多杀性巴氏杆菌荚膜的生物合成及其调控机制研究进展

doi:10.3864/j.issn.0578-1752.2020.03.016

Abstract

Abstract:

Pasteurella multocida can be widely infected with a variety of animals, causing hemorrhagic septicemia or infectious pneumonia. P. multocida possess a viscous capsular polysaccharide on the cell surface, which is a critical structural component and virulence factor, and plays an important role in the interaction between bacteria and the host, promoting the adhesion of bacteria to the host surface and enhancing the virulence of the bacteria. The molecular structure of the P. multocida capsule is similar to that of vertebrate glycosaminoglycan, which is polymerized by repeated disaccharide units to form a linear polysaccharide chain, which is an important immunological material basis for molecular mimicry, resistance to phagocytosis, and immune evasion during the infection of the host. In recent years, a series of important research advances have been made in the biosynthesis and regulation mechanism aspects of P. multocida capsule, providing a certain basic knowledge for the molecular pathogenesis of P. multocida capsule, and supplying a theoretical basis for the development of the capsular polysaccharide vaccine of P. multocida. This paper systematically illuminates the biosynthesis and expression regulation mechanisms of P. multocida capsule, including the serotyping of the capsule, the composition and structure of the capsular polysaccharide, the biosynthesis gene cluster and function of the capsule, the molecular synthesis mechanism of capsular polysaccharide, the expression regulation mechanism of capsular biosynthesis gene cluster, a total of five aspects. According to the capsular antigen, P. multocida is divided into five capsular serogroups of A, B, D, E, and F. The type A capsule GAG component is hyaluronic acid; the type D is heparosan; the type F is chondroitin, which is repeatedly composed of its corresponding disaccharide unit [β-GlcUA/β-GlcNAc], [β-GlcUA/α-GlcNAc], [β-GlcUA/β-GalNAc], respectively; the type B capsular polysaccharide is composed of arabinose, mannose and galactose in a certain structural form, and the composition and chemical structure of type E capsular polysaccharide are uncertain. Genes related to the biosynthesis of A, B, D, E and F capsules of P. multocida exist in the form of gene clusters and are divided into three distinct functional regions, R1, R2 and R3; the R1 region is responsible for transporting the capsular polysaccharide, the R2 region is responsible for the activation of the monosaccharide and the assembly of the capsular polysaccharide, and the R3 region is responsible for the modification of capsular polysaccharide (phospholipid replacement); according to the structure and the number of genes of the R2 region, the biosynthetic gene clusters of the five capsules can be divided into two categories: type A, D and F are Class I, and R2 contains 4 genes; types B and E are Class II, and R2 contains 9 genes, and using the specific gene in the R2 region to design primers, the capsular serotype of P. multocida can be rapidly identified by PCR. The capsular GAG of P. multocida is synthesized in the cytoplasm, and then exported to the cell surface via the ABC transporter formed by the protein encoded by the R1 region, and tightly bound to the cell surface by covalent attachment to the phospholipid; during the biosynthesis of the P. multocida capsular GAG, the glycosyltransferase gene located in the R2 region determines the type of activated monosaccharide and the type of capsular polysaccharide after assembly. In the biosynthetic gene cluster of the P. multocida capsule, the R1 and R2 regions form an operon with the same transcriptional direction, while the R3 transcription direction is opposite, and the promoter regions of both are located on the DNA sequence between the R2 and R3 regions; the transcriptional process of the P. multocida capsular biosynthesis gene cluster is positive regulated by the Fis protein, and the translation process is mainly positive regulated by Hfq protein.

Key words: Pasteurella multocida, capsule, glycosaminoglycan, biosynthesis, expression regulation

GUAN LiJun,XUE Yun,DING WenWen,ZHAO ZhanQin. Advances in Mechanisms of Biosynthesis and Regulation of Pasteurella multocida Capsule[J].Scientia Agricultura Sinica, 2020, 53(3): 658-668.

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References 79

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功能区 Region	开放阅读框（ORF）			A型/D型/F型编码的蛋白质 Protein encoded by type A/D/F	功能 Function
功能区 Region	A型 Type A	D型 Type B	F型 Type D	A型/D型/F型编码的蛋白质 Protein encoded by type A/D/F	功能 Function
R1	hexA	hexA	hexA	HexA	ATP结合蛋白 ATP-binding protein
	hexB	hexB	hexB	HexB	内膜蛋白 Inner membrane protein
	hexC	hexC	hexC	HexC	具有周质结构域的内膜蛋白 Inner membrane protein with a periplasmic domain
	hexD	hexD	hexD	HexD	外膜蛋白（脂蛋白） Outer membrane protein (lipoprotein)
R2	hyaB	dcbB	fcbB	HyaB/DcbB/FcbB	糖基转移酶 Glycosyltransferase
	hyaC	dcbC	fcbC	HyaC/DcbC/FcbC	UDP-葡萄糖脱氢酶,催化UDP-葡萄糖转化为UDP-葡糖醛酸 UDP-glucose dehydrogenase, catalyzing the conversion of UDP-glucose to UDP-glucuronic acid
	hyaD	dcbF	fcbD	HyaD/DcbF/FcbD	透明质酸合成酶/肝素合成酶/软骨素合成酶 PmHAS/PmHS/PmCS
	hyaE	dcbE	fcbE	HyaE/ DcbE / FcbE	起始酶,将重复单元的第一个残基转移至 Poly-KDO linker Initiates enzyme, which transfers from the first residue of the repeat-unit to the poly-KDO linker
R3	phyA	phyA	phyA	PhyA	β-KDO糖基转移酶 β-Kdo glycosyltransferase
R3	phyB	phyB	phyB	PhyB	β-KDO糖基转移酶 β-Kdo glycosyltransferase

功能区 Region	开放阅读框（ORF）		B型/E型编码的蛋白质 Protein encoded by type B/E	功能 Function
功能区 Region	B型 Type B	E型 Type F	B型/E型编码的蛋白质 Protein encoded by type B/E	功能 Function
R1	cexA	cexA	CexA	ATP结合蛋白 ATP-binding protein
	cexB	cexB	CexB	内膜蛋白 Inner membrane protein
	cexC	cexC	CexC	具有周质结构域的内膜蛋白 Inner membrane protein with a periplasmic domain
	cexD	cexD	CexD	外膜蛋白 Outer membrane protein
R2	bcbA	ecbA	BcbA/EcbA	UDP-N-乙酰葡糖胺2-差向异构酶 UDP-N-acetylglucosamine 2-epimerase
	bcbB	ecbB	BcbB/EcbB	UDP-N-乙酰-D-甘露糖胺糖醛酸脱氢酶 UDP-N-acetyl-D-mannosaminuronic acid dehydrogenase
	bcbC	ecbK	BcbC/EcbK	假定的糖基转移酶,可能参与活化单糖的聚合 Putative glycosyltransferase, which likely involved in polymerization of the activated sugar
	bcbDEFGI	ecbDEFGI	BcbDEFGI/EcbDEFGI	未知 Unknown
	bcbH	ecbJ	BcbH/EcbJ	未知,是B型或E型的独有基因 Unknown, unique to serogroup B or E
R3	lipA	lipA	LipA	参与多糖的磷脂替换 Involved in phospholipid substitution of GAG
R3	lipB	lipB	LipB	参与多糖的磷脂替换 Involved in phospholipid substitution of GAG

Advances in Mechanisms of Biosynthesis and Regulation of Pasteurella multocida Capsule

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