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中国生物工程杂志

CHINA BIOTECHNOLOGY
中国生物工程杂志
中国生物工程杂志  2022, Vol. 42 Issue (1/2): 46-57    DOI: 10.13523/j.cb.2110034
工业微生物的设计、改造与应用专题     
黄原胶生物合成及分子调控*
戢传富1,王璐2,苟敏1,宋文枫3,夏子渊1,**(),汤岳琴1
1 四川大学建筑与环境学院 四川省环境保护有机废弃物资源化利用重点实验室 成都 610065
2 中国石油勘探开发研究院 提高石油采收率国家重点实验室 北京 100083
3 中国石油勘探开发研究院 北京 100083
The Review of Biosynthesis and Molecular Regulation of Xanthan Gum
JI Chuan-fu1,WANG Lu2,GOU Min1,SONG Wen-feng3,XIA Zi-yuan1,**(),TANG Yue-qin1
1 Sichuan Provincial Key Lab for Resource Utilization of Organic Waste, College of Architecture and Environment, Sichuan University, Chengdu 610065, China
2 State Key Laboratory of Enhanced Oil Recovery, Research Institute of Petroleum Exploration and Development, CNPC, Beijing 100083, China
3 Research Institute of Petroleum Exploration and Development, CNPC, Beijing 100083, China
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摘要：

黄原胶(xanthan gum)是由黄单胞菌属(Xanthomonas)产生的一种胞外多糖,其凭借优越的流变性和稳定性被广泛应用于食品、石油、农业等行业。目前黄原胶的合成途径已经被阐明,其研究主要集中在如何通过分子调控影响其合成及改性以适应于不同行业需求。通过对黄原胶的一级和二级结构、流变性及稳定性、生物合成途径的介绍,综述了黄单胞菌属黄原胶的生物合成分子调控研究进展。主要结论如下:现有的分子调控研究集中于对黄原胶合成途径中各阶段关键基因、信号分子及其他因子的调控;在黄原胶前体合成阶段,可针对葡萄糖转化为磷酸糖、磷酸糖前体转化为二磷酸核苷糖、细胞内碳水化合物利用及转运等过程的相关基因对黄原胶的合成进行调控;在黄原胶的组装及分泌阶段,主要可通过调节gum基因簇的结构蛋白及启动子调控因子等调控;信号分子调控与环二鸟苷酸(c-di-GMP)信号网络系统及群体感应(quorum sensing,QS)系统中的信号分子水平等相关;其他因子包括脂多糖改性O-抗原相关基因,蛋白质/金属转运和分泌相关基因,肽聚糖和聚羟基脂肪酸酯(polyhydroxyalkanoates,PHA)的底物竞争途径,以及血红蛋白基因等。未来应进一步挖掘新的黄原胶生物合成调控因子及完善分子调控机制。
关键词： 黄原胶;  黄单胞菌;  生物合成;  分子调控;  环二鸟苷酸;  群体感应    
Abstract:

Xanthan gum is an extracellular polysaccharide produced by genus Xanthomonas. It is widely used in food, petroleum, agriculture and other industries because of its superior rheology and stability. At present, the synthetic pathway of xanthan gum has been clarified, and its research mainly focuses on how to affect its synthesis and modification through molecular regulation to meet the needs of different industries.By introducing the primary and secondary structure, rheology, stability and biosynthetic pathway of xanthan gum, this paper summarizes the research progress on the molecular regulation of xanthan gum biosynthesis in Xanthomonas sp. The main conclusions are as follows: the existing research on molecular regulation focuses on the regulation of key genes in each stage of xanthan gum synthesis pathway, signal molecules and other factors; in the synthesis stage of xanthan gum precursor, xanthan gum production can be regulated by changing the expression of related genes involved in the conversion of glucose to phosphate sugar, the conversion of phosphate sugar precursor to nucleoside diphosphate, utilization and transport of intracellular carbohydrate; in the assembly and secretion stage of xanthan gum, the synthesis of xanthan gum can be regulated by regulating the structural proteins and promoter regulators of gum gene cluster; the regulation of signal molecule level in c-di-GMP signal network system and quorum sensing (QS) system can affect the synthesis and secretion of xanthan gum; other factors can also regulate the synthesis of xanthan gum, including genes related to lipopolysaccharide modified O-antigen, genes related to protein/metal transport and secretion, substrate competition pathways of peptidoglycan and polyhydroxyalkanoate (PHA), and hemoglobin genes. In the future, we will further explore new regulatory factors for xanthan gum biosynthesis and reveal the molecular regulatory mechanism.
Key words: Xanthan gum    Xanthomonas    Biosynthesis    Molecular regulation    c-di-GMP    Quorum sensing
收稿日期: 2021-10-22 出版日期: 2022-03-03
ZTFLH:  Q819  
基金资助: * 国家重点研发计划资助项目(2018YFA0902100)
通讯作者: 夏子渊     E-mail: ziyuanxia@scu.edu.cn
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引用本文:

