植物油体表达体系的建立及Profilin2维管束特异表达启动子的区段缺失分析
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摘要
本论文的目的是：（1）建立在植物油体中表达外源蛋白的表达体系，以简化
目的蛋白的分离纯化过程；（2）对维管束特异表达的Profilin2启动子进行区段分
析，寻找控制维管束特异表达的区段和元件，为在抗维管束病害的基因工程中
使抗病基因产物直接在维管束中特异表达并对病原体建立防御体系，以节省能
量，同时增加转基因作物的安全性。
为建立油体表达体系，本论文进行了以下研究：（1）分离和克隆了油菜
Oleosin基因的启动子及编码芝麻Oleosin蛋白的结构基因。（2）克隆了鲑鱼降钙
素的突变基因msCT[Ser~6，des-Leu~(19)]，将其插在芝麻油体蛋白Oleosin基因的C
端。据文献报道，鲑鱼降钙素（sCT）在治疗缺钙症等中有明显的疗效。（3）构
建了由油菜Oleosin基因启动子驱动的Oleosin-Calcitonin融合蛋白植物表达载
体，转化油菜和棉花，获得了转基因植株。转基因油菜经PCR检测，证明降钙
素基因已整合到油菜基因组中。转基因棉花经PCR-Southern和Western检测，
证明msCT基因在棉花中整合并在油体中表达。目前已得到T_2代转基因棉花株
系44个，由于在田间对T_1、T_2代植株的叶片用高浓度（5000 ppm）的卡那霉素
溶液涂抹，故筛选出的Kan~R株理论上应为高表达的个体。用植物油体表达体系
在转基因棉籽油中表达鲑鱼降钙素，在文献中尚未见报道。
在Profilin2维管束特异表达启动子的区段分析中，根据Pfn全长启动子
（Pfn1.7，-1667～-1bp）及其5’端4种不同长度（Pfn1.4、Pfn1.2、Pfn1.0、Pfn0.6）
的缺失分析，检测gus基因在转基因伽蓝菜中的表达，可将Pfn全长启动子分
成三部分：区段A，-1667～-1380 bp，缺失该区段（Pfn1.4），由维管束特异表
达变为组成型表达，由此推测在该区段内存在维管束特异表达的元件。区段B，
-1153～-597bp，强烈抑制gus基因的表达，推测在该区段中存在负调控因子。
区段C，-597～-1bp，即Pfn0.6，除在维管束中表达较强外，还在薄壁细胞中
表达，故可认为该区段是Profilin2的基本启动子。进一步分析发现，Profilin2
启动子在-1391～-1388 bp处及-565～-562bp处各有一个bZIP蛋白结合位点
的核心序列ACGT（bZIP蛋白是植物中最丰富的一类转录因子）；-1647～-1640bp
处有一个与控制菜豆苯丙氨酸氨裂合酶2（PAL2）维管束特异表达的序列AC-I
（CCCACCTACC）相类似的序列（CCACCTAC）。为弄清ACGT及AC-I序列在
Profilin2启动子中是否调控维管束的特异表达，我们又分别构建了P1（-1667～-
刃 县
1403，含 AC－l）、PZ（如27一羽80，含 ACGT）、P3（－1667～羽80，含 ACGT
和AC－1）、P4（－1667一1627，含AC－l）、PS（－1403一1376，含ACGT）等五
个不同区段与80 hp CaMV35S最小启动子（P35Sm）连接的融合启动子，构建
植物表达载体，转化烟草和伽蓝菜。在烟草短暂表达中，PZ（含ACGT）启动子
的表达强度明显高于n（含AC0，推测bZIP蛋白结合序列ACGT在调控表
达强度方面可能起更重要的作用。转基因烟草的稳定表达结果表明，P135Smfus
构建物（含AC工序列）仅在茎、叶维管束中检测到GUS表达。鉴于355最小启
动子在植物的所有发育阶段都不能表达，故说明AC．I作为一个顺式作用元件
k七lementX 确能调控维管柬的特异表达。其余构建物因目前仅得到转基因
的小芽，尚不能进行检测，故其对稳定表达的影响尚待进一步研究。据作者所
知，这是首次对profilinZ启动子进行的区段缺失和基元序列分析。
The main objectives of this study are (1) to establish an oil
body-based
expression system, by which the isolation and purification
process of a target protein
can be simplified; (2) to analyze fragments of a
vascular-specific Arabidopsis
proflhin2 promoter, aiming at identifying domains and motifs that
regulate genes of
interest specifically expressed in plant vascular bundles. This
is particularly important
in the genetic engineering of crops with enhanced resistance to
vascular diseases,
since the gene product specifically expressed in the vascular
system may directly
provide a defense mechanism to vascular-resided pathogens.
