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Article(id=1152687440251765384, tenantId=1146029695717560320, journalId=1149652044408987649, issueId=1152687434774000221, articleNumber=null, orderNo=null, doi=10.19812/j.cnki.jfsq11-5956/ts.20250120006, pmid=null, cstr=null, oa=null, hot=null, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=null, receivedDate=1737302400000, receivedDateStr=2025-01-20, revisedDate=null, revisedDateStr=null, acceptedDate=null, acceptedDateStr=null, onlineDate=1752751701652, onlineDateStr=2025-07-17, pubDate=1747238400000, pubDateStr=2025-05-15, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1752751701652, onlineIssueDateStr=2025-07-17, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1752751701652, creator=13701087609, updateTime=1752751701652, updator=13701087609, issue=Issue{id=1152687434774000221, tenantId=1146029695717560320, journalId=1149652044408987649, year='2025', volume='16', issue='9', pageStart='1', pageEnd='324', issueExtLink='null', onlineDate='null', pubDate='null', beforeIssueId=null, nextIssueId=null, price=null, status=1, issueComplete=1, articleOrder=1, issueType=-1, specialIssue=0, createTime=1752751700342, creator=13701087609, updateTime=1756708585928, updator=13701087609, preIssue=null, nextIssue=null, ext={EN=IssueExt(id=1169283815848555430, tenantId=1146029695717560320, journalId=1149652044408987649, issueId=1152687434774000221, language=EN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=), CN=IssueExt(id=1169283815848555431, tenantId=1146029695717560320, journalId=1149652044408987649, issueId=1152687434774000221, language=CN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=)}, issueFiles=null}, startPage=33, endPage=39, ext={EN=ArticleExt(id=1152687441304535719, articleId=1152687440251765384, tenantId=1146029695717560320, journalId=1149652044408987649, language=EN, title=Effects of glnA gene knockout on the production of available curdlan by Agrobacterium, columnId=1152687436237812322, journalTitle=Journal of Food Safety & Quality, columnName=Special Topic: Application of Fermentation Technology in Food, runingTitle=null, highlight=null, articleAbstract=

Objective To explore the effects of glnA gene on the production of curdlan by Agrobacterium. Methods A glnA gene knockout strain ΔglnA was constructed in this paper. The growth and metabolism of ΔglnA strain and the changes in yield, gel properties and infrared structure of the obtained gum were analyzed, and the expression of genes related to the synthesis of curdlan was explored by quantitative real-time polymerase chain reaction. Results The amino nitrogen consumption of the ΔglnA strain during fermentation was consistent with that of CGMCC11546. In terms of sucrose consumption, there was no significant difference before 12 h, but there was a significant difference between the ΔglnA strain and CGMCC11546 after 15 h, and sucrose consumption was significantly reduced compared to CGMCC11546. The ΔglnA strain could produce approximately 4 g/L of natural gum, which was 60% lower than CGMCC11546. Conclusion The glnA gene does not affect bacterial growth or the structure of collagen, but it can reduce the synthesis of collagen. This study provides insights into the role of glnA in curdlan production and lays a molecular biological foundation for further efforts to enhance curdlan yield.

, correspAuthors=Bo-Wei YAO, Xue-Xia YANG, authorNote=null, correspAuthorsNote=null, copyrightStatement=null, copyrightOwner=null, extLink=null, articleAbsUrl=null, sourceXml=null, magXml=null, pdfUrl=null, pdf=null, pdfFileSize=null, pdfExtLink=null, richHtmlUrl=null, mobilePdfUrl=null, reviewReport=null, pdfFirstPage=null, abstractGraph=null, abstractGraphContent=null, abstractVideo=null, citation=null, cebUrl=null, magXmlContent=null, mapNumber=null, authorCompany=null, fund=null, authors=null, authorsList=Bo-Wei YAO, Shi-Qiang WU, Hong-Liang GAO, Xue-Xia YANG), CN=ArticleExt(id=1152687464369013027, articleId=1152687440251765384, tenantId=1146029695717560320, journalId=1149652044408987649, language=CN, title=glnA基因敲除对土壤杆菌产可得然胶的影响, columnId=1152687436393001573, journalTitle=食品安全质量检测学报, columnName=本期专题：发酵技术在食品中的应用, runingTitle=null, highlight=null, articleAbstract=

目的 探究glnA基因对土壤杆菌产可得然胶的影响。方法 构建glnA基因敲除株ΔglnA菌株, 分析ΔglnA菌株生长代谢情况和产生的可得然胶在产量、凝胶性质和红外结构的变化, 并通过实时荧光定量聚合酶链式反应的方法探究可得然胶合成相关基因的表达量。结果 ΔglnA菌株在发酵过程中氨基氮消耗情况与CGMCC11546一致, 在蔗糖消耗方面, ΔglnA菌株与CGMCC11546在12 h之前无明显差别, 但在15 h之后出现明显差异, 蔗糖消耗比CGMCC11546明显降低。ΔglnA菌株可得然胶产量约4 g/L, 相对于CGMCC11546降低了60%。结论 glnA基因不影响菌体生长和可得然胶结构, 但可以使可得然胶的合成减少。本研究通过对glnA基因进行敲除, 研究其对可得然胶合成的影响, 为后续提高可得然胶产量提供一些分子生物学基础。

