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Article(id=1226460578919658394, tenantId=1146029695717560320, journalId=1192105938417971205, issueId=1226460576751206672, articleNumber=null, orderNo=null, doi=10.13343/j.cnki.wsxb.20250057, pmid=null, cstr=null, oa=null, hot=null, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=research-article, receivedDate=1737302400000, receivedDateStr=2025-01-20, revisedDate=null, revisedDateStr=null, acceptedDate=1740931200000, acceptedDateStr=2025-03-03, onlineDate=1770340588550, onlineDateStr=2026-02-06, pubDate=1754236800000, pubDateStr=2025-08-04, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1770340588550, onlineIssueDateStr=2026-02-06, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1770340588550, creator=13701087609, updateTime=1770340588550, updator=13701087609, issue=Issue{id=1226460576751206672, tenantId=1146029695717560320, journalId=1192105938417971205, year='2025', volume='65', issue='8', pageStart='1', pageEnd='3812', issueExtLink='null', onlineDate='null', pubDate='null', beforeIssueId=null, nextIssueId=null, price=null, status=1, issueComplete=1, articleOrder=1, issueType=-1, specialIssue=null, createTime=1770340588033, creator=13701087609, updateTime=1770363610188, updator=13701087609, preIssue=null, nextIssue=null, ext={EN=IssueExt(id=1226557138735117113, tenantId=1146029695717560320, journalId=1192105938417971205, issueId=1226460576751206672, language=EN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=), CN=IssueExt(id=1226557138735117114, tenantId=1146029695717560320, journalId=1192105938417971205, issueId=1226460576751206672, language=CN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=)}, issueFiles=null}, startPage=3657, endPage=3670, ext={EN=ArticleExt(id=1226460579263591331, articleId=1226460578919658394, tenantId=1146029695717560320, journalId=1192105938417971205, language=EN, title=Yeast asci lysate as an immunostimulatory dietary supplement, columnId=1192149543992045670, journalTitle=Acta Microbiologica Sinica, columnName=Research Article, runingTitle=null, highlight=null, articleAbstract=

Yeast β-glucan has been widely used as a dietary supplement for its multiple biological activities, including immune stimulation. Previous studies in our laboratory have shown that yeast chs3Δ spores exposed in the glucan layer can activate immune effects more effectively than trophoblasts. However, chs3Δ spores are formed within the ascospores and are encapsulated by the ascospore cell wall, which greatly limits their direct application. [Objective] To explore whether the lysates of chs3Δ asci have the potential to be used as novel immune-stimulating dietary supplements. [Methods] We prepared chs3Δ asci lysates by ultrasonic freeze drying, enzymatic freeze drying, and enzymatic-assisted ultrasonic freeze drying and then explored their immune effects in depth. [Results] The chs3Δ asci lysate prepared by ultrasonic freeze drying induced the highest level of inflammatory cytokines compared with the vegetative cell lysate and chs3Δ spores, and the asci lysate achieved immunostimulatory responses through activation of Dectin-1 receptor. Further studies showed that the chs3Δ asci lysate also exhibited immunostimulatory activity and trained immunity-inducing ability in mice. [Conclusion] The chs3Δ asci lysate has great potential as a highly effective immunostimulatory dietary supplement.

, correspAuthors=Nakanishi Hideki, authorNote=null, correspAuthorsNote=
*E-mail:
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酵母β-葡聚糖具有免疫刺激等多种生物学活性，已被广泛用作膳食补充剂。我们前期研究已表明，葡聚糖层外露的酵母chs3Δ孢子比营养细胞能更有效地激活免疫效应。然而，chs3Δ孢子是在子囊内形成的，且被子囊细胞壁所包裹，这极大地限制了其直接应用。 【目的】 通过裂解chs3Δ子囊，探究其子囊裂解物是否具有作为新型免疫刺激性膳食补充剂的潜力。 【方法】 采用超声冻干、酶解冻干、酶解辅助超声冻干3种方法制备chs3Δ子囊裂解物，并对其免疫效应进行深入探究。 【结果】 与营养细胞裂解物和chs3Δ孢子相比，采用超声冻干法制备的chs3Δ子囊裂解物诱导的炎性细胞因子水平最高，且该裂解物通过激活C型凝集素1 (Dectin-1)受体实现免疫刺激反应。进一步研究显示，chs3Δ子囊裂解物在小鼠体内也展现出免疫刺激活性及训练免疫诱导能力。 【结论】 chs3Δ子囊裂解物具备作为高效免疫刺激性膳食补充剂的巨大潜力。

