Ahansaz, N., Tarrah, A., Pakroo, S., Corich, V., Giacomini, A., 2023. Lactic acid bacteria in dairy foods: prime sources of antimicrobial compounds. Fermentation 9 (11), 964. https://doi.org/10.3390/fermentation9110964. Azzini, E., Raguzzini, A., Polito, A., 2021. A brief review on vitamin B12 deficiency looking at some case study reports in adults. Int. J. Mol. Sci. 22 (18), 9694. https:// doi.org/10.3390/ijms22189694. Bordiga, M., Travaglia, F., Locatelli, M., 2019. Valorisation of grape pomace: an approach that is increasingly reaching its maturity - a review. Int. J. Food Sci. Technol. 54 (4), 933-942. https://doi.org/10.1111/ijfs.14118. Calvillo, A., ´ Pellicer, T., Carnicer, M., Planas, A., 2022. Bioprocess strategies for vitamin B12 production by microbial fermentation and its market applications. Bioengineering 9 (8), 365. https://doi.org/10.3390/bioengineering9080365. Canon, F., Nidelet, T., Guédon, E., Thierry, A., Gagnaire, V., 2020. Understanding the mechanisms of positive microbial interactions that benefit lactic acid bacteria co cultures. Front. Microbiol. 11, 2088. https://doi.org/10.3389/fmicb.2020.02088. Chamlagain, B., Deptula, P., Edelmann, M., Kariluoto, S., Grattepanche, F., Lacroix, C., Varmanen, P., Piironen, V., 2016. Effect of the lower ligand precursors on vitamin B12 production by food-grade Propionibacteria. LWT-Food Sci. Technol. 72, 117-124. https://doi.org/10.1016/j.lwt.2016.04.023. Chamlagain, B., Sugito, T.A., Deptula, P., Edelmann, M., Kariluoto, S., Varmanen, P., Piironen, V., 2018. In situ production of active vitamin B12 in cereal matrices using Propionibacterium freudenreichii. Food Sci. Nutr. 6 (1), 67-76. https://doi.org/ 10.1002/fsn3.528. Chamlagain, B., Rautio, S., Edelmann, M., Ollilainen, V., Piironen, V., 2020. Niacin contents of cereal-milling products in food-composition databases need to be updated. J. Food Compos. Anal. 91, 103518. https://doi.org/10.1016/j. jfca.2020.103518. Chamlagain, B., Peltonen, L., Edelmann, M., Ramos-Diaz, J.M., Kemppinen, A., Jouppila, K., Varmanen, P., Piironen, V., 2021. Bioaccessibility of vitamin B12 synthesized by Propionibacterium freudenreichii and from products made with fermented wheat bran extract. Curr. Res. Food Sci. 4, 499-502. https://doi.org/ 10.1016/j.crfs.2021.07.009. Chamlagain, B., Loivamaa, I., Zhang, R., Hovilehto, E., Deptula, P., Edelmann, M., in vitamin B12 