植物中C-糖基转移酶的结构与应用

doi:10.13560/j.cnki.biotech.bull.1985.2021-1572

摘要/Abstract

摘要：

糖基转移酶是一类专门催化糖基化反应的后修饰酶，根据与糖供体连接的受体分子不同，分为O-、C-、N-、S-四种糖基转移酶。近年来，伴随着结构生物学的不断发展，关于C-糖基转移酶的结构解析及定向改造的报道日益增多，为复杂天然产物修饰提供理论依据。本文综述了植物C-糖基转移酶的研究进展，从其底物杂泛性、糖供体结合的空间结构与生物技术应用的角度进行讨论，探究造成底物催化特征的结构基础，为后续通过合成生物学手段异源合成植物天然产物提供参考。

关键词: C-糖基转移酶, 生物合成途径, 催化机制, 晶体结构, 合成生物学

Abstract:

Glycosyltransferases are a class of post-modification enzymes that specifically catalyse glycosylation reactions and are classified into 4 types of glycosyltransferases，O-，C-，N- and S-，depending on the acceptor molecule attached to the sugar donor. In recent years，along with the vigorous development of structural biology，reports on the structural elucidation and targeted modification of C-glycosyltransferases have gradually increased，providing theoretical guidance for the modification of complex natural products. This paper reviews the research progress of plant C-glycosyltransferases，discusses them from the perspectives of their substrate specificity，spatial structure of sugar donor binding and biotechnological applications，and explores the structural basis of the substrate catalytic characteristics，which may provide a reference for the subsequent heterologous synthesis of plant natural products by means of synthetic biology.

Key words: C-glycosyltransferase, biosynthetic pathway, catalytic mechanism, crystal structure, synthetic biology

赵玉雪, 王芸, 余璐瑶, 刘京晶, 斯金平, 张新凤, 张磊. 植物中C-糖基转移酶的结构与应用[J]. 生物技术通报, 2022, 38(10): 18-28.

ZHAO Yu-xue, WANG Yun, YU Lu-yao, LIU Jing-jing, SI Jin-ping, ZHANG Xin-feng, ZHANG Lei. Structure and Application of C-glycosyltransferases in Plants[J]. Biotechnology Bulletin, 2022, 38(10): 18-28.