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作者贡献声明

杜聪聪：实验设计、数据核查、论文撰写；李雨晴：辅助动物实验；刘国玉：辅助突变菌株构建、菌株筛选；周小满：论文核查；中西秀树：数据核查、论文核查、提供基金支持。

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Fish & Shellfish Immunology, 2006, 20(5): 682-692., articleTitle=Immunostimulatory nature of β-glucans and baker’s yeast in gnotobiotic Artemia challenge tests, refAbstract=null), Reference(id=1226596300423082040, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226460578919658394, doi=null, pmid=null, pmcid=null, year=2017, volume=104, issue=null, pageStart=367, pageEnd=376, url=null, language=null, rfNumber=[30], rfOrder=31, authorNames=ISHIMOTO Y, ISHIBASHI KI, YAMANAKA D, ADACHI Y, KANZAKI K, OKITA K, IWAKURA Y, OHNO N, journalName=International Journal of Biological Macromolecules, refType=null, unstructuredReference=ISHIMOTO Y, ISHIBASHI KI, YAMANAKA D, ADACHI Y, KANZAKI K, OKITA K, IWAKURA Y, OHNO N. Modulation of an innate immune response by soluble yeast β-glucan prepared by a heat degradation method[J]. International Journal of Biological Macromolecules, 2017, 104: 367-376., articleTitle=Modulation of an innate immune response by soluble yeast β-glucan prepared by a heat degradation method, refAbstract=null), Reference(id=1226596300544716860, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226460578919658394, doi=null, pmid=null, pmcid=null, year=2022, volume=100, issue=null, pageStart=100, pageEnd=null, url=null, language=null, rfNumber=[31], rfOrder=32, authorNames=DAVIES SJ, ESPOSITO G, MAPIYE C, VILLOT C, CHEVAUX E, RAFFRENATO E, journalName=Journal of Animal Science, refType=null, unstructuredReference=DAVIES SJ, ESPOSITO G, MAPIYE C, VILLOT C, CHEVAUX E, RAFFRENATO E. 225 effects of live yeast supplement in pre-weaning and finishing diets of Holstein bull calves on meat production and quality[J]. Journal of Animal Science, 2022, 100(Supplement_3): 100., articleTitle=225 effects of, refAbstract=supplementinga), Reference(id=1226596300670545987, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226460578919658394, doi=null, pmid=null, pmcid=null, year=2018, volume=236, issue=null, pageStart=149, pageEnd=158, url=null, language=null, rfNumber=[32], rfOrder=33, authorNames=SOUSA DO, OLIVEIRA CA, VELASQUEZ AV, SOUZA JM, CHEVAUX E, MARI LJ, SILVA LFP, journalName=Animal Feed Science and Technology, refType=null, unstructuredReference=SOUSA DO, OLIVEIRA CA, VELASQUEZ AV, SOUZA JM, CHEVAUX E, MARI LJ, SILVA LFP. Live yeast supplementation improves rumen fibre degradation in cattle grazing tropical pastures throughout the year[J]. Animal Feed Science and Technology, 2018, 236: 149-158., articleTitle=Live yeast supplementation improves rumen fibre degradation in cattle grazing tropical pastures throughout the year, refAbstract=null)], funds=[Fund(id=1226596295364752049, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226460578919658394, awardId=32071467, language=EN, fundingSource=National Natural Science Foundation of China(32071467), fundOrder=null, country=null), Fund(id=1226596295465415359, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226460578919658394, awardId=32071467, language=CN, fundingSource=国家自然科学基金(32071467), fundOrder=null, country=null)], companyList=[AuthorCompany(id=1226596286422495400, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226460578919658394, xref=1., ext=[AuthorCompanyExt(id=1226596286443466921, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226460578919658394, companyId=1226596286422495400, language=EN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=1.School of Biotechnology, Jiangnan University, Wuxi, Jiangsu, China), AuthorCompanyExt(id=1226596286485409964, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226460578919658394, companyId=1226596286422495400, language=CN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=1.江南大学 生物工程学院，江苏 无锡)]), AuthorCompany(id=1226596286598656177, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226460578919658394, xref=2., ext=[AuthorCompanyExt(id=1226596286636404915, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226460578919658394, companyId=1226596286598656177, language=EN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=2.Beijing Key Laboratory of the Innovative Development of Functional Staple and Nutritional Intervention for Chronic Disease, China National Research Institute of Food and Fermentation Industries Co. , Ltd. , Beijing, China), AuthorCompanyExt(id=1226596286669959350, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226460578919658394, companyId=1226596286598656177, language=CN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=2.中国食品发酵工业研究院有限公司，功能主食创制与慢病营养干预北京市重点实验室，北京)])], figs=[ArticleFig(id=1226596291644404147, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226460578919658394, language=EN, label=Figure 1, caption=Preparation of chs3Δ asci lysate by ultrasonic freeze drying and analysis of their immunostimulatory effects. A: Schematic diagram of the preparation of the asci lysate; B: ELISA to detect that chs3Δ asci lysate prepared under different conditions stimulated the secretion of TNF-α by murine-derived macrophages; C: ELISA to detect that chs3Δ asci lysate prepared under different conditions stimulated the secretion of IL-6 by murine-derived macrophages; D: ELISA to detect that chs3Δ asci lysate and chs3Δ spore stimulate the secretion of TNF-α level by murine-derived macrophages, respectively; E: ELISA to detect the secretion of IL-6 level by murine-derived macrophages stimulated by chs3Δ asci lysate and chs3Δ spores, respectively. *: P<0.05; **: P<0.01; ***: P<0.001; ns: No significant differences. The same as below., figureFileSmall=HZDDTCu5iiUqioMpudyw2Q==, figureFileBig=QixaW2dGE4Xtk1AUzmO+Xw==, tableContent=null), ArticleFig(id=1226596291795399100, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226460578919658394, language=CN, label=图1, caption=超声冻干法制备 chs3Δ子囊裂解物及其免疫刺激作用。A：子囊裂解物制备示意图；B：ELISA检测不同条件下制备的chs3Δ子囊裂解物刺激鼠源巨噬细胞分泌TNF-α；C：ELISA检测不同条件下制备的chs3Δ子囊裂解物刺激鼠源巨噬细胞分泌IL-6水平；D：ELISA检测chs3Δ子囊裂解物和chs3Δ孢子分别刺激鼠源巨噬细胞分泌TNF-α水平；E：ELISA检测chs3Δ子囊裂解物和chs3Δ孢子分别刺激鼠源巨噬细胞分泌IL-6水平。*：P<0.05；**：P<0.01；***：P<0.001；ns：无显著性差异。下同。