植物超分子纤维素合酶复合体的结构与组装研究进展

doi:10.3969/j.issn.1004-1524.20231227

浙江农业学报 ›› 2024, Vol. 36 ›› Issue (10): 2402-2415.DOI: 10.3969/j.issn.1004-1524.20231227

植物超分子纤维素合酶复合体的结构与组装研究进展

任世坦¹^,²(), 郑林², 姜廷波¹, 周博如¹^,^*(), 王宏芝²^,^*()

1.东北林业大学林学院,黑龙江哈尔滨 150040
2.北京市农林科学院生物技术研究所,北京 100097

收稿日期:2023-10-31 出版日期:2024-10-25 发布日期:2024-10-30
作者简介:任世坦(1999—),男,河南濮阳人,硕士研究生,研究方向为森林保护学。E-mail:rst0428@foxmail.com
通讯作者:
^*周博如,E-mail:896099005@qq.com;王宏芝,E-mail:wanghongzhi@baafs.net.cn
基金资助:
北京市农林科学院优秀青年基金项目(YXQN202402);北京市农林科学院科技创新能力建设专项(KJCX20230203)

Research progress on the structure and assembly of plant supramolecular cellulose synthase complexes

REN Shitan¹^,²(), ZHENG Lin², JIANG Tingbo¹, ZHOU Boru¹^,^*(), WANG Hongzhi²^,^*()

1. College of Forestry, Northeast Forestry University, Harbin 150040, China
2. Institute of Biotechnology, Beijing Academy of Agriculture and Forestry Sciences, Beijing 100097, China).

Received:2023-10-31 Online:2024-10-25 Published:2024-10-30

摘要/Abstract

摘要：

纤维素是构成植物细胞壁的主要成分,也是自然界最丰富的可再生资源。植物纤维素由β-1,4-葡聚糖链组成,其合成是由位于质膜上的纤维素合酶复合体(cellulose synthase complex, CSC)催化完成。文章从生物化学、遗传学和进化角度分析了植物CSC结构及组装方面的最新研究成果,包括纤维素合酶(cellulose synthase, CesA)亚基的蛋白结构特征和生物学功能、CesA组装成玫瑰花环CSC核心复合体的机制,以及CSC结构与纤维素特性的关系。进一步探讨了细胞骨架等辅助蛋白和蛋白翻译后修饰在超分子CSC组装与纤维素合成调控中的作用。最后,文章展望了利用单碱基编辑技术突变植物保守结构域(plant-conserved region, PCR)、N端结构域(N-terminal domain, NTD)或跨膜结构域(transmembrane domain, TM)等结构域的关键氨基酸位点,改变纤维素合酶复合体稳定性和功能,为培育生物量增加、抗逆性增强、适合特定工业应用的新型植物提供理论基础,为植物纤维素基因工程遗传改良提供了新策略。

关键词: 纤维素合酶(CesA), 纤维素合酶复合体(CSC), CSC结构, CSC组装, 纤维素

Abstract:

Cellulose is the main component of plant cell wall and the most abundant renewable resource in nature. Plant cellulose is composed of β-1,4-glucan chains, and its synthesis is catalyzed by the cellulose synthase complex (CSC) located on the plasma membrane. In this paper, the latest research findings on the structure and assembly of plant CSC were analyzed from the perspectives of biochemistry, genetics and evolution, these included examining the protein structure characteristics and biological functions of cellulose synthase (CesA) subunits, the mechanism by which CesAs assembly into CSC rosettes core complex, and the relationship between CSC structure and cellulose properties. The roles of auxiliary proteins such as cytoskeleton and post-translational modification of proteins in the assembly of supramolecular CSC and the regulation of cellulose synthesis was further explored. This paper prospected the use of single base editing technology to mutate key amino acid sites in domains such as plant-conserved region (PCR), N-terminal domain (NTD) or transmembrane domain (TM). By regulating the stability and function of CSC, it is expected to cultivate new plants with increased biomass, enhanced stress resistance and suitable for specific industrial applications, which provides a novel strategy for genetic improvement of plant cellulose genetic engineering.

Key words: cellulose synthase (CesA), cellulose synthase complex (CSC), CSC structure, CSC assembly, cellulose

中图分类号:

任世坦, 郑林, 姜廷波, 周博如, 王宏芝. 植物超分子纤维素合酶复合体的结构与组装研究进展[J]. 浙江农业学报, 2024, 36(10): 2402-2415.

