植物顶端组织高效诱导型启动子的创建与表达调控研究
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摘要
目前，启动子的研究是国内外的一个研究热点，现已发现了大
量组织特异性启动子和诱导型启动子。在仔细研究了这些特异性启
动子的结构特点，以及其中起决定作用的顺式作用调控元件的基础
上，本文从中选择了两个研究背景比较清楚、序列结构具有共性的
植物顶端新生组织特异性调控元件B1元件和水杨酸诱导型调控元
件as-1元件进行了进一步研究。
以35S启动子的基本框架结构（-90至+1区间的序列）为基础，
利用人工合成的这两个调控元件进行重新组装、构建了九个分别含
有一个、两个、四个B1元件和as-1元件的系列启动子和带有gfp
基因的质粒，将它们分别命名为pUC191B1a.GFP、pUC191B2a.GFP、
pUC191B4a.GFP、pUC192B1a.GFP、pUC192B2a.GFP、pUC192B4a.GFP、
PUC194B1a.GFP、pUC194B2a.GFP和pUC194B4a.GFP。并进一步
分别构建出九个不同的植物表达载体，分别命名为pBI1B1a.GFP、
pBI1B2a.GFP、pBI1B4a.GFP、pBI2B1a.GFP、pBI2B2a.GFP、pBI2B4a.GFP、
PBI4B1a.GFP、PBI4B2a.GFP、pBI4B4a.GFP。利用农杆菌介导的叶
盘转化法转化烟草，通过高浓度的卡那霉素连续筛选，最终获得186
株转基因烟草植株。经过荧光显微镜初步筛选，获得含有1B1a启
动子的植株9株、1B2a启动子的植株12株、1B4a启动子的植株14
株、2B1a启动子植株8株、2B2a启动子的植株11株、2B4a启动子
的植株9株、4B1a启动子的植株15株、4B2a启动子的植株15株
和4B4a启动子的植株19株。经PCR扩增和Southern blot分析表明
各种表达载体已分别整合进转基因烟草植株的基因组中。
以每种启动子的转基因烟草植株为一个小群体，分别利用荧光
显微镜在蓝色激发光下观察各种转基因烟草植株中各组织部位的
GFP表达水平，并利用荧光分光光度计在460nm激发光和509nm
发射光下对不同部位叶片中GFP的表达量进行了定量分析。两种不
同的分析方法一致证明：（1）含有B1顺式作用元件的启动子具有在
植株顶端叶片和茎尖中表达的组织特异性，其在茎中的表达也主要
行 男 二
集中在茎的外表皮和韧皮部组织。仅旧 元件具有累加效应，启动
子中含有BI 元件越多，该启动子在顶端组织特异性表达就越强，
含有四个 田 元件的启动子的顶端组织特异性最显著。在转基因烟
草植株的同一部位，含有四个 ＊ 元件启动子的表达水平是含有两
个田元件启动子表达水平的2．9倍，是含有单个BI元件启动子表
达水平的3．9倍，含有两个BI元件启动子的表达水平是含有单个BI
元件启动子表达水平的 1．3倍。o）含有相同 BI元件而 asJ元件数
目不同的三种启动子的表达水平没有明显差异，说明正常条件下含
有 BI 元件和 as1元件启动子的表达水平主要由 BI 元件的多少决
定，而不受asJ元件的影响。
在应用ZmM、20mM和50mM水杨酸溶液喷洒转基因烟草植株
叶片后，在不同时间段内选取同一部位的叶片，通过荧光显微镜观
察和荧光分光光度计测量两种方法，定性、定量地分析了不同浓度
的水杨酸诱导下含有不同启动子的转基因烟草植株中GFP表达水平
的变化。两种方法一致证明：（1）as－1 元件能赋予启动子水杨酸诱
导特性。（2）水杨酸的诱导效果与所喷洒水杨酸的浓度及 的值密切
相关，供试溶液中以50mM PH6．8 的水杨酸溶液诱导效果最佳。（3）
水杨酸诱导后基因表达水平的增长幅度与启动子中as－1元件的数目
密切相关，含有单个asJ 元件的启动子在诱导后表达水平比诱导前
提高2．2～2．8倍，含有两个asJ元件的启动子在诱导后表达水平比
诱导前提高3～3．5倍，含有四个asA元件的启动子在诱导后表达水
平比诱导前提高 13～17倍。O）增加 BI顺式作用元件数目有协同增
强as八元件表达的作用，含有四个BI元件和四个as八元件的4B4a
启动子，其不但具有很高的顶端叶片和茎尖组织特异性，而且在水
杨酸诱导后 24小时左右，其表达水平比诱导前可提高 17．3倍。如
此高效表达目前未见类似报道。
本研究结果有力地证明了来自启动子中的组织特异性BI元件
和诱导性as1元件，可被直接用于改造或构建人工融合启动子，
并赋予启动子组织特异性表达的特性和水杨酸诱导特性，这一研
MK 3
究结果为今后构建组织特异性或诱导型启动子提供了依据。
将BI和as－1顺式作用调控元件串联叠加起来构建成新型的、
既有顶端组织特异性又有水杨酸诱导性并能高效表达的启动子，
目前尚未见到类似报道。
Based on the study of published cis-elements of tissue-specific
and inducible
promoters. Salicylic acid (SA) inducible as-I element and the
green tissue-
specific 81 element, derived from the cauliflower mosaic virus
35S promoter,
were selected. Because both of the two promoters have well-known
research
background and more commonness than other cis-elements.