, figureFileSmall=HZDDTCu5iiUqioMpudyw2Q==, figureFileBig=QixaW2dGE4Xtk1AUzmO+Xw==, tableContent=null), ArticleFig(id=1226596291996725706, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226460578919658394, language=EN, label=Figure 2, caption=chs3Δ asci lysate activate murine-derived macrophages immune system. A: ELISA for chs3Δ asci lysate and vegetative cell lysate to stimulate murine-derived macrophages to secrete TNF-α levels; B: ELISA for chs3Δ asci lysate and vegetative cell lysate to stimulate murine-derived macrophages to secrete IL-6 levels; C: ELISA for each yeast asci lysate to stimulate murine-derived macrophages to secrete TNF-α levels, respectively; D: ELISA for each yeast asci lysate stimulates IL-6 secretion by murine-derived macrophages, respectively., figureFileSmall=WxyW7OmHbkNzfaL9lHSIfg==, figureFileBig=S931QyVs8AOsFLbCkozYZw==, tableContent=null), ArticleFig(id=1226596292126749146, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226460578919658394, language=CN, label=图2, caption=chs3Δ子囊裂解物激活鼠源巨噬细胞免疫系统。A：ELISA检测chs3Δ子囊裂解物和营养细胞裂解物刺激鼠源巨噬细胞分泌TNF-α水平；B：ELISA检测chs3Δ子囊裂解物和营养细胞裂解物刺激鼠源巨噬细胞分泌IL-6水平；C：ELISA检测各酵母子囊裂解物分别刺激鼠源巨噬细胞分泌TNF-α水平；D：ELISA检测各酵母子囊裂解物分别刺激鼠源巨噬细胞分泌IL-6水平。, figureFileSmall=WxyW7OmHbkNzfaL9lHSIfg==, figureFileBig=S931QyVs8AOsFLbCkozYZw==, tableContent=null), ArticleFig(id=1226596292252578277, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226460578919658394, language=EN, label=Figure 3, caption=chs3Δ asci lysate induce cytokines production of murine-derived macrophages via Dectin-1 signal. A: RT-qPCR to detect the relative expression of Dectin-1 mRNA; B: ELISA to detect the level of TNF-α secreted by murine-derived macrophages stimulated by chs3Δ asci lysate to Dectin-1 knockdown; C: ELISA to detect the level of IL-6 secreted by murine-derived macrophages stimulated by chs3Δ asci lysate to Dectin-1 knockdown., figureFileSmall=1Q5Pk84uoA6JUq8L7PY5PA==, figureFileBig=LlOMRGxxQnmiw6OCMcIlmw==, tableContent=null), ArticleFig(id=1226596292370018802, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226460578919658394, language=CN, label=图3, caption=chs3Δ子囊裂解物通过Dectin-1激活鼠源巨噬细胞分泌细胞因子。A：采用RT-qPCR检测Dectin-1 mRNA的相对表达量；B：利用ELISA检测chs3Δ子囊裂解物刺激Dectin-1敲低的鼠源巨噬细胞分泌TNF-α的水平；C：借助ELISA检测chs3Δ子囊裂解物刺激Dectin-1敲低的鼠源巨噬细胞分泌IL-6的水平。, figureFileSmall=1Q5Pk84uoA6JUq8L7PY5PA==, figureFileBig=LlOMRGxxQnmiw6OCMcIlmw==, tableContent=null), ArticleFig(id=1226596292479070722, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226460578919658394, language=EN, label=Figure 4, caption=chs3Δ asci lysate activate mouse peritoneal primary macrophage cell immune system. A: ELISA to detect the level of TNF-α secretion by primary macrophages stimulated by chs3Δ asci lysate and vegetative cell lysate; B: ELISA to detect the level of IL-6 secretion by primary macrophages stimulated by chs3Δ asci lysate and vegetative cell lysate; C: ELISA to detect the level of TNF-α secretion by primary macrophages stimulated by yeast asci lysate between them; D: ELISA for stimulation of primary macrophage secretion of IL-6 levels between yeast asci lysates., figureFileSmall=V+gZ0/i/nwfzRQfbYLFiNA==, figureFileBig=MoUqWG1XhMOW/4+zTaC3KA==, tableContent=null), ArticleFig(id=1226596292600705551, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226460578919658394, language=CN, label=图4, caption=chs3Δ子囊裂解物激活小鼠腹腔原代巨噬细胞免疫系统。A：利用ELISA检测chs3Δ子囊裂解物和营养细胞裂解物刺激原代巨噬细胞分泌TNF-α的水平；B：借助ELISA检测chs3Δ子囊裂解物和营养细胞裂解物刺激原代巨噬细胞分泌IL-6的水平；C：采用ELISA检测酵母子囊裂解物之间刺激原代巨噬细胞分泌TNF-α的水平；D：通过ELISA检测酵母子囊裂解物之间刺激原代巨噬细胞分泌IL-6的水平。, figureFileSmall=V+gZ0/i/nwfzRQfbYLFiNA==, figureFileBig=MoUqWG1XhMOW/4+zTaC3KA==, tableContent=null), ArticleFig(id=1226596292743311901, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226460578919658394, language=EN, label=Figure 5, caption=chs3Δ asci lysate induce cytokines production of mouse peritoneal primary macrophage cells via Dectin-1 signal. A: RT-qPCR to detect the relative expression of Dectin-1 mRNA; B: ELISA to detect the level of TNF-α secreted by Dectin-1 knockdown primary macrophages stimulated by chs3Δ asci lysate; C: ELISA to detect the level of IL-6 secreted by chs3Δ asci lysate stimulated by Dectin-1 knockdown primary macrophages., figureFileSmall=CE2K5Q+pwclP7jgbCFeI9Q==, figureFileBig=fJPSdeLGkNSqqNLv/aE9vw==, tableContent=null), ArticleFig(id=1226596292860752426, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226460578919658394, language=CN, label=图5, caption=chs3Δ子囊裂解物通过Dectin-1激活小鼠腹腔原代巨噬细胞分泌细胞因子。A：采用RT-qPCR检测Dectin-1 mRNA的相对表达量；B：利用ELISA检测chs3Δ子囊裂解物刺激Dectin-1敲低的原代巨噬细胞分泌TNF-α的水平；C：借助ELISA检测chs3Δ子囊裂解物刺激Dectin-1敲低的原代巨噬细胞分泌IL-6的水平。, figureFileSmall=CE2K5Q+pwclP7jgbCFeI9Q==, figureFileBig=fJPSdeLGkNSqqNLv/aE9vw==, tableContent=null), ArticleFig(id=1226596293020135991, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226460578919658394, language=EN, label=Figure 6, caption=chs3Δ asci lysate activate cells to secrete cytokines in vivo of mice. A: Values of body weight changes in mice during one week of gavage of chs3Δ asci lysate; B: ELISA for TNF-α levels in serum of chs3Δ asci lysate stimulated mice; C: ELISA for IL-6 levels in serum of chs3Δ asci lysate stimulated mice., figureFileSmall=L2YYPVTSuU04krUZ9QbHvw==, figureFileBig=eSLCVqetsEkHb7Wo8vcFpg==, tableContent=null), ArticleFig(id=1226596293141770820, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226460578919658394, language=CN, label=图6, caption=chs3Δ子囊裂解物在小鼠体内激活细胞分泌细胞因子。A：在灌胃chs3Δ子囊裂解物的一周内，小鼠的体重变化值；B：ELISA检测chs3Δ子囊裂解物刺激小鼠血清中的TNF-α水平；C：ELISA检测chs3Δ子囊裂解物刺激小鼠血清中的IL-6水平。, figureFileSmall=L2YYPVTSuU04krUZ9QbHvw==, figureFileBig=eSLCVqetsEkHb7Wo8vcFpg==, tableContent=null), ArticleFig(id=1226596293284377175, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226460578919658394, language=EN, label=Figure 7, caption=chs3Δ asci lysate induce trained immunity. A: Schematic of chs3Δ asci lysate training immunity; B: ELISA to detect TNF-α levels in serum from mice in activated and inactivated states; C: ELISA to detect IL-6 levels in serum from mice in activated and inactivated states; D: ELISA to detect TNF-α levels secreted by LPS-stimulated primary macrophages in mice; E: ELISA to detect IL-6 levels secreted by LPS-stimulated mouse primary macrophages., figureFileSmall=/m95FCvCsZFkXtuiiE5dFQ==, figureFileBig=QjF9EoiDlpi3Uf6IfEbuVA==, tableContent=null), ArticleFig(id=1226596293385040483, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226460578919658394, language=CN, label=图7, caption=chs3Δ子囊裂解物诱导训练免疫。A：chs3Δ子囊裂解物训练免疫示意图；B：ELISA检测小鼠激活状态和非激活状态下血清中的TNF-α水平；C：ELISA检测小鼠激活状态和非激活状态下血清中的IL-6水平；D：ELISA检测LPS刺激小鼠原代巨噬细胞分泌的TNF-α水平；E：ELISA检测LPS刺激小鼠原代巨噬细胞分泌的IL-6水平。, figureFileSmall=/m95FCvCsZFkXtuiiE5dFQ==, figureFileBig=QjF9EoiDlpi3Uf6IfEbuVA==, tableContent=null), ArticleFig(id=1226596293598950001, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226460578919658394, language=EN, label=Table 1, caption=

