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Article(id=1226460584309342941, tenantId=1146029695717560320, journalId=1192105938417971205, issueId=1226460576751206672, articleNumber=null, orderNo=null, doi=10.13343/j.cnki.wsxb.20250100, pmid=null, cstr=null, oa=null, hot=null, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=research-article, receivedDate=1739376000000, receivedDateStr=2025-02-13, revisedDate=null, revisedDateStr=null, acceptedDate=1741881600000, acceptedDateStr=2025-03-14, onlineDate=1770340589834, onlineDateStr=2026-02-06, pubDate=1754236800000, pubDateStr=2025-08-04, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1770340589834, onlineIssueDateStr=2026-02-06, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1770340589834, creator=13701087609, updateTime=1770340589834, 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=3721, endPage=3730, ext={EN=ArticleExt(id=1226460584791687942, articleId=1226460584309342941, tenantId=1146029695717560320, journalId=1192105938417971205, language=EN, title=An efficient chitinolytic bacterium Bacillus cereus BSF-CH1: isolation, identification, and application in puparium biotransformation of Hermetia illucens, columnId=1192149543992045670, journalTitle=Acta Microbiologica Sinica, columnName=Research Article, runingTitle=null, highlight=null, articleAbstract=

The puparium of Hermetia illucens is rich in chitin and protein, while efficient and environmentally friendly utilization methods remain to be developed. [Objective] To isolate chitinolytic bacteria from the puparium pile of H. illucens and explore their potential in puparium biotransformation. [Methods] Strains were isolated by the plate screening method and identified by 16S rRNA gene sequencing. The 3,5-dinitrosalicylic acid (DNS) method was employed to determine the chitinase activity. Whole genome sequencing by PacBio HiFi was conducted to elucidate the degradation mechanism. The application potential of the strain was explored by puparium fermentation experiments. [Results] Among the seven isolated strains, Bacillus cereus BSF-CH1 showed the highest chitinase activity, reaching a maximum chitinase activity of 0.48 U/mL on the second day of fermentation. The genome of BSF-CH1 contained three chitinase genes, seven chitin deacetylase genes, and four chitodextrinase genes. Puparium biotransformation experiments showed that BSF-CH1 could degrade 47.2% of puparium mass within 7 days, with degradation rates of 64.6% and 59.1% for chitin and protein, respectively. [Conclusion] This study reports an efficient chitinolytic bacterium isolated from the puparium of H. illucens, providing new insights into the puparium biotransformation and having important implications for promoting the sustainable development of the H. illucens industry.

, correspAuthors=Yunsheng WANG, authorNote=null, correspAuthorsNote=
*E-mail:
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黑水虻(Hermetia illucens)养殖产生的蛹壳富含几丁质和蛋白质，但目前缺乏高效环保的利用方法。 【目的】 从黑水虻蛹壳堆中分离筛选几丁质降解菌，探索其在蛹壳生物转化中的应用潜力。 【方法】 使用平板筛选和16S rRNA基因测序进行分离和鉴定，采用3,5-二硝基水杨酸(3,5-dinitrosalicylic acid, DNS)法检测其酶活力。使用PacBio HiFi技术进行全基因组测序，以阐明其降解机制。通过蛹壳发酵实验探索其应用潜力。 【结果】 共鉴定得到7株分离菌，其中活性最强的菌株为蜡样芽孢杆菌(Bacillus cereus)BSF-CH1，该菌株在发酵第2天时几丁质酶活力达到最高值0.48 U/mL。全基因组测序结果显示BSF-CH1含有3个几丁质酶(chitinase)基因、7个几丁质脱乙酰酶(chitin deacetylase)基因及4个壳糊精酶基因(chitodextrinase)。蛹壳生物转化实验表明，BSF-CH1能在7 d内将蛹壳质量降解47.2%，几丁质和蛋白质的降解率分别达到64.6%和59.1%。 【结论】 本研究报道了从黑水虻蛹壳中分离的高效几丁质降解菌，为蛹壳的生物转化利用提供了新思路，对促进黑水虻产业的可持续发展具有重要意义。