Based on a truncation of?0 355 promoter (-90 to +1), nine
different promoters
were constructed with artificially synthesized as-I sequence and
BI sequence.
Each one of them respectively has one copy, two copies or four
copies of as-I
sequence and 81 sequence, and were subsequently designated as
IBIa promoter,
182a promoter, lB4a promoter, 2BIa promoter, 2B2a promoter, 2B4a
promoter,
4Bta promoter, 4B2a promoter and 4B4a promoter. And all the
constructs were
confirmed by sequencing. Then nine plant expression vectors
containing these
corresponding promoters were constructed, in which the GFP gene
was
downstreamed as a reporter gene. These plant expression vectors
were
subsequently designated as p811 B 1 a.GFP, p811 B I a.GFP, p811 B
I a.GFP.
pBI2Bla.GFP, pBl4BIa.GFP, pBIIB2a.GFP, pBl2B2a.GFP, pBl4B2a.GFP,
pBIlB4a.GFP, pBI2B4a.GFP and pBl4B4a.GFP. Tobacco leaf discs were
transformed with Agrobacterium tumefaciences Conn LBA4404
containing these
expression vectors, and 186 trangenic plants were obtained. Among
of them, 106
pass muster transgenic plants were firstly screened out by
Fluorescent
Microscope.
Results of testing transgenic plants with PCR, Southern blot
methods confirmed
the successful integration of exogenous gene into tobacco plants
genome.
The GFP expressing level of different tissues located on
different parts of
respective trangenic plants transformed by different promoters
were observed by
the Fluorescent Microscopy, and the expressing level of different
leaves located
5
on different parts of transgenic plants were quantitatively
analyzed by Fluorescent
Spectrophotometer. The results of two methods were highly
consistent with each
other and confirmed following conclusions:
?The promoters containing B! element show a tissue-specific
expression
principally in new leaves and the stem apex.
?The B I element shows a accumulative characteristic when adding
the its copy
number in promoter. The expression level of promoters containing
four
copies of B! sequence is 2.9 fold the level of promoters
containing two
copies of B1 sequence, is 3.9 fold the level of promoters
containing one
copy of 81 sequence. The expression level of promoters containing
two
copies of B I sequence is I .3 fold the level of promoters
containing one copy
of 81 sequence.
?No difference has been observed between the different transgenic
plants of
three kinds of promoters that containing same copy number of BI
element
and different copy number of as-I element. The results mean that
the
expression level of promoter is principally dependent on the
number of 81
sequence contained in the promoter, and the number of as-I
sequence has no
obvious influence on it under the general condition.
After sprayed by salicylic acid solutions at various
concentration, such as 2mM,
20mM and 50mM, the transgenic tobacco plants carrying different
promoters were
tested for their response to the SA treatment by Fluorescent
Microscopy and
Fluorescent Spectrophotometer both qualitatively and
quantitatively. The results
of two methods were highly consistent with each other and
confirmed following
conclusions:
?The as-I element shows a positive response to the SA signal
transduction,
and the promoter containing as-I element show a trait of
transcription
activation by SA inducement.
