Article(id=1280817534341465067, tenantId=1146029695717560320, journalId=1192105938417971205, issueId=1280817479555462000, articleNumber=null, orderNo=null, doi=10.13343/j.cnki.wsxb.20250819, pmid=null, cstr=null, oa=null, hot=null, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=research-article, receivedDate=1761840000000, receivedDateStr=2025-10-31, revisedDate=null, revisedDateStr=null, acceptedDate=1770912000000, acceptedDateStr=2026-02-13, onlineDate=1783300296625, onlineDateStr=2026-07-06, pubDate=1783094400000, pubDateStr=2026-07-04, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1783300296625, onlineIssueDateStr=2026-07-06, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1783300296625, creator=13701087609, updateTime=1783300296625, updator=13701087609, issue=Issue{id=1280817479555462000, tenantId=1146029695717560320, journalId=1192105938417971205, year='2026', volume='66', issue='7', pageStart='3121', pageEnd='3677', issueExtLink='null', onlineDate='null', pubDate='1783094400000', pubDateStr='2026-07-04', beforeIssueId=null, nextIssueId=null, price=null, status=1, issueComplete=1, articleOrder=1, issueType=1, specialIssue=null, createTime=1783300283564, creator='13701087609', updateTime=1783326087324, updator='13701087609', preIssue=null, nextIssue=null, articleTotal=null, ext={EN=IssueExt(id=1280925708813832745, tenantId=1146029695717560320, journalId=1192105938417971205, issueId=1280817479555462000, language=EN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=), CN=IssueExt(id=1280925708813832746, tenantId=1146029695717560320, journalId=1192105938417971205, issueId=1280817479555462000, language=CN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=)}, issueFiles=null, downloadFileDto=null}, startPage=3324, endPage=3336, ext={EN=ArticleExt(id=1280817534710563820, articleId=1280817534341465067, tenantId=1146029695717560320, journalId=1192105938417971205, language=EN, title=High-level expression and activity identification of chicken NK-lysin in Pichia pastoris, columnId=1192149543992045670, journalTitle=Acta Microbiologica Sinica, columnName=Research Article, runingTitle=null, highlight=null, articleAbstract=

Objective To achieve high-level expression and purification of the recombinant 9 kDa and 15 kDa chicken NK-lysin proteins (designated as cNKL9 and cNKL15) and evaluate their in vitro antibacterial and immunomodulatory activities. Methods The target genes—cNKL9 and cNKL15—were individually cloned into the pPIC9K vector to construct recombinant expression plasmids for heterologous expression in Pichia pastoris. After confirming the correct expression of the recombinant proteins via SDS-PAGE and Western blotting, we used nickel affinity chromatography to obtain highly purified recombinant cNKL9 and cNKL15 proteins. The in vitro antibacterial activitiesof recombinant proteins against Salmonella typhimurium and Escherichia coli O157:H7 were assessed by colony forming unit (CFU) assays. We measured the mRNA levels by RT-qPCR and analyzed the transcriptional changes of relevant cytokines after applying the recombinant proteins to the chicken macrophage cell line HD11. High-level expression of the recombinant proteins was achieved in a 15 L fermenter for scale-up culture via optimization of the high-density fermentation process. Results The P. pastoris expression plasmids pPIC9K-cNKL9 andpPIC9K-cNKL15 were successfully constructed. SDS-PAGE and Western blotting results revealed specific bands of recombinant cNKL9 and cNKL15 proteins in the culture supernatant, confirming the successful secretory expression of the target proteins. Antibacterial activity assay demonstrated that the recombinant cNKL9 protein exhibited significant inhibitory activities against both S. typhimurium and E. coli O157:H7, whereas the recombinant cNKL15 protein showed no obvious inhibitory effects on either pathogenic strain. RT-qPCR analysis showed that the recombinant cNKL15 protein significantly upregulated the transcription levels of cytokines such as CCL4 and CCL20 in HD11 cells, while the recombinant cNKL9 protein did not exhibit immunomodulatory activity. Furthermore, in the 15 L fermenter, the heterologous expression yields of cNKL9 and cNKL15 reached 0.64 g/L and 0.53 g/L, respectively. Conclusion For the first time, we achieved high-level expression of recombinant cNKL9 and cNKL15 proteins in the P. pastoris system. Specifically, cNKL9 exhibited inhibitory activities against S. typhimurium and E. coli O157:H7, while cNKL15 exerted immunomodulatory effects by upregulating the mRNA levels of cytokines in the chicken macrophage cell line HD11. These findings provide experimental evidence for the development of cNKL9 and cNKL15 as antibacterial or immunomodulatory agents, indicating their considerable application potential.