, correspAuthors=王运生, authorNote=null, correspAuthorsNote=null, copyrightStatement=null, copyrightOwner=null, extLink=null, articleAbsUrl=null, sourceXml=fRV9CTVbKp21EDdUAVemaw==, magXml=HACgQ8ANjCMAA9wmoNLtnA==, pdfUrl=null, pdf=h/tEi9+ehZaj+NeCI1J71Q==, pdfFileSize=2334137, pdfExtLink=null, richHtmlUrl=null, mobilePdfUrl=null, reviewReport=null, pdfFirstPage=null, abstractGraph=SAvKYzpdByALdKlM2pHn4w==, abstractGraphContent=null, abstractVideo=null, citation=null, cebUrl=null, magXmlContent=dYPzQGM6yxLepp3fV11S5g==, mapNumber=null, authorCompany=null, fund=null, authors=

作者贡献声明

姚祥：研究构思和设计、实验操作、论文撰写与修改；刘倩：协助数据分析；王胜辉：协助几丁质降解菌筛选鉴定；杨健：协助蛋白质含量的检测；彭才望：指导蛋白质检测；陈武：指导几丁质降解菌筛选；尹丽娟：指导酶活力检测；戴良英：指导氨基酸含量检测；王运生：研究构思和设计、论文指导与修改。

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A: Colony morphology of BSF-CH1 on chitin plates; B: Gram staining results of BSF-CH1 (100×); C: Phylogenetic tree constructed based on the 16S rRNA gene sequence of BSF-CH1. Scale bar represents 0.01 nucleotide substitutions per site, the numbers in parentheses are the accession numbers of these strains’ 16S rRNA gene sequences in GenBank., figureFileSmall=qEDWjrhS23CR4i3nN66ESw==, figureFileBig=aYIVeJQssG9e5iHOcaI3Qg==, tableContent=null), ArticleFig(id=1226596293712196217, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226460584309342941, language=CN, label=图1, caption=几丁质降解菌BSF-CH1的分离与鉴定。A：BSF-CH1在几丁质平板的菌落形态；B：BSF-CH1的革兰氏染色结果(100×)；C：基于BSF-CH1 16S rRNA基因序列构建的系统发育树。标尺为每个位点有0.01个核苷酸取代，括号中的数字是这些菌株的16S rRNA基因序列在GenBank中的登录号。, figureFileSmall=qEDWjrhS23CR4i3nN66ESw==, figureFileBig=aYIVeJQssG9e5iHOcaI3Qg==, tableContent=null), ArticleFig(id=1226596293812859529, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226460584309342941, language=EN, label=Figure 2, caption=The changes in enzyme activity and pH during the fermentation process of BSF-CH1. A: The changes in enzyme activity during the fermentation process of BSF-CH1; B: The changes in pH during the fermentation process of BSF-CH1. Different lowercase letters indicate significant differences among treatments at P<0.05., figureFileSmall=TFj9BKaL9V4MoMI85/OLsw==, figureFileBig=A3sEApGvswDkOfk2p+8E9Q==, tableContent=null), ArticleFig(id=1226596293884162709, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226460584309342941, language=CN, label=图2, caption=BSF-CH1发酵过程中的酶活力和pH变化情况。A：BSF-CH1发酵过程中的酶活力变化情况；B：BSF-CH1发酵过程中的pH变化情况。, figureFileSmall=TFj9BKaL9V4MoMI85/OLsw==, figureFileBig=A3sEApGvswDkOfk2p+8E9Q==, tableContent=null), ArticleFig(id=1226596295251505826, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226460584309342941, language=EN, label=Figure 3, caption=Circular maps of BSF-CH1 genome. A: The circular map of the chromosome (chr) genome; B: The circular map of the plasmid 1 genome; C: The circular map of the plasmid 2 genome., figureFileSmall=4U/OCdG14ArPPd0iiz/Qqw==, figureFileBig=sjCKl5Mfhmyu/76ENj1cdg==, tableContent=null), ArticleFig(id=1226596295373140659, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226460584309342941, language=CN, label=图3, caption=BSF-CH1基因组圈图。A：染色体(chr)基因组圈图；B：质粒1 (plasmid 1)基因组圈图；C：质粒2 (plasmid 2)基因组圈图。, figureFileSmall=4U/OCdG14ArPPd0iiz/Qqw==, figureFileBig=sjCKl5Mfhmyu/76ENj1cdg==, tableContent=null), ArticleFig(id=1226596295477998273, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226460584309342941, language=EN, label=Figure 4, caption=Changes in various indicators during the biotransformation process of pupal cases. A: Changes in reducing sugar content; B: Changes in free amino acid content; C: Changes in protein/chitin content. Different lowercase letters indicate significant differences among treatments at P<0.05., figureFileSmall=p01cVaeyfbwiGlBiXF/IgQ==, figureFileBig=Ru7omB2mT08k/qOmXhu6Ag==, tableContent=null), ArticleFig(id=1226596295595438796, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226460584309342941, language=CN, label=图4, caption=蛹壳生物转化过程中各指标的变化。A：还原糖含量变化情况；B：游离氨基酸含量变化情况；C：蛋白质/几丁质含量变化情况。, figureFileSmall=p01cVaeyfbwiGlBiXF/IgQ==, figureFileBig=Ru7omB2mT08k/qOmXhu6Ag==, tableContent=null), ArticleFig(id=1226596295704490707, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226460584309342941, language=EN, label=Table 1, caption=