6
?Inductive efficiency is closely dependent on concentration and
pH value of
SA solution. The results suggest that the inducti
量组织特异性启动子和诱导型启动子。在仔细研究了这些特异性启
动子的结构特点，以及其中起决定作用的顺式作用调控元件的基础
上，本文从中选择了两个研究背景比较清楚、序列结构具有共性的
植物顶端新生组织特异性调控元件B1元件和水杨酸诱导型调控元
件as-1元件进行了进一步研究。
以35S启动子的基本框架结构（-90至+1区间的序列）为基础，
利用人工合成的这两个调控元件进行重新组装、构建了九个分别含
有一个、两个、四个B1元件和as-1元件的系列启动子和带有gfp
基因的质粒，将它们分别命名为pUC191B1a.GFP、pUC191B2a.GFP、
pUC191B4a.GFP、pUC192B1a.GFP、pUC192B2a.GFP、pUC192B4a.GFP、
PUC194B1a.GFP、pUC194B2a.GFP和pUC194B4a.GFP。并进一步
分别构建出九个不同的植物表达载体，分别命名为pBI1B1a.GFP、
pBI1B2a.GFP、pBI1B4a.GFP、pBI2B1a.GFP、pBI2B2a.GFP、pBI2B4a.GFP、
PBI4B1a.GFP、PBI4B2a.GFP、pBI4B4a.GFP。利用农杆菌介导的叶
盘转化法转化烟草，通过高浓度的卡那霉素连续筛选，最终获得186
株转基因烟草植株。经过荧光显微镜初步筛选，获得含有1B1a启
动子的植株9株、1B2a启动子的植株12株、1B4a启动子的植株14
株、2B1a启动子植株8株、2B2a启动子的植株11株、2B4a启动子
的植株9株、4B1a启动子的植株15株、4B2a启动子的植株15株
和4B4a启动子的植株19株。经PCR扩增和Southern blot分析表明
各种表达载体已分别整合进转基因烟草植株的基因组中。
以每种启动子的转基因烟草植株为一个小群体，分别利用荧光
显微镜在蓝色激发光下观察各种转基因烟草植株中各组织部位的
GFP表达水平，并利用荧光分光光度计在460nm激发光和509nm
发射光下对不同部位叶片中GFP的表达量进行了定量分析。两种不
同的分析方法一致证明：（1）含有B1顺式作用元件的启动子具有在
植株顶端叶片和茎尖中表达的组织特异性，其在茎中的表达也主要
行 男 二
集中在茎的外表皮和韧皮部组织。仅旧 元件具有累加效应，启动
子中含有BI 元件越多，该启动子在顶端组织特异性表达就越强，
含有四个 田 元件的启动子的顶端组织特异性最显著。在转基因烟
草植株的同一部位，含有四个 ＊ 元件启动子的表达水平是含有两
个田元件启动子表达水平的2．9倍，是含有单个BI元件启动子表
达水平的3．9倍，含有两个BI元件启动子的表达水平是含有单个BI
元件启动子表达水平的 1．3倍。o）含有相同 BI元件而 asJ元件数
目不同的三种启动子的表达水平没有明显差异，说明正常条件下含
有 BI 元件和 as1元件启动子的表达水平主要由 BI 元件的多少决
定，而不受asJ元件的影响。
在应用ZmM、20mM和50mM水杨酸溶液喷洒转基因烟草植株
叶片后，在不同时间段内选取同一部位的叶片，通过荧光显微镜观
察和荧光分光光度计测量两种方法，定性、定量地分析了不同浓度
的水杨酸诱导下含有不同启动子的转基因烟草植株中GFP表达水平
的变化。两种方法一致证明：（1）as－1 元件能赋予启动子水杨酸诱
导特性。（2）水杨酸的诱导效果与所喷洒水杨酸的浓度及 的值密切
相关，供试溶液中以50mM PH6．8 的水杨酸溶液诱导效果最佳。（3）
水杨酸诱导后基因表达水平的增长幅度与启动子中as－1元件的数目
密切相关，含有单个asJ 元件的启动子在诱导后表达水平比诱导前
提高2．2～2．8倍，含有两个asJ元件的启动子在诱导后表达水平比
诱导前提高3～3．5倍，含有四个asA元件的启动子在诱导后表达水
平比诱导前提高 13～17倍。O）增加 BI顺式作用元件数目有协同增
强as八元件表达的作用，含有四个BI元件和四个as八元件的4B4a
启动子，其不但具有很高的顶端叶片和茎尖组织特异性，而且在水
杨酸诱导后 24小时左右，其表达水平比诱导前可提高 17．3倍。如
此高效表达目前未见类似报道。
本研究结果有力地证明了来自启动子中的组织特异性BI元件
和诱导性as1元件，可被直接用于改造或构建人工融合启动子，
并赋予启动子组织特异性表达的特性和水杨酸诱导特性，这一研
MK 3
究结果为今后构建组织特异性或诱导型启动子提供了依据。
将BI和as－1顺式作用调控元件串联叠加起来构建成新型的、
既有顶端组织特异性又有水杨酸诱导性并能高效表达的启动子，
目前尚未见到类似报道。
Based on the study of published cis-elements of tissue-specific
and inducible
promoters. Salicylic acid (SA) inducible as-I element and the
green tissue-
specific 81 element, derived from the cauliflower mosaic virus
35S promoter,
were selected. Because both of the two promoters have well-known
research
background and more commonness than other cis-elements.