Double-stranded siRNA sequences

, figureFileSmall=null, figureFileBig=null, tableContent=
NamesiRNA sequences (5′→3′)
Dectin-1 siRNA (mouse)

GCUGUUACCUAUUUAGCUUdTdT;

AAGCUAAAUAGGUAACAGCdTdT

Control siRNA (mouse)

UUCUCCGAACGUGUCACGUtt;

ACGUGACACGUUCGGAGAAtt

GAPDH siRNA (mouse)

CACUCAAGAUUGUCAGCAAdTdT;

UUGCUGACAAUCUUGAGUGdTdT

), ArticleFig(id=1226596293716390522, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226460578919658394, language=CN, label=表1, caption=

双链siRNA的序列

, figureFileSmall=null, figureFileBig=null, tableContent=
NamesiRNA sequences (5′→3′)
Dectin-1 siRNA (mouse)

GCUGUUACCUAUUUAGCUUdTdT;

AAGCUAAAUAGGUAACAGCdTdT

Control siRNA (mouse)

UUCUCCGAACGUGUCACGUtt;

ACGUGACACGUUCGGAGAAtt

GAPDH siRNA (mouse)

CACUCAAGAUUGUCAGCAAdTdT;

UUGCUGACAAUCUUGAGUGdTdT

), ArticleFig(id=1226596293791888005, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226460578919658394, language=EN, label=Table 2, caption=

Target gene primer sequences

, figureFileSmall=null, figureFileBig=null, tableContent=
Primers namePrimer sequences (5′→3′)
mRNA-M-GAPDH-FGCCAAAAGGGTCATCATCTC
mRNA-M-GAPDH-RGTAGAGGCAGGGATGATGTTC
mRNA-M-Dectin-1-FGACTTCAGCACTCAAGACATCC
mRNA-M-Dectin-1-RTTGTGTCGCCAAAATGCTAGG
mRNA-M-β-actin-FGGCTGTATTCCCCTCCATCG
mRNA-M-β-actin-RCCAGTTGGTAACAATGCCATGT
), ArticleFig(id=1226596293892551319, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226460578919658394, language=CN, label=表2, caption=

目的基因的引物序列

, figureFileSmall=null, figureFileBig=null, tableContent=
Primers namePrimer sequences (5′→3′)
mRNA-M-GAPDH-FGCCAAAAGGGTCATCATCTC
mRNA-M-GAPDH-RGTAGAGGCAGGGATGATGTTC
mRNA-M-Dectin-1-FGACTTCAGCACTCAAGACATCC
mRNA-M-Dectin-1-RTTGTGTCGCCAAAATGCTAGG
mRNA-M-β-actin-FGGCTGTATTCCCCTCCATCG
mRNA-M-β-actin-RCCAGTTGGTAACAATGCCATGT
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酵母子囊裂解物作为免疫刺激性膳食的补充剂
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杜聪聪 1 , 李雨晴 1 , 刘国玉 1, 2 , 周小满 1 , 中西秀树 1, *
微生物学报 | 研究报告 2025,65(8): 3657-3670
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微生物学报 | 研究报告 2025, 65(8): 3657-3670
酵母子囊裂解物作为免疫刺激性膳食的补充剂
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杜聪聪1, 李雨晴1, 刘国玉1, 2, 周小满1, 中西秀树1, *
作者信息
  • 1.江南大学 生物工程学院，江苏 无锡
  • 2.中国食品发酵工业研究院有限公司，功能主食创制与慢病营养干预北京市重点实验室，北京
Yeast asci lysate as an immunostimulatory dietary supplement
Congcong DU1, Yuqing LI1, Guoyu LIU1, 2, Xiaoman ZHOU1, Nakanishi Hideki1, *
Affiliations
  • 1.School of Biotechnology, Jiangnan University, Wuxi, Jiangsu, China
  • 2.Beijing Key Laboratory of the Innovative Development of Functional Staple and Nutritional Intervention for Chronic Disease, China National Research Institute of Food and Fermentation Industries Co. , Ltd. , Beijing, China
出版时间: 2025-08-04 doi: 10.13343/j.cnki.wsxb.20250057
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摘要
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酵母β-葡聚糖具有免疫刺激等多种生物学活性，已被广泛用作膳食补充剂。我们前期研究已表明，葡聚糖层外露的酵母chs3Δ孢子比营养细胞能更有效地激活免疫效应。然而，chs3Δ孢子是在子囊内形成的，且被子囊细胞壁所包裹，这极大地限制了其直接应用。 【目的】 通过裂解chs3Δ子囊，探究其子囊裂解物是否具有作为新型免疫刺激性膳食补充剂的潜力。 【方法】 采用超声冻干、酶解冻干、酶解辅助超声冻干3种方法制备chs3Δ子囊裂解物，并对其免疫效应进行深入探究。 【结果】 与营养细胞裂解物和chs3Δ孢子相比，采用超声冻干法制备的chs3Δ子囊裂解物诱导的炎性细胞因子水平最高，且该裂解物通过激活C型凝集素1 (Dectin-1)受体实现免疫刺激反应。进一步研究显示，chs3Δ子囊裂解物在小鼠体内也展现出免疫刺激活性及训练免疫诱导能力。 【结论】 chs3Δ子囊裂解物具备作为高效免疫刺激性膳食补充剂的巨大潜力。

酿酒酵母  /  酵母β-葡聚糖  /  膳食补充剂  /  免疫刺激  /  C型凝集素1

Yeast β-glucan has been widely used as a dietary supplement for its multiple biological activities, including immune stimulation. Previous studies in our laboratory have shown that yeast chs3Δ spores exposed in the glucan layer can activate immune effects more effectively than trophoblasts. However, chs3Δ spores are formed within the ascospores and are encapsulated by the ascospore cell wall, which greatly limits their direct application. [Objective] To explore whether the lysates of chs3Δ asci have the potential to be used as novel immune-stimulating dietary supplements. [Methods] We prepared chs3Δ asci lysates by ultrasonic freeze drying, enzymatic freeze drying, and enzymatic-assisted ultrasonic freeze drying and then explored their immune effects in depth. [Results] The chs3Δ asci lysate prepared by ultrasonic freeze drying induced the highest level of inflammatory cytokines compared with the vegetative cell lysate and chs3Δ spores, and the asci lysate achieved immunostimulatory responses through activation of Dectin-1 receptor. Further studies showed that the chs3Δ asci lysate also exhibited immunostimulatory activity and trained immunity-inducing ability in mice. [Conclusion] The chs3Δ asci lysate has great potential as a highly effective immunostimulatory dietary supplement.