, authors=Sixuan CHEN, Meiyu SHI, Tao TU, Huoqing HUANG, Huiying LUO, Bin YAO, Yuan WANG, Honglian ZHANG, authorsList=Sixuan CHEN, Meiyu SHI, Tao TU, Huoqing HUANG, Huiying LUO, Bin YAO, Yuan WANG, Honglian ZHANG, authorCompany=null, correspAuthors=Yuan WANG, Honglian ZHANG, authorNote=null, correspAuthorsNote=, 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, fund=null), CN=ArticleExt(id=1280817535876580341, articleId=1280817534341465067, tenantId=1146029695717560320, journalId=1192105938417971205, language=CN, title=鸡NK-lysin在毕赤酵母中的高效表达及活性鉴定, columnId=1192149544164012138, journalTitle=微生物学报, columnName=研究报告, runingTitle=null, highlight=null, articleAbstract=

目的 高效表达并纯化9 kDa和15 kDa鸡NK-lysin重组蛋白(cNKL9和cNKL15)，评估其体外抗菌活性及免疫调节活性。 方法 将cNKL9和cNKL15目的基因分别克隆至pPIC9K载体中，构建重组表达质粒进行异源表达。经SDS-PAGE分析及Western blotting特异性验证，确认重组蛋白正确表达后，通过镍亲和层析法获得高纯度的cNKL9和cNKL15重组蛋白。以鼠伤寒沙门氏菌和大肠埃希氏菌O157:H7为靶标菌株，采用平板活菌计数法(CFU法)评估重组蛋白的体外抗菌活性；通过RT-qPCR技术检测mRNA表达水平，分析重组蛋白处理HD11鸡巨噬细胞系后相关细胞因子的转录水平变化。在15 L发酵罐放大培养体系中，通过优化高密度发酵工艺实现重组蛋白的高效表达。 结果 成功构建毕赤酵母重组表达质粒pPIC9K-cNKL9和pPIC9K-cNKL15。SDS-PAGE及Western blotting验证结果显示，培养上清中可检测到重组cNKL9和cNKL15蛋白的特异性条带，证实目标蛋白已实现正确分泌表达。抑菌活性检测结果表明，重组cNKL9蛋白对鼠伤寒沙门氏菌及大肠埃希氏菌O157:H7均具有显著抗菌活性，而重组cNKL15蛋白对上述2株病原菌无明显抑制作用。RT-qPCR结果显示，重组cNKL15蛋白显著上调HD11细胞中CCL4、CCL20等细胞因子的基因转录水平，重组cNKL9蛋白则无此免疫调节作用。此外，在15 L发酵罐放大培养体系中，cNKL9和cNKL15的异源表达量分别可达0.64 g/L和0.53 g/L。 结论 本研究在毕赤酵母系统中实现了cNKL9和cNKL15蛋白的高效重组表达。其中，cNKL9蛋白对鼠伤寒沙门氏菌和大肠埃希氏菌O157:H7具有抗菌活性，cNKL15蛋白能够通过上调鸡巨噬细胞HD11中细胞因子的mRNA表达水平发挥免疫调节功能。上述结果为cNKL9和cNKL15蛋白作为抗菌或免疫调节产品的开发提供了实验依据，具有良好的应用前景。

