Epsilon-变形菌趋化信号通路中偶联蛋白CheV与趋化受体蛋白的CZB结构域的功能研究

doi:10.11978/2020055

热带海洋学报 ›› 2021, Vol. 40 ›› Issue (2): 27-38.doi: 10.11978/2020055CSTR: 32234.14.2020055

Epsilon-变形菌趋化信号通路中偶联蛋白CheV与趋化受体蛋白的CZB结构域的功能研究

刘煜耿^1,²(), 毛颖津^1,², 章灿川^1,², 高贝乐¹()

1.中国科学院热带海洋生物资源与生态重点实验室, 广东省海洋药物重点实验室, 中国科学院南海海洋研究所, 广东广州 510301
2.中国科学院大学, 北京 100049

收稿日期:2020-05-26 修回日期:2020-06-29 出版日期:2021-03-10 发布日期:2020-07-29
通讯作者: 高贝乐
作者简介:刘煜耿(1994—), 男, 广东省揭阳市人, 硕士研究生, 研究方向为微生物功能基因组学。email: yugengliu@163.com
基金资助:
国家自然科学基金(31870064)

Functional study of coupling protein CheV and CZB domain of chemoreceptors in the Epsilon-proteobacteria chemotaxis signaling pathway

LIU Yugeng^1,²(), MAO Yingjin^1,², ZHANG Canchuan^1,², GAO Beile¹()

1. CAS Key Laboratory of Tropical Marine Bio-resources and Ecology, Guangdong Key Laboratory of Marine Materia Medica, South China Sea Institute of Oceanology, Chinese Academy of Sciences, Guangzhou 510301, China
2. University of Chinese Academy of Sciences, Beijing 100049, China

Received:2020-05-26 Revised:2020-06-29 Online:2021-03-10 Published:2020-07-29
Contact: GAO Beile
Supported by:
National Science Foundation of China(31870064)

摘要/Abstract

摘要：

细菌通过趋化系统感知和响应外界环境变化并进行趋利避害, 因此很多细菌进化出了非常复杂多样的趋化系统以适应不同的生态位。Epsilon-变形菌广泛存在于自然界中, 能适应不同的生态环境, 特别是深海Epsilon-变形菌能适应深海热液口和冷泉等极端环境, 其趋化系统可能有其独特之处。通过BlastP和MIST数据库分析, 我们发现大部分深海Epsilon-变形菌拥有F3类型趋化系统且具有单拷贝的双功能域融合蛋白CheV。此外, 一种特殊结构域CZB (C-terminal Zinc-Binding) 存在于深海Epsilon-变形菌的趋化受体蛋白中, 而且在部分深海Epsilon-变形菌的趋化受体中还发现并鉴定了一种新的结构域CZB-like结构域。文章以Epsilon-变形菌的模式菌株空肠弯曲杆菌(Campylobacter jejuni 81-176)为研究对象, 用细菌双杂交试验证实了CheV能与所有含MA结构域的趋化受体相互作用。用电感耦合等离子体质谱(ICP-Mass)技术证实了CZB-like结构域不能结合Zn离子, 但细菌双杂交试验证实它能介导趋化受体Tlp9与CheV的互作。

关键词: Epsilon-变形菌, 趋化系统, 融合蛋白CheV, CZB结构域

Abstract:

Epsilon-proteobacteria is widely distributed, from deep-sea hydrothermal vent to surface sea water, from free-living environment to host-associated one. Chemotaxis plays an important role in bacteria survival through sensing and responding to environmental changes. Thus, some bacteria have evolved into many complex and diverse chemotaxis systems. Epsilon-proteobacteria can adapt to different environments; especially, some species can survive in the deep-sea extreme environments including hydrothermal vent and cold seep, whose chemotaxis system may have special characteristics. Bioinformatics analyses by BlastP and MIST database revealed that most Epsilon-proteobacteria species in the deep-sea hydrothermal vent have F3 type chemotaxis system. They all contain a single copy of CheV, which is a double domain fusion protein. Besides, a unique domain, CZB (C-terminal Zinc-Binding) domain, exists in chemoreceptors of deep-sea Epsilon- proteobacteria. A CZB-like domain is also identified in Epsilon-proteobacteria. Using the model strain Campylobacter jejuni 81-176, we confirmed that CheV can interact with all chemoreceptors with MA domain by bacterial two-hybrid experiments. Additionally, we demonstrated that CZB-like domain cannot bind Zn by ICP-Mass, but it can promote the interaction between chemoreceptor Tlp9 and CheV.

Key words: Epsilon-proteobacteria, chemotaxis, CheV, CZB domain

中图分类号:

P735.4

刘煜耿, 毛颖津, 章灿川, 高贝乐. Epsilon-变形菌趋化信号通路中偶联蛋白CheV与趋化受体蛋白的CZB结构域的功能研究[J]. 热带海洋学报, 2021, 40(2): 27-38.