, correspAuthors=姚博伟, 杨雪霞, authorNote=null, correspAuthorsNote=
* 姚博伟(1993—), 女, 硕士, 工程师, 主要研究方向为应用微生物及食品质量安全。E-mail:
* 杨雪霞(1971—), 女, 博士, 副教授, 主要研究方向为应用微生物与酶学研究。E-mail:
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Molecular cloning: A laboratory manual[M]. 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School of Life Sciences, East China Normal University, Shanghai 200241, China), AuthorCompanyExt(id=1169272761055715689, tenantId=1146029695717560320, journalId=1149652044408987649, articleId=1152687440251765384, companyId=1169272761038938471, language=CN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=3.华东师范大学生命科学学院, 上海 200241)])], figs=[ArticleFig(id=1169272762548887942, tenantId=1146029695717560320, journalId=1149652044408987649, articleId=1152687440251765384, language=EN, label=Fig.1, caption=Vector and result of double digestion, figureFileSmall=bVlvJtb4AfCx1WOgoNAjFw==, figureFileBig=jRsyiJ4WRFgUwYH71jud2g==, tableContent=null), ArticleFig(id=1169272762603413895, tenantId=1146029695717560320, journalId=1149652044408987649, articleId=1152687440251765384, language=CN, label=图1, caption=质粒和双酶切结果

注: A. pT-glnA-UD质粒图谱; B. pEXG2-glnA-UD质粒图谱; C. pEXG2-glnA-UD双酶切电泳结果; 图C中M. 10000 DNA ladder; 1. glnA基因上下游同源臂融合序列; 2. pEXG2-glnA-UD双酶切电泳结果。

, figureFileSmall=bVlvJtb4AfCx1WOgoNAjFw==, figureFileBig=jRsyiJ4WRFgUwYH71jud2g==, tableContent=null), ArticleFig(id=1169272762704077192, tenantId=1146029695717560320, journalId=1149652044408987649, articleId=1152687440251765384, language=EN, label=Fig.2, caption=Result of colony PCR, figureFileSmall=m9nISZlhyFifx7WNdJ8RtQ==, figureFileBig=MhMht/8uaxEpVwkOntDMMg==, tableContent=null), ArticleFig(id=1169272762766991753, tenantId=1146029695717560320, journalId=1149652044408987649, articleId=1152687440251765384, language=CN, label=图2, caption=菌落PCR鉴定结果

注: A. 同源臂融合片段扩增结果; B. glnA基因片段扩增结果; M. DL2000 marker; 1~10. 菌株编号; ck. 土壤杆菌CGMCC11546。

, figureFileSmall=m9nISZlhyFifx7WNdJ8RtQ==, figureFileBig=MhMht/8uaxEpVwkOntDMMg==, tableContent=null), ArticleFig(id=1169272762855072138, tenantId=1146029695717560320, journalId=1149652044408987649, articleId=1152687440251765384, language=EN, label=Fig.3, caption=Curdlan yield after fermentation for 96 h, figureFileSmall=pP4ZprTWAmKM6xjPvojqog==, figureFileBig=pR9qJPMdF3BrwCHej9874w==, tableContent=null), ArticleFig(id=1169272762917986699, tenantId=1146029695717560320, journalId=1149652044408987649, articleId=1152687440251765384, language=CN, label=图3, caption=发酵96 h可得然胶粗产量

注: ***表示差异极显著(P<0.001)。

, figureFileSmall=pP4ZprTWAmKM6xjPvojqog==, figureFileBig=pR9qJPMdF3BrwCHej9874w==, tableContent=null), ArticleFig(id=1169272763006067084, tenantId=1146029695717560320, journalId=1149652044408987649, articleId=1152687440251765384, language=EN, label=Fig.4, caption=Growth and metabolic status of ΔglnA strain and Agrobacterium sp. CGMCC11546 strain 24 hours before fermentation., figureFileSmall=3VXLo8uvZNsgXL9TSfIx9w==, figureFileBig=H1pG8hW/Luju+mYcUrcYVw==, tableContent=null), ArticleFig(id=1169272763127701901, tenantId=1146029695717560320, journalId=1149652044408987649, articleId=1152687440251765384, language=CN, label=图4, caption=发酵前24 h ΔglnA菌株与土壤杆菌CGMCC11546菌株生长代谢情况

