浓香型白酒发酵体系中己酸菌研究进展

摘要/Abstract

摘要： 浓香型白酒发酵体系中己酸生成菌（以下简称“己酸菌”）的己酸合成代谢对提高浓香型白酒的发酵质量非常重要。因此，有必要深入全面了解浓香型白酒发酵体系中己酸菌的种类及其己酸合成代谢特征。本综述介绍了目前浓香型白酒发酵体系中已经分离的己酸菌株的种类多样性、系统进化关系、生理代谢特征、己酸合成代谢机制以及其与己酸菌、非己酸菌之间的协同代谢关系。本文为理解己酸菌群在浓香型白酒发酵体系中的原位己酸合成代谢规律提供参考，为将来靶向提高己酸菌群在浓香型白酒发酵和生物质转化高附加值己酸工艺中进行己酸合成培养工程中的应用提供理论依据。

关键词: 浓香型白酒发酵体系, 己酸菌株, 代谢特征, 己酸合成机制, 己酸菌群, 协同代谢

Abstract: In Chinese strong-flavor Baijiu (CSFB) fermentation ecosystem, the caproic acid-synthesis metabolism of caproic acid-producing bacteria (CPBs) are very important for improving the fermentation quality of CSFB. Therefore, it is necessary to thoroughly understand the types of CPBs and their caproic acid-synthesis metabolic characteristics. This minireview introduced the diversity of CPB strains isolated from the CSFB fermentation ecosystem, their phylogenetic relationships, physiological metabolic characteristics, caproic acid-synthesis metabolic mechanism, and their synergistic metabolic relationships with other CPBs or non-CPBs. This paper provides a reference for understanding the in-situ caproic acid-synthesis metabolic pattern of CPBs from CSFB fermentation ecosystem, and further provides a theoretical basis for the future targeted application of CPBs in the CSFB fermentation and the high value-added caproic acid synthetic-culture engineering for biomass conversion.

Key words: Chinese strong-flavor Baijiu fermentation ecosystem, Caproic acid-producing bacteria, Metabolic characteristics, Caproic acid-synthesis Mechanism, CPBs microbiota, Cooperative metabolism

中图分类号:

TS201.3

张会敏邢新会王越崔磊王秀本常强孙伟席鲜会薛正莲. 浓香型白酒发酵体系中己酸菌研究进展[J]. 食品科学.

参考文献

[1] LIU M, TANG Y, GUO X, et al. Deep sequencing reveals high bacterial diversity and phylogenetic novelty in pit mud from Luzhou Laojiao cellars for Chinese strong-flavor Baijiu[J]. Food research international, 2017, 102: 68-76. DOI: 10.1016/j.foodres.2017.09.075.
[2] WEI Y, ZOU W, SHEN C, et al. Basic flavor types and component characteristics of Chinese traditional liquors: A review[J]. Journal of Food Science, 2020, 85(12): 4096-4107. DOI: 10.1111/1750-3841.15536.
[3] XU Y, ZHAO J, LIU X, et al. Flavor mystery of Chinese traditional fermented baijiu: the great contribution of ester compounds[J]. Food Chemistry, 2022, 369: 130920. DOI: 10.1016/j.foodchem.2021.130920.
[4] 何培新, 胡晓龙, 郑燕, 等. 中国浓香型白酒 “增己降乳” 研究与应用进展[J]. 轻工学报, 2018, 33(4): 1-12. DOI: 10.3969/j.issn.2096-1553.2018.04.001.
[5] HUANG Z, ZENG Y, SUN Q, et al. Insights into the mechanism of flavor compound changes in strong flavor baijiu during storage by using the density functional theory and molecular dynamics simulation[J]. Food Chemistry, 2022, 373: 131522. DOI: 10.1016/j.foodchem.2021.131522.
[6] ZOU W, YE G, ZHANG K. Diversity, function, and application of Clostridium in Chinese strong flavor baijiu ecosystem: a review[J]. Journal of food science, 2018, 83(5): 1193-1199. DOI: 10.1111/1750-3841.14134.
[7] WANG X, DU H, XU Y. Source tracking of prokaryotic communities in fermented grain of Chinese strong-flavor liquor[J]. International Journal of Food Microbiology, 2017, 244: 27-35. DOI: 10.1016/j.ijfoodmicro.2016.12.018.