REN Shitan, ZHENG Lin, JIANG Tingbo, ZHOU Boru, WANG Hongzhi. Research progress on the structure and assembly of plant supramolecular cellulose synthase complexes[J]. Acta Agriculturae Zhejiangensis, 2024, 36(10): 2402-2415.

图/表 3

图1 纤维素合酶结构与玫瑰花环结构的形成[15] A,PttCESA8纤维素合酶二级结构示意图。TM,跨膜结构域;NTD,N端结构域,包括RING结构域和可变结构域1(VR1);PCR,植物保守结构域;CSR,类型特异结构域;IF,细胞质螺旋结构。B,PttCESA8核心单体三维结构图。C,PttCESA8核心三聚体组装示意图。D,PttCESA8的低分辨率三维结构图。Stalk是由NTD组成的叶柄状结构。E,纤维素合酶18聚体组装示意图。F,玫瑰花环结构示意图。CSC的玫瑰花环结构具有6个花瓣,每个花瓣由CESA三聚体组成。

Fig.1 Structure of cellulose synthase and model of rosette formation A, Cellulose synthase secondary structure diagram of PttCESA8. TM, Transmembrane domain; NTD, N-terminal domain, including RING domain and variable domain 1 (VR1); PCR, Plant conserved domain; CSR, Class specific domain; IF, Cytoplasmic helical structure. B, Three-dimensional structure diagram of PttCESA8 core monomer. C, PttCESA8 core homotrimer assembly diagram. D, Low-resolution three-dimensional structure diagram of full-length PttCESA8. Stalk is a petiole-mounted structure composed of NTD. E, Assembly diagram of cellulose synthase 18 polymers. F, Schematic diagram of the rosette structure. CSC’s rosette structure has six petals, each of which is composed of a CesA homotrimer.

图2 拟南芥、水稻、杨树的CesA蛋白质序列对比拟南芥、水稻和杨树CesA蛋白序列来源于 https://phytozome-next.jgi.doe.gov/,利用MEGA6和GeneDoc软件进行序列对比。根据杨树PtCESA8结构特征对10个CesA主要功能结构域和相关突变位点分别进行了标记:RING结构域标记为藏青色;可变结构域标记为绿色;植物保守结构域(PCR)标记为蓝色;类型特异性结构域(CSR)标记为粉色;跨膜结构域(TM)标记为灰色;小红色方框标记突变位点;红色箭头标记突变体蛋白翻译提前终止位置;蓝色和黑色直线分别标记CesA的β折叠链和α螺旋结构;蓝色方框标记保守的D, D, D, QxxRW基序;红色和黑色圆圈分别标记CesA三聚体聚合和纤维素配位相关的氨基酸残基。

Fig.2 Alignment of CesA protein sequences from Arabidopsis thaliana, Oryza sativa and Populus The CesAs protein sequences of Arabidopsis, rice and poplar were obtained from https://phytozome-next.jgi.doe.gov/, and the sequence alignment was performed by MEGA6 and GeneDoc software. According to the structural characteristics of poplar PtCESA8, functionally important regions of 10 CesAs and related mutation sites were marked: RING domain is marked in navy blue; Variable region is marked in green; plant conserved domain (PCR) is marked in blue; class-specific domain (CSR) is marked in pink; transmembrane domain (TM) is marked in gray; small red box marks the mutation site; red arrow marks the premature termination position of mutant protein translation; blue and black straight lines indicate β-stands and α-helices of CesAs, respectively; blue boxes mark the conserved D, D, D, QxxRW motifs; red and black circles indicate residues implicated in CesAs trimerization and cellulose coordination, respectively.

图3 超分子纤维素合酶复合体模式图 CC,纤维素合酶伴侣蛋白;PM,质膜;MT,微管;CesA,纤维素合酶;CSI,纤维素合酶和微管的连接蛋白。

Fig.3 Model of supramolecular cellulose synthase complex CC, Companion of cellulose synthase; PM, Plasma membrane; MT, Microtubule; CesA, Cellulose synthase; CSI, POM-POM2/cellulose synthase interacting 1.

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植物超分子纤维素合酶复合体的结构与组装研究进展

Research progress on the structure and assembly of plant supramolecular cellulose synthase complexes

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