图/表 6

参考文献 66

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序号 Number	基因名称 Gene name	基因ID Gene ID	植物来源 Species	糖受体 Sugar receptor	糖供体 Sugar donor	参考文献 Reference
1	OsCGT	CAQ77160	水稻Oryza sativa	黄酮	UDP-Glc UDP-Ara	[35]
2	UGT708A6	NP_001132650.1	玉米Zea mays	黄酮二氢黄酮	UDP-Glc	[36]
3	FeCGTa（UGT708C1）	BAP90360	荞麦Fagopyrum esculentum	二氢查耳酮二氢黄酮	UDP-Glc UDP-Xyl	[37]
4	FeCGTb（UGT708C2）	BAP90361	荞麦F. esculentum	二氢查耳酮二氢黄酮	UDP-Glc UDP-Xyl	[37]
5	UGT708D1	BAR73279	大豆Glycine max	黄烷酮黄酮	UDP-Glc	[38]
6	GtUF6CGT1	BAQ19550	三花龙胆Gentiana triflora	黄酮	UDP-Glc	[39]
7	MiCGT	ALD83754	芒果Mangifera indica	苯丙素	UDP-Glc	[40]
8	MiCGTb	AMM73095	芒果M. indica	苯丙素	UDP-Glc	[40]
9	PlUGT43	A0A172J2G3.2	葛根Pueraria lobata	异黄酮	UDP-Glc	[41]
10	FcCGT（UGT708G1）	BBA18062	金柑Fortunella crassifolia	二氢查耳酮二氢黄酮	UDP-Glc UDP-Xyl UDP-Gal	[42]
11	CuCGT（UGT708G2）	BBA18063	温州蜜柑Citrus unshiu	二氢查耳酮二氢黄酮	UDP-Glc UDP-Xyl UDP-Gal	[42]
12	TcCGT1	MK644229	中国金莲花Trollius chinensis	黄酮黄酮醇二氢查尔酮	UDP-Xyl UDP-Gal UDP-Ara	[43]
13	WjGT1	LC465149	块茎山萮菜Eutrema japonicum	黄酮黄酮醇二氢查耳酮	UDP-Glc	[44]
14	GgCGT	MH998596	光果甘草Glycyrrhiza glabra	二氢查耳酮异黄酮	UDP-Glc UDP-Xyl UDP-Gal UDP-Ara	[45]
15	OsUGT708A4	Q5VMG8	水稻O. sativa	二氢黄酮二氢查耳酮	UDP-Glc	[46]
16	OsUGT708A2	Q5VME5	水稻O. sativa	二氢黄酮二氢查耳酮	UDP-Glc	[46]
17	OsUGT708A1	A2YBW7	水稻O. sativa	二氢黄酮二氢查耳酮	UDP-Glc UDP-Ara	[46]
18	OsUGT708A39	A2YBW8	水稻O. sativa	二氢黄酮二氢查耳酮	UDP-Glc UDP-Ara	[46]
19	OsUGT708A40	A2YBX1	水稻O. sativa	二氢黄酮二氢查耳酮	UDP-Glc UDP-Ara	[46]
20	ZmUGT708A5	B6SWX3	玉米Z. mays	二氢查耳酮	UDP-Glc UDP-Ara	[46]
21	ZmUGT708A11	A0A1D6LXE6	玉米Z. mays	二氢黄酮二氢查耳酮	UDP-Glc UDP-Ara	[46]
22	ZmUGT708A41	B4FAN3	玉米Z. mays	二氢黄酮二氢查耳酮	UDP-Glc	[46]
23	ZmUGT708A42	B4FM13	玉米Z. mays	二氢黄酮二氢查耳酮	UDP-Glc	[46]
24	SbUGT708A36	C5Z8Y8	高粱Sorghum bicolor	二氢黄酮二氢查耳酮	UDP-Glc UDP-Ara	[46]
25	SbUGT708A35	C5Z8Y5	高粱S. bicolor	二氢黄酮二氢查耳酮	UDP-Glc UDP-Ara	[46]
26	SbUGT708A34	C5Z8Y2	高粱S. bicolor	二氢黄酮二氢查耳酮	UDP-Glc UDP-Ara	[46]
27	SbUGT708A38	A0A194YGR9	高粱S. bicolor	二氢黄酮二氢查耳酮	UDP-Glc UDP-Ara	[46]
28	TaUGT708A14	A0A3B6RGT4	小麦Triticum aestivum	二氢黄酮二氢查耳酮	UDP-Glc UDP-Ara	[46]
29	TaUGT708A52	A0A3B6SEV6	小麦T. aestivum	二氢黄酮二氢查耳酮	UDP-Glc UDP-Ara	[46]
30	TaUGT708A53	A0A3B6TPG6	小麦T. aestivum	二氢黄酮二氢查耳酮	UDP-Glc UDP-Ara	[46]
31	TaUGT708A15	A0A3B6RGR9	小麦T. aestivum	二氢黄酮二氢查耳酮	UDP-Glc	[46]
32	TaUGT708A54	A0A3B6SB95	小麦T. aestivum	二氢黄酮二氢查耳酮	UDP-Glc	[46]
33	TaUGT708A55	A0A3B6TNL8	小麦T. aestivum	二氢黄酮	UDP-Glc	[46]
34	BdUGT708A7	I1GYZ6	二穗短柄草 Brachypodium distachyon	二氢黄酮二氢查耳酮	UDP-Glc	[46]
35	BdUGT708A8	I1GZD7	二穗短柄草B. distachyon	二氢黄酮二氢查耳酮	UDP-Glc UDP-Ara	[46]
36	SiUGT708A31	K3XWN4	小米Setaria italica	二氢黄酮二氢查耳酮	UDP-Glc UDP-Ara	[46]
37	SiUGT708A32	K3XWJ1	小米S. italica	二氢黄酮二氢查耳酮	UDP-Glc	[46]
38	SiUGT708A33	K3Y181	小米S. italica	二氢黄酮二氢查耳酮	UDP-Glc UDP-Ara	[46]
39	PhUGT708A43	MK616588	毛竹Phyllostachys heterocycla	二氢黄酮二氢查耳酮	UDP-Glc UDP-Ara	[46]
40	PhUGT708A46	MK616589	毛竹P. heterocycla	二氢黄酮二氢查耳酮	UDP-Glc	[46]
41	PhUGT708A48	MK616590	毛竹P. heterocycla	二氢黄酮二氢查耳酮	UDP-Glc UDP-Ara	[46]
42	PhUGT708A50	MK616591	毛竹P. heterocycla	二氢查耳酮	UDP-Glc UDP-Ara	[46]
43	PgUGT708A44	MK616592	淡竹P. heterocycla	二氢黄酮二氢查耳酮	UDP-Glc UDP-Ara	[46]
44	PpUGT708A45	MK616593	高节竹Phyllostachys prominens	二氢查耳酮	UDP-Glc	[46]
45	DcaCGT	MT452646	铁皮石斛Dendrobium catenatum	二氢黄酮二氢查耳酮	UDP-Glc	[47]
46	NnCGT1（UGT708N1）	LOC104598527	莲Nelumbo nucifera	黄酮	UDP-Glc	[48]
47	NnCGT2（UGT708N2）	LOC104603347	莲N. nucifera	黄酮	UDP-Ara UDP-Xyl	[48]
48	SbCGT	MK894443	黄芩Scutellaria baicalensis	黄酮	UDP-Glc UDP-Ara	[49]
49	LpCGT	MK894451	兰氏萍Landoltia punctata	黄酮	UDP-Glc UDP-Ara	[49]
50	AbCGT	MN747045	芦荟Aloe barbadensis	二氢查尔酮酚	UDP-Glc	[50]