Strains of isolated bacteria and enzyme activity

, figureFileSmall=null, figureFileBig=null, tableContent=

Serial

numbers

StrainsEnzyme activity (U/mL)
BSF-CH1Bacillus cereus0.48
BSF-2Virgibacillus salarius0.31
BSF-3Bacillus thuringiensis0.31
BSF-4Bacillus licheniformis0.37
BSF-5Streptomyces diastaticus0.23
BSF-6Klebsiella pneumoniaeND
BSF-7Brevibacterium aviumND
), ArticleFig(id=1226596295847097058, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226460584309342941, language=CN, label=表1, caption=

分离菌株的菌种及酶活力

, figureFileSmall=null, figureFileBig=null, tableContent=

Serial

numbers

StrainsEnzyme activity (U/mL)
BSF-CH1Bacillus cereus0.48
BSF-2Virgibacillus salarius0.31
BSF-3Bacillus thuringiensis0.31
BSF-4Bacillus licheniformis0.37
BSF-5Streptomyces diastaticus0.23
BSF-6Klebsiella pneumoniaeND
BSF-7Brevibacterium aviumND
), ArticleFig(id=1226596295947760365, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226460584309342941, language=EN, label=Table 2, caption=

Distribution and functional annotation of genes related to chitin metabolism in the genome of BSF-CH1

, figureFileSmall=null, figureFileBig=null, tableContent=
Gene numberFunctional annotationSequence numberStart positionEnd positionAmino acid size (bp)
A1_01415Chitin deacetylaseChr192 314193 397360
A1_01730ChitodextrinaseChr267 606269 631675
A1_08430Chitin deacetylaseChr1 539 6161 540 321234
A1_13285ChitodextrinaseChr2 509 1152 510 483455
A1_14040Chitin deacetylaseChr2 630 4352 631 263275
A1_15185Chitin deacetylaseChr2 878 6182 879 461280
A1_17890ChitinaseChr3 421 2713 422 354360
A1_18280Chitin deacetylaseChr3 510 2163 511 116299
A1_25140Chitin deacetylaseChr4 779 7674 780 505245
A1_25160Chitin deacetylaseChr4 785 2524 785 957234
A1_27870ChitodextrinasePlasmid 167 71369 954746
A1_27875ChitinasePlasmid 170 08672 453788
A1_28005ChitinasePlasmid 1100 866102 870667
A1_28425ChitodextrinasePlasmid 1194 267195 644458
), ArticleFig(id=1226596296056812281, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1226460584309342941, language=CN, label=表2, caption=

