百子莲脱水素基因ApSK3对逆境与激素信号的应答模式与调控机制

doi:10.16420/j.issn.0513-353x.2020-0660

摘要/Abstract

摘要：

克隆了百子莲（Agapanthus praecox）脱水素基因ApSK₃上游2 195 bp的启动子序列,对百子莲不同组织和多种非生物胁迫与ABA处理的样品进行基因定量分析,构建多个ApSK₃不同长度缺失的启动子片段与GUS基因融合的表达载体并转化拟南芥,以揭示ApSK₃基因对不同逆境与激素信号的应答模式及顺式作用元件的调控功能。结果表明：ApSK₃启动子序列包含多个与植物逆境、激素应答及生长发育相关的顺式作用元件,且ApSK₃的表达具有组织特异性,果实中表达量最高,叶、根次之,花中最低;ApSK₃对ABA信号与盐胁迫最敏感,其次为干旱、高渗和低温胁迫,对高温胁迫响应不明显。ApSK₃-P::GUS 融合表达载体转化拟南芥,ApSK₃启动子在拟南芥的整个生长发育过程中均具有较强的表达活性,幼苗根部的表达活性强于叶片,且随着果实的发育成熟启动子的活性明显增强。ApSK₃不同长度缺失的启动子片段分析结果表明-2 175 ~-950 bp片段对启动子的活性起到重要的调控作用;多个顺式作用元件响应干旱胁迫;ApSK₃-P通过两个ABRE元件共同响应ABA信号;-526 ~-533 bp的 ERE元件参与响应乙烯信号;-561 ~-567 bp的P-box元件响应了赤霉素信号,-2 175 ~-1 167 bp区域可以增强对GA的响应。研究结果证明ApSK₃启动子可积极响应干旱、渗透、盐、低温、ABA、GA和乙烯信号;启动子上存在多个顺式作用元件响应干旱胁迫,ApSK₃-P通过两个ABRE、ERE与P-box元件分别响应ABA、乙烯与赤霉素信号。

关键词: 百子莲, 脱水素, 启动子, 非生物胁迫, 顺式作用元件

Abstract:

This study cloned the dehydrin ApSK₃（Agapanthus praecox）2 195 bp promoter sequence of ApSK₃-P using real-time PCR technology to analyze tissue specificity of ApSK₃ gene,and gene expression pattern response to various abiotic stress and ABA signal. Expression vectors with five promoter deletion fragments of ApSK₃-P were constructed and transformed into Arabidopsis thaliana,in order to reveal the response pattern of ApSK₃ gene to different stress and hormone signals and the regulatory function of cis-acting elements. Bio-informatic analysis showed that the ApSK₃-P contained many typical cis-acting elements related to plant stress,hormone response and plant development. ApSK₃ gene was tissue-specific expression,the highest expression was found in fruits,followed by leaves and roots,and the lowest value was shown in flowers. ApSK₃ is more sensitive to ABA and salt signals,followed by drought and low temperature,and no obvious response to high temperature. ApSK₃-P::GUS were transformed into A. thaliana and it showed that ApSK₃-P presented strong activity in seedlings of A. thaliana,GUS activity of roots was stronger than that of leaves,and the activity gradually increased in the developing fruits. Different promoter deletion fragments of ApSK₃-P test results showed that-2 175 to-950 bp fragment played an important role in the regulation of promoter activity,multiple cis-acting elements responsed to drought stress,two tandem ABRE elements participated in response to ABA signals,and ApSK₃-P can be activated by ERE（-526 to-533 bp）and P-box（-561 to-567 bp）under ethylene and gibberellin signal,respectively. These results indicated that ApSK₃-P can be activated by drought,cold,salt,ABA,GA and ethylene,and ABRE,ERE and P-box elements of ApSK₃-P play important roles in response to abiotic stress and hormone signals.

Key words: Agapanthus praecox, dehydrin, promoter, abiotic stress, cis-acting element

中图分类号:

S682.1

杨天宸, 陈晓童, 吕可, 张荻. 百子莲脱水素基因ApSK₃对逆境与激素信号的应答模式与调控机制[J]. 园艺学报, 2021, 48(8): 1565-1578.