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Blood, 2010, 116(18): 3465-3474., articleTitle=Granulysin activates antigen-presenting cells through TLR4 and acts as an immune alarmin, refAbstract=null), Reference(id=1280925232189916027, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1280817534341465067, doi=null, pmid=null, pmcid=null, year=2020, volume=161, issue=3, pageStart=245, pageEnd=258, url=null, language=null, rfNumber=[33], rfOrder=32, authorNames=Sparrow EL, Fowler DW, Fenn J, Caron J, Copier J, Dalgleish AG, Bodman-Smith MD, journalName=Immunology, refType=null, unstructuredReference=Sparrow EL, Fowler DW, Fenn J, Caron J, Copier J, Dalgleish AG, Bodman-Smith MD. The cytotoxic molecule granulysin is capable of inducing either chemotaxis or fugetaxis in dendritic cells depending on maturation: a role for Vδ2⁺ γδ T cells in the modulation of immune response to tumour[J]. Immunology, 2020, 161(3): 245-258., articleTitle=The cytotoxic molecule granulysin is capable of inducing either chemotaxis or fugetaxis in dendritic cells depending on maturation: a role for Vδ2⁺ γδ T cells in the modulation of immune response to tumour, refAbstract=null)], funds=[Fund(id=1280925227815256914, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1280817534341465067, awardId=2022YFD1300701, language=EN, fundingSource=The National Key Research and Development Program of China(2022YFD1300701), fundOrder=null, country=null), Fund(id=1280925227886560083, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1280817534341465067, awardId=2022YFD1300701, language=CN, fundingSource=国家重点研发计划(2022YFD1300701), fundOrder=null, country=null), Fund(id=1280925227949474644, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1280817534341465067, awardId=32372915, language=EN, fundingSource=The National Natural Science Foundation of China(32372915), fundOrder=null, country=null), Fund(id=1280925228020777813, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1280817534341465067, awardId=32372915, language=CN, fundingSource=国家自然科学基金(32372915), fundOrder=null, country=null), Fund(id=1280925228087886678, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1280817534341465067, awardId=CARS-41, language=EN, fundingSource=The China Agriculture Research System(CARS-41), fundOrder=null, country=null), Fund(id=1280925228146606935, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1280817534341465067, awardId=CARS-41, language=CN, fundingSource=现代农业产业技术体系项目(CARS-41), fundOrder=null, country=null)], companyList=[AuthorCompany(id=1280925220273898254, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1280817534341465067, xref=null, ext=[AuthorCompanyExt(id=1280925220282286863, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1280817534341465067, companyId=1280925220273898254, language=EN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=State Key Laboratory of Animal Nutrition and Feeding, Institute of Animal Science, Chinese Academy of Agricultural Sciences, Beijing, China), AuthorCompanyExt(id=1280925220294869776, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1280817534341465067, companyId=1280925220273898254, language=CN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=中国农业科学院北京畜牧兽医研究所，畜禽营养与饲养全国重点实验室，北京)])], figs=[ArticleFig(id=1280925225088959300, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1280817534341465067, language=EN, label=Figure 1, caption=Construction of the pPIC9K-cNKL9 and pPIC9K-cNKL15 expression plasmids. A: Schematic diagram of pPIC9K-cNKL9 and pPIC9K-cNKL15 expression plasmids construction; B: Agarose gel electrophoresis analysis of PCR-amplified fragments of cNKL9 and cNKL15 genes; C: Agarose gel electrophoresis analysis for validation of recombinant plasmids pPIC9K-cNKL9/cNKL15 construction. The recombinant plasmids were verified by PCR amplification with the 5′AOX and 3′AOX primers derived from the pPIC9K vector. Lane M: DNA marker., figureFileSmall=qmcCQ6a377opWZI/PwGP1A==, figureFileBig=nyiFYL6yhgC7udQNGhKP/w==, tableContent=null), ArticleFig(id=1280925225160262469, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1280817534341465067, language=CN, label=图1, caption=pPIC9K-cNKL9/cNKL15 表达质粒的构建, figureFileSmall=qmcCQ6a377opWZI/PwGP1A==, figureFileBig=nyiFYL6yhgC7udQNGhKP/w==, tableContent=null), ArticleFig(id=1280925225256731462, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1280817534341465067, language=EN, label=Figure 2, caption=Expression and purification of recombinant cNKL9 and cNKL15 proteins in Pichia pastoris. A: SDS-PAGE analysis of the expression of recombinant cNKL9 and cNKL15 proteins in the culture supernatants from two positive clones; B: Western blotting verification of recombinant proteins in the culture supernatant of two positive clones using an anti-His antibody; C: SDS-PAGE detection of purified recombinant cNKL9 and cNKL15 proteins. Lanes 1-2: The culture supernatant of two transformants; Lane 1×: One volume of protein sample; Lane 2×: Two volumes of protein sample., figureFileSmall=O3kUofuqFnyQTH5Epd5qeQ==, figureFileBig=KbJbWbqeLhLGk2tPOuRagg==, tableContent=null), ArticleFig(id=1280925225336423239, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1280817534341465067, language=CN, label=图2, caption=重组蛋白cNKL9和cNKL15在毕赤酵母中的表达及纯化, figureFileSmall=O3kUofuqFnyQTH5Epd5qeQ==, figureFileBig=KbJbWbqeLhLGk2tPOuRagg==, tableContent=null), ArticleFig(id=1280925227035116360, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1280817534341465067, language=EN, label=Figure 3, caption=Antibacterial and immunomodulatory activities of recombinant cNKL9 and cNKL15 proteins. A: Relative survival rates of Salmonella typhimurium and Escherichia coli O157:H7 treated withrecombinant cNKL9 and cNKL15 proteins at different concentrations, compared with untreated control; B: mRNA expression levels of cytokines in HD11 cells treated with different concentrations of cNKL9 and cNKL15 proteins, detected by RT-qPCR. All data present the mean±SD from three independent experiments., figureFileSmall=n1i4ciImwEO+YxSZCln8gA==, figureFileBig=OPO6Dwpa2t24GyZJoVYBXw==, tableContent=null), ArticleFig(id=1280925227119002441, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1280817534341465067, language=CN, label=图3, caption=cNKL9和cNKL15重组蛋白的抗菌活性及免疫调节活性, figureFileSmall=n1i4ciImwEO+YxSZCln8gA==, figureFileBig=OPO6Dwpa2t24GyZJoVYBXw==, tableContent=null), ArticleFig(id=1280925227202888522, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1280817534341465067, language=EN, label=Figure 4, caption=Optimization of high-density fermentation process and expression analysis of recombinant Pichia pastoris strains for cNKL9 and cNKL15 proteins. A: Changes in wet cell weight of recombinant strains during methanol induction; B: Changes in crude protein content in the high-density fermentation supernatants of recombinant strains at different induction times; C: SDS-PAGE analysis of target protein expression levels in the fermentation supernatants of recombinant strains at different induction times., figureFileSmall=6ldjKy6P/zUbTsfe0o2l1g==, figureFileBig=5jrj85vPlgcbt/8mmaVTDw==, tableContent=null), ArticleFig(id=1280925227282580299, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1280817534341465067, language=CN, label=图4, caption=毕赤酵母重组cNKL9和cNKL15蛋白菌株的高密度发酵工艺优化与表达分析, figureFileSmall=6ldjKy6P/zUbTsfe0o2l1g==, figureFileBig=5jrj85vPlgcbt/8mmaVTDw==, tableContent=null), ArticleFig(id=1280925227349689164, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1280817534341465067, language=EN, label=Table 1, caption=