[8] ZHANG H, MENG Y, WANG Y, et al. Prokaryotic communities in multidimensional bottom-pit-mud from old and young pits used for the production of Chinese Strong-Flavor Baijiu[J]. Food Chemistry, 2020, 312: 126084. DOI: 10.1016/j.foodchem.2019.126084.
[9] GAO J, LIU G, LI A, et al. Domination of pit mud microbes in the formation of diverse flavour compounds during Chinese strong aroma-type Baijiu fermentation [J]. LWT, 2021, 137: 110442. DOI: 10.1016/j.lwt.2020.110442.
[10] GAO Z, WU Z, ZHANG W. Effect of Pit Mud on Bacterial Community and Aroma Components in Yellow Water and Their Changes during the Fermentation of Chinese Strong-Flavor Liquor[J]. Foods, 2020, 9(3): 372. DOI: 10.3390/foods9030372.
[11] LI H, HUANG J, LIU X, et al. Characterization of interphase microbial community in Luzhou-flavored liquor manufacturing pits of various ages by polyphasic detection methods[J]. Journal of Microbiology and Biotechnology, 2017, 27(1): 130-140. DOI: 10.4014/jmb.1605.05036.
[12] LIU M, LIU C, TIAN X, et al. Bioremediation of degraded pit mud by indigenous microbes for Baijiu production[J]. Food Microbiology, 2022, 108: 104096. DOI: 10.1016/j.fm.2022.104096.
[13] LIU M, TANG Y, GUO X, et al. Structural and Functional Changes in Prokaryotic Communities in Artificial Pit Mud during Chinese Baijiu Production[J]. mSystems, 2020, 5(2): e00829-19. DOI: 10.1128/msystems.00829-19.
[14] WU Q, JIANG Y, CHEN Y, et al. Opportunities and challenges in microbial medium chain fatty acids production from waste biomass[J]. Bioresource Technology, 2021, 340: 125633. DOI: 10.1016/j.biortech.2021.125633.
[15] WU Q, GUO W, BAO X, et al. Upgrading liquor-making wastewater into medium chain fatty acid: Insights into co-electron donors, key microflora, and energy harvest[J]. Water Research, 2018, 145: 650-659. DOI: 10.1016/j.watres.2018.08.046.
[16] YU P, WU M, BAO W, et al. Performance of a mixed inoculum of sludge and pit mud for short and medium-chain fatty acids production: Insight into key microbiome and functional potential in anaerobic fermentation inoculum[J]. Chemical Engineering Journal, 2023, 466: 143142. DOI: 10.1016/j.cej.2023.143142
[17] GAO M, LIN Y, WANG P, et al. Production of medium-chain fatty acid caproate from Chinese liquor distillers’ grain using pit mud as the fermentation microbes[J]. Journal of Hazardous Materials, 2021, 417: 126037. DOI: 10.1016/j.jhazmat.2021.126037.
[18] STAMATOPOULOU P, MALKOWSKI J, CONRADO L, et al. Fermentation of organic residues to beneficial chemicals: a review of medium-chain fatty acid production[J]. Processes, 2020, 8(12): 1571. DOI: 10.3390/pr8121571.
[19] FLAIZ M, BAUR T, BRAHNER S, et al. Caproicibacter fermentans gen. nov., sp. nov., a new caproate-producing bacterium and emended description of the genus Caproiciproducens[J]. International Journal of Systematic and Evolutionary Microbiology, 2020, 70(7): 4269-4279. DOI: 10.1099/ijsem.0.004283.
[20] WANG H, GU Y, ZHAO D, et al. Caproicibacterium lactatifermentans sp. nov., isolated from pit clay used for the production of Chinese strong aroma-type liquor[J]. International Journal of Systematic and Evolutionary Microbiology, 2022, 72(1): 005206. DOI: 10.1099/ijsem.0.004283.
[21] LIU C, DU Y, ZHENG J, et al. Production of caproic acid by Rummeliibacillus suwonensis 3B-1 isolated from the pit mud of strong-flavor baijiu[J]. Journal of Biotechnology, 2022, 358: 33-40. DOI: 10.1016/j.jbiotec.2022.08.017.