YANG Tianchen, CHEN Xiaotong, LÜ Ke, ZHANG Di. Expression Pattern and Regulation Mechanism of ApSK₃ Dehydrin （Agapanthus praecox）Response to Abiotic Stress and Hormone Signals[J]. Acta Horticulturae Sinica, 2021, 48(8): 1565-1578.

图/表 9

表1 引物序列

Table 1 Corresponding primer sequences

引物名称	序列（5′-3′）	退火温度/℃
Primer name	Primer sequence	T_m
Ap-Actin-S	CAGTGTCTGGATTGGAGG	50.0
Ap-Actin-A	TAGAAGCACTTCCTGTG	50.0
RT-ApSK₃-S	AAGAGCCAAGAGGAGGTT	55.0
RT-ApSK₃-A	CTTCTTCTCGCCGTCTTC	55.0
ApSK₃-SP1	TGTGGCCGGGGAGCTTCTGTTT	61.4
ApSK₃-SP2	ACGTCCTGTTCGGTTACCACGG	61.4
ApSK₃-SP3	CCACTTCCTCTTCGTCGCTCGA	61.4
ApSK₃-SP2-1	AGCTAGAGAAACGAATTATCACATCCCTC	59.6
ApSK₃-SP2-2	ACTTTCAAGCTCTCGACTCCGG	59.5
ApSK₃-SP2-3	CCCTCTCTCACTCACCTCCACA	61.4
Sp-2175-S	ATGACCATGATTACGCCAAGCTTGTGTTGCTATTTGTTAGAGAA	56.0
Sp-1167-S	ATGACCATGATTACGCCAAGCTTAAAGTAAAAAGAGCCAACACTTG	55.0
Sp-950-S	ATGACCATGATTACGCCAAGCTTCCACATCCGTCCATTAGTGC	57.0
Sp-646-S	ATGACCATGATTACGCCAAGCTTGGCAACATAATCATTAAGATACTGT	57.0
Sp-291-S	ATGACCATGATTACGCCAAGCTTGCTGATGGATAAGTAAAGAATAA	55.0
Sp-A	ACTGACCACCCGGGGATCCTTTTTTAATTAATTATAAACTTCAATG	59.0
pBI121-S	CGGCTCGTATGTTGTGTGGAATTG	60.0
pBI121-A	CGTTGGGGTTTCTACAGGACGTAA	58.0
Kana-S	TGGATTGCACGCAGGTTCTC	58.0
Kana-A	CTCGATGCGATGTTTCGCTT	58.0

图1 ApSK3启动子的序列及顺式作用元件分布起始密码子“ATG”的“A”被指定为“+ 1”。红色标注的为逆境胁迫及激素相关的元件,水平箭头表示方向。序列上方的垂直箭头表示不同缺失片段的起始点;蓝色核序列代表用于扩增缺失片段的特殊引物（Sp-S）。

Fig. 1 DNA sequence analysis of ApSK3 promoter and the description of cis-elements The“A”of the translation initiation code“ATG”of ApSK3 is designated as“+ 1”. These important cis-elements related with stress and hormone are red markerd. The horizontal arrows show their directions. The vertical arrows above the sequence indicate the start point of different deletion fragments;the blue nucleotide sequences represent special primers for amplifying deletion fragments（Sp-S）.

表2 ApSK3启动子中的顺式作用元件预测结果及功能列表

Table 2 Identification of cis-acting elements and functions in the ApSK3 promoter

