泡菜為典型之乳酸發酵蔬菜食品，其發酵品質、風味形成及功能性成分與微生物群落變化密切相關。鹽類除具有調味與抑菌功能外，其礦物質組成與理化特性亦可能影響發酵過程中之菌相演替與代謝表現。本研究以海鹽、岩鹽及食鹽作為不同鹽類處理，探討其對高麗菜發酵過程中微生物群落結構與胞外多醣（exopolysaccharides, EPS）生成之影響。本研究利用全長 16S rRNA 基因定序分析不同發酵階段之菌相組成，並以 alpha diversity、beta diversity、heatmap、ternary plot 與 Venn diagram 等方式比較不同鹽類處理間之微生物分布差異。結果顯示，隨發酵進行，菌相多樣性逐漸下降，群落逐漸由少數優勢菌群主導。不同鹽類處理間雖未形成完全分離之菌群結構，但仍呈現一定程度之分布差異，其中食鹽組之菌相分布相對集中，而海鹽組與岩鹽組之間則具有較高相似性。菌相組成分析顯示，Weissella、Leuconostoc 與 Lactiplantibacillus 為主要優勢乳酸菌，其中 Weissella cibaria 於多項分析中皆呈現高度優勢，顯示其可能為本研究發酵系統中的核心菌群。不同鹽類處理雖共享部分核心菌群，但於 ASV 層級仍存在差異，顯示不同鹽源可能對特定微生物具有選擇性影響。此外，部分 Enterobacter-related taxa 較常出現在岩鹽組，而多種乳酸菌則較集中於海鹽組與食鹽組。EPS 分析結果顯示，不同鹽類處理之 EPS 產量皆於發酵中期達到較高累積量，其中岩鹽組整體呈現較高之 EPS 生成趨勢。綜合菌相分析推測，岩鹽中較複雜之礦物質組成可能與 EPS-producing lactic acid bacteria 之代謝活性具有關聯。整體而言，本研究顯示不同鹽類來源可能透過影響發酵微生物群落結構，進一步改變泡菜發酵過程中之菌相演替與 EPS 生成趨勢，並可作為未來功能性發酵食品開發與發酵條件優化之參考依據。

Pickle is a typical lactic acid-fermented vegetable food whose quality, flavor, and functional properties are closely associated with microbial community dynamics. In addition to seasoning and antimicrobial effects, different salt sources may influence microbial succession and metabolic activities through differences in mineral composition and physicochemical properties. In this study, marine salt, halite, and edible salt were used to investigate their effects on microbial community structure and exopolysaccharide (EPS) production during cabbage fermentation. The results showed that microbial diversity gradually decreased during fermentation, and the microbial community became progressively dominated by a limited number of taxa. Although microbial communities among salt treatments were not completely separated, distinct distribution tendencies were still observed. Community composition analysis revealed that Weissella, Leuconostoc, and Lactiplantibacillus were the predominant lactic acid bacteria during fermentation, with Weissella cibaria consistently exhibiting high abundance across multiple analyses. Differences among salt treatments were also observed at the ASV level, indicating that different salt sources may selectively influence specific microbial populations. EPS analysis demonstrated that EPS accumulation increased during the middle stage of fermentation under all salt treatments, with the halite group generally exhibiting higher EPS production. Combined with microbial community analysis, the results suggest that the more complex mineral composition of halite may be associated with the metabolic activities of EPS-producing lactic acid bacteria. Overall, this study demonstrates that different salt sources may influence microbial succession and EPS production during pickle fermentation by altering microbial community structure, providing useful information for the development of functional fermented foods and optimization of fermentation conditions.

Bautista-Gallego, J., Rantsiou, K., Garrido-Fernández, A., Cocolin, L., & Arroyo-López, F. N. (2013). Salt reduction in vegetable fermentation: reality or desire? Journal of Food Science, 78, R1095-R1100.
Cheigh, H. S., Park, K. Y., & Lee, C. Y. (1994). Biochemical, microbiological, and nutritional aspects of kimchi (Korean fermented vegetable products). Critical Reviews in Food Science and Nutrition, 34, 175-203.
Cocolin, L., Alessandria, V., Dolci, P., Gorra, R., & Rantsiou, K. (2013). Culture independent methods to assess the diversity and dynamics of microbiota during food fermentation. International Journal of Food Microbiology, 167, 29-43.
de Filippis, F., Parente, E., & Ercolini, D. (2018). Recent past, present, and future of the food microbiome. Annual Review of Food Science and Technology, 9, 589-608.
de Filippis, F., Parente, E., & Ercolini, D. (2017). Metagenomics insights into food fermentations. Microbial Biotechnology, 10, 91-102.
de Vuyst, L., & Degeest, B. (1999). Heteropolysaccharides from lactic acid bacteria. FEMS Microbiology Reviews, 23, 153-177.