[22] LUO H, LI T, ZHENG J, et al. Isolation, Identification, and Fermentation Medium Optimization of a Caproic Acid?Producing Enterococcus casseliflavus Strain from Pit Mud of Chinese Strong Flavor Baijiu Ecosystem[J]. Polish Journal of Microbiology, 2022, 71(4): 563-575. DOI: 10.33073/pjm-2022-052.
[23] LI K, CHEN Y, LIU T, et al. Analysis of spatial distribution of bacterial community associated with accumulation of volatile compounds in Jiupei during the brewing of special-flavor liquor[J]. LWT, 2020, 130: 109620. DOI: 10.1016/j.lwt.2020.109620.
[24] LU M, ZHOU W, JI F, et al. Profiling prokaryotic community in pit mud of Chinese strong-aroma type liquor by using oligotrophic culturing[J]. International Journal of Food Microbiology, 2020, 337: 108951. DOI: 10.1016/j.ijfoodmicro.2020.108951.
[25] CHAI L, QIAN W, ZHONG X, et al. Mining the factors driving the evolution of the pit mud microbiome under the impact of long-term production of strong-flavor baijiu[J]. Applied and Environmental Microbiology, 2021, 87(17): e00885-21. DOI: 10.1128/AEM.00885-21.
[26] XU Y, EU M, ZHAO D, et al. Simulated fermentation of strong-flavor baijiu through functional microbial combination to realize the stable synthesis of important flavor chemicals[J]. Foods, 2023, 12(3): 00644. DOI: 10.3390/foods12030644.
[27] LIU M, ZHAO K, TANG Y, et al. Analysis of Clostridium cluster I community diversity in pit mud used in manufacture of Chinese Luzhou-flavor liquor[J]. Food Science and Biotechnology, 2015, 24(3): 995-1000. DOI: 10.1007/s10068-015-0127-7.
[28] ZHU X, ZHOU Y, WANG Y, et al. Production of high-concentration n-caproic acid from lactate through fermentation using a newly isolated Ruminococcaceae bacterium CPB6[J]. Biotechnology for biofuels, 2017, 10: 102. DOI: 10.3390/pr8121571.
[29] WANG H, GU Y, ZHOU W, et al. Adaptability of a caproate-producing bacterium contributes to its dominance in an anaerobic fermentation system[J]. Applied and Environmental Microbiology, 2021, 87(20): e01203-21. DOI: 10.1128/AEM.01203-21.
[30] TINDALL B J. The names Hungateiclostridium Zhang et al. 2018, Hungateiclostridium thermocellum (Viljoen et al. 1926) Zhang et al. 2018, Hungateiclostridium cellulolyticum (Patel et al. 1980) Zhang et al. 2018, Hungateiclostridium aldrichii (Yang et al. 1990) Zhang et al. 2018, Hungateiclostridium alkalicellulosi (Zhilina et al. 2006) Zhang et al. 2018, Hungateiclostridium clariflavum (Shiratori et al. 2009) Zhang et al. 2018, Hungateiclostridium straminisolvens (Kato et al. 2004) Zhang et al. 2018 and Hungateiclostridium saccincola (Koeck et al. 2016) Zhang et al. 2018 contravene Rule 51b of the International Code of Nomenclature of Prokaryotes and require replacement names in the genus Acetivibrio Patel et al. 1980[J]. International Journal of Systematic and Evolutionary Microbiology, 2019, 69(12): 3927-3932. DOI: 10.1099/ijsem.0.003685
[31] WU L, FAN J, CHEN J, et al. Chemotaxis of Clostridium strains isolated from pit mud and its application in baijiu fermentation[J]. Foods, 2022, 11(22): 3639. DOI: 10.3390/foods11223639.
[32] ZHANG C, GUO M, LIU J, et al. A new method for screening and culture of Clostridium from pit mud under non-anaerobic conditions [J]. Journal of Microbiological Methods, 2022, 200: 106559.
[33] ZOU W, YE G, LIU C, et al. Comparative genome analysis of Clostridium beijerinckii strains isolated from pit mud of Chinese strong flavor baijiu ecosystem[J]. G3, 2021, 11(11): jkab317. DOI: 10.1093/g3journal/jkab317.