序号 Number	基因名称 Gene name	基因ID Gene ID	植物来源 Species	糖受体 Sugar receptor	糖供体 Sugar donor	参考文献 Reference
1	OsCGT	CAQ77160	水稻Oryza sativa	黄酮	UDP-Glc UDP-Ara	[35]
2	UGT708A6	NP_001132650.1	玉米Zea mays	黄酮二氢黄酮	UDP-Glc	[36]
3	FeCGTa（UGT708C1）	BAP90360	荞麦Fagopyrum esculentum	二氢查耳酮二氢黄酮	UDP-Glc UDP-Xyl	[37]
4	FeCGTb（UGT708C2）	BAP90361	荞麦F. esculentum	二氢查耳酮二氢黄酮	UDP-Glc UDP-Xyl	[37]
5	UGT708D1	BAR73279	大豆Glycine max	黄烷酮黄酮	UDP-Glc	[38]
6	GtUF6CGT1	BAQ19550	三花龙胆Gentiana triflora	黄酮	UDP-Glc	[39]
7	MiCGT	ALD83754	芒果Mangifera indica	苯丙素	UDP-Glc	[40]
8	MiCGTb	AMM73095	芒果M. indica	苯丙素	UDP-Glc	[40]
9	PlUGT43	A0A172J2G3.2	葛根Pueraria lobata	异黄酮	UDP-Glc	[41]
10	FcCGT（UGT708G1）	BBA18062	金柑Fortunella crassifolia	二氢查耳酮二氢黄酮	UDP-Glc UDP-Xyl UDP-Gal	[42]
11	CuCGT（UGT708G2）	BBA18063	温州蜜柑Citrus unshiu	二氢查耳酮二氢黄酮	UDP-Glc UDP-Xyl UDP-Gal	[42]
12	TcCGT1	MK644229	中国金莲花Trollius chinensis	黄酮黄酮醇二氢查尔酮	UDP-Xyl UDP-Gal UDP-Ara	[43]
13	WjGT1	LC465149	块茎山萮菜Eutrema japonicum	黄酮黄酮醇二氢查耳酮	UDP-Glc	[44]
14	GgCGT	MH998596	光果甘草Glycyrrhiza glabra	二氢查耳酮异黄酮	UDP-Glc UDP-Xyl UDP-Gal UDP-Ara	[45]
15	OsUGT708A4	Q5VMG8	水稻O. sativa	二氢黄酮二氢查耳酮	UDP-Glc	[46]
16	OsUGT708A2	Q5VME5	水稻O. sativa	二氢黄酮二氢查耳酮	UDP-Glc	[46]
17	OsUGT708A1	A2YBW7	水稻O. sativa	二氢黄酮二氢查耳酮	UDP-Glc UDP-Ara	[46]
18	OsUGT708A39	A2YBW8	水稻O. sativa	二氢黄酮二氢查耳酮	UDP-Glc UDP-Ara	[46]
19	OsUGT708A40	A2YBX1	水稻O. sativa	二氢黄酮二氢查耳酮	UDP-Glc UDP-Ara	[46]
20	ZmUGT708A5	B6SWX3	玉米Z. mays	二氢查耳酮	UDP-Glc UDP-Ara	[46]
21	ZmUGT708A11	A0A1D6LXE6	玉米Z. mays	二氢黄酮二氢查耳酮	UDP-Glc UDP-Ara	[46]
22	ZmUGT708A41	B4FAN3	玉米Z. mays	二氢黄酮二氢查耳酮	UDP-Glc	[46]
23	ZmUGT708A42	B4FM13	玉米Z. mays	二氢黄酮二氢查耳酮	UDP-Glc	[46]
24	SbUGT708A36	C5Z8Y8	高粱Sorghum bicolor	二氢黄酮二氢查耳酮	UDP-Glc UDP-Ara	[46]
25	SbUGT708A35	C5Z8Y5	高粱S. bicolor	二氢黄酮二氢查耳酮	UDP-Glc UDP-Ara	[46]
26	SbUGT708A34	C5Z8Y2	高粱S. bicolor	二氢黄酮二氢查耳酮	UDP-Glc UDP-Ara	[46]
27	SbUGT708A38	A0A194YGR9	高粱S. bicolor	二氢黄酮二氢查耳酮	UDP-Glc UDP-Ara	[46]
28	TaUGT708A14	A0A3B6RGT4	小麦Triticum aestivum	二氢黄酮二氢查耳酮	UDP-Glc UDP-Ara	[46]
29	TaUGT708A52	A0A3B6SEV6	小麦T. aestivum	二氢黄酮二氢查耳酮	UDP-Glc UDP-Ara	[46]
30	TaUGT708A53	A0A3B6TPG6	小麦T. aestivum	二氢黄酮二氢查耳酮	UDP-Glc UDP-Ara	[46]
31	TaUGT708A15	A0A3B6RGR9	小麦T. aestivum	二氢黄酮二氢查耳酮	UDP-Glc	[46]
32	TaUGT708A54	A0A3B6SB95	小麦T. aestivum	二氢黄酮二氢查耳酮	UDP-Glc	[46]
33	TaUGT708A55	A0A3B6TNL8	小麦T. aestivum	二氢黄酮	UDP-Glc	[46]
34	BdUGT708A7	I1GYZ6	二穗短柄草 Brachypodium distachyon	二氢黄酮二氢查耳酮	UDP-Glc	[46]
35	BdUGT708A8	I1GZD7	二穗短柄草B. distachyon	二氢黄酮二氢查耳酮	UDP-Glc UDP-Ara	[46]
36	SiUGT708A31	K3XWN4	小米Setaria italica	二氢黄酮二氢查耳酮	UDP-Glc UDP-Ara	[46]
37	SiUGT708A32	K3XWJ1	小米S. italica	二氢黄酮二氢查耳酮	UDP-Glc	[46]
38	SiUGT708A33	K3Y181	小米S. italica	二氢黄酮二氢查耳酮	UDP-Glc UDP-Ara	[46]
39	PhUGT708A43	MK616588	毛竹Phyllostachys heterocycla	二氢黄酮二氢查耳酮	UDP-Glc UDP-Ara	[46]
40	PhUGT708A46	MK616589	毛竹P. heterocycla	二氢黄酮二氢查耳酮	UDP-Glc	[46]
41	PhUGT708A48	MK616590	毛竹P. heterocycla	二氢黄酮二氢查耳酮	UDP-Glc UDP-Ara	[46]
42	PhUGT708A50	MK616591	毛竹P. heterocycla	二氢查耳酮	UDP-Glc UDP-Ara	[46]
43	PgUGT708A44	MK616592	淡竹P. heterocycla	二氢黄酮二氢查耳酮	UDP-Glc UDP-Ara	[46]
44	PpUGT708A45	MK616593	高节竹Phyllostachys prominens	二氢查耳酮	UDP-Glc	[46]
45	DcaCGT	MT452646	铁皮石斛Dendrobium catenatum	二氢黄酮二氢查耳酮	UDP-Glc	[47]
46	NnCGT1（UGT708N1）	LOC104598527	莲Nelumbo nucifera	黄酮	UDP-Glc	[48]
47	NnCGT2（UGT708N2）	LOC104603347	莲N. nucifera	黄酮	UDP-Ara UDP-Xyl	[48]
48	SbCGT	MK894443	黄芩Scutellaria baicalensis	黄酮	UDP-Glc UDP-Ara	[49]
49	LpCGT	MK894451	兰氏萍Landoltia punctata	黄酮	UDP-Glc UDP-Ara	[49]
50	AbCGT	MN747045	芦荟Aloe barbadensis	二氢查尔酮酚	UDP-Glc	[50]

