增总带数在16-58之间，多态检出率在40％-88％之间。从中选择了四对扩增总带
数多，多态性检出率高的引物组合用作分析。
3采用AFLP的荧光标记检测技术对135份材料进行了遗传多样性分析。选用SM系数作为相似性系数
的计算方法，选用UPGMA作为聚类方法。
4聚类结果表明：芜菁和所有的白菜类蔬菜各聚为两类，在白菜类蔬菜类群内部，薹菜和大白菜单独
聚为一类。这表明，芜菁与大白菜、小白菜的相似性程度较低，大白菜与小白菜的相似性程度较高，
大白菜在所有的白菜类蔬菜当中是比较特殊的类群。同意曹家树等提出的将芜菁和白菜（包括大白
菜和小白菜）归为Brassica campestris的两个不同的亚种，大白菜则为白菜亚种中的一个特殊的
变种的观点。
5在小白菜的不同类型中，薹菜与大白菜聚为一类，相似性程度较高，表明薹菜与大白菜的起源可能
存在密切的关系。
6小白菜的AFLP M结果与其综合园艺学分类不符，说明小白菜各类型在DNA M上的差异与其形
态学差异并不存在完全对应的关系。小白菜各类型间的椰以仕程度较大白菜各类型间的＋引以性程度
低，分化程度高，说明小白菜的起源较大白菜要早。
7大白菜的AFLP聚类结果表明，目前生产中所用的大白菜材料之间榔以性矗缠舀交高，而两个大白菜农
家品种与它可l］之间的相似性稷图脚氏。这反映了大ewhIS．＊ru＊品种的遗传多样性较为狭窄，有必要从
大白菜的原始类型中挖掘新的种质资源。
第H部分，利用InRNAgn显示技术研究大白菜结球过程中的基因表达
IAN了4种G、Cng高的锚定引物和3种随机弓1物，共作了12个l）Iyl，nl，反应。在大白菜莲座期和
包吧溯叶片的 mRNA差异显示电泳图谱中，选择了 23个差异 cM ）17ling＋Y了 Reverse NOrthe。检验，
从中得到了两个在包心期表达量增加的阳性cDNA片段。
2对这两个cM片 了了核酌辙拓Uop性分析，发现它呵门u与 申因子egnl和呛删泪肽
转移酶的编匝归丐ung同源。根据核音酸序列阶即叮U，序列比较的结果表明在不同植物
中这两种蛋白质的氨基酸序列 叵强的保守性。
在国际上对大白菜的分子生物学研穷也是刚＠瞒涉，捌〕的工作尚处于初始阶段，有关基因的结构
和功8铱抹进行研究，所以在此不对这些基因溯沽细讨论。相信随着我士门工作进一步的深入，将为大
白菜的分子＋打下重要的基础。
This work is part of our research towards the molecular breeding
of Chinese cabbage (Brassica
calnpestirs). The work has been done in two steps: the analysis
of genetic diversity in vegetable
crops of Brassica campestris using AFLP technique and the mRNA
differential display of the
heading process of Chinese cabbage.
The vegetable crops of Chinese cabbage originated from China,
which comprised non-reading
Chinese cabbage and heading Chinese cabbage. It is the unique
species with special tastes among
all kinds of vegetable crops in the world and one of the earliest
cultivated vegetables in China
Its gennplasm resources are plentiful and have a wide-tange of
distribution. Understanding the
relationships of different types of Chinese cabbage and their
genetic diversity will benefit
the effective collection, documentation and utilization of the
gennplasm and further improve
the breeding work.
The leafy head of heading Chinese cabbage (Brassica campestrisL.
ssp. pekinensis) is typical
an organ of morphological alteration. The vegetative growth of
heading Chinese cabbage involves
four successive stages characterized by definite types of
juvenile, rosette, folding and head
leaves. From rosette stage to folding stage, morphological and
physiological changes occur in
the leaves before the heading process initiates. Studay on the
heading process of Chinese cabbage
will help understanding its unique development mechanism and
metabolism pathways. It also can
serve as the basis for the molecular breeding of Chinese cabbage.
In this study, AFLP technique, followed by the detection using
the fluorescent and sliver
staining systems, has been used to explore the genetic diversity
and taxonomy of 135 accessions
of Brassica campestris collected from different parts of China
and forei~ countries. SM
coefficient was chosen to generate similarity coefficient matrix
and the dendrogram constructed
using UPGMA method. mRNA differential display analysis was then
carried out to see the difference
in gene expression at rosette stage and folding stage of the
heading Chinese cabbage.