The sequences of the optimized cNKL9 and cNKL15 genes

, figureFileSmall=null, figureFileBig=null, tableContent=

Gene names	Sequences (5′→3′)
cNKL9	ATCAAGTGCAGATTTTGTGTTTCTTTGGTTAAGAAAGTCCAGAAAA TTGTTGGTGACGATCCAGACGAAGACGCTATCAACAACGCATTGAA CAAGGTTTGTTCCACTGGTAGAAGACAAAGATCTATTTGTAAGCAA CTTCTGAAGAAGTTGAGACAACAGTTGTCCGATGCCTTGCAGAACA ATGACGATCCTAGAGATGTTTGTACTACATTGGGTCTGTGC
cNKL15	TTCTCTGGCTTGACCCCTGAGCATTCTGCTCTGGCTAGAGCTCATCCT TGTGACGGCGAGCAATTTTGTCAGAACTTGGCACCAGAAGACCCAC AAGGAGATCAATTACTTCAGAGAGAAGAACTTGGTTTGATCTGTGAG TCTTGTAGAAAAATTATCCAAAAGTTGGAAGATATGGTTGGCCCTCAA CCTAATGAGGATACCGTTACTCAGGCCGCTTCTAGAGTTTGTGATAAG ATGAAGATTCTGAGAGGTGTTTGTAAAAAGATAATGAGAACCTTCTTG AGAAGAATCTCCAAGGACATCTTGACTGGTAAAAAGCCACAAGCTAT CTGTGTTGACATCAAGATCTGCAAGGAAAAGACCGGTTTGATT

), ArticleFig(id=1280925227412603725, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1280817534341465067, language=CN, label=表1, caption=

优化后的 cNKL9 和 cNKL15 基因序列

, figureFileSmall=null, figureFileBig=null, tableContent=

Gene names	Sequences (5′→3′)
cNKL9	ATCAAGTGCAGATTTTGTGTTTCTTTGGTTAAGAAAGTCCAGAAAA TTGTTGGTGACGATCCAGACGAAGACGCTATCAACAACGCATTGAA CAAGGTTTGTTCCACTGGTAGAAGACAAAGATCTATTTGTAAGCAA CTTCTGAAGAAGTTGAGACAACAGTTGTCCGATGCCTTGCAGAACA ATGACGATCCTAGAGATGTTTGTACTACATTGGGTCTGTGC
cNKL15	TTCTCTGGCTTGACCCCTGAGCATTCTGCTCTGGCTAGAGCTCATCCT TGTGACGGCGAGCAATTTTGTCAGAACTTGGCACCAGAAGACCCAC AAGGAGATCAATTACTTCAGAGAGAAGAACTTGGTTTGATCTGTGAG TCTTGTAGAAAAATTATCCAAAAGTTGGAAGATATGGTTGGCCCTCAA CCTAATGAGGATACCGTTACTCAGGCCGCTTCTAGAGTTTGTGATAAG ATGAAGATTCTGAGAGGTGTTTGTAAAAAGATAATGAGAACCTTCTTG AGAAGAATCTCCAAGGACATCTTGACTGGTAAAAAGCCACAAGCTAT CTGTGTTGACATCAAGATCTGCAAGGAAAAGACCGGTTTGATT