Saccharomyces cerevisiae  /  yeast β-glucan  /  dietary supplement  /  immune stimulation  /  Dectin-1
杜聪聪, 李雨晴, 刘国玉, 周小满, 中西秀树. 酵母子囊裂解物作为免疫刺激性膳食的补充剂. 微生物学报, 2025 , 65 (8) : 3657 -3670 . DOI: 10.13343/j.cnki.wsxb.20250057
Congcong DU, Yuqing LI, Guoyu LIU, Xiaoman ZHOU, Nakanishi Hideki. Yeast asci lysate as an immunostimulatory dietary supplement[J]. Acta Microbiologica Sinica, 2025 , 65 (8) : 3657 -3670 . DOI: 10.13343/j.cnki.wsxb.20250057
正文
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β-葡聚糖是一类由β-d-葡萄糖单元组成的直链多糖，其主链通过β-1,3糖苷键连接构成，侧链则由β-1,4或β-1,6键连接构成，其中酵母及大多数真菌衍生的β-葡聚糖在1、6位存在支链[1-2]。研究表明，酵母β-葡聚糖展现出多样化的生物活性特征，在激活细胞免疫应答、抗氧化性能、促进伤口愈合进程以及癌症治疗方面展现出潜在效用[3-4]。鉴于这些广泛的生物学效应，酵母β-葡聚糖在食品领域得到了广泛应用，在开发具有保健功能的食品和食品添加剂方面体现了其作为天然活性成分的重要价值[5-6]。
C型凝集素1 (Dectin-1)作为C型凝集素家族中的模式识别受体，在巨噬细胞和树突状细胞中高度表达[7-8]。作为β-葡聚糖的特异性识别受体，Dectin-1通过介导一系列生物学反应，有效抵御细菌、寄生虫及病毒等多种感染性疾病，在细胞免疫调节过程中发挥核心作用[9-10]。此外，有研究发现，酵母β-葡聚糖对细胞的免疫刺激效应尤为突出，不仅能够激活体内的免疫系统，还具备诱导并训练免疫应答的独特功能[11-12]。训练免疫是一种先天免疫细胞的长期功能重编程现象，由外源性或内源性刺激激活，并在恢复到非激活状态后对第二次相同的或不同的刺激产生更为快速和强烈的激活效应[13-14]。
酿酒酵母又称面包酵母，是一种公认的安全微生物。目前，酵母细胞及源自酵母细胞壁的β-葡聚糖成分均被广泛应用于膳食补充剂领域[15]。酵母细胞壁的β-葡聚糖由β-1,3糖苷键连接的主链与β-1,6糖苷键形成的分支共同构成[16]。此外，酵母细胞壁组成成分还包括甘露聚糖和微量的几丁质。甘露聚糖和β-葡聚糖构成了酵母细胞壁的双层结构，而β-葡聚糖层被甘露聚糖层所覆盖[17]。酿酒酵母的二倍体细胞在营养缺乏的情况下分化为休眠的单倍体孢子[18]。在产孢过程中，母细胞通过减数分裂产生4个单倍体核，随后这些核被孢子质膜和孢子壁分别包裹起来，形成4个独立的孢子，最终母细胞成为一个包裹着4个孢子的子囊。孢子壁由4层结构组成，从内到外依次是甘露聚糖层、β-葡聚糖层、壳聚糖层和二酪氨酸层[19]。在孢子壁的合成过程中，各层结构是按照一定顺序由内向外逐层构建的。其中，Dit1是合成二酪氨酸层的关键酶，因此突变体(dit1Δ孢子)无法形成二酪氨酸层，细胞壁最外层是壳聚糖层，其次是β-葡聚糖层[20]。此外，Chs3作为组装合成壳聚糖层的关键酶，被敲除后壳聚糖层缺失，进一步导致了二酪氨酸层形成受阻。因此，在chs3Δ孢子中，由于缺少了壳聚糖层和二酪氨酸层的覆盖，β-葡聚糖层会直接暴露于孢子表面[21]。
先前的研究已经揭示葡聚糖层外露的chs3Δ孢子具备免疫刺激活性，能够在体外通过Dectin-1受体介导有效激活细胞免疫反应[22]。然而，鉴于孢子形成于子囊内部并被子囊壁所包裹，且子囊壁的最内层同样富含β-葡聚糖。因此，推测chs3Δ子囊裂解物可能具有作为新型免疫刺激性膳食补充剂的潜力。为了验证这一假设，本研究采用超声冻干、酶解冻干以及酶解辅助超声冻干3种技术成功制备了具有生物活性的chs3Δ子囊裂解物；此外，进一步评估了chs3Δ子囊裂解物的免疫刺激活性及作为Dectin-1受体激活剂的潜力，以期为chs3Δ子囊裂解物应用于膳食补充剂增强免疫功能的研究提供参考依据。
1 材料与方法
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1.1 主要试剂和仪器
所有培养基配制原料及常用化合物购自国药集团化学试剂有限公司；酵母破壁酶，西格玛奥德里奇(上海)贸易有限公司；Lipo8000TM染试剂、青霉素-链霉素溶液、脂多糖，碧云天生物技术股份有限公司；DMEM高糖培养基、胎牛血清、Opti-MEM减血清培养基、PBS溶液，赛默飞仕尔科技有限公司；ELISA试剂盒，南京森贝伽生物科技有限公司；RNAiso PLUS，宝日医生物技术(北京)有限公司；三氯甲烷、异丙醇，生工生物工程(上海)股份有限公司；反转录试剂盒，爱博泰克生物科技有限公司。
台式水平恒温摇床、超净工作台，上海三发科学仪器有限公司；高速低温离心机，HITACHI公司；超声波细胞粉碎机，宁波新芝生物科技股份有限公司；冷冻干燥机，EYELA公司；多功能酶标仪，BioTek公司；细胞培养箱、实时荧光定量PCR仪，赛默飞仕尔科技有限公司。
1.2 培养基
YPAD培养基：酵母提取物10 g/L，胰蛋白胨20 g/L，腺嘌呤30 mg/L，葡萄糖20 g/L；YPAce培养基：酵母提取物10 g/L，胰蛋白胨20 g/L，腺嘌呤30 mg/L，乙酸钾20 g/L；产孢培养基：乙酸钾20 g/L。
1.3 实验酵母的培养与产孢