注: A. 生长曲线; B. 发酵前24 h可得然胶粗产量; C. 发酵前24 h氮含量; D. 发酵前24 h蔗糖含量。

, figureFileSmall=3VXLo8uvZNsgXL9TSfIx9w==, figureFileBig=H1pG8hW/Luju+mYcUrcYVw==, tableContent=null), ArticleFig(id=1169272763178033550, tenantId=1146029695717560320, journalId=1149652044408987649, articleId=1152687440251765384, language=EN, label=Fig.5, caption=Real-time fluorescence quantification PCR results of genes related to ccurdlan synthesis obtained from the ΔglnA strain, figureFileSmall=Wjq1X2zNRE+a3o5F0qjd0w==, figureFileBig=2RBInL0Y4m+B4+Xl0ZdoFw==, tableContent=null), ArticleFig(id=1169272763228365199, tenantId=1146029695717560320, journalId=1149652044408987649, articleId=1152687440251765384, language=CN, label=图5, caption=ΔglnA菌株可得然胶合成相关基因实时荧光定量PCR结果, figureFileSmall=Wjq1X2zNRE+a3o5F0qjd0w==, figureFileBig=2RBInL0Y4m+B4+Xl0ZdoFw==, tableContent=null), ArticleFig(id=1169272763299668368, tenantId=1146029695717560320, journalId=1149652044408987649, articleId=1152687440251765384, language=EN, label=Table 1, caption=

Primers of the experiment

, figureFileSmall=null, figureFileBig=null, tableContent=
组别 引物名称 序列(5'→3')
同源重组基因敲除所用引物 gUF CGGGATCCAGCCCGACCAGCAGTTTCAG
gUR AGAAACACAGACCCGGTGATACGGAACCACGAGCATGACATGGC
gDF GCCATGTCATGCTCGTGGTTCCGTATCACCGGGTCTGTGTTTCT
gDR CCAAGCTTGCGCCCATTTTCGGGATGGA
gF AAGGTGGCGTAGATGCCGTG
gR CTGGAAGCCGGTCGTAGCAC
实时荧光定量PCR所用引物 q-fixN-F CACGTAGAAGGATGTGGCGA
q-fixN-R TGGGGTGTTGTCGGTTTTCT
q-ppx-F TCGGCTATCGGAACAATCGG
q-ppx-R CATGTACCACCGACCGCATA
q-exoC-F GCAACGGAAATCAGATCGGC
q-exoC-R GTCGACCAGTCTGTCAGCAA
q-galU-F CAAGTCGCTTCATGCCATCG
q-galU-R ATCACGGCTTTTCCGTCACT
q-crdR-F TCCTCAATGTGCCCGTTTCC
q-crdR-R GATCTTCACGAAAGCCCGGT
q-crdS-F CCAGACCATAGACAGCCTGC
q-crdS-R GGCAATGGCTAAACCGTTCG
q-ntrB-F TCGGCTATCGGAACAATCGG
q-ntrB-R CATGTACCACCGACCGCATA
q-ntrC-F AGACCGATGTGACGCTGATG
q-ntrC-R TTCCGATTCGATCAGGTCGC
), ArticleFig(id=1169272763383554449, tenantId=1146029695717560320, journalId=1149652044408987649, articleId=1152687440251765384, language=CN, label=表1, caption=

实验所需要的引物

, figureFileSmall=null, figureFileBig=null, tableContent=
组别 引物名称 序列(5'→3')
同源重组基因敲除所用引物 gUF CGGGATCCAGCCCGACCAGCAGTTTCAG
gUR AGAAACACAGACCCGGTGATACGGAACCACGAGCATGACATGGC
gDF GCCATGTCATGCTCGTGGTTCCGTATCACCGGGTCTGTGTTTCT
gDR CCAAGCTTGCGCCCATTTTCGGGATGGA
gF AAGGTGGCGTAGATGCCGTG
gR CTGGAAGCCGGTCGTAGCAC
实时荧光定量PCR所用引物 q-fixN-F CACGTAGAAGGATGTGGCGA
q-fixN-R TGGGGTGTTGTCGGTTTTCT
q-ppx-F TCGGCTATCGGAACAATCGG
q-ppx-R CATGTACCACCGACCGCATA
q-exoC-F GCAACGGAAATCAGATCGGC
q-exoC-R GTCGACCAGTCTGTCAGCAA
q-galU-F CAAGTCGCTTCATGCCATCG
q-galU-R ATCACGGCTTTTCCGTCACT
q-crdR-F TCCTCAATGTGCCCGTTTCC
q-crdR-R GATCTTCACGAAAGCCCGGT
q-crdS-F CCAGACCATAGACAGCCTGC
q-crdS-R GGCAATGGCTAAACCGTTCG
q-ntrB-F TCGGCTATCGGAACAATCGG
q-ntrB-R CATGTACCACCGACCGCATA
q-ntrC-F AGACCGATGTGACGCTGATG
q-ntrC-R TTCCGATTCGATCAGGTCGC
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glnA基因敲除对土壤杆菌产可得然胶的影响
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姚博伟 1, * , 武世强 2 , 高红亮 3 , 杨雪霞 2, *
食品安全质量检测学报 | 本期专题：发酵技术在食品中的应用 2025,16(9): 33-39
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食品安全质量检测学报 | 本期专题：发酵技术在食品中的应用 2025, 16(9): 33-39
glnA基因敲除对土壤杆菌产可得然胶的影响
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姚博伟1, * , 武世强2, 高红亮3, 杨雪霞2, *
作者信息
  • 1.宁夏计量质量检验检测研究院, 银川 750000
  • 2.东华大学生物与医学工程学院, 上海 201620
  • 3.华东师范大学生命科学学院, 上海 200241