The results obtained are:
A the analysis of genetic diversity in vegetable crops of Brassica
cawpestiris using AFLP
technique
1 The amplified bands by the two kinds primer combinations
E+3/M+2 and E+3/M+3, respectively,
were calculated and compared. It was found that the fonner primer
combination produced
3
too many bands, while the later produced a proper number of bands
and the AFLP
fingerprinting was very clear after the sliver-staining.
2 It was found that the number of amplified bands, with the 64
pairs of E+3/M+3 primer
combinations, was between 16 and 58, the percentage of
polymorphic bands was between
40%?8%. Four E+3/M+3 primer combinations with the higher number
of polymorphic bands
were therefore, selected.
3 Clustering results showed that all the accessions were grouped
into two separate groups,
one included all turnip accessions and another all Chinese
cabbage accessions. The heading
Chinese cabbage and the “taicai” a type of non-heading Chinese
cabbage, formed one unique
subgroup in the group of Chinese cabbage. This suggested that
turnip and Chinese cabbage
belong to two different subspecies of Brassica ca'npestris and
the heading Chinese cabbage
is a special variety within the subspecies.
4 That "Taicai" was clustered together with heading Chinese
cabbage into one group suggested
that the "taicai" have a close relationship with the heading
Chinese cabbage and may be
the ancestor of the heading Chinese cabbage.
6 Clustering results of non-heading Chinese cabbage were complex
and showed much deviation
from those of horticultural taxonomy. This indicates that the
difference at the DNA level
is not the same as that at the morphological level in the
non-heading Chinese cabbage.
The similarity amon
分是利用AFLP分子标记技术研究白菜类蔬菜的遗传多样性，第二部分是利用mRNA差异显示技术研究大
白菜结球过程中的基因表达。
白菜类蔬菜包括大白菜和小白菜，原产于我国，是我国在世界各国蔬菜品种和蔬菜栽培中独具风格
的种类。白菜类蔬菜是我国最早的栽培植物之一，它们分布广泛，品种资源非常丰富。对其进行遗传多
样性分析，明确白菜类蔬菜之间的分类关系，有助于人们进一步开发利用其种质资源和更加有效的选择
育种材料。
大白菜的叶球是典型的变态器官。根据叶片发育的形态特征，大白菜的营养生长可分为幼苗期、莲
座期、包心期和结球期四个时期。从莲座期至包心期，是大白菜叶片形态特征和生理功能发生转变的关
键时期，它标志着叶球发生的开始。对大白菜结球过程的研究不仅可以揭示其独特的发育机制和代谢途
径，而且能够为白菜和其它芸薹属蔬菜的分子育种打下基础。
本实验采用AFLP的荧光标记检测技术和银染检测技术，对135份大白菜、小白菜、芜菁材料进行
了遗传多样性和分类研究。在此基础上 进一步以大白菜“北京桔红心”为材料，对其莲座期和包心期
的叶片进行了DD-PCR分析，为揭示由莲座期至包心期发育过程中大白菜叶片基因表达的变化作了一些初
步的尝试。现将主要结果总结如下：
第一部分，利用AFLP分子标记技术研究白菜类蔬菜的遗传多样性
1采用银染检测方法，比较了E+3/M+2、E+3/M+3引物组合对白菜类蔬菜DNA的扩增条带数，发现前者
的扩增条带过于密集，而后者扩增条带数目适中，电泳图谱清晰。
2采用银染检测方法，比较了64对E+3/M+3引物组合对白菜类蔬菜DNA的扩增总带数和多态性检出率，
发现不同引物的扩增总带数在16-58之间，多态检出率在40％-88％之间。从中选择了四对扩增总带
数多，多态性检出率高的引物组合用作分析。
3采用AFLP的荧光标记检测技术对135份材料进行了遗传多样性分析。选用SM系数作为相似性系数
的计算方法，选用UPGMA作为聚类方法。
4聚类结果表明：芜菁和所有的白菜类蔬菜各聚为两类，在白菜类蔬菜类群内部，薹菜和大白菜单独
聚为一类。这表明，芜菁与大白菜、小白菜的相似性程度较低，大白菜与小白菜的相似性程度较高，
大白菜在所有的白菜类蔬菜当中是比较特殊的类群。同意曹家树等提出的将芜菁和白菜（包括大白
菜和小白菜）归为Brassica campestris的两个不同的亚种，大白菜则为白菜亚种中的一个特殊的
变种的观点。
5在小白菜的不同类型中，薹菜与大白菜聚为一类，相似性程度较高，表明薹菜与大白菜的起源可能
存在密切的关系。
6小白菜的AFLP M结果与其综合园艺学分类不符，说明小白菜各类型在DNA M上的差异与其形
态学差异并不存在完全对应的关系。小白菜各类型间的椰以仕程度较大白菜各类型间的＋引以性程度
低，分化程度高，说明小白菜的起源较大白菜要早。
7大白菜的AFLP聚类结果表明，目前生产中所用的大白菜材料之间榔以性矗缠舀交高，而两个大白菜农
家品种与它可l］之间的相似性稷图脚氏。这反映了大ewhIS．＊ru＊品种的遗传多样性较为狭窄，有必要从
大白菜的原始类型中挖掘新的种质资源。
第H部分，利用InRNAgn显示技术研究大白菜结球过程中的基因表达
IAN了4种G、Cng高的锚定引物和3种随机弓1物，共作了12个l）Iyl，nl，反应。在大白菜莲座期和
包吧溯叶片的 mRNA差异显示电泳图谱中，选择了 23个差异 cM ）17ling＋Y了 Reverse NOrthe。检验，
从中得到了两个在包心期表达量增加的阳性cDNA片段。
2对这两个cM片 了了核酌辙拓Uop性分析，发现它呵门u与 申因子egnl和呛删泪肽
转移酶的编匝归丐ung同源。根据核音酸序列阶即叮U，序列比较的结果表明在不同植物
中这两种蛋白质的氨基酸序列 叵强的保守性。
在国际上对大白菜的分子生物学研穷也是刚＠瞒涉，捌〕的工作尚处于初始阶段，有关基因的结构
和功8铱抹进行研究，所以在此不对这些基因溯沽细讨论。相信随着我士门工作进一步的深入，将为大
白菜的分子＋打下重要的基础。
This work is part of our research towards the molecular breeding
of Chinese cabbage (Brassica
calnpestirs). The work has been done in two steps: the analysis
of genetic diversity in vegetable