本研究所用菌株为酵母SK1 (高产孢型)背景的野生型菌株，在此基础上，通过同源重组的方法将chs3基因、dit1基因整个开放阅读框替换为包含标记基因his5+的DNA序列，成功构建出chs3Δ、dit1Δ突变体菌株[21-22]。酵母产孢参考文献[23]的方法进行制备：从培养酵母的YPAD平板上挑取单克隆酵母细胞，接种于5 mL YPAD培养基中，30 ℃、220 r/min培养10 h。取2 mL YPAD培养菌液接种于100 mL YPAce培养基中，30 ℃、220 r/min培养24 h后，7 000×g离心2 min收集菌体，随后用100 mL乙酸钾重悬菌体，30 ℃、220 r/min培养24 h进行产孢，产孢结束后7 000×g离心2 min收集沉淀为子囊，无菌水清洗2次用于后续实验。
1.4 子囊裂解物的制备
1.4.1 超声冻干法获取子囊裂解物
称取湿重约100 mg的子囊，使用3 mL预冷的无菌水重悬子囊，在48%功率下进行超声处理5 min，每个循环处理时间5 s、间歇时间2 s。超声结束后冷冻干燥处理24 h，-20 ℃冻存备用。
1.4.2 酶解冻干法获取子囊裂解物
称取湿重约100 mg的子囊，使用3 mL预冷的无菌水重悬子囊，吹散均匀。随后加入2 μL酵母破壁酶，37 ℃、220 r/min处理1 h，吸取2 μL混合液在显微镜下观察子囊壁已破裂。酶解结束后冷冻干燥处理24 h，-20 ℃冻存备用。
1.4.3 酶解辅助超声处理获取子囊裂解物
先采用上述酶解的方式处理子囊，待子囊壁破裂后，采用1.4.1节中的超声方式继续处理。最后冷冻干燥处理24 h，-20 ℃冻存备用。
1.5 孢子纯化
称取湿重约60 mg子囊于1.5 mL EP管中，加入1 mL山梨醇缓冲溶液，吹散均匀。随后加入1 μL酵母破壁酶，37 ℃、220 r/min处理1 h，吸取2 μL混合液在显微镜下观察子囊壁已破裂，4 ℃、7 000×g离心2 min收集沉淀。在此基础上，去除子囊壁碎片：首先使用0.5%吐温-20低速洗涤3次，随后无菌水低速洗除吐温-20，收集孢子沉淀，最后-20 ℃冻存备用。
1.6 哺乳动物细胞培养及刺激
本研究所用哺乳动物细胞为鼠源巨噬细胞RAW264.7 (中国科学院细胞库)。将-80 ℃冻存的鼠源巨噬细胞复苏，传代3代后进行实验。接种106个/mL细胞于12孔板培养，待贴壁后更换新鲜DMEM培养基。随后刺激细胞：每孔滴加7.2 mg/mL超声冻干的样品，混合均匀，5% CO2、37 ℃环境下静置培养24 h。最后收集细胞上清液，检测刺激后哺乳动物细胞的细胞因子分泌水平。
1.7 酶联免疫法检测蛋白表达
采用ELISA试剂盒测定超声冻干的样品刺激鼠源巨噬细胞以及小鼠原代腹腔巨噬细胞分泌的细胞因子水平，包括肿瘤坏死因子-α (tumor necrosis factor-α, TNF-α)和白介素-6 (interleukin-6, IL-6)；测定灌胃小鼠超声冻干的样品后小鼠血清中分泌的细胞因子水平，具体操作方法参照试剂盒说明书进行。本实验方案及实验过程在江南大学伦理与动物福利审查委员会的监管下进行，伦理审核批准号：JN.No20240630c0641101[314]。
1.8 siRNA转染细胞
基于Dectin-1基因序列，使用网络工具箱(http://biodev.extra.cea.fr/DSIR/DSIR.html)设计用于敲低靶向基因的22 bp的小干扰RNA (small interfering RNA, siRNA)序列，具体序列信息如表1所示。细胞转染前接种7×105个/mL巨噬细胞于12孔板培养，待细胞密度达到70%以备转染。转染前，每孔更换1 mL新鲜的DMEM高糖双抗培养基。随后准备无菌EP管，依次加入50 μL Opti-MEM® Medium、40 pmol Dectin-1 siRNA、40 pmol Control siRNA以及1.6 μL Lipo8000TM，用移液枪轻轻地吹打均匀，逐滴滴加于细胞上清中，随后提取细胞总RNA，取适量采用RT-qPCR检测Dectin-1 mRNA的相对表达量，其余-20 ℃冻存备用。
1.9 实时荧光定量PCR
采用实时荧光定量PCR (quantitative real-time PCR, RT-qPCR)测定哺乳动物细胞中目的基因的相对转录水平。在NCBI网站 (https://www.ncbi.nlm.nih.gov/)上查找鼠源相关目的基因，从而设计引物序列并委托天霖生物科技(上海)有限公司进行合成，具体引物信息如表2所示。
1.10 动物实验设计
1.10.1 动物实验分组及实验设计
选用从斯贝福(北京)生物技术有限公司订购的品系为C57BL/6N雌性小鼠，鼠龄6周，体重为17-22 g。本研究严格遵守国家标准实验动物福利伦理要求(实验方案及实验过程在江南大学伦理与动物福利审查委员会的监管下进行，伦理审核批准文号：JN.No20240630c0641101[314])。在实验期间小鼠自由进水并喂食标准饲料，饲养条件：温度20-26 ℃，相对湿度40%-70%，噪声≤60 dB，动物照度15-20 lx。经1周适应喂养后，将小鼠随机分为对照组(每天分别灌胃0.9%生理盐水)和实验组(每天分别灌胃营养细胞、营养细胞裂解物、chs3Δ孢子和chs3Δ子囊裂解物，灌胃浓度为12 mg/kg，体积均为200 μL)。持续一周实验后，对小鼠进行吸入麻醉，眼眶取血收集小鼠全血。血液静置1.5 h，在4 ℃、3 000×g条件下离心15 min，吸取血清后置于-80 ℃冻存备用。
1.10.2 小鼠原代巨噬细胞提取和免疫刺激
实验前连续3 d对小鼠腹腔注射1 mL 3%的巯基乙酸盐，随后将其腹腔剪开小口，皮肤拉向两侧，暴露腹膜，注射1 mL预冷PBS溶液，吹吸3次，最后吸取腹腔液分装于2 mL Ep管，3 000×g离心15 min收集原代巨噬细胞。在此基础上，使用1 mL DMEM高糖双抗培养基重悬细胞，并对细胞计数，随后接种2.5×106个/mL细胞于6 cm板，5% CO2、37 ℃环境下静置培养4 h后，待细胞贴壁更换新鲜培养基。随后刺激细胞：每孔滴加7.2 mg/mL野生型子囊裂解物、chs3Δ孢子、dit1Δ子囊裂解物和chs3Δ子囊裂解物，混合均匀，5% CO2、37 ℃环境下静置培养24 h。最后收集细胞上清液，检测刺激后哺乳动物细胞的细胞因子分泌水平。
1.10.3 脂多糖刺激原代巨噬细胞