戢传富,王璐,苟敏,宋文枫,夏子渊,汤岳琴. 黄原胶生物合成及分子调控*[J]. 中国生物工程杂志, 2022, 42(1/2): 46-57.

JI Chuan-fu,WANG Lu,GOU Min,SONG Wen-feng,XIA Zi-yuan,TANG Yue-qin. The Review of Biosynthesis and Molecular Regulation of Xanthan Gum. China Biotechnology, 2022, 42(1/2): 46-57.

链接本文:

https://manu60.magtech.com.cn/biotech/CN/10.13523/j.cb.2110034    或    https://manu60.magtech.com.cn/biotech/CN/Y2022/V42/I1/2/46

图1  黄原胶的一级结构
图2  Xcc菌中黄原胶的合成途径(根据文献[18,19]制作)
基因名称 基因功能 对黄原胶合成的调控作用 调控机制 参考文献
磷酸糖前体合成阶段
eddH 编码磷酸葡萄糖酸脱水酶 负调控 限制前体6-磷酸葡萄糖的细胞内利用率限制黄原胶合成 [20]
xoo2314 编码葡萄糖-6-磷酸脱氢酶 负调控 改变中心碳代谢的碳流量以增强黄原胶合成 [21]
oprB 编码碳水化合物转运蛋白 缺失后,黄原胶产量及黏度增加 改变中心碳代谢的碳流量以增强黄原胶合成 [22]
α-淀粉酶基因 编码嗜热α-淀粉酶 可生产耐高温的黄原胶,且产量提高 [23]
NDP-sugars合成阶段
pgi 编码黄原胶合成前体相关的葡萄糖- 6-磷酸异构酶 正调控 黄原胶合成前体合成的关键酶 [26]
pig 编码菌黄素合成相关的酶 敲除pig基因簇,黄原胶产量下降;敲除pig基因簇中的糖基转移酶基因xanK后,黄原胶产量提升 菌黄素合成与黄原胶合成存在前体竞争 [27-28]
rmd 编码尿苷二磷酸葡萄糖差向异构酶 外源导入该基因,正调控黄原胶合成 [29]
ugdH 编码UDP-葡萄糖脱氢酶 正调控 黄原胶合成前体UDP-葡糖醛酸合成的关键酶 [30]
表1  黄原胶前体合成阶段关键基因调控
基因名称 基因功能 对黄原胶合成的调控作用 调控机制 参考文献
gumD 编码糖基转移酶 高表达后,黄原胶产量、黏度、均分子质量、乙酰基含量均提高 黄原胶合成途径上的关键基因,直接影响黄原胶的合成 [31]
gumB、gumC 编码黄原胶的链长及外排相关的蛋白质 高表达后,黄原胶黏度、剪切性能值提高 黄原胶合成途径上的关键基因,直接影响黄原胶的合成 [32]
hpaR1 编码结合gum基因簇启动子的HpaR1转录蛋白 正调控 转录调控,结合黄原胶合成的gum基因簇的启动子区域 [33]
vemR 编码接收结构域调控信号的VemR调控因子 正调控 作为一种仅含一个结构接受域的响应调节器,转录调控黄原胶合成基因的转录 [33]
cysB 编码结合gum基因簇启动子的CysB蛋白 正调控 转录调控,直接结合gum基因启动子区域 [34]
soxS 编码结合gum基因簇启动子的氧化应激蛋白 正调控 转录调控,直接结合gum基因启动子区域 [35]
detR 编码与防御系统和调节毒力相关的DetR蛋白 正调控 转录调控,影响其合成途径的关键基因gumD及DSF等信号通路 [36]
表2  黄原胶合成组装阶段关键基因调控