Di Cagno, R., Coda, R., De Angelis, M., & Gobbetti, M. (2013). Exploitation of vegetables and fruits through lactic acid fermentation. Food Microbiology, 33, 1-10.
Ercolini, D. (2013). High-throughput sequencing and metagenomics: moving forward in the culture-independent analysis of food microbial ecology. Applied and Environmental Microbiology, 79, 3148-3155.
Ferrocino, I., Bellio, A., Romano, A., Macori, G., Rantsiou, K., Decastelli, L., & Cocolin, L. (2018). RNA-based amplicon sequencing reveals active microbial communities in food ecosystems. Food Microbiology, 73, 295-303.
Fusco, V., Quero, G. M., Cho, G. S., Kabisch, J., Meske, D., Neve, H., Bockelmann, W., & Franz, C. M. A. P. (2015). The genus Weissella: taxonomy, ecology and biotechnological potential. Frontiers in Microbiology, 6, 155.
Hutkins, R. W. (2018). Microbiology and Technology of Fermented Foods (2nd ed.). Hoboken, NJ: Wiley-Blackwell.
Jeong, S. H., Lee, S. H., Jung, J. Y., Choi, E. J., & Jeon, C. O. (2013). Microbial succession and metabolite changes during long-term storage of kimchi. Journal of Food Science, 78, M763-M769.
Jung, J. Y., Lee, S. H., & Jeon, C. O. (2011). Kimchi microflora: history, current status, and perspectives for industrial kimchi production. Applied Microbiology and Biotechnology, 98, 2385-2393.
Jung, J. Y., Lee, S. H., Jin, H. M., Hahn, Y., Madsen, E. L., & Jeon, C. O. (2014). Metatranscriptomic analysis of lactic acid bacterial gene expression during kimchi fermentation. International Journal of Food Microbiology, 163, 171-179.
Kim, D. W., Kim, B. M., Lee, H. J., Jang, G. J., Song, S. H., Lee, J. I., Lee, S. B., Shim, J. M., Lee, K. W., Kim, J. H., Ham, K. S., Chen, F., & Kim, H. J. (2017). Effects of different salt treatments on the fermentation metabolites and bacterial profiles of kimchi. Journal of Food Science, 82, 1124-1131.
Lee, D. Y., Park, S. E., Kim, E. J., Kim, H. W., Cho, K. M., Kwon, S. J., Roh, S. W., Kwak, S., Whon, T. W., & Son, H. S. (2024). A comparative study of the physicochemical, microbial, and metabolic profiling of kimchi during long-term fermentation under varying salinity conditions. LWT - Food Science and Technology, 196, 115838.
Lee, M. E., Jang, J. Y., Lee, J. H., Park, H. W., Choi, H. J., & Kim, T. W. (2005). Starter cultures for kimchi fermentation. Journal of Microbiology and Biotechnology, 25, 559-568.
Lee, M. A., Choi, Y. J., Kim, Y. S., Chon, S. Y., Chung, Y. B., Park, S. H., Yun, Y. R., Min, S. G., Yang, H. C., & Seo, H. Y. (2022). Effects of salt type on the metabolites and microbial community in kimchi fermentation. Heliyon, 8, e11360.
Lee, M. A., Choi, Y. J., Lee, H., Hwang, S., Lee, H. J., Park, S. J., Chung, Y. B., Yun, Y. R., Park, S. H., Min, S., Kwon, L. S., & Seo, H. Y. (2021). Influence of salinity on the microbial community composition and metabolite profile in kimchi. Fermentation, 7, 308.
Leech, J., Cabrera-Rubio, R., Walsh, A. M., Macori, G., Walsh, C. J., Barton, W., Finnegan, L., Crispie, F., O’Sullivan, O., Claesson, M. J., Cotter, P. D., & Hill, C. (2020). Fermented-food metagenomics reveals substrate-associated differences in taxonomy and health-associated and antibiotic resistance determinants. mSystems, 5, e00522-19.
Leroy, F., & De Vuyst, L. (2004). Lactic acid bacteria as functional starter cultures for the food fermentation industry. Trends in Food Science & Technology, 15, 67-78.
Lin, X., Tang, Y., Hu, Y., Lu, Y., Sun, Q., Lv, Y., Zhang, Q., Wu, C., Zhu, M., He, Q., & Chi, Y. (2021). Sodium reduction in traditional fermented foods: challenges, strategies, and perspectives. Journal of Agricultural and Food Chemistry, 69, 8065-8080.
Lynch, K. M., Zannini, E., Coffey, A., & Arendt, E. K. (2018). Lactic acid bacteria exopolysaccharides in foods and beverages: isolation, properties, characterization, and health benefits. Annual Review of Food Science and Technology, 9, 155-176.
Lynch, K. M., McSweeney, P. L. H., & Sheehan, J. J. (2018). Exopolysaccharide production by lactic acid bacteria and its application in dairy products. International Dairy Journal, 86, 34-45.