待原代巨噬细胞贴壁，更换新鲜DMEM培养基。称取5 mg脂多糖粉末溶于5 mL PBS缓冲液中，配制成1 mg/mL母液于-20 ℃冻存。使用2 μg/mL脂多糖刺激细胞，混合均匀，5% CO2、37 ℃环境下静置培养24 h。最后收集细胞上清液，采用ELISA检测细胞因子水平。
1.11 数据处理
每个实验重复3次，取平均值。所有数据用GraphPad Prism 8.0软件统计和绘图。数据用平均数±标准差表示，采用单因素方差分析检测多组间数据。P<0.05表示差异具有统计学意义。*表示P<0.05，**表示P<0.01，***表示P<0.001，****表示P<0.000 1，ns代表无显著性差异。
2 结果与分析
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2.1 子囊裂解物最优制备方式
通过研究比较超声冻干、酶解冻干和酶解辅助超声冻干3种用于破裂子囊壁的手段，获得免疫活性最强的chs3Δ子囊裂解物。第1种方案是超声处理chs3Δ子囊使细胞壁破裂，随后将其裂解液冷冻干燥(图1A)，随后置于-20 ℃冻存备用；第2种方案是采用酵母溶壁酶处理chs3Δ子囊使其细胞壁破裂，随后将其裂解液冷冻干燥；第3种方案是使用酵母溶壁酶处理chs3Δ子囊，随后继续采用超声处理，最后将其裂解液冷冻干燥。3种方案对比结果如图1B、1C所示，采用超声冻干处理的chs3Δ子囊刺激鼠源巨噬细胞分泌TNF-α和IL-6水平最高(P<0.001)。此外，溶壁酶通常用于裂解酵母细胞壁，具有降解细胞壁上的β-葡聚糖的作用[24]。因此，酶解冻干和酶解辅助超声冻干获得的子囊裂解物免疫活性降低很可能是由于酶处理后减少了β-葡聚糖的含量。此外，本研究发现相较于chs3Δ孢子，超声冻干的chs3Δ子囊裂解物的免疫刺激性更强(图1D、1E)。上述结果表明通过超声冻干法获取的子囊裂解物能显著激活鼠源巨噬细胞免疫效应，且该方法简单及高效。因此本研究选择超声冻干法获取子囊裂解物进行深入研究。
根据营养细胞壁内层的β-葡聚糖能刺激细胞免疫反应[3]，本研究利用超声冻干手段获取chs3Δ营养细胞裂解物，探究其免疫刺激作用。如图2A、2B所示，营养细胞裂解物能够激活免疫反应(P<0.05)，但是其免疫刺激反应显著低于chs3Δ子囊裂解物的刺激作用(P<0.05)。分析可能是从chs3Δ子囊内释放的chs3Δ孢子对细胞产生了免疫刺激作用。此外，孢子随着子囊壁破裂而释放出来，由于野生型和突变体孢子的最外层细胞壁成分不同[19]，因此可能对细胞的免疫刺激效果不同。chs3Δ孢子最外层是β-葡聚糖，而dit1Δ孢子的β-葡聚糖被壳聚糖所覆盖，因此推测chs3Δ子囊裂解物的免疫活性最强。如图2C、2D所示，相较于dit1Δ子囊裂解物，chs3Δ子囊裂解物刺激鼠源巨噬细胞分泌TNF-α和IL-6的水平显著升高(P<0.01)。上述结果成功验证了推测，即chs3Δ子囊裂解物的免疫刺激作用最强。
2.2 chs3Δ子囊裂解物在体外通过Dectin-1激活鼠源巨噬细胞免疫反应
虽然上述结果表明chs3Δ子囊裂解物能激活鼠源巨噬细胞的免疫反应，但其具体机制尚不清晰。鉴于chs3Δ子囊裂解物含有β-葡聚糖，本研究尝试采用siRNA干扰方式降低鼠源巨噬细胞的β-葡聚糖受体Dectin-1f表达水平，随后通过检测鼠源巨噬细胞分泌细胞因子水平的变化，进而探究chs3Δ子囊裂解物对其免疫激活的潜在机制。如图3A所示，使用靶向性siRNA处理受体细胞后，其Dectin-1的表达水平显著下降(P<0.05)。结果发现，随着鼠源巨噬细胞Dectin-1表达水平的降低，chs3Δ子囊裂解物刺激细胞分泌TNF-α和IL-6的水平也随之显著降低(图3B、3C)。上述结果表明，chs3Δ子囊裂解物是通过Dectin-1介导激活鼠源巨噬细胞免疫反应的。
2.3 chs3Δ子囊裂解物在体内通过Dectin-1激活小鼠原代巨噬细胞免疫系统
本研究已发现chs3Δ子囊裂解物能在体外条件显著激活细胞免疫反应，为进一步探究其在体内是否同样对免疫有刺激作用，设计了小鼠原代细胞实验。首先探究chs3Δ子囊裂解物对原代巨噬细胞的免疫激活作用。如图4A、4B所示，与chs3Δ营养细胞裂解物相比，chs3Δ子囊裂解物刺激原代巨噬细胞分泌的TNF-α和IL-6水平显著升高(P<0.01)。在此基础上，与野生型子囊裂解物及突变体dit1Δ子囊裂解物相比，chs3Δ子囊裂解物刺激原代巨噬细胞分泌TNF-α和IL-6的水平也显著升高(P<0.01)，此外chs3Δ子囊裂解物的免疫刺激活性高于chs3Δ孢子(图4C、4D)，这很可能是因为子囊壁上的β-葡聚糖具有较高的免疫活性。此结果与上述体外结果一致，表明chs3Δ子囊裂解物是一种具有生物活性的免疫刺激剂，并且能激活小鼠原代巨噬细胞免疫反应。
为探究chs3Δ子囊裂解物在体内激活小鼠原代巨噬细胞免疫反应是否同样与Dectin-1信号激活有关，本研究首先敲低原代巨噬细胞Dectin-1的表达水平。如图5A所示，Dectin-1的表达水平显著下降(P<0.05)。随着Dectin-1表达水平降低(图5A)，chs3Δ子囊裂解液刺激原代巨噬细胞分泌TNF-α和IL-6水平也随之显著降低(图5B、5C)。上述结果表明，chs3Δ子囊裂解液刺激原代巨噬细胞产生免疫反应是由Dectin-1信号介导的。
2.4 chs3Δ子囊裂解物在体内对小鼠免疫系统激活的影响
为探究chs3Δ子囊裂解物对小鼠整体免疫系统的影响，对品系为C57BL/6N小鼠口服chs3Δ子囊裂解物。如图6A所示，口服酵母子囊裂解物与生理盐水组相比，小鼠的体重变化量无显著性差异，表明chs3Δ子囊裂解物对小鼠的健康无显著影响。采用ELISA检测小鼠血清中的细胞因子分泌水平，结果如图6B、6C所示，chs3Δ子囊裂解物的小鼠血清中检测到TNF-α和IL-6水平均显著增加，这是因为与chs3Δ孢子相比，位于chs3Δ子囊壁上的β-葡聚糖在小鼠体内也具有高度的免疫活性(P<0.01)。上述结果表明，chs3Δ子囊裂解物作为免疫刺激剂激活了小鼠体内的免疫反应。
2.5 chs3Δ子囊裂解物诱导训练免疫