蛋白质ID PDB ID	描述Description	植物来源Species	参考文献Reference
6JTD	TcCGT1与UDP复合的晶体结构	中国金莲花T. chinensis	[43]
6L5P	GgCGT与UDP-Glu复合物的晶体结构	光果甘草G. glabra	[45]
6LLW	UGT708C1与UDP复合的晶体结构	荞麦F. esculentum	[52]
6LF6	ZmCGTa与UDP复合的晶体结构	玉米Z. mays	[49]
6LG0	SbCGTa与UDP复合的晶体结构	黄芩S. baicalensis	[49]
6LG1	LpCGTa与UDP复合物的晶体结构	兰氏萍L. punctata	[49]

蛋白质ID PDB ID	描述Description	植物来源Species	参考文献Reference
6JTD	TcCGT1与UDP复合的晶体结构	中国金莲花T. chinensis	[43]
6L5P	GgCGT与UDP-Glu复合物的晶体结构	光果甘草G. glabra	[45]
6LLW	UGT708C1与UDP复合的晶体结构	荞麦F. esculentum	[52]
6LF6	ZmCGTa与UDP复合的晶体结构	玉米Z. mays	[49]
6LG0	SbCGTa与UDP复合的晶体结构	黄芩S. baicalensis	[49]
6LG1	LpCGTa与UDP复合物的晶体结构	兰氏萍L. punctata	[49]