[34] LI C, WANG Y, XIE G, et al. Complete genome sequence of Clostridium butyricum JKY6D1 isolated from the pit mud of a Chinese flavor liquor-making factory[J]. Journal of Biotechnology, 2016, 220: 23-24. DOI: 10.1016/j.jbiotec.2016.01.003.
[35] CANDRY P, RADI? L, FAVERE J, et al. Mildly acidic pH selects for chain elongation to caproic acid over alternative pathways during lactic acid fermentation[J]. Water Research, 2020, 186: 116396. DOI: 10.1016/j.watres.2020.116396.
[36] LIU C, HUANG D, LIU L, et al. Clostridium swellfunianum sp. nov., a novel anaerobic bacterium isolated from the pit mud of Chinese Luzhou-flavor liquor production[J]. Antonie van Leeuwenhoek, 2014, 106(3): 817-825. DOI: 10.1007/s10482-014-0251-z.
[37] XU P, CHAI L, QIU T, et al. Clostridium fermenticellae sp. nov., isolated from the mud in a fermentation cellar for the production of the Chinese liquor, baijiu[J]. International Journal of Systematic and Evolutionary Microbiology, 2019, 69(3): 859-865. DOI: 10.1099/ijsem.0.003254.
[38] TAO Y, LI J, RUI J, et al. Prokaryotic communities in pit mud from different-aged cellars used for the production of Chinese strong-flavored liquor[J]. Applied and Environmental Microbiology, 2014, 80(7): 2254-2260. DOI: 10.1128/AEM.04070-13.
[39] HU X, DU H, REN C, et al. Illuminating anaerobic microbial community and cooccurrence patterns across a quality gradient in Chinese liquor fermentation pit muds[J]. Applied and Environmental Microbiology, 2016, 82(8): 2506-2515. DOI: 10.1128/AEM.03409-15.
[40] XU J, SUN L, XING X, et al. Culturing bacteria from fermentation pit muds of Baijiu with culturomics and amplicon-based metagenomic approaches[J]. Frontiers in microbiology, 2020, 11: 1223. DOI: 10.3389/fmicb.2020.01223.
[41] WANG J, HAO S, REN Q. Analysis of Bacterial Diversity in Fermented Grains of Baijiu Based on Culturomics and Amplicon Sequencing[J]. Fermentation, 2023, 9(3): 260. DOI: 10.3390/fermentation9030260.
[42] WATTERSON W, TANYERI M, WATSON A, et al. Droplet-based high-throughput cultivation for accurate screening of antibiotic resistant gut microbes[J]. eLife, 2020, 9: e56998. DOI: 10.7554/eLife.56998.
[43] TAO Y, ZHU X, WANG H, et al. Complete genome sequence of Ruminococcaceae bacterium CPB6: A newly isolated culture for efficient n-caproic acid production from lactate[J]. Journal of Biotechnology, 2017, 259: 91-94. DOI: 10.1016/j.jbiotec.2017.07.036.
[44] ZHU X, TAO Y, LIANG C, et al. The synthesis of n-caproate from lactate: a new efficient process for medium-chain carboxylates production[J]. Scientific reports, 2015, 5: 14360. DOI: 10.1038/srep14360.
[45] GU Y, ZHU X, LIN F, et al. Caproicibacterium amylolyticum gen. nov., sp. nov., a novel member of the family Oscillospiraceae isolated from pit clay used for making Chinese strong aroma-type liquor[J]. International Journal of Systematic and Evolutionary Microbiology, 2021, 71(4): 004789. DOI: 10.1099/ijsem.0.004789.
[46] HU X, DU H, XU Y. Identification and quantification of the caproic acid-producing bacterium Clostridium kluyveri in the fermentation of pit mud used for Chinese strong-aroma type liquor production[J]. International Journal of Food Microbiology, 2015, 214: 116-122. DOI: 10.1016/j.ijfoodmicro.2015.07.032.
[47] 胡晓龙. 浓香型白酒窖泥中梭菌群落多样性与窖泥质量关联性研究[D]. 无锡: 江南大学, 2015: 70.
[48] WANG Y, LI B, DONG H, et al. Complete genome sequence of Clostridium kluyveri JZZ applied in Chinese strong-flavor liquor production[J]. Current microbiology, 2018, 75(11): 1429-1433. DOI: 10.1007/s00284-018-1539-4.