Meanwhile, the vascular-
specific expression, compared with the constitutive expression,
may save energy and
reduce risks of transgenic crops.
For the establishment of an oil body-based expression system, the
following
experiments were conducted: (1) Cloning of a rape seed (Brassica
napus) oleosin
promoter and a sesame structure gene coding for oleosin protein
which inserts in the
surface of the oil body. (2) Cloning of a mutated msCT (Ser6,
des-leu'9) gene coding
for salmon calcitonin (sCT) protein, an efficient drug used for
curing calcium
deficiency, etc. The msCT gene was inserted in the C'-end of the
sesame oleosin gene
driven by the rape seed oleosin promoter. (3) Construction of a
plant expression vector
and transformation of B. napus by Agrobacterium-mediated gene
transfer and of
cotton (Gossypium babardense) by pollen-tube pathway. PCR
analysis showed that
the msCT gene was integrated in the rape seed genome.
PCR-Southern hybridization
of transgenic cotton was positive and Western analysis of
oleosin-calcitonin fusion
protein extracted from cotton-seed oil showed an expected ?9 kD
band indicating
that the msCT gene was specifically expressed in the cotton-seed
oil body. At present,
44 transgenic cotton lines of 12 generation have been obtained.
Since a high
concentration of kanamycin (5,000 ppm) was applied on leaf in the
field testing, the
transgenic plant selected should be the one with higher gene
expression. As far as we
know, the establishment of an oil body-based system to produce
calcitonin protein in
cotton-seed oil has not been previously reported in the
literature.
Based on the transient and stable gus gene expression driven by
different 5'-
deletions of Pfn promoter (Pflil .4, Pfnl .2, Pfhl .0, PfliO.6),
the full-length promoter
(Pfril.7, -1667 -1 bp) can be divided into 3 parts: (1) Domain A,
-1667 ?1380 bp.
Deletion of this part, i.e. from Pfnl .7 to Pfnl .4, resulted in
the transition of gus gene
3
expression from vascular-specific to constitutive. It suggested
that element(s)
responsible for vascular-specific expression might be existed in
this region. (2)
Domain B, located at -1153 -597 bp, strongly inhibited gus gene
expression
indicating there might be negative regulatory element(s)
presented. (3) Domain C, -
597 -1 bp (PfhO.6) was considered as a basic domain of profilin2,
since GUS
activity was not only relatively strong in vascular system but
also in parenchyma cells.