Jung, M. Y., Kim, T. W., Lee, C., Kim, J. Y., Song, H. S., Kim, Y. B., Ahn, S. W., Kim, J. S., Roh, S. W., & Lee, S. H. (2018). Role of jeotgal, a Korean traditional fermented fish sauce, in microbial dynamics and metabolite profiles during kimchi fermentation. Food Chemistry, 265, 135-143.
Marco, M. L., Heeney, D., Binda, S., Cifelli, C. J., Cotter, P. D., Foligné, B., Gänzle, M., Kort, R., Pasin, G., Pihlanto, A., Smid, E. J., & Hutkins, R. W. (2017). Health benefits of fermented foods: microbiota and beyond. Current Opinion in Biotechnology, 44, 94-102.
Papadimitriou, K., Alegría, Á., Bron, P. A., de Angelis, M., Gobbetti, M., Kleerebezem, M., Lemos, J. A., Linares, D. M., Ross, P., Stanton, C., Turroni, F., van Sinderen, D., Varmanen, P., Ventura, M., Zúñiga, M., Tsakalidou, E., & Kok, J. (2016). Stress physiology of lactic acid bacteria. Microbiology and Molecular Biology Reviews, 80, 837-890.
Park, E. J., Chun, J., Cha, C. J., Park, W. S., Jeon, C. O., & Bae, J. W. (2012). Bacterial community analysis during fermentation of ten representative kinds of kimchi with barcoded pyrosequencing. Food Microbiology, 30, 197-204.
Park, K. Y., Jeong, J. K., Lee, Y. Y., & Daily, J. W. (2014). Health benefits of kimchi (Korean fermented vegetables) as a probiotic food. Journal of Medicinal Food, 17, 6-20.
Patel, A., Prajapati, J. B., Holst, O., & Ljungh, Å. (2012). Determining probiotic potential of exopolysaccharide producing lactic acid bacteria isolated from vegetables and traditional Indian fermented foods. International Journal of Fermented Foods, 1, 87-101.
Quince, C., Walker, A. W., Simpson, J. T., Loman, N. J., & Segata, N. (2017). Shotgun metagenomics, from sampling to analysis. Nature Biotechnology, 35, 833-844.
Ruas-Madiedo, P., & de los Reyes-Gavilán, C. G. (2005). Methods for the screening, isolation, and characterization of exopolysaccharides produced by lactic acid bacteria. Journal of Dairy Science, 88, 843-856.
Ruas-Madiedo, P., & de los Reyes-Gavilán, C. G. (2005). Invited review: methods for the screening, isolation, and characterization of exopolysaccharides produced by lactic acid bacteria. Journal of Dairy Science, 88, 843-856.
Ruijssenaars, H. J., Stingele, F., & Hartmans, S. (2000). Biopolymer production by bacterial strains from carbohydrate-rich waste products. Applied Microbiology and Biotechnology, 64, 1-10.
Seo, B. J., Bajpai, V. K., Rather, I. A., & Park, Y. H. (2018). Partially purified exopolysaccharide from lactobacillus plantarum YML009 with total phenolic content, antioxidant and free radical scavenging efficacy. BMC Microbiology, 18, 35.
Seo, S. H., Park, S. E., Kim, E. J., Lee, K. I., Na, C. S., & Son, H. S. (2018). A GC-MS based metabolomics approach to determine the effect of salinity on kimchi. Food Research International, 105, 492-498.
Stewart, E. J. (2012). Growing unculturable bacteria. Journal of Bacteriology, 194, 4151-4160.
Tamang, J. P., Watanabe, K., & Holzapfel, W. H. (2016). Diversity of microorganisms in global fermented foods and beverages. Frontiers in Microbiology, 7, 377.
Walsh, A. M., Crispie, F., Kilcawley, K. N., O’Sullivan, O., & Cotter, P. D. (2017). Microbial succession and flavor production in the fermented dairy ecosystem. Current Opinion in Biotechnology, 49, 100-108.
Welman, A. D., & Maddox, I. S. (2003). Exopolysaccharides from lactic acid bacteria: perspectives and challenges. Trends in Biotechnology, 21, 269-274.
Wolfe, B. E., & Dutton, R. J. (2015). Fermented foods as experimentally tractable microbial ecosystems. Cell, 161, 49-55.
Wolfe, B. E., Button, J. E., Santarelli, M., & Dutton, R. J. (2014). Cheese rind communities provide tractable systems for in situ and in vitro studies of microbial diversity. Cell, 158, 422-433.
Zannini, E., Waters, D. M., Coffey, A., & Arendt, E. K. (2016). Production, properties, and industrial food application of lactic acid bacteria-derived exopolysaccharides. Applied Microbiology and Biotechnology, 100, 1121-1135.
Zotta, T., Parente, E., & Ricciardi, A. (2017). Aerobic metabolism in the genus Lactobacillus: impact on stress response and potential applications in the food industry. Journal of Applied Microbiology, 122, 857-869.