BSF-CH1基因组中几丁质代谢相关基因的分布及功能注释

, figureFileSmall=null, figureFileBig=null, tableContent=
Gene numberFunctional annotationSequence numberStart positionEnd positionAmino acid size (bp)
A1_01415Chitin deacetylaseChr192 314193 397360
A1_01730ChitodextrinaseChr267 606269 631675
A1_08430Chitin deacetylaseChr1 539 6161 540 321234
A1_13285ChitodextrinaseChr2 509 1152 510 483455
A1_14040Chitin deacetylaseChr2 630 4352 631 263275
A1_15185Chitin deacetylaseChr2 878 6182 879 461280
A1_17890ChitinaseChr3 421 2713 422 354360
A1_18280Chitin deacetylaseChr3 510 2163 511 116299
A1_25140Chitin deacetylaseChr4 779 7674 780 505245
A1_25160Chitin deacetylaseChr4 785 2524 785 957234
A1_27870ChitodextrinasePlasmid 167 71369 954746
A1_27875ChitinasePlasmid 170 08672 453788
A1_28005ChitinasePlasmid 1100 866102 870667
A1_28425ChitodextrinasePlasmid 1194 267195 644458
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一株高效几丁质降解菌蜡样芽孢杆菌BSF-CH1的分离鉴定及其在黑水虻蛹壳生物转化中的应用
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姚祥 1 , 刘倩 1 , 王胜辉 1 , 杨健 1 , 彭才望 2 , 陈武 1 , 尹丽娟 1 , 戴良英 1 , 王运生 1, *
微生物学报 | 研究报告 2025,65(8): 3721-3730
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微生物学报 | 研究报告 2025, 65(8): 3721-3730
一株高效几丁质降解菌蜡样芽孢杆菌BSF-CH1的分离鉴定及其在黑水虻蛹壳生物转化中的应用
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姚祥1, 刘倩1, 王胜辉1, 杨健1, 彭才望2, 陈武1, 尹丽娟1, 戴良英1, 王运生1, *
作者信息
  • 1.湖南农业大学 植物保护学院，湖南 长沙
  • 2.湖南农业大学 机电工程学院，湖南 长沙
An efficient chitinolytic bacterium Bacillus cereus BSF-CH1: isolation, identification, and application in puparium biotransformation of Hermetia illucens
Xiang YAO1, Qian LIU1, Shenghui WANG1, Jian YANG1, Caiwang PENG2, Wu CHEN1, Lijuan YIN1, Liangying DAI1, Yunsheng WANG1, *
Affiliations
  • 1.College of Plant Protection, Hunan Agricultural University, Changsha, Hunan, China
  • 2.College of Mechanical and Electrical Engineering, Hunan Agricultural University, Changsha, Hunan, China
出版时间: 2025-08-04 doi: 10.13343/j.cnki.wsxb.20250100
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摘要
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黑水虻(Hermetia illucens)养殖产生的蛹壳富含几丁质和蛋白质，但目前缺乏高效环保的利用方法。 【目的】 从黑水虻蛹壳堆中分离筛选几丁质降解菌，探索其在蛹壳生物转化中的应用潜力。 【方法】 使用平板筛选和16S rRNA基因测序进行分离和鉴定，采用3,5-二硝基水杨酸(3,5-dinitrosalicylic acid, DNS)法检测其酶活力。使用PacBio HiFi技术进行全基因组测序，以阐明其降解机制。通过蛹壳发酵实验探索其应用潜力。 【结果】 共鉴定得到7株分离菌，其中活性最强的菌株为蜡样芽孢杆菌(Bacillus cereus)BSF-CH1，该菌株在发酵第2天时几丁质酶活力达到最高值0.48 U/mL。全基因组测序结果显示BSF-CH1含有3个几丁质酶(chitinase)基因、7个几丁质脱乙酰酶(chitin deacetylase)基因及4个壳糊精酶基因(chitodextrinase)。蛹壳生物转化实验表明，BSF-CH1能在7 d内将蛹壳质量降解47.2%，几丁质和蛋白质的降解率分别达到64.6%和59.1%。 【结论】 本研究报道了从黑水虻蛹壳中分离的高效几丁质降解菌，为蛹壳的生物转化利用提供了新思路，对促进黑水虻产业的可持续发展具有重要意义。

几丁质降解菌  /  黑水虻蛹壳  /  蜡样芽孢杆菌  /  生物转化  /  基因组分析

The puparium of Hermetia illucens is rich in chitin and protein, while efficient and environmentally friendly utilization methods remain to be developed. [Objective] To isolate chitinolytic bacteria from the puparium pile of H. illucens and explore their potential in puparium biotransformation. [Methods] Strains were isolated by the plate screening method and identified by 16S rRNA gene sequencing. The 3,5-dinitrosalicylic acid (DNS) method was employed to determine the chitinase activity. Whole genome sequencing by PacBio HiFi was conducted to elucidate the degradation mechanism. The application potential of the strain was explored by puparium fermentation experiments. [Results] Among the seven isolated strains, Bacillus cereus BSF-CH1 showed the highest chitinase activity, reaching a maximum chitinase activity of 0.48 U/mL on the second day of fermentation. The genome of BSF-CH1 contained three chitinase genes, seven chitin deacetylase genes, and four chitodextrinase genes. Puparium biotransformation experiments showed that BSF-CH1 could degrade 47.2% of puparium mass within 7 days, with degradation rates of 64.6% and 59.1% for chitin and protein, respectively. [Conclusion] This study reports an efficient chitinolytic bacterium isolated from the puparium of H. illucens, providing new insights into the puparium biotransformation and having important implications for promoting the sustainable development of the H. illucens industry.