crops of Brassica campestris using AFLP technique and the mRNA
differential display of the
heading process of Chinese cabbage.
The vegetable crops of Chinese cabbage originated from China,
which comprised non-reading
Chinese cabbage and heading Chinese cabbage. It is the unique
species with special tastes among
all kinds of vegetable crops in the world and one of the earliest
cultivated vegetables in China
Its gennplasm resources are plentiful and have a wide-tange of
distribution. Understanding the
relationships of different types of Chinese cabbage and their
genetic diversity will benefit
the effective collection, documentation and utilization of the
gennplasm and further improve
the breeding work.
The leafy head of heading Chinese cabbage (Brassica campestrisL.
ssp. pekinensis) is typical
an organ of morphological alteration. The vegetative growth of
heading Chinese cabbage involves
four successive stages characterized by definite types of
juvenile, rosette, folding and head
leaves. From rosette stage to folding stage, morphological and
physiological changes occur in
the leaves before the heading process initiates. Studay on the
heading process of Chinese cabbage
will help understanding its unique development mechanism and
metabolism pathways. It also can
serve as the basis for the molecular breeding of Chinese cabbage.
In this study, AFLP technique, followed by the detection using
the fluorescent and sliver
staining systems, has been used to explore the genetic diversity
and taxonomy of 135 accessions
of Brassica campestris collected from different parts of China
and forei~ countries. SM
coefficient was chosen to generate similarity coefficient matrix
and the dendrogram constructed
using UPGMA method. mRNA differential display analysis was then
carried out to see the difference
in gene expression at rosette stage and folding stage of the
heading Chinese cabbage.
The results obtained are:
A the analysis of genetic diversity in vegetable crops of Brassica
cawpestiris using AFLP
technique
1 The amplified bands by the two kinds primer combinations
E+3/M+2 and E+3/M+3, respectively,
were calculated and compared. It was found that the fonner primer
combination produced
3
too many bands, while the later produced a proper number of bands
and the AFLP
fingerprinting was very clear after the sliver-staining.
2 It was found that the number of amplified bands, with the 64
pairs of E+3/M+3 primer
combinations, was between 16 and 58, the percentage of
polymorphic bands was between
40%?8%. Four E+3/M+3 primer combinations with the higher number
of polymorphic bands
were therefore, selected.
3 Clustering results showed that all the accessions were grouped
into two separate groups,
one included all turnip accessions and another all Chinese
cabbage accessions. The heading
Chinese cabbage and the “taicai” a type of non-heading Chinese
cabbage, formed one unique
subgroup in the group of Chinese cabbage. This suggested that
turnip and Chinese cabbage
belong to two different subspecies of Brassica ca'npestris and
the heading Chinese cabbage
is a special variety within the subspecies.
4 That "Taicai" was clustered together with heading Chinese
cabbage into one group suggested
that the "taicai" have a close relationship with the heading
Chinese cabbage and may be
the ancestor of the heading Chinese cabbage.
6 Clustering results of non-heading Chinese cabbage were complex
and showed much deviation
from those of horticultural taxonomy. This indicates that the
difference at the DNA level
is not the same as that at the morphological level in the
non-heading Chinese cabbage.
The similarity amon
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