通讯作者:

* 姚博伟(1993—), 女, 硕士, 工程师, 主要研究方向为应用微生物及食品质量安全。E-mail:
* 杨雪霞(1971—), 女, 博士, 副教授, 主要研究方向为应用微生物与酶学研究。E-mail:
Effects of glnA gene knockout on the production of available curdlan by Agrobacterium
Bo-Wei YAO1, * , Shi-Qiang WU2, Hong-Liang GAO3, Xue-Xia YANG2, *
Affiliations
  • 1. NingXia Academy of Metrology & Quality Inspection, Yinchuan 750000, China
  • 2. College of Biological Science and Medical Engineering, Donghua University, Shanghai 201620, China
  • 3. School of Life Sciences, East China Normal University, Shanghai 200241, China
出版时间: 2025-05-15 doi: 10.19812/j.cnki.jfsq11-5956/ts.20250120006
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摘要
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目的 探究glnA基因对土壤杆菌产可得然胶的影响。方法 构建glnA基因敲除株ΔglnA菌株, 分析ΔglnA菌株生长代谢情况和产生的可得然胶在产量、凝胶性质和红外结构的变化, 并通过实时荧光定量聚合酶链式反应的方法探究可得然胶合成相关基因的表达量。结果 ΔglnA菌株在发酵过程中氨基氮消耗情况与CGMCC11546一致, 在蔗糖消耗方面, ΔglnA菌株与CGMCC11546在12 h之前无明显差别, 但在15 h之后出现明显差异, 蔗糖消耗比CGMCC11546明显降低。ΔglnA菌株可得然胶产量约4 g/L, 相对于CGMCC11546降低了60%。结论 glnA基因不影响菌体生长和可得然胶结构, 但可以使可得然胶的合成减少。本研究通过对glnA基因进行敲除, 研究其对可得然胶合成的影响, 为后续提高可得然胶产量提供一些分子生物学基础。

可得然胶  /  土壤杆菌  /  基因敲除  /  glnA基因

Objective To explore the effects of glnA gene on the production of curdlan by Agrobacterium. Methods A glnA gene knockout strain ΔglnA was constructed in this paper. The growth and metabolism of ΔglnA strain and the changes in yield, gel properties and infrared structure of the obtained gum were analyzed, and the expression of genes related to the synthesis of curdlan was explored by quantitative real-time polymerase chain reaction. Results The amino nitrogen consumption of the ΔglnA strain during fermentation was consistent with that of CGMCC11546. In terms of sucrose consumption, there was no significant difference before 12 h, but there was a significant difference between the ΔglnA strain and CGMCC11546 after 15 h, and sucrose consumption was significantly reduced compared to CGMCC11546. The ΔglnA strain could produce approximately 4 g/L of natural gum, which was 60% lower than CGMCC11546. Conclusion The glnA gene does not affect bacterial growth or the structure of collagen, but it can reduce the synthesis of collagen. This study provides insights into the role of glnA in curdlan production and lays a molecular biological foundation for further efforts to enhance curdlan yield.