Detailed sequence analysis indicated that there was a core ACGT
motif, the bZIP
protein binding site, both at -1391 ?-1388 bp and -565 ?-562 bp
in the profllin2
promoter. It is known that bZIP proteins are the most abundant
transcription factors in
plants. Analysis also shown that there was an AC-I (CCCACCTACC)
like sequence at
-1647 ?-1640 bp (CCACCTAC) of the profihin2 promoter. The AC-I
motif, also as a
protein-binding site, was known to be very important in
controlling vascular-specific
expression of the bean phenylalanine ammonia-lyase2 (PAL2)
promoter. To clarify
whether the ACGT or AC-I like sequence in profilin2 promoter
plays a similar
regulatory role in vascular-specific expression, we further
constructed following plant
expression vectors contai
目的蛋白的分离纯化过程；（2）对维管束特异表达的Profilin2启动子进行区段分
析，寻找控制维管束特异表达的区段和元件，为在抗维管束病害的基因工程中
使抗病基因产物直接在维管束中特异表达并对病原体建立防御体系，以节省能
量，同时增加转基因作物的安全性。
为建立油体表达体系，本论文进行了以下研究：（1）分离和克隆了油菜
Oleosin基因的启动子及编码芝麻Oleosin蛋白的结构基因。（2）克隆了鲑鱼降钙
素的突变基因msCT[Ser~6，des-Leu~(19)]，将其插在芝麻油体蛋白Oleosin基因的C
端。据文献报道，鲑鱼降钙素（sCT）在治疗缺钙症等中有明显的疗效。（3）构
建了由油菜Oleosin基因启动子驱动的Oleosin-Calcitonin融合蛋白植物表达载
体，转化油菜和棉花，获得了转基因植株。转基因油菜经PCR检测，证明降钙
素基因已整合到油菜基因组中。转基因棉花经PCR-Southern和Western检测，
证明msCT基因在棉花中整合并在油体中表达。目前已得到T_2代转基因棉花株
系44个，由于在田间对T_1、T_2代植株的叶片用高浓度（5000 ppm）的卡那霉素
溶液涂抹，故筛选出的Kan~R株理论上应为高表达的个体。用植物油体表达体系
在转基因棉籽油中表达鲑鱼降钙素，在文献中尚未见报道。
在Profilin2维管束特异表达启动子的区段分析中，根据Pfn全长启动子
（Pfn1.7，-1667～-1bp）及其5’端4种不同长度（Pfn1.4、Pfn1.2、Pfn1.0、Pfn0.6）
的缺失分析，检测gus基因在转基因伽蓝菜中的表达，可将Pfn全长启动子分
成三部分：区段A，-1667～-1380 bp，缺失该区段（Pfn1.4），由维管束特异表
达变为组成型表达，由此推测在该区段内存在维管束特异表达的元件。区段B，
-1153～-597bp，强烈抑制gus基因的表达，推测在该区段中存在负调控因子。
区段C，-597～-1bp，即Pfn0.6，除在维管束中表达较强外，还在薄壁细胞中
表达，故可认为该区段是Profilin2的基本启动子。进一步分析发现，Profilin2
启动子在-1391～-1388 bp处及-565～-562bp处各有一个bZIP蛋白结合位点
的核心序列ACGT（bZIP蛋白是植物中最丰富的一类转录因子）；-1647～-1640bp
处有一个与控制菜豆苯丙氨酸氨裂合酶2（PAL2）维管束特异表达的序列AC-I
（CCCACCTACC）相类似的序列（CCACCTAC）。为弄清ACGT及AC-I序列在
Profilin2启动子中是否调控维管束的特异表达，我们又分别构建了P1（-1667～-
刃 县
1403，含 AC－l）、PZ（如27一羽80，含 ACGT）、P3（－1667～羽80，含 ACGT
和AC－1）、P4（－1667一1627，含AC－l）、PS（－1403一1376，含ACGT）等五
个不同区段与80 hp CaMV35S最小启动子（P35Sm）连接的融合启动子，构建
植物表达载体，转化烟草和伽蓝菜。在烟草短暂表达中，PZ（含ACGT）启动子
的表达强度明显高于n（含AC0，推测bZIP蛋白结合序列ACGT在调控表
达强度方面可能起更重要的作用。转基因烟草的稳定表达结果表明，P135Smfus
构建物（含AC工序列）仅在茎、叶维管束中检测到GUS表达。鉴于355最小启
动子在植物的所有发育阶段都不能表达，故说明AC．I作为一个顺式作用元件
k七lementX 确能调控维管柬的特异表达。其余构建物因目前仅得到转基因
的小芽，尚不能进行检测，故其对稳定表达的影响尚待进一步研究。据作者所
知，这是首次对profilinZ启动子进行的区段缺失和基元序列分析。
The main objectives of this study are (1) to establish an oil
body-based
expression system, by which the isolation and purification
process of a target protein
can be simplified; (2) to analyze fragments of a
vascular-specific Arabidopsis
proflhin2 promoter, aiming at identifying domains and motifs that
regulate genes of
interest specifically expressed in plant vascular bundles. This
is particularly important
in the genetic engineering of crops with enhanced resistance to
vascular diseases,
since the gene product specifically expressed in the vascular
system may directly
provide a defense mechanism to vascular-resided pathogens.
