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中国生物工程杂志

China Biotechnology
China Biotechnology
China Biotechnology  2022, Vol. 42 Issue (1/2): 46-57    DOI: 10.13523/j.cb.2110034
Orginal Article     
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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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     
Received: 22 October 2021      Published: 03 March 2022
ZTFLH:  Q819  
Corresponding Authors: Zi-yuan XIA     E-mail: ziyuanxia@scu.edu.cn
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Chuan-fu JI
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Cite this article:

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.

URL:

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

Fig.1 Primary structure of xanthan gum
Fig.2 Synthesis pathway of xanthan gum in Xcc PGI: Glucose-6-phosphate isomerase; PGM and PMM: Glucose phosphate mutase and mannose phosphate mutase; PMI and GMP: Mannose-6-phosphate isomerase and mannose-1-phosphate aminotransferase; UGP: Utp-glucose-1-phosphate uridine transferase; UGD: UDP glucose 6-dehydrogenase
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]
Table 1 Regulation of key genes in xanthan gum precursor synthesis stage
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]
Table 2 Regulation of key genes in the process of xanthan synthesis and assembly
Fig.3 Regulation of xanthan gum synthesis and other virulence factors by c-di-GMP regulatory network DGC: Diguanylate cyclase; PDE: Phosphodiesterase; c-di-GMP: Cyclic diglucoside; GTP: Guanosine triphosphate; GMP: Guanosine monophosphate
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]
Table 3 Effects of signaling molecules regulation on xanthan gum synthesis
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]
Table 4 Effects of other regulatory factors on xanthan gum synthesis
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