chitinolytic bacteria  /  puparium of Hermetia illucens  /  Bacillus cereus  /  biotransformation  /  genome analysis
姚祥, 刘倩, 王胜辉, 杨健, 彭才望, 陈武, 尹丽娟, 戴良英, 王运生. 一株高效几丁质降解菌蜡样芽孢杆菌BSF-CH1的分离鉴定及其在黑水虻蛹壳生物转化中的应用. 微生物学报, 2025 , 65 (8) : 3721 -3730 . DOI: 10.13343/j.cnki.wsxb.20250100
Xiang YAO, Qian LIU, Shenghui WANG, Jian YANG, Caiwang PENG, Wu CHEN, Lijuan YIN, Liangying DAI, Yunsheng WANG. An efficient chitinolytic bacterium Bacillus cereus BSF-CH1: isolation, identification, and application in puparium biotransformation of Hermetia illucens[J]. Acta Microbiologica Sinica, 2025 , 65 (8) : 3721 -3730 . DOI: 10.13343/j.cnki.wsxb.20250100
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随着全球人口增长和环境问题日益突出，如何实现有机废弃物的资源化利用已成为当前亟待解决的问题。黑水虻(Hermetia illucens)因其高效转化有机废物的能力而受到广泛关注[1]。研究表明，黑水虻幼虫可将餐厨垃圾等有机废弃物转化为富含蛋白质和脂肪的生物质[2-3]，是理想的饲用蛋白源。然而，黑水虻产业发展面临的一个重要问题是蛹壳副产物的处理与利用。
黑水虻蛹壳含有丰富的几丁质和蛋白质[4]。其中几丁质是由N-乙酰氨基葡萄糖通过β-1,4糖苷键连接而成的线性多糖，是自然界中含量仅次于纤维素的第二丰富的生物大分子[5]。几丁质及其衍生物壳聚糖因具有生物相容性、可降解性和抗菌活性等特点，在农业、食品、医药和环境保护等领域具有广泛应用前景[6]。
目前，蛹壳中几丁质的提取主要采用化学法，即通过强酸去除矿物质、强碱脱蛋白以获得粗几丁质[7]。该方法不仅污染环境、能耗高，而且产物应用受限。微生物发酵法作为一种绿色环保的替代方案近年来备受关注[8]。已有研究表明，某些微生物可通过分泌几丁质酶、蛋白酶等水解酶来降解甲壳类废弃物[9-11]。然而，目前关于专门针对黑水虻蛹壳的降解菌研究较少，其生物转化过程中的分子机制也尚未阐明。
几丁质降解菌通常具有复杂的几丁质代谢系统，包括几丁质酶、几丁质脱乙酰酶和多糖单加氧酶等[12]。这些酶的协同作用可将几丁质逐步降解为N-乙酰氨基葡萄糖单体或寡聚物。近年来，随着基因组测序技术的发展，对几丁质降解菌的分子机制研究取得了重要进展[13-14]。然而，从黑水虻蛹壳中分离的几丁质降解菌的基因组特征及其降解机制仍有待深入研究。基于此，本研究旨在：(1) 从黑水虻蛹壳堆中分离筛选高效几丁质降解菌；(2) 通过全基因组测序解析其分子特征和代谢潜力；(3) 评估其在蛹壳生物转化中的应用效果。本研究将为黑水虻蛹壳的绿色高值化利用提供新思路，对推动昆虫养殖产业的可持续发展具有重要意义。
1 材料与方法
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1.1 材料
1.1.1 样品来源
黑水虻蛹壳样品采集自湖南农业大学植保基地黑水虻养殖室(28°11′N, 113°04′E)。采样时选取堆放时间为1-2周的化蛹之后的蛹壳，用无菌水洗净、晾干后于-20 ℃保存备用。
1.1.2 主要试剂和仪器
3,5-二硝基水杨酸(3,5-dinitrosalicylic acid, DNS)、革兰氏染色试剂盒均购自北京索莱宝科技有限公司；刚果红购自上海麦克林生化科技股份有限公司；细菌基因组DNA提取试剂盒购自天根生化科技(北京)有限公司。