Meanwhile, the vascular-
specific expression, compared with the constitutive expression,
may save energy and
reduce risks of transgenic crops.
For the establishment of an oil body-based expression system, the
following
experiments were conducted: (1) Cloning of a rape seed (Brassica
napus) oleosin
promoter and a sesame structure gene coding for oleosin protein
which inserts in the
surface of the oil body. (2) Cloning of a mutated msCT (Ser6,
des-leu'9) gene coding
for salmon calcitonin (sCT) protein, an efficient drug used for
curing calcium
deficiency, etc. The msCT gene was inserted in the C'-end of the
sesame oleosin gene
driven by the rape seed oleosin promoter. (3) Construction of a
plant expression vector
and transformation of B. napus by Agrobacterium-mediated gene
transfer and of
cotton (Gossypium babardense) by pollen-tube pathway. PCR
analysis showed that
the msCT gene was integrated in the rape seed genome.
PCR-Southern hybridization
of transgenic cotton was positive and Western analysis of
oleosin-calcitonin fusion
protein extracted from cotton-seed oil showed an expected ?9 kD
band indicating
that the msCT gene was specifically expressed in the cotton-seed
oil body. At present,
44 transgenic cotton lines of 12 generation have been obtained.
Since a high
concentration of kanamycin (5,000 ppm) was applied on leaf in the
field testing, the
transgenic plant selected should be the one with higher gene
expression. As far as we
know, the establishment of an oil body-based system to produce
calcitonin protein in
cotton-seed oil has not been previously reported in the
literature.
Based on the transient and stable gus gene expression driven by
different 5'-
deletions of Pfn promoter (Pflil .4, Pfnl .2, Pfhl .0, PfliO.6),
the full-length promoter
(Pfril.7, -1667 -1 bp) can be divided into 3 parts: (1) Domain A,
-1667 ?1380 bp.
Deletion of this part, i.e. from Pfnl .7 to Pfnl .4, resulted in
the transition of gus gene
3
expression from vascular-specific to constitutive. It suggested
that element(s)
responsible for vascular-specific expression might be existed in
this region. (2)
Domain B, located at -1153 -597 bp, strongly inhibited gus gene
expression
indicating there might be negative regulatory element(s)
presented. (3) Domain C, -
597 -1 bp (PfhO.6) was considered as a basic domain of profilin2,
since GUS
activity was not only relatively strong in vascular system but
also in parenchyma cells.
Detailed sequence analysis indicated that there was a core ACGT
motif, the bZIP
protein binding site, both at -1391 ?-1388 bp and -565 ?-562 bp
in the profllin2
promoter. It is known that bZIP proteins are the most abundant
transcription factors in
plants. Analysis also shown that there was an AC-I (CCCACCTACC)
like sequence at
-1647 ?-1640 bp (CCACCTAC) of the profihin2 promoter. The AC-I
motif, also as a
protein-binding site, was known to be very important in
controlling vascular-specific
expression of the bean phenylalanine ammonia-lyase2 (PAL2)
promoter. To clarify
whether the ACGT or AC-I like sequence in profilin2 promoter
plays a similar
regulatory role in vascular-specific expression, we further
constructed following plant
expression vectors contai
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