水稻胚乳淀粉合成相关酶的结构、功能及其互作研究进展

doi:10.16819/j.1001-7216.2017.6132

摘要/Abstract

摘要：

淀粉作为稻米最主要的储藏物质,其生物合成过程直接影响水稻的产量和品质。水稻淀粉的生物合成在造粉体中通过一系列酶促反应完成。本文综述了淀粉合成过程中ADP-葡萄糖焦磷酸化酶(AGPase)、淀粉合酶(SSs)、淀粉分支酶(BE)及淀粉脱支酶(DBE)的结构、功能以及各酶之间的相互作用的最新研究进展,以期为稻米品质改良提供理论参考依据。

关键词: 水稻, 淀粉合成, 胚乳

Abstract:

Starch is the predominant rice seed reserves. Rice yield and eating quality are influenced by the process of starch biosynthesis. In endosperm amyloplast, starch biosynthesis is a result of complex network of many enzymes. This report summarized the structures, functions and interactions of ADP-glucose pyrophosphorylase (AGPase), starch synthases (SSs), starch branching enzymes (BEs) and starch debranching enzymes (DBEs). It will provide helpful information to breed high yield and good quality rice.

Key words: rice, starch synthesis, endosperm

中图分类号:

陈雅玲, 包劲松. 水稻胚乳淀粉合成相关酶的结构、功能及其互作研究进展[J]. 中国水稻科学, 2017, 31(1): 1-12.

Yaling CHEN, Jinsong BAO. Progress in Structures, Functions and Interactions of Starch Synthesis Related Enzymes in Rice Endosperm[J]. Chinese Journal OF Rice Science, 2017, 31(1): 1-12.

图/表 4

图1 光合作用组织和非光合作用组织中淀粉合成途径和相关代谢[7] TPT-磷酸丙糖转运蛋白; GPT-G6P转运蛋白; G1PT-假定的G1P转运蛋白[8]; BT1-Brittle1内包膜蛋白。A-叶片中瞬时淀粉合成; B-造粉体中储藏淀粉合成; C-淀粉合成途径。

Fig. 1. Schematic representation of the starch biosynthetic pathway and the related metabolism in photosynthetic and non-photosynthetic tissues[7]. TPT, Triose phosphate translocator; GPT, G6P translocator; G1PT, Putative G1P transporter[8]; BT1, Brittle-1 protein. A, The synthesis of assimilatory starch in the leaf; B, The synthesis of reserve starch in storage tissues; C, The starch synthetic pathway from G1P.

表1 水稻胚乳中淀粉合成相关基因、功能及表达特征

Table 1 Function and expression pattern of starch synthesis genes in rice endosperm.

酶 Enzyme	基因 Gene	染色体 Chromosome	最高表达时期 Highest expression stage(DAF)	突变体 Mutant	功能 Function
AGPase	AGPLSU	5,1,3,7	6~15	osagpl2	酶的调控中心Regulation center of the enzyme
	AGPSSU	9,8	6	osagps2	酶的活性中心 Activity center of the enzyme
SSs	GBSSⅠ	6	6,15	waxy	合成直连淀粉或形成超长链 Synthesis of amylose or formation of extra-long chain
	SSⅠ	6	10	ssⅠ	延伸DP 6-7的链形成DP 8-12的链 Formation of the DP 8-12 chain through elongated the DP 6-7 chain
	SSⅡa/SSⅡ-3	6	无明显表达高峰 No obvious peaks	Most japonica	延伸DP≤10的短链形成 DP 12-24的中链 Formation of the DP 12-24 chain through elongated the DP≤10 short chain
	SSⅢa/SSⅢ-2	8	无明显表达高峰 No obvious peak	flo5	形成 DP>30的长链Formation of DP>30 long chain
	SSⅣa	1	无明显表达高峰 No obvious peak	-	涉及淀粉颗粒的形成及控制颗粒数量 Involve formation of starch granules and control the number of starch granules
	SSⅣb	5	10	ss4b	涉及淀粉颗粒的形成及控制颗粒数量 Involve formation of starch granules and control the number of starch granules
BEs	BEⅠ	6	10	sbe1	分支多聚葡萄糖链,形成B链簇状结构 Branch glucose chain and form the cluster structure of b chain
	BEⅡb	2	10	ae	分支多聚葡萄糖链,形成A链 Branch glucose chain and form A chain
DBEs	ISA1	8	10	sugary1	形成同源复合物,去除不当分支 Cleavage improper branch through homologous complexes of ISA1
	ISA2	5	10	-	与ISA1形成异源复合物 Form the heterogeneous complexes with ISA1
	PUL	4	10	pul	补偿ISA功能 Compensate the function of ISA

图2 水稻淀粉合成相关酶3D结构 A-水稻AGPase的最小分子动力学结构[13]。蓝色的a链-LS; 绿色的c链-LS; 蓝绿色的b链-SS; 红色的d链-SS。B-水稻GBSSⅠ的同源模型[21]。蓝色为α螺旋,棕色为β片层,橙色为无规则卷曲。C-日本晴和93-11的SSⅠ三维结构[35]。蓝色为淀粉合酶催化区域; 黄色为葡萄糖转移酶区域。D-水稻BEⅠ结构[53]。紫色为N-末端,淡蓝色为CBM48区域,蓝绿色为α淀粉酶区域,橙色为C-末端。

Fig. 2. 3D structure of starch synthesis related enzymes in rice. A, Molecular dynamics minimized structure of the rice AGPase[13]. Blue chain a, LS; Green chain c, LS; Cyan chain b, SS; Red chain d, SS. B, Homology model of GBSSⅠ, alpha helices are in blue, beta plated sheets in orange and coils in brown. C, Computational models of the 3D structure of SSⅠ of Nipponbare and 93-11. The blue color represents the starch synthase catalytic domain, and the yellow color represents the glycosyl transferase group. D, Structure of BEⅠ from Oryza sativa L. The purple color represents N-domain; The nattier blue represents CBM48; The blue-green color represents α-amylase; The orange represents C-domain.

图3 水稻中淀粉合成相关酶之间的关联网络 BE-淀粉分支酶Ⅰ; SBE-淀粉分支酶Ⅱb; Osl-14800-普鲁蓝酶;Osl-33746-淀粉合酶Ⅱc。

Fig. 3. Potential association networks of starch synthesis enzymes in rice endosperm. BE, Branching enzyme Ⅰ; BEⅡb, Branching enzyme Ⅱb; Osl-14800, Pullulanase; Osl-33746, Starch synthase Ⅱc.

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