curdlan  /  Agrobacterium  /  gene knockout  /  glnA gene
姚博伟, 武世强, 高红亮, 杨雪霞. glnA基因敲除对土壤杆菌产可得然胶的影响. 食品安全质量检测学报, 2025 , 16 (9) : 33 -39 . DOI: 10.19812/j.cnki.jfsq11-5956/ts.20250120006
Bo-Wei YAO, Shi-Qiang WU, Hong-Liang GAO, Xue-Xia YANG. Effects of glnA gene knockout on the production of available curdlan by Agrobacterium[J]. Journal of Food Safety & Quality, 2025 , 16 (9) : 33 -39 . DOI: 10.19812/j.cnki.jfsq11-5956/ts.20250120006
正文
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0 引言
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可得然胶, 又称为可得然胶多糖, 是一种微生物胞外多糖, 不溶于水, 可溶于NaOH、二甲基亚砜(dimethyl sulfoxide, DMSO)等碱性溶剂中。可得然胶最早发现于粪产碱杆菌(Alcaligenes faecalis var. myxogens) 10C3K[1], 后陆续发现根瘤菌(Rhizobium)[2]、土壤杆菌(Agrobacterium)[3]等菌株也可以合成可得然胶。由于其加热成胶性、持水性、无毒性、可降解性等性能受到食品、生命医学、新型材料、环境保护等行业的广泛关注[4-7]。可得然胶是一种β-D-1,3-葡聚糖[8-9], 可得然胶分子式为(C6H10O5)n, 在碱溶液中有单链螺旋结构、三螺旋结构以及松散环状结构3种构象。三螺旋结构在普遍存在, 是可得然胶的主要结构[10]。60 ℃时可形成低固胶(可逆凝胶); 80 ℃时可形成高固胶(不可逆凝胶)[11]。可得然胶具有抗脱水性、稳定性、分散性、抗冻融性和抗消化性[12], 这些性质使得可得然胶在诸多领域中都有着广泛的应用。可得然胶水悬浮液高温和低温下会形成两种结构不同的凝胶, 分别称为高凝固热不可逆凝胶和低凝固热可逆凝胶[13-14]。相较于其他亲水胶体, 可得然胶具有独特的加热流变性和保水性能, 作为稳定剂、持水剂、增稠剂、乳化剂等在面制品[15]、肉制品[16]、乳制品[17]、速冻食品[18]、复合食品添加剂[19]和食品保鲜材料[20]中广泛应用。
可得然胶在氮源限制的条件下合成, 主要包括两个方面: 底物合成和能量供给。可得然胶合成的底物是尿苷二磷酸(uridine diphosphate, UDP)-葡萄糖, 尿苷三磷酸(uridine triphosphate, UTP)-葡萄糖-1-磷酸尿苷转移酶(由galU基因编码)催化UTP和葡萄糖-1-磷酸生成UDP-葡萄糖, 在不同氧浓度下galU基因转录水平有些显著变化, 因此galU是可得然胶合成过程中的一个关键酶[21]。可得然胶合成需要大量的能量, 转录组分析表明, 在高氧浓度下可得然胶有着很高产量, 同时三羧酸循环和电子传递链中的关键基因以及galU基因均呈现显著上调。研究发现, 参与半胱氨酸和蛋氨酸代谢的metZ和metH基因在胶原蛋白的合成中起着重要作用。敲除metZ和metH基因会降低土壤杆菌对氮源和糖的利用, 减少腺苷三磷酸(adenosine triphosphate, ATP)合成, 抑制L-甲流氨酸合成, 导致可得然胶合成过程中糖利用率降低, 能量供应不足, 导致减产[22]。细菌在不缺少营养物质, 但是碳源过剩时可以产生聚-3-羟基烷酸(poly-3-hydroxyalkanoate, PHA), 碳源不足时PHA亦可作碳源供细菌使用[23-24]。聚-3-羟基丁酸(poly-3-hydroxybutyrate, PHB)是PHA中研究最为广泛的一种。PHB的合成涉及多个基因[25], phbC基因编码的聚羟基丁酸合酶, 可催化3-羟基丁酸合成PHB。土壤杆菌CGMCC11546的phbC基因敲除后可得然胶产量降低约46%[26]。
研究发现crdASC操纵子(包含crdA、crdS、crdC)和crdR对于可得然胶的生物合成至关重要, 在crdASC操纵子中, crdS编码一种糖基转移酶, 将UDP-葡萄糖的葡萄糖残基转移到可得然胶链中[27]。CrdS与糖基转移酶(glycosyltransferase, GT)家族II具有相似性, 其介导细胞膜内测葡萄糖基聚合。crdS的突变导致可得然胶合成障碍, 通过回补crdS基因可消除该基因突变带来的影响, 从而证明crdS对于可得然胶生物合成是必需的[28]。基于转录组分析, 在氮饥饿时crdASC操纵子上调近100倍, 证明了crdASC操纵子在土壤杆菌ATCC31749菌株的可得然胶生物合成中的突出作用[29]。YU等[30]已经确定, 作为正调节剂的CrdR可以改善土壤杆菌ATCC31749中的可得然胶生物合成, 并且crdR位于crdA起始密码子的上游。还推断CrdR通过调节crdASC操纵子的表达来调节可得然胶的生物合成。