图3  c-di-GMP调控网络系统影响黄原胶和其他毒力因子的合成
基因名称 基因功能 对黄原胶合成的调控作用 调控机制 参考文献
clp 作为c-di-GMP的效应子及全局转录调节剂 正调控和负调控均有 作为全局转录因子,通过转录调控影响黄原胶合成。存在调控级联 [38]
xpsE 二型分泌系统的结构蛋白组成部分,为蛋白质分泌提供能量 正调控 转录后调控,通过为二型分泌系统结构蛋白提供能量影响黄原胶分泌过程 [39]
gdpX1 编码GdpX1蛋白,作为DGCs控制c-di-GMP的合成 正调控 作为DGCs控制c-di-GMP的合成,转录调控下游基因的表达,影响黄原胶的合成。该系统存在调控级联 [40]
dgcA 编码DgcA1蛋白,作为DGCs控制c-di-GMP的合成 正调控 作为DGCs控制c-di-GMP的合成,转录调控下游基因的表达,影响黄原胶的合成。该系统存在调控级联 [41]
edpX1 编码EdpX1蛋白,作为PDEs控制c-di-GMP的降解 负调控 作为DGCs控制c-di-GMP的降解,转录调控下游基因的表达,影响黄原胶的合成。该系统存在调控级联 [42]
vieAxoo 编码VieAxoo蛋白,作为PDEs控制c-di-GMP的降解 负调控 作为DGCs控制c-di-GMP的降解,转录调控下游基因的表达,影响黄原胶的合成。该系统存在调控级联 [42]
rpf基因簇 调节群感效应QS的信号分子DSF的水平 正调控和负调控 (1)转录调控,激活自身PDEs活性以调节c-di-GMP水平影响黄原胶合成。(2)与黄原胶竞争前体葡糖醛酸 [45-48]
ravS/ravR 组成低氧感应双组分系统,与DSF介导的QS相互作用 正调控 影响全局转录因子clp及c-di-GMP的表达水平,在转录水平调控黄原胶合成 [49]
pdeR 编码PdeR蛋白,作为PDEs控制c-di-GMP的降解 负调控 作为DGCs控制c-di-GMP的降解,转录调控下游基因的表达,影响黄原胶的合成。该系统存在调控级联 [50]
表3  信号分子调控对黄原胶合成的影响
基因名称 基因功能 对黄原胶合成的调控作用 调控机制 参考文献
wxcA 与脂多糖改性O-抗原相关,影响LPS的生物合成 正调控 转录调控,影响黄原胶合成糖基转移的反应以及影响相关蛋白质的组装分泌和信号转导 [51]
wxoD 编码假定的o-抗原乙酰化酶,影响LPS的生物合成 负调控 减少竞争代谢LPS的合成,以增加黄原胶合成的代谢资源 [52]
wxcB 与脂多糖改性O-抗原相关,影响LPS的生物合成 负调控 减少竞争代谢LPS的合成,以增加黄原胶合成的代谢资源。 [53]
tatC 双精氨酸转运系统中的编码基因 正调控 [54]
zur 调控细菌的锌离子摄入能力 正调控 [55]
phaR 影响细菌合成聚羟基脂肪酸酯的能力 正调控 通过碳水化合物代谢影响黄原胶合成途径上的碳通量 [57]
vgb 血红蛋白编码基因,与菌体的摄氧能力相关 正调控 通过影响细菌的氧摄取能力,增强菌体体内氧代谢水平促进黄原胶的合成 [58]
xc_4097 编码脂质酰基转移酶 该基因缺失后,可产生
无色黄原胶		 通过脂质代谢影响黄色素的合成,进而生产无色黄原胶 [59]
表4  其他调控因子对黄原胶合成的影响
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