), ArticleFig(id=1280925227479712590, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1280817534341465067, language=EN, label=Table 2, caption=

Primers used for vector construction

, figureFileSmall=null, figureFileBig=null, tableContent=

Primer names	Primer sequences (5′→3′)
cNKL9-F	GAAAAAAGAGAGGCTGAAGCTATCAAGTGCAGATTTTGTGTTTC
cNKL9-R	CTTAATGGTGATGATGATGATGGCACAGACCCAATGTAGTAC
cNKL15-F	GAAAAAAGAGAGGCTGAAGCTGAACCACCTAGAGATGATCAC
cNKL15-R	CTTAATGGTGATGATGATGATGACCTTTGCACAGACCCAATGTAGTAC
5′AOX	GACTGGTTCCAATTGACAAGC
3′AOX	GCAAATGGCATTCTGACATCC

), ArticleFig(id=1280925227542627151, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1280817534341465067, language=CN, label=表2, caption=

载体构建所用引物

, figureFileSmall=null, figureFileBig=null, tableContent=

Primer names	Primer sequences (5′→3′)
cNKL9-F	GAAAAAAGAGAGGCTGAAGCTATCAAGTGCAGATTTTGTGTTTC
cNKL9-R	CTTAATGGTGATGATGATGATGGCACAGACCCAATGTAGTAC
cNKL15-F	GAAAAAAGAGAGGCTGAAGCTGAACCACCTAGAGATGATCAC
cNKL15-R	CTTAATGGTGATGATGATGATGACCTTTGCACAGACCCAATGTAGTAC
5′AOX	GACTGGTTCCAATTGACAAGC
3′AOX	GCAAATGGCATTCTGACATCC

), ArticleFig(id=1280925227618124624, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1280817534341465067, language=EN, label=Table 3, caption=

Primers used for RT-qPCR

, figureFileSmall=null, figureFileBig=null, tableContent=

Primer names	Primer sequences (5′→3′)
CCL4-F	GCTGCCCTTCAGCTTTG
CCL4-R	TCAGTTCAGTTCCATCTTGTTCATGTA
CCL20-F	GGCTTGAGCACCAAGAGTTTG
CCL20-R	CCCTTAATGACCTTCCGAGGC
IL-1β-F[23]	TGGGCATCAAGGGCTACA
IL-1β-R[23]	TCGGGTTGGTTGGTGATG
IL-8-F[23]	GGCTTGCTAGGGGAAATGA
IL-8-R[23]	AGCTGACTCTGACTAGGAAACTGT
TNF-α-F	TCAGGACAGCCTATGCCAAC
TNF-α-R	CTTGGTTGCTTTGTGCCTGG
GAPDH-F[23]	GGTGGTGCTAAGCGTGTTAT
GAPDH-R[23]	ACCTCTGTCATCTCTCCACA

), ArticleFig(id=1280925227689427793, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1280817534341465067, language=CN, label=表3, caption=

RT-qPCR所用引物

, figureFileSmall=null, figureFileBig=null, tableContent=

Primer names	Primer sequences (5′→3′)
CCL4-F	GCTGCCCTTCAGCTTTG
CCL4-R	TCAGTTCAGTTCCATCTTGTTCATGTA
CCL20-F	GGCTTGAGCACCAAGAGTTTG
CCL20-R	CCCTTAATGACCTTCCGAGGC
IL-1β-F[23]	TGGGCATCAAGGGCTACA
IL-1β-R[23]	TCGGGTTGGTTGGTGATG
IL-8-F[23]	GGCTTGCTAGGGGAAATGA
IL-8-R[23]	AGCTGACTCTGACTAGGAAACTGT
TNF-α-F	TCAGGACAGCCTATGCCAAC
TNF-α-R	CTTGGTTGCTTTGTGCCTGG
GAPDH-F[23]	GGTGGTGCTAAGCGTGTTAT
GAPDH-R[23]	ACCTCTGTCATCTCTCCACA

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