NtrC-NtrB是在氮源耗尽时参与氮信号级联的双组分系统, 研究表明土壤杆菌ATCC31749中ntrBC突变体在酵母提取物-葡萄糖培养基上可以产生可得然胶。ntrBC操纵子是可得然胶生物合成的关键调节机制, ntrBC基因可用于在氮耗尽下启动氮同化[31], 此时糖类物质便不再供应细胞生长, 而是被用于合成可得然胶, 所以氮元素的利用情况是影响可得然胶生物合成的主要因素[32-34]。在土壤杆菌 CGMCC11546中也存在类似的双组分系统Gene0716-Gene0717, 其中起主要作用的是转录因子Gene0716, 但是通过增强gene0717的表达可以提高野生株的可得然胶产量[35]。
国内对于可得然胶的研究起步较晚, 但发展迅速, 在高产菌株诱变选育及改造[36-37]、发酵工艺优化[38-39]、合成机制调控[40]等方面报道了诸多研究成果, 为我国可得然胶产业发展奠定了基础。但是, 目前我国可得然胶仍未实现完全自主化生产, 究其原因主要是存在产量低/凝胶强度差等一系列瓶颈问题[41]。为进一步了解土壤杆菌可得然胶合成的分子机制, 通过基因改造提高可得然胶产量, 华东师范大学生命科学学院微生物实验室在前期研究中发现glnA基因突变能够明显降低可得然胶产量[40]。本研究通过构建glnA基因敲除株, 并研究glnA基因对土壤杆菌CGMCC11546产可得然胶的影响, 以期从分子生物学角度为提高可得然胶产量及工业化应用奠定基础。
1 材料与方法
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1.1 材料与试剂
土壤杆菌CGMCC11546、大肠埃希氏菌(Escherichia coli) DH5α菌株以及质粒pMD-19T、pEXG2均为华东师范大学生命科学学院微生物实验室保存。
细菌基因组DNA提取试剂盒、质粒小提试剂盒和琼脂糖凝胶回收试剂盒(北京天根生化科技有限公司); DNA片段纯化试剂盒、限制性核酸内切酶、T4 DNA连接酶(大连宝生物工程有限公司); 2×Taq Master Mix、2×Fast Pfu Master Mix、DNA marker(美国赛默飞世尔科技有限公司); 反转录试剂盒(北京普洛麦格生物技术有限公司); 荧光定量聚合酶链式反应(polymerase chain reaction, PCR)试剂盒、Gold View核酸染液(上海翊圣生物科技有限公司); DMSO、NaOH、KH2PO4、K2HPO4·3H2O、(NH4)2HPO4、NH4Cl、蔗糖、MgSO4·7H2O、玉米浆、CaCO3、95%乙醇、甘油、庆大霉素(国药集团化学试剂有限公司); PCR引物委托南京金斯瑞生物科技有限公司合成(表1)。
1.2 仪器与设备
GNP型隔水式恒温培养箱、DHG-9246A型电热恒温鼓风干燥箱(上海精宏实验设备有限公司); UV-2700型分光光度计(日本岛津仪器有限公司); SorvallRC-6plus型冷冻离心机、Nanodrop型微量分光光度计(美国赛默飞世尔科技有限公司); PB-10型pH酸度计、SQP型精密电子天平(精度0.0001 g)(德国赛多利斯仪器有限公司); MLS-378L-PC型立式压力蒸汽灭菌器(日本松下电器有限公司); sw-cj-2FD型洁净工作台(苏净安泰空气技术有限公司); Sub-Cell GT型电泳槽、PowerEase型稳压电泳仪、cfx opus 96型实时荧光PCR仪(伯乐生命医学产品有限公司); Mastercycler型PCR仪(德国艾本德股份有限公司); SCIENTZ-18N型真空冷冻干燥机(宁波新之生物科技有限公司)。
1.3 培养基配制
LB培养基: 胰蛋白胨1.0 g/L、酵母粉0.5 g/L、NaCl 1.0 g/L、pH 7.2。配制固体培养基时, 加入2%琼脂。
LB-蔗糖培养基: 蔗糖8.0 g/L、胰蛋白胨1.0 g/L、酵母粉0.5 g/L、NaCl 1.0 g/L、pH 7.2。配制固体培养基加入2%琼脂。
种子培养基: 蔗糖2.0 g/L、(NH4)2HPO4 1.5 g/L、KH2PO4 0.15 g/L、MgSO4·7H2O 0.1 g/L、玉米浆0.4 g/L、CaCO3 0.3 g/L、pH 7.2。
发酵培养基: 蔗糖9.0 g/L、(NH4)2HPO4 0.2 g/L、KH2PO4 0.2 g/L、MgSO4·7H2O 0.1 g/L、玉米浆0.3 g/L、CaCO3 0.2 g/L、pH 7.2。
1.4 实验方法
1.4.1 感受态细胞制备及转化
按照《分子克隆指南》进行制备和转化步骤进行[42]。
1.4.2 glnA基因敲除菌株的获得
参考文献[26], 以CGMCC11546基因组DNA为模板, PCR扩增得到glnA上下游同源臂融合片段glnA-UD。先将glnA-UD连接到pMD-19T载体上, 构建克隆载体pT-glnA-UD(图1A), 经酶切、测序鉴定无误后回收glnA-UD。再将glnA-UD与pEXG2质粒连接构建基因打靶载体pEXG2-glnA-UD(图1B), 鉴定无误后电转进CGMCC11546菌株进行同源重组。将菌液稀释涂布在含有50 μg/mL庆大霉素的LB固体培养基上并挑选存活的菌落接种在含有8%蔗糖的LB平板上培养, 验证无误后保存。
1.4.3 ΔglnA菌株可得然胶产量验证
按照按照文献[35,38]所述方法进行可得然胶产量的测定。