LIU Yugeng, MAO Yingjin, ZHANG Canchuan, GAO Beile. Functional study of coupling protein CheV and CZB domain of chemoreceptors in the Epsilon-proteobacteria chemotaxis signaling pathway[J]. Journal of Tropical Oceanography, 2021, 40(2): 27-38.

图/表 11

图1

表1

表2

表3

表4

引物序列"

引物名称	序列(5’→ 3’)
pKNT25+cheV_F	CTCTAGAGGATCCCCGGGTAATGTTTGATGAAAATATCGT
pKNT25+cheV_R	GTCATTGAATTCGAGCTCGGTTACCCCTGTTCTTGAGATT
pKNT25+cheW_F	CAGGTCGACTCTAGAGGATCCCCGGGTAATGAGTAATGAAAAATTAGAGCAAATTTTGC
pKNT25+cheW_R	CTGCATGGTCATTGAATTCGAGCTCGGAAATTCGCGCTTAAGTAAAGCTTCTACTTTG
pUT18C+tlp1_F	ACTGCAGGTCGACTCTAGATGAAATTAAAAAGATGCTTTTGGCTT
pUT18C+tlp1_R	ATATCGATGAATTGCTCGAGTTAAAATCTTTTTTTACTCACATCTTCAAG
pUT18C+tlp4_F	ACGCCACTGCAGGTCGACTCTAGATCTCTCCCCACTTGCAGCTATCCAAACAGGT
pUT18C+tlp4_R	TAGTTATATCGATGAATTGCTCGAGTTAAAACCTTTTCTTCTTAACATCTTCTAA
pUT18C+tlp6_F	ACGCCACTGCAGGTCGACTCTAGATATGTTTGGAAGTAAAATAAACCATTCTGAT
pUT18C+tlp6_R	TAGTTATATCGATGAATTGCTCGAGTTAATGATCTGACTCATCAAGCATTTCTTT
pUT18C+tlp7_F	ACTGCAGGTCGACTCTAGATCAATTTATCGAAAAAACTCATAAGGCAGTT
pUT18C+tlp7_R	ATATCGATGAATTGCTCGAGTTAAATTTGAAATTGGTTAAGTTCGCTTTC
pUT18C+tlp8_F	ACGCCACTGCAGGTCGACTCTAGATATGTTTGGTGCTAAGAAAAATAATACTGAA
pUT18C+tlp8_R	TAGTTATATCGATGAATTGCTCGAGTTATGACATCGCTTTAGCAACTTCAGCAGAGCT
pUT18C+tlp9_F	ACGCCACTGCAGGTCGACTCTAGATTTAGCAAACATAGAAGTGACAGCAAGATCT
pUT18C+tlp9_R	TAGTTATATCGATGAATTGCTCGAGTTATATTTTTAATTTTGCTAAGATTTCAGC
pUT18C+tlp10_F	ACTGCAGGTCGACTCTAGATTCAAGTCATCAAAATTCACAAAAACTCAA
pUT18C+tlp10_R	ATATCGATGAATTGCTCGAGTTACTGAAAGCTACTTAATTTTTCGGAGAG
pUT18C+aer1_F	ACTGCAGGTCGACTCTAGATAAAGAAATAGTTTTGTCTGAAAATGCTTTA
pUT18C+aer1_R	ATATCGATGAATTGCTCGAGTTAATTATTTTCTTGTAAGTTAAAAATAAGTTCAT
pUT18C+aer2_F	ACTGCAGGTCGACTCTAGATTCAAGAGAAATTTTTTTACAAGAAGATAGT
pUT18C+aer2_R	ATATCGATGAATTGCTCGAGTTATTTAGCTTCTTGAAGAGAAAAGATTAGC
pUT18C+tlp9 (no CZB-like)_F	ACTGCAGGTCGACTCTAGATTTAGCAAACATAGAAGTGACAGCAAGATCT
pUT18C+tlp9 (no CZB-like)_R	ATATCGATGAATTGCTCGAGTTAATAAAGAATATGATCGATTTTAACCAC
pCH363+tlp9_F	ACAGCTATGACCATGATTACGCCATTAGCAAACATAGAAGTGACAGCAAGATCT
pCH363+tlp9_R	TGGCCTCGCTGGCGGCTGAATTCGATATTTTTAATTTTGCTAAGATTTCAGCACT
pGEX-6P+tlp6_F	TCTGTTCCAGGGGCCCCTGGGATCCTTGCTTAGACAACACAAAGATGAGC
pGEX-6P+tlp6_R	GCTCGAGTCGACCCGGGAATTCCGGTTATATTTTTAATTTTGCTAAGATTTCAGCAC
pGEX-6P+tlp9_F	TCTGTTCCAGGGGCCCCTGGGATCCATGTTTGGAAGTAAAATAAACCATTCTGATCTTC
pGEX-6P+tlp9_R	GCTCGAGTCGACCCGGGAATTCCGGTTAATGATCTGACTCATCAAGCATTTC
pGEX-6P+tlp9 (no CZB-like)_F	TCTGTTCCAGGGGCCCCTGGGATCCATGTTTGGAAGTAAAATAAACCATTCTGATCTTC
pGEX-6P+tlp9 (no CZB-like)_R	GCTCGAGTCGACCCGGGAATTCCGGTTAATCGATTTTAACCACAGACAAAATCAATC