[49] 彭兵, 祝熙, 李忠奎, 等. 窖泥高产己酸菌分离鉴定及培养条件优化的研究[J]. 中国酿造, 2016, 35(05): 43-46. DOI: 10.11882/j.issn.0254-5071.2016.05.009.
[50] 张会敏, 孟雅静, 王艳丽, 等. 新老窖池黄水的差异性及静置培养对其影响[J]. 食品科学, 2020, 41(2): 215-222. DOI: 10.7506/spkx1002-6630-20190531-382.
[51] ZHANG M, WU X, MU D, et al. Profiling the influence of physicochemical parameters on the microbial community and flavor substances of zaopei[J]. Journal of the Science of Food and Agriculture, 2021, 101(15): 6300-6310. DOI: 10.1002/jsfa.11299.
[52] KANG J, SUN Y, HUANG X, et al. Unraveling the microbial compositions, metabolic functions, and antibacterial properties of Huangshui, a byproduct of Baijiu fermentation[J]. Food Research International, 2022, 157: 111320. DOI: 10.1016/j.foodres.2022.111320.
[53] DING X, WU C, HUANG J, et al. Interphase microbial community characteristics in the fermentation cellar of Chinese Luzhou-flavor liquor determined by PLFA and DGGE profiles[J]. Food Research International, 2015, 72: 16-24. DOI: 10.1016/j.foodres.2015.03.018.
[54] SCARBOROUGH M, LAWSON C, HAMILTON J, et al. Metatranscriptomic and thermodynamic insights into medium-chain fatty acid production using an anaerobic microbiome[J]. mSystems, 2018, 3(6): e00221-18. DOI: 10.1128/msystems.00221-18.
[55] HAN W, HE P, SHAO L, et al. Metabolic interactions of a chain elongation microbiome[J]. Applied and Environmental Microbiology, 2018, 84(22): e01614-18. DOI: 10.1128/AEM.01614-18.
[56] WANG H, LI X, WANG Y, et al. Improvement of n-caproic acid production with Ruminococcaceae bacterium CPB6: selection of electron acceptors and carbon sources and optimization of the culture medium[J]. Microbial Cell Factories, 2018, 17: 99. DOI: 10.1186/s12934-018-0946-3.
[57] TAO Y, WANG X, LI X, et al. The functional potential and active populations of the pit mud microbiome for the production of Chinese strong‐flavour liquor[J]. Microbial biotechnology, 2017, 10(6): 1603-1615. DOI: 10.1111/1751-7915.12729.
[58] ZHU X, HUANG H, HE Y, et al. A preliminary study on the feasibility of industrialization for n-caproic acid recovery from food wastewater: From lab to pilot[J]. Bioresource Technology, 2022, 366: 128154. DOI: 10.1016/j.biortech.2022.128154.
[59] FANG G, CHAI L, ZHONG X, et al. Comparative Genomics Unveils the Habitat Adaptation and Metabolic Profiles of Clostridium in an Artificial Ecosystem for Liquor Production [J]. mSystems, 2022, 7(3): e00297-22. DOI: 10.1128/msystems.00297-22.
[60] SHI X, WU L, WEI W, et al. Insights into the microbiomes for medium-chain carboxylic acids production from biowastes through chain elongation[J]. Critical Reviews in Environmental Science and Technology, 2022, 52(21): 3787-3812. DOI: 10.1080/10643389.2021.1957342.
[61] QIAN W, LU Z, CHAI L, et al. Cooperation within the microbial consortia of fermented grains and pit mud drives organic acid synthesis in strong-flavor Baijiu production [J]. Food Research International, 2021, (147): 110449.
[62] CHAI L, XU P, QIAN W, et al. Profiling the Clostridia with butyrate-producing potential in the mud of Chinese liquor fermentation cellar[J]. International Journal of Food Microbiology, 2019, 297: 41-50. DOI: 10.1016/j.ijfoodmicro.2019.02.023.
[63] SUN H, CHAI L, FANG G, et al. Metabolite-based mutualistic interaction between two novel Clostridial species from pit mud enhances butyrate and caproate production[J]. Applied and Environmental Microbiology, 2022, 88(13): e00484-22. DOI: 10.1128/aem.00484-22.