Based on a truncation of?0 355 promoter (-90 to +1), nine
different promoters
were constructed with artificially synthesized as-I sequence and
BI sequence.
Each one of them respectively has one copy, two copies or four
copies of as-I
sequence and 81 sequence, and were subsequently designated as
IBIa promoter,
182a promoter, lB4a promoter, 2BIa promoter, 2B2a promoter, 2B4a
promoter,
4Bta promoter, 4B2a promoter and 4B4a promoter. And all the
constructs were
confirmed by sequencing. Then nine plant expression vectors
containing these
corresponding promoters were constructed, in which the GFP gene
was
downstreamed as a reporter gene. These plant expression vectors
were
subsequently designated as p811 B 1 a.GFP, p811 B I a.GFP, p811 B
I a.GFP.
pBI2Bla.GFP, pBl4BIa.GFP, pBIIB2a.GFP, pBl2B2a.GFP, pBl4B2a.GFP,
pBIlB4a.GFP, pBI2B4a.GFP and pBl4B4a.GFP. Tobacco leaf discs were
transformed with Agrobacterium tumefaciences Conn LBA4404
containing these
expression vectors, and 186 trangenic plants were obtained. Among
of them, 106
pass muster transgenic plants were firstly screened out by
Fluorescent
Microscope.
Results of testing transgenic plants with PCR, Southern blot
methods confirmed
the successful integration of exogenous gene into tobacco plants
genome.
The GFP expressing level of different tissues located on
different parts of
respective trangenic plants transformed by different promoters
were observed by
the Fluorescent Microscopy, and the expressing level of different
leaves located
5
on different parts of transgenic plants were quantitatively
analyzed by Fluorescent
Spectrophotometer. The results of two methods were highly
consistent with each
other and confirmed following conclusions:
?The promoters containing B! element show a tissue-specific
expression
principally in new leaves and the stem apex.
?The B I element shows a accumulative characteristic when adding
the its copy
number in promoter. The expression level of promoters containing
four
copies of B! sequence is 2.9 fold the level of promoters
containing two
copies of B1 sequence, is 3.9 fold the level of promoters
containing one
copy of 81 sequence. The expression level of promoters containing
two
copies of B I sequence is I .3 fold the level of promoters
containing one copy
of 81 sequence.
?No difference has been observed between the different transgenic
plants of
three kinds of promoters that containing same copy number of BI
element
and different copy number of as-I element. The results mean that
the
expression level of promoter is principally dependent on the
number of 81
sequence contained in the promoter, and the number of as-I
sequence has no
obvious influence on it under the general condition.
After sprayed by salicylic acid solutions at various
concentration, such as 2mM,
20mM and 50mM, the transgenic tobacco plants carrying different
promoters were
tested for their response to the SA treatment by Fluorescent
Microscopy and
Fluorescent Spectrophotometer both qualitatively and
quantitatively. The results
of two methods were highly consistent with each other and
confirmed following
conclusions:
?The as-I element shows a positive response to the SA signal
transduction,
and the promoter containing as-I element show a trait of
transcription
activation by SA inducement.
6
?Inductive efficiency is closely dependent on concentration and
pH value of
SA solution. The results suggest that the inducti
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