类别	名称	描述	核心序列	数量	位置/bp
Category	Name	Description	Core Sequence	Number	Position
结构元件 Structure elements	TATA-Box	转录起始-30核心启动子元件 Core promoter element around-30 of transcription start	TATATA (5)	16	123 ~-128,-1314 ~-1319, -1704 ~-1709, -1928 ~-1933, -1973 ~-1978
			TATAAA (1)		-595 ~-600
			TATA (7)		36 ~-39,-192~-195,-478 ~-481,-507 ~-510,-1208 ~-1211,-1753 ~-1756, -1778 ~-1781
			TTTATA (3)		-15 ~-20,-1385 ~ -1390,-1459 ~-1464
	CAAT-Box	启动子和增强子区域调控元件 Common cis-acting element in promoter and enhancer regions	CAAT (9)	15	-89 ~-92,-822 ~-825, -991 ~-994,-1002 ~ -1005,-1661 ~-1664, -1713 ~-1716,-1840 ~ -1843,-1965 ~-1968, -2010 ~-2013
	CAAT-Box		CAAAT (6)	15	-340 ~-344,-492 ~ -496,-1359 ~-1363, -1367 ~-1370,-1558 ~ -1561,-2068 ~-2072
胁迫响应元件Abiotic stress responsive elements	ARE	厌氧顺式调控元件 cis-Acting regulatory element essential for the anaerobic induction	AACCA	2	-584 ~-588, -1534 ~-1538
	MBS	干旱诱导MYB结合位点 MYB binding site involved in drought-inducibility	CAACTG CAACAG	2	-1512 ~-1517, -319 ~-324
	circadian	昼夜顺式调控元件 Circadian-control element	CAAAGTTATC	1	-1628 ~-1637
	TC-rich repeats	防卫和胁迫响应顺式调控元件 cis-Acting element involved in defense and stress responsiveness	GTTTTCTAA ATTCTCTAAC	2	-1127 ~-1135 -2159 ~-2168
激素响应元件Phytohor- mone responsive element	ABRE	脱落酸响应顺式调控元件 Abscisic acid responsive element	CACGTG	2	-977 ~-982, -881 ~-886
	CGTCA-motif	茉莉酸甲酯响应顺式调控元件 MeJA-responsive element	CGTCA	2	-165 ~-169, -1370 ~-1374
	ERE	乙烯响应顺式调控元件 Ethylene-responsive element	ATTTCAAA	1	-526 ~-533
	P-box	赤霉素响应顺式调控元件 Gibberellin-responsive element	CCTTTTG	1	-561 ~-567
	TGA-box	生长素响应顺式调控元件 Auxin-responsive element	CGGTGCAGT	1	-170 ~-178
生长发育调控元件 Development-related elements	Skn-1 motif	胚乳表达调控顺式作用元件 cis-Acting element involved in endosperm expression	GTCAT	5	-164 ~-168,-1005 ~-1009,-1785 ~-1789, -1956 ~-1960, -2023 ~-2027
生长发育调控元件 Development-related elements	GCN4 motif	胚乳表达调控顺式作用元件 cis-Acting element involved in endosperm expression	CAAGCCA	1	-1650 ~-1656

表2 ApSK3启动子中的顺式作用元件预测结果及功能列表

Table 2 Identification of cis-acting elements and functions in the ApSK3 promoter

类别	名称	描述	核心序列	数量	位置/bp
Category	Name	Description	Core Sequence	Number	Position
结构元件 Structure elements	TATA-Box	转录起始-30核心启动子元件 Core promoter element around-30 of transcription start	TATATA (5)	16	123 ~-128,-1314 ~-1319, -1704 ~-1709, -1928 ~-1933, -1973 ~-1978
			TATAAA (1)		-595 ~-600
			TATA (7)		36 ~-39,-192~-195,-478 ~-481,-507 ~-510,-1208 ~-1211,-1753 ~-1756, -1778 ~-1781
			TTTATA (3)		-15 ~-20,-1385 ~ -1390,-1459 ~-1464
	CAAT-Box	启动子和增强子区域调控元件 Common cis-acting element in promoter and enhancer regions	CAAT (9)	15	-89 ~-92,-822 ~-825, -991 ~-994,-1002 ~ -1005,-1661 ~-1664, -1713 ~-1716,-1840 ~ -1843,-1965 ~-1968, -2010 ~-2013
	CAAT-Box		CAAAT (6)	15	-340 ~-344,-492 ~ -496,-1359 ~-1363, -1367 ~-1370,-1558 ~ -1561,-2068 ~-2072
胁迫响应元件Abiotic stress responsive elements	ARE	厌氧顺式调控元件 cis-Acting regulatory element essential for the anaerobic induction	AACCA	2	-584 ~-588, -1534 ~-1538
	MBS	干旱诱导MYB结合位点 MYB binding site involved in drought-inducibility	CAACTG CAACAG	2	-1512 ~-1517, -319 ~-324
	circadian	昼夜顺式调控元件 Circadian-control element	CAAAGTTATC	1	-1628 ~-1637
	TC-rich repeats	防卫和胁迫响应顺式调控元件 cis-Acting element involved in defense and stress responsiveness	GTTTTCTAA ATTCTCTAAC	2	-1127 ~-1135 -2159 ~-2168
激素响应元件Phytohor- mone responsive element	ABRE	脱落酸响应顺式调控元件 Abscisic acid responsive element	CACGTG	2	-977 ~-982, -881 ~-886
	CGTCA-motif	茉莉酸甲酯响应顺式调控元件 MeJA-responsive element	CGTCA	2	-165 ~-169, -1370 ~-1374
	ERE	乙烯响应顺式调控元件 Ethylene-responsive element	ATTTCAAA	1	-526 ~-533
	P-box	赤霉素响应顺式调控元件 Gibberellin-responsive element	CCTTTTG	1	-561 ~-567
	TGA-box	生长素响应顺式调控元件 Auxin-responsive element	CGGTGCAGT	1	-170 ~-178
生长发育调控元件 Development-related elements	Skn-1 motif	胚乳表达调控顺式作用元件 cis-Acting element involved in endosperm expression	GTCAT	5	-164 ~-168,-1005 ~-1009,-1785 ~-1789, -1956 ~-1960, -2023 ~-2027
生长发育调控元件 Development-related elements	GCN4 motif	胚乳表达调控顺式作用元件 cis-Acting element involved in endosperm expression	CAAGCCA	1	-1650 ~-1656