[64] CHEN S, HUANG J, QIN H, et al. Characterizing the interaction relationship of the microbial communities between Zaopei and pit mud disturbing by Daqu[J]. Food Science and Biotechnology, 2021, 30(10): 1357–1367. DOI: 10.1007/s10068-021-00975-z.
[65] MU Y, HUANG J, ZHOU R, et al. Exploring the response patterns of strong-flavor baijiu brewing microecosystem to fortified Daqu under different pit ages[J]. Food Research International, 2022, 155: 111062. DOI: 10.1016/j.foodres.2022.111062.
[66] GAO L, XIE F, REN X, et al. Correlation between microbial diversity and flavor metabolism in Huangshui: A by-product of solid-state fermentation Baijiu[J]. LWT, 2023, 181: 114767. DOI: 10.1016/j.lwt.2023.114767.
[67] 孟雅静, 王艳丽, 丁峰, 等. 浓香型白酒新、老窖池分层池底窖泥菌群总氢代谢与乳酸含量之间的关系[J]. 食品科学, 2021, 42(18): 171-177. DOI: 10.7506/spkx1002-6630-20200514-161.
[68] YUAN S, JIN Z, ALI A, et al. Caproic acid producing bacteria in Chinese Baijiu brewing[J]. Frontiers in Microbiology, 2022, 13: 1651. DOI: 10.3389/fmicb.2022.883142.
[69] FU J, CHEN L, YANG S, et al. Metagenome and analysis of metabolic potential of the microbial community in pit mud used for Chinese strong-flavor liquor production[J]. Food Research International, 2021, 143: 110294. DOI: 10.1016/j.foodres.2021.110294.
[70] FERNáNDEZ-BLANCO C, VEIGA M C, KENNES C. Efficient production of n-caproate from syngas by a co-culture of Clostridium aceticum and Clostridium kluyveri [J]. Journal of Environmental Management, 2022, 302: 113992.
[71] BAUMLER M, SCHNEIDER M, EHRENREICH A, et al. Synthetic co‐culture of autotrophic Clostridium carboxidivorans and chain elongating Clostridium kluyveri monitored by flow cytometry[J]. Microbial Biotechnology, 2022, 15(5): 1471-1485. DOI: 10.1111/1751-7915.13941.
[72] OTTEN J, ZOU Y, PAPOUTSAKIS E. The potential of caproate (hexanoate) production using Clostridium kluyveri syntrophic cocultures with Clostridium acetobutylicum or Clostridium saccharolyticum[J]. Frontiers in Bioengineering and Biotechnology, 2022, 10: 965614. DOI: 10.3389/fbioe.2022.965614.
[73] ZHANG C, LIU H, WU P, et al. Clostridium kluyveri enhances caproate production by synergistically cooperating with acetogens in mixed microbial community of electro-fermentation system[J]. Bioresource Technology, 2023, 369: 128436. DOI: 10.1016/j.biortech.2022.128436.
[74] PARERA I, SOUSA D. Upgrading dilute ethanol to odd-chain carboxylic acids by a synthetic co-culture of Anaerotignum neopropionicum and Clostridium kluyveri[J]. Biotechnology for Biofuels and Bioproducts, 2023, 16(1): 1-17. DOI: 10.1186/s13068-023-02336-w.
[75] CUI Y, YANG K, ZHOU K. Using co-culture to functionalize Clostridium fermentation[J]. Trends in Biotechnology, 2021, 39(9): 914-926. DOI: 10.1016/j.tibtech.2020.11.016.
[76] LOUCA S, POLZ M, MAZEL F, et al. Function and functional redundancy in microbial systems[J]. Nature ecology & evolution, 2018, 2(6): 936-943. DOI: 10.1038/s41559-018-0519-1.
[77] AULAKH S K, SELLéS VIDAL L, SOUTH E J, et al. Spontaneously established syntrophic yeast communities improve bioproduction[J]. Nature Chemical Biology, 2023. DOI: 10.1038/s41589-023-01341-2.
[78] DIENDER M, OLM I P, SOUSA D Z. Synthetic co-cultures: novel avenues for bio-based processes[J]. Current Opinion in Biotechnology, 2021, 67: 72-79. DOI: 10.1016/j.copbio.2021.01.006.