高压灭菌锅购自上海申安医疗器械厂；生化培养箱购自上海一恒科学仪器有限公司；超净工作台购自苏州安泰空气技术有限公司；高速离心机购自Eppendorf公司；PCR仪购自Bio-Rad公司。
1.1.3 培养基
富集培养基(g/L)：胶体几丁质20.0，K2HPO4 0.7，KH2PO4 0.3，MgSO4·7H2O 0.5， FeSO4·7H2O 0.02，NaCl 1.5，pH 7.0。
筛选培养基(g/L)：在富集培养基基础上添加琼脂20.0和刚果红0.2。
几丁质培养基：在富集培养基基础上添加琼脂20.0 g/L和胰蛋白胨5.0 g/L。
种子培养基(g/L)：胶体几丁质20.0，K2HPO4 0.7，KH2PO4 0.3，MgSO4·7H2O 0.5，FeSO4·7H2O 0.02，NaCl 5.0，pH 7.0。
发酵培养基：在种子培养基基础上添加胰蛋白胨10.0 g/L。
所有培养基121 ℃灭菌20 min。
1.2 几丁质降解菌的分离与筛选
称取2 g蛹壳样品于无菌研钵中研磨，加入10 mL无菌水制成悬浮液。取1 mL接种于富集培养基中，30 ℃、180 r/min振荡培养6 h。将培养液进行10-3-10-6梯度稀释，每个稀释度取100 μL涂布于筛选培养基平板，37 ℃培养5 d。挑选具有明显透明圈的菌落进行纯化，获得单菌落后保存于-80 ℃甘油管中。
1.3 菌株鉴定
1.3.1 形态学观察
将纯化菌株接种于几丁质培养基平板，37 ℃培养7 d后观察菌落形态。挑取单菌落进行革兰氏染色，光学显微镜下观察细胞形态。
1.3.2 16S rRNA基因测序
使用细菌基因组DNA提取试剂盒提取菌株DNA。采用通用引物27F (5′-AGAGTTTGATCC TGGCTCAG-3′)和1492R (5′-GGTTACCTTGTT ACGACTT-3′)[15]扩增16S rRNA基因。PCR反应体系(50 μL)：模板DNA 2 μL，正、反向引物(10 μmol/L)各1 μL，2×Taq PCR Mix 25 μL，ddH2O 21 μL。PCR反应程序：95 ℃预变性5 min；95 ℃变性30 s，55 ℃退火30 s，72 ℃延伸90 s，35个循环；72 ℃终延伸10 min。PCR产物送至北京擎科生物科技股份有限公司测序，结果通过NCBI BLAST进行比对分析。
1.4 几丁质酶活力测定
1.4.1 DNS标准曲线的制作
配制10 mg/mL N-乙酰氨基葡萄糖溶液，取0-80 μL于2 mL离心管中，加入蒸馏水至500 μL。加入500 μL DNS试剂，100 ℃水浴10 min后冷却，12 000 r/min离心10 min。取200 μL上清于96孔板中，测定OD540值。以N-乙酰氨基葡萄糖浓度为横坐标，OD540为纵坐标绘制标准曲线。
1.4.2 酶活力测定
DNS法测酶活[16]，将菌株接种于种子培养基中，30 ℃预培养24 h后，按1%接种量转接至发酵培养基中培养。每隔24 h取样，12 000 r/min离心10 min收集上清作为粗酶液。取250 μL粗酶液与等体积2%胶体几丁质混合，50 ℃反应30 min。加入500 μL DNS试剂终止反应，100 ℃水浴10 min后测定OD540。以灭活的粗酶液为对照。酶活力单位(U)定义为每分钟产生1 μmol还原糖所需的酶量。
1.5 全基因组测序与分析
选取酶活最高的菌株BSF-CH1进行全基因组测序。使用PacBio HiFi技术进行测序(西安浩瑞基因技术有限公司)。采用PGAP流程[17]进行基因组注释，使用antiSMASH[18]预测次级代谢产物基因簇，使用KEGG[19]分析代谢通路。
1.6 蛹壳生物转化实验
将蛹壳洗净烘干后研磨成粉末(粒径≤0.5 mm)。在500 mL三角瓶中加入5 g蛹壳粉末和100 mL无菌水。将BSF-CH1种子液按1%接种量接种，30 ℃、160 r/min振荡培养7 d。每隔24 h取样测定还原糖含量(DNS法)、游离氨基酸含量(茚三酮比色法[20])、pH值、蛹壳降解率、几丁质含量(改良酸碱法[21])、蛋白质含量(凯氏定氮法[22])。
1.7 数据分析