图2 百子莲不同组织中脱水素基因ApSK3的表达量虚线表示Ap-actin（内参）表达水平。

Fig. 2 Expression level of ApSK3 in different tissues of Agapanthus praecox The dotted line indicated the expression level of Ap-actin（internal reference gene）.

图3 ApSK3 -P::GUS在转基因拟南芥不同组织中的GUS组织化学染色

Fig. 3 GUS histochemical staining of different tissues of ApSK3-P::GUS transgenic Arabidopsis Bar = 1 cm.

图4 百子莲在不同胁迫及ABA处理条件下根和叶片中ApSK3的表达分析 Ap-actin为内参基因。大写和小写字母分别表示在相同处理下,根和叶中不同时刻基因表达量间的差异显著性（P < 0.05）。

Fig. 4 Quantitative real-time PCR analysis of ApSK3 gene transcripts in Agapanthus praecox roots and leaves in response to various abiotic stresses and ABA treatment Ap-actin was used as internal reference gene. Values with different uppercase and lowercase letters are significantly different among samples in the roots and leaves,respectively.（P < 0.05）.

图5 不同胁迫、ABA、GA和乙烯处理后ApSK3-P转基因及空载拟南芥（35S启动子）的GUS活性分析 * 表示不同植物生长调节剂、胁迫处理组与对照间的差异显著（P < 0.05,α = 0.05）

Fig. 5 GUS activity analysis of transgenic Arabidopsis seedlings containing ApSK3-P or 35S promoter under abiotic,ABA,GA and Ethylene treatments Asterisks（*）indicate significant differences between ApSK3-P under different treatments and control.（P < 0.05,α = 0.05）. Ap-actin was used as internal reference gene. Values with different uppercase and lowercase letters are significantly different among samples in the roots and leaves,respectively.

图6 ApSK3启动子及其缺失片段的载体构建

Fig. 6 Construction of vector of ApSK3 promoter and its deletion fragments

图7 转ApSK3启动子不同5'缺失片段的拟南芥在非生物胁迫（A）及外源ABA、GA、乙烯（B）处理下的GUS活性不同大写字母代表不同启动子缺失片段在相同处理下的差异显著（P < 0.05）;不同小写字母代表相同启动子缺失片段在不同处理下的差异显著（P < 0.05）;柱上数字代表相同启动子缺失片段在不同处理下与对照的比值。

Fig. 7 GUS activity of transgenic Arabidopsis seedlings containing ApSK3 promoter and its 5' deletion constructs under abiotic（A）and ABA,GA,Ethylene（B）treatments Capital letters represent the significance of difference between the differernt ApSK3 promoters in the same treated condition;lower-case letters represent the significance among the same ApSK3 promoter under different treatments（P < 0.05,least significant difference test）;numbers represent the fold of the same ApSK3 promoter under different treatments/Control.

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百子莲脱水素基因ApSK₃对逆境与激素信号的应答模式与调控机制

Expression Pattern and Regulation Mechanism of ApSK₃ Dehydrin （Agapanthus praecox）Response to Abiotic Stress and Hormone Signals

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