Xin等[25]和Vetvicka等[26]发现酵母β-葡聚糖能够诱导训练免疫反应，从而增强对二次感染疾病的抵抗力。研究表明，酵母β-葡聚糖对小鼠产生训练免疫后，脂多糖(lipopolysaccharides, LPS)二次刺激小鼠原代巨噬细胞后，诱导其产生更强烈的免疫反应和高水平的细胞因子[27-28]。因此，本研究探究chs3Δ子囊裂解物是否能对小鼠产生训练免疫反应，具体实验流程如图7A所示，首先小鼠连续7 d口服chs3Δ子囊裂解物以激活小鼠免疫反应，随后间歇7 d，待小鼠的免疫反应恢复至非激活状态。在此基础上，提取小鼠腹腔原代巨噬细胞，并使用LPS刺激进行二次刺激。
如图7B、7C所示，在灌胃7 d后检测小鼠血清中的细胞因子水平，发现chs3Δ子囊裂解物显著激活了小鼠免疫反应，这与上述结果一致(图6B、6C)。在7 d间歇后，采用ELISA法检测小鼠血清中的细胞因子水平，发现口服chs3Δ子囊裂解物的小鼠免疫反应已恢复至非激活状态(图7B、7C)。随后采用LPS二次刺激原代巨噬细胞，结果如图7D、7E所示，在chs3Δ子囊裂解物刺激的小鼠体内提取的原代巨噬细胞在LPS的二次刺激后，其分泌的TNF-α和IL-6水平均显著提高。上述结果表明，chs3Δ子囊裂解物能够诱导训练免疫反应。
3 讨论与结论
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酵母β-葡聚糖被广泛用于膳食补充剂，其具有免疫刺激性[15]。研究表明，酵母β-葡聚糖作为免疫刺激剂能够激活细胞免疫系统，诱导鼠源巨噬细胞分泌促炎细胞因子TNF-α和IL-6[29-30]。目前，市场上商业化酵母β-葡聚糖主要来源于酿酒酵母营养细胞壁，该细胞壁结构由葡聚糖层和甘露糖层构成，其中葡聚糖层被甘露糖层覆盖[17]。然而，Liu等[22]研究发现葡聚糖层外露的chs3Δ孢子比营养细胞能更有效地激活免疫效应。遗憾的是，这些孢子是在子囊内形成的，且被富含β-葡聚糖的子囊壁紧紧包裹[18]，这极大地限制了chs3Δ孢子的直接应用潜力。因此，本研究旨在探索出几种用于破裂子囊壁的手段，根据比较不同方式获取的chs3Δ子囊裂解物对鼠源巨噬细胞免疫刺激性的强弱，以筛选出最优的制备方案。发现通过超声冻干法获取的子囊裂解物激活鼠源巨噬细胞免疫效应最强，这是因为超声冻干法不仅能有效裂解chs3Δ孢子的子囊壁，释放出大量的β-葡聚糖，同时还能保持β-葡聚糖结构的完整性。这一发现不仅为chs3Δ孢子的高效利用提供了新思路，也为开发新型免疫刺激剂奠定了坚实基础。
研究表明酵母营养细胞和其细胞壁内层的β-葡聚糖作为膳食补充剂，能有效提高机体免疫能力[31-32]。目前，酵母chs3Δ孢子作为一种无毒性的β-葡聚糖微球，在体外展现出了比营养细胞更为强大的免疫激活能力[22]。本研究通过小鼠口服chs3Δ孢子实验设计，进一步确认了其不仅安全无毒，而且在体内展现出的免疫刺激性也超越了营养细胞。鉴于chs3Δ孢子被子囊壁包裹，子囊壁富含β-葡聚糖这一免疫刺激剂的特点[17]，本研究采取了破除子囊壁的策略，以深入探究chs3Δ子囊裂解物的免疫刺激活性。先前的研究已经揭示chs3Δ孢子能够在体外通过Dectin-1受体介导，有效激活细胞免疫反应[22]。此外，它能通过Dectin-1受体显著激活细胞免疫效应，诱导细胞因子的分泌，并通过训练免疫机制增强小鼠的免疫防御能力。因此，推测chs3Δ子囊裂解物具有作为一种新型免疫刺激性膳食补充剂的潜力。
综上所述，本研究通过采用超声冻干这一简便且高效的技术手段，成功制备了具有免疫刺激活性的chs3Δ子囊裂解物。在体外实验与体内模型中均验证了它作为有效免疫刺激剂的潜能，其能够显著激活细胞免疫反应。进一步的研究揭示，chs3Δ子囊裂解物是Dectin-1受体的激动剂，并能通过训练免疫机制增强小鼠对抗感染的能力。值得注意的是，口服chs3Δ子囊裂解物对小鼠未显示出明显的毒性效应，且其免疫刺激活性相较于已报道的营养细胞更为突出。此外，chs3Δ子囊裂解物的制备流程简易、成本效益高，具备良好的工业化生产潜力。基于上述研究发现，chs3Δ子囊裂解物在作为膳食补充剂以提升人体免疫力方面展现出了广阔的应用前景。这不仅为免疫调节剂的开发提供了新的思路，也为利用微生物资源促进健康提供了科学依据和实践路径。
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  • 国家自然科学基金(32071467)
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doi: 10.13343/j.cnki.wsxb.20250057
  • 接收时间：2025-01-20
  • 首发时间：2026-02-06
  • 出版时间：2025-08-04
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  • 收稿日期：2025-01-20
  • 录用日期：2025-03-03
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National Natural Science Foundation of China(32071467)
国家自然科学基金(32071467)
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    1.江南大学 生物工程学院，江苏 无锡
    2.中国食品发酵工业研究院有限公司，功能主食创制与慢病营养干预北京市重点实验室，北京

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Number of
species		 占总种数比例
Percentage of
total species (%)		 属
Genus		 种数
Number of
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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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