所有实验设3个重复。使用SPSS 25.0进行统计分析，数据以平均值±标准差表示。采用单因素方差分析(one-way ANOVA)和Duncan多重比较检验显著性差异(P<0.05)。使用Origin 2021绘制图表。
2 结果与分析
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2.1 几丁质降解菌的分离与鉴定
从黑水虻蛹壳堆中共分离获得7株能在几丁质筛选培养基上形成透明圈的细菌。通过测定各菌株的几丁质酶活力，发现它们具有不同程度的几丁质降解能力(表1)。其中3株属于芽孢杆菌属，活性最高的BSF-CH1菌株在发酵第2天时达到0.48 U/mL，显著高于其他菌株(P<0.05)。链霉菌属菌株表现出较弱的降解活性，而克雷伯氏菌属和短杆菌属菌株未检测到明显的酶活力。选择酶活力最高的BSF-CH1进行后续研究。
2.2 菌株BSF-CH1的生物学特征
2.2.1 BSF-CH1的形态特征与16S rRNA基因鉴定
如图1A所示，BSF-CH1在几丁质平板上形成较大菌落，菌落整体呈圆形，灰白色，边缘不规则，表面粗糙，似融蜡状，菌落周围存在较大几丁质降解圈，显示出优异的几丁质降解能力。革兰氏染色呈阳性，细胞杆状，长2.0-3.5 μm，宽0.5-0.8 μm，单个或成链排列(图1B)。16S rRNA基因序列分析表明，BSF-CH1与B. cereus ATCC 14579相似度最高。用MEGA 7.0软件构建系统发育树(图1C)，BSF-CH1与蜡样芽孢杆菌亲缘关系最近。结合菌落形态和革兰氏染色结果，将其鉴定为蜡样芽孢杆菌(Bacillus cereus)。
2.2.2 BSF-CH1发酵液几丁质酶活及pH值变化
BSF-CH1的几丁质酶活力在发酵初期快速上升，第2天达到最高值0.48 U/mL，随后缓慢下降(图2A)。pH值从初始的7.0逐渐升高到8.5左右(图2B)，这可能与蛋白质降解产生氨基化合物有关。
2.3 基因组特征分析
2.3.1 基因组总体特征
BSF-CH1基因组测序获得5 596 kb完整序列，包含1个环状染色体(5 255 kb)和2个质粒(274 kb和67 kb)，基因组圈图如图3所示。总G+C含量为35.2%，共预测到5 582个蛋白质编码基因，其中5 573个(99.84%)获得功能注释。
2.3.2 几丁质代谢相关基因
基因组分析发现了多个与几丁质代谢相关的基因(表2)，包括3个几丁质酶基因(chitinase)、7个几丁质脱乙酰酶基因(chitin deacetylase)和4个壳糊精酶基因(chitodextrinase)。这些基因的存在有力地支持了BSF-CH1具有高效降解几丁质的能力。此外，基因组中还存在多个编码几丁质结合蛋白和纤维连接蛋白的基因，这些蛋白可能参与了几丁质的识别和结合过程。
2.4 蛹壳生物转化效果
2.4.1 降解产物分析
在7 d发酵过程中，还原糖含量从0.75 μmol/mL增加至1.48 μmol/mL (图4A)，游离氨基酸含量从1.86 μmol/mL升至11.26 μmol/mL (图4B)，表明BSF-CH1能有效降解蛹壳中的几丁质和蛋白质。
2.4.2 降解效率评估
经过7 d发酵处理，蛹壳总质量降解率达(47.2±2.1)%，几丁质含量从15.30%降至10.25%，降解率为(64.6±3.2)%，蛋白质含量从17.80%降至13.80%，降解率为(59.1±2.8)% (图4C)。这些初步研究表明，BSF-CH1具有较好的蛹壳降解能力。
3 讨论与结论
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本研究从黑水虻蛹壳中分离到的BSF-CH1菌株被鉴定为蜡样芽孢杆菌，这与之前报道的几丁质降解菌种类相似。廖会君等[23]和王敏等[24]分别从雪灵芝和黄粉虫虫蜕中分离到具有较强几丁质降解能力的芽孢杆菌，表明芽孢杆菌可能是自然界中广泛分布的几丁质降解菌。BSF-CH1来源于蛹壳堆，说明其已适应该生态位，这可能是其具有较强降解能力的重要原因。