表4

图2

表5

图3

图4

表6

图5

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物种名称	菌株	生物样品序列号	生物项目序列号	基因组编号	纯培养	呼吸类型	参考文献
Caminibacter mediatlanticus	TB-2	SAMN10884411	PRJNA521379	GCA_006459125.1	是	厌氧	Giovannelli 等(2011)
Cetia pacifica	TB6	SAMN03737945	PRJNA284962	GCA_003346815.1	是	厌氧	Grosche等(2015)
Hydrogenimonas sp.	MAG	SAMN00210774	PRJNA543187	GCA_005886055.1	否	兼性厌氧	Takai等(2004)
Lebetimonas sp.^#	JS138	SAMN03737948	PRJNA66817	GCA_003660105.1	否	厌氧	Meyer等(2014)
Nautilia profundicola	AmH	SAMN03737951	PRJNA284966	GCA_003544915.1	是	厌氧	Smith等(2008)
Nautilia sp.	PV-1	SAMN03737952	PRJNA284967	GCA_003346755.1	是	厌氧	Smith等(2008)
Nitratiruptor sp.	SB155-2	SAMN03737954	PRJNA66821	GCA_003544935.1	是	兼性厌氧	Nakagawa等(2005)
Sulfurimonas autotrophica	DSM 16294	SAMN03737955	PRJNA66823	GCA_003346775.1	是	好氧	Inagaki等(2003)
Campylobacter jejuni^*	81-176	SAMEA4030738	PRJEB6403	GCA_900637395.1	是	微好氧	Snelling等(2005)

质粒名称	质粒描述
pGEX-6P	用于构建GST融合蛋白表达载体
pKNT25	细菌双杂交T25融合蛋白表达载体
pUT18C	细菌双杂交T18(在N端)融合蛋白表达载体
pCH363	细菌双杂交T18(在C端)融合蛋白表达载体

基因	基因编号	蛋白结构域示意图
cheV	CJJ81176_0311
cheW	CJJ81176_0309
tlp1	CJJ81176_1498
tlp2	CJJ81176_0180
tlp3^#	CJJ81176_1548
tlp3^#	CJJ81176_1549
tlp4	CJJ81176_0289
tlp5^*	CJJ81176_0271-4	Pseudogene
tlp6	CJJ81176_0473
tlp7	CJJ81176_0975
tlp8	CJJ81176_1128
tlp9	CJJ81176_1205
tlp10	CJJ81176_0046
aer1	CJJ81176_1204
aer2	CJJ81176_1206

菌株	F类型	CheV蛋白编号	菌株	F类型	CheV蛋白编号
Campylobacter jejuni^*	1 (F3)	WP_002857365.1	Nautilia profundicola	1 (F3)	WP_041361491.1
Caminibacter mediatlanticus	1 (F3)	WP_007474092.1	Nautilia sp. PV-1	1 (F3)	WP_127679461.1
Cetia pacifica	1 (F3)	WP_123351600.1	Nitratiruptor sp. SB155-2	1 (F14)	WP_012081979.1
Hydrogenimonas sp. MAG	1 (F3)	BBG65199.1	Sulfurimonas autotrophica	2 (F3/F8)	WP_013326429.1 WP_013327173.1^#
Lebetimonas sp. JS138	1 (F3)	WP_024787070.1	Sulfurimonas autotrophica	2 (F3/F8)	WP_013326429.1 WP_013327173.1^#

菌株	CZB个数	含CZB的蛋白编号	CZB-like个数	含CZB-like的蛋白编号
Campylobacter jejuni^*	1	WP_002857440.1	1	WP_002790076.1
Caminibacter mediatlanticus	2	WP_138323537.1, WP_138323100.1	1	WP_138323156.1
Cetia pacifica	2	WP_123352714.1, WP_123352123.1	1	WP_123351565.1
Hydrogenimonas sp. MAG	5	BBG65387.1, BBG65955.1, BBG65459.1, BBG65926.1, BBG65424.1	0	0
Lebetimonas sp. JS138	1	WP_024791915.1	0	0
Nautilia profundicola	2	WP_012663710.1, WP_015902667.1	0	0
Nautilia sp. PV-1	2	WP_127679641.1, WP_127679872.1	0	0
Nitratiruptor sp. SB155-2	2	WP_012081746.1, WP_012081977.1	0	0
Sulfurimonas autotrophica	3	WP_013327874.1, WP_041675273.1, WP_013327332.1	0	0

Epsilon-变形菌趋化信号通路中偶联蛋白CheV与趋化受体蛋白的CZB结构域的功能研究

Functional study of coupling protein CheV and CZB domain of chemoreceptors in the Epsilon-proteobacteria chemotaxis signaling pathway

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