1.4.4 ΔglnA菌株生长曲线测定
参考文献[26], 将活化菌液按2%的接种量接入种子培养基(250 mL锥形瓶装液量50 mL), 30 ℃、250 r/min培养, 每隔2 h取样, 测OD600值。
1.4.5 氮含量测定
按照HJ 536—2009《水质 氨氮的测定 水杨酸分光光度法》标准检测氮含量。
1.4.6 蔗糖含量测定
按照GB 5009.8—2023《食品安全国家标准 食品中果糖、葡萄糖、蔗糖、麦芽糖、乳糖的测定》标准检测蔗糖含量。
1.4.7 可得然胶合成相关基因的定量分析
以CGMCC11549基因表达量为参考, 用实时荧光定量PCR引物分别检测发酵6 h和12 h的fixN、ppx、exoC、galU、crdR、crdS、ntrB、ntrC和glnA基因表达量。
1.5 数据处理
本研究中采用Excel 2013、SPSS 22.0统计软件进行数据处理, 采用GraphPad prism 8.0进行图表绘制, 每组实验数据均测定3次。
2 结果与讨论
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2.1 glnA基因同源重组基因敲除载体构建
将鉴定无误的glnA上下游同源臂融合片段glnA-UD与pEXG2质粒构建基因打靶载体pEXG2-glnA-UD, 酶切鉴定, 如图1C所示。酶切结果显示, 酶切产物有两条条带, 较小的片段在1000 bp左右, 较大的片段在5000 bp左右, 条带大小正确, 基因打靶载体pEXG2-glnA-UD准确无误。
2.2 ΔglnA菌株的验证
利用基因重组方法构建glnA敲除株。将PEXG2-glnA-UD电转化到土壤杆菌CGMCC11546菌株中, 并进行菌落PCR鉴定, 结果如图2所示。10个选定菌株均扩增为同源臂融合片段未扩增为glnA片段。取编号为1的菌株表示为ΔglnA, 发酵96 h后可以获得可得然胶的粗产量, 如图3所示。从发酵结果可以看出, 经过96 h的发酵后, ΔglnA菌株和土壤杆菌CGMCC11546的可得然胶粗产量存在极显著差异, ΔglnA菌株可得然胶产量约4 g/L, 相对于CGMCC11546降低了60%
2.3 ΔglnA菌株生长代谢情况
为了消除glnA基因敲除对菌体生长的影响, 本研究测定了ΔglnA菌株的生长曲线、发酵24 h可得然胶粗产量、24 h后的氮含量和蔗糖含量。从生长曲线看(图4A), ΔglnA菌株和土壤杆菌CGMCC11546之间没有差异。两种菌株均在6 h进入指数生长期期, 在18 h进入稳定期。
可得然胶粗产量方面(图4B), 发酵9 h后, ΔglnA菌株的产量开始低于土壤杆菌CGMCC11546菌株。12 h后ΔglnA菌株产量稳定。而土壤杆菌CGMCC11546菌株产量继续增长, 二者在12 h后出现明显差异。
氮消耗方面(图4C), 无论是ΔglnA菌株还是土壤杆菌 CGMCC11546菌株, 氮元素在约15 h后基本被消耗, 21 h后无法检测到氮元素。这表明ΔglnA菌株和土壤杆菌CGMCC11546菌株的生长没有差异, 这与生长结果一致。
蔗糖消耗方面(图4D), ΔglnA菌株和土壤杆菌CGMCC11546菌株在12 h前的蔗糖消耗量没有明显差异, 但在15 h后, ΔglnA菌株蔗糖消耗少, 这与ΔglnA菌株可用粗胶生产减少结果一致。
上述结果表明, ΔglnA菌株和土壤杆菌CGMCC11546菌株的生长正常无差别。然而, ΔglnA菌株可得然胶合成和蔗糖消耗明显低于土壤杆菌 CGMCC11546菌株, 这说明glnA基因的敲除会降低土壤杆菌对糖的消耗效率, 减少糖向可得然胶的转化。
2.4 不同时期可得然胶合成相关基因的表达
实时荧光定量PCR检测发现, 在发酵6 h时, ΔglnA菌株中fixN基因表达量上调3.8倍, ntrC基因表达量上调10倍。NtrC是一个转录调控因子, 可以正向调控glnA基因的表达, glnA基因的作用便是催化L-谷氨酸和NH3合成L-谷氨酰胺, 从而细菌快速利用氮源[35]。而在ΔglnA菌株中NtrC的高表达或许是因为glnA的敲除解除了谷氨酰胺对NtrC的反馈抑制, 提高了NtrC的表达。发酵12 h时, ΔglnA菌株中, galU、crdR、crdS基因以及ntrB、ntrC基因表达量明显降低, galU基因下调了3.8倍, crdR基因下调了1.6倍, crdS基因下调了33倍, ntrB和ntrC也分别下调了3倍和4倍。
从实时荧光定量PCR结果(图5)来看, glnA基因敲除能够提高了fixN基因、exoC基因、以及ntrC基因表达, 其中ntrC基因编码的NtrC是NtrBC中的核心蛋白质, 而NtrBC参与氮代谢, 且NtrC调控的基因正是glnA, 在ΔglnA菌株中, crdR和crdS基因表达量同样是降低的, 且galU基因表达量也是降低的, GalU的产物UDP-葡萄糖是可得然胶合成的底物, galU基因表达降低, 导致UDP-葡萄糖减少, 从而影响了可得然胶合成。综合来看, glnA基因可以影响UDP-葡萄糖的合成和crdR和crdS的表达, 两者共同作用使得ΔglnA菌株中可得然胶产量减少。
3 结论
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产可得然胶土壤杆菌CGMCC11546菌株中, 基因glnA被敲除之后可得然胶产量降低了60%。土壤杆菌CGMCC11546菌株、ΔglnA菌株在发酵过程中生长代谢情况均正常, 可得然胶产量降低并不是因为菌体生长不好、受损引起的。在发酵过程中氨基氮消耗情况是一致的, ΔglnA菌株和土壤杆菌CGMCC11546生长一致。糖消耗方面, ΔglnA菌株与土壤杆菌CGMCC11546在12 h之前无明显差异, 12 h之后, ΔglnA菌株糖的消耗开始小于CGMCC11546菌株, 在15 h之后ΔglnA菌株与土壤杆菌CGMCC11546之前出现明显差异, 土壤杆菌CGMCC11546蔗糖消耗比ΔglnA菌株明显增加。综合生长曲线、可得然胶粗产量、氨基氮和糖结果, 初步得到glnA基因的敲除不影响菌体生长, 但是影响了可得然胶的合成。从定量结果来看, glnA基因敲除能够提高fixN基因的表达, 同时在产胶期exoC基因表达增加, glnA基因可以影响细胞中能量的合成这也与之前预测的基因功能吻合。但是在ΔglnA菌株中, 编码可得然胶合酶催化亚基的crdS基因与调控基因crdR表达量均减少, 这就可以解释ΔglnA菌株可得然胶产量降低是因为crdR和crdS基因表达量下调引起的。在ΔglnA菌株中, crdR和crdS基因表达量同样是降低的, 且galU基因表达量也是降低的, GalU的产物UDP-葡萄糖是可得然胶合成的底物, galU基因表达降低, 导致UDP-葡萄糖减少, 从而影响了可得然胶合成。综合来看, glnA基因可以降低UDP-葡萄糖的合成和crdR和crdS的表达, 两者共同作用使得ΔglnA菌株中可得然胶产量减少, 这与已经报道的结果相吻合[40]。本研究表明glnA基因敲除后, 与氮代谢相关的基因表达水平均下调, 氮代谢受阻而引起可得然胶合成受限, 最终导致可得然胶合成量减小。glnA基因参与氮代谢, 其编码产物为谷氨酰胺合成酶, 催化L-谷氨酸和NH3合成L-谷氨酰胺。前期研究中发现glnA基因突变可得然胶产量明显下降(数据未发表), 本研究通过对glnA基因进行敲除, 初步研究其对可得然胶合成的影响, 为后续提高可得然胶产量提供分子生物学基础。
基金
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  • 宁夏自然科学基金项目(2023AAC03735)
  • 2023年度宁夏回族自治区青年科技托举人才培养项目(宁科协发组字〔2024〕6号)
  • 宁夏科技惠民项目(2024CMG03049)
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doi: 10.19812/j.cnki.jfsq11-5956/ts.20250120006
  • 接收时间：2025-01-20
  • 首发时间：2025-07-17
  • 出版时间：2025-05-15
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  • 收稿日期：2025-01-20
基金
宁夏自然科学基金项目(2023AAC03735)
2023年度宁夏回族自治区青年科技托举人才培养项目(宁科协发组字〔2024〕6号)
宁夏科技惠民项目(2024CMG03049)
作者信息
    1.宁夏计量质量检验检测研究院, 银川 750000
    2.东华大学生物与医学工程学院, 上海 201620
    3.华东师范大学生命科学学院, 上海 200241

通讯作者:

* 姚博伟(1993—), 女, 硕士, 工程师, 主要研究方向为应用微生物及食品质量安全。E-mail:
* 杨雪霞(1971—), 女, 博士, 副教授, 主要研究方向为应用微生物与酶学研究。E-mail:
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https://castjournals.cast.org.cn/joweb/spaq/CN/10.19812/j.cnki.jfsq11-5956/ts.20250120006
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2种不同金属材料的力学参数
科
Family		 属数
Number of
genus		 种数
Number of
species		 占总种数比例
Percentage of
total species (%)		 属
Genus		 种数
Number of
species		 占总种数比例
Percentage of total
species (%)
鹅膏菌科Amanitaceae 2 11 5.26 鹅膏菌属 Amanita 10 4.78
小菇科 Mycenaceae 2 12 5.74 丝盖伞属 Inocybe 5 2.39
多孔菌科 Polyporaceae 8 14 6.70 蜡蘑属 Laccaria 5 2.39
红菇科 Russulaceae 3 23 11.00 小皮伞属 Marasmius 6 2.87
小菇属 Mycena 11 5.26
光柄菇属 Pluteus 5 2.39
红菇属 Russula 17 8.13
栓菌属 Trametes 5 2.39
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