基因组分析揭示BSF-CH1具有完整的几丁质降解系统，包含3个几丁质酶基因、7个几丁质脱乙酰酶基因和4个壳糊精酶基因。BSF-CH1拥有较多的几丁质脱乙酰酶基因，这可能是其具有较高降解效率的分子基础。壳糊精酶基因的存在确保了降解产物的进一步水解，避免了中间产物的积累抑制。这种多酶协同的降解机制使BSF-CH1能够高效降解复杂的几丁质结构。
本研究发现，BSF-CH1的几丁质酶活力在发酵第2天时达到峰值(0.48 U/mL)，这与培养基pH值变化趋势相吻合。pH值的升高可能与以下因素有关：(1) 几丁质降解释放N-乙酰氨基葡萄糖，进一步脱乙酰产生氨基化合物；(2) 蛋白质降解产生氨基酸和氨基化合物；(3) 菌体代谢产生碱性物质。pH值的变化可能会影响酶的活性和稳定性，这提示在工业应用中需要考虑pH调控策略。
BSF-CH1对蛹壳的降解效率(7 d总降解率47.2%)与Dhole等[25]筛选出的降解效率最高的嗜麦芽寡养单胞菌(Stenotrophomonas maltophilia)相近，同时显著优于其筛选出的其他菌株。BSF-CH1表现出的高效降解能力可能源于：(1) 菌株来源于蛹壳环境，具有良好的适应性；(2) 具有完整的几丁质降解酶系；(3) 能同时降解几丁质和蛋白质，降低了底物复杂性的影响。
蛹壳生物转化实验结果表明，BSF-CH1能够有效降解蛹壳中的几丁质和蛋白质，同时产生还原糖和氨基酸等小分子物质。这些小分子物质可作为生物肥料、饲料添加剂或微生物培养基的原料，实现蛹壳的高值化利用。与传统的化学提取法相比，微生物发酵法具有环境友好、条件温和、成本较低等优势，更具可持续性。
然而要实现该技术的工业化应用，仍需进一步优化发酵条件(如pH、温度、通气量、底物浓度等)和产物分离纯化工艺。此外，还可以通过基因工程手段改造BSF-CH1，进一步提高其几丁质酶和蛋白酶的产量，或引入新的代谢途径，生产更多高附加值产品。
本研究从黑水虻蛹壳堆肥环境中分离筛选到一株高效几丁质降解菌BSF-CH1，该菌株属于蜡样芽孢杆菌，具有优异的几丁质和蛋白质降解能力。全基因组分析揭示了其几丁质降解的潜在分子机制。BSF-CH1发酵转化蛹壳技术为黑水虻蛹壳的高值化利用提供了一种新颖、环保、可持续的途径，有助于推动黑水虻养殖业的绿色发展。
基金
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  • 湖南省自然科学基金面上项目(2023JJ30310)
  • 湖南省教育厅资助科研项目(22A0169)
文献
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2025年第65卷第8期
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doi: 10.13343/j.cnki.wsxb.20250100
  • 接收时间：2025-02-13
  • 首发时间：2026-02-06
  • 出版时间：2025-08-04
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  • 收稿日期：2025-02-13
  • 录用日期：2025-03-14
基金
General Program of the Natural Science Foundation of Hunan Province(2023JJ30310)
湖南省自然科学基金面上项目(2023JJ30310)
Scientific Research Project Funded by the Education Department of Hunan Province(22A0169)
湖南省教育厅资助科研项目(22A0169)
作者信息
    1.湖南农业大学 植物保护学院，湖南 长沙
    2.湖南农业大学 机电工程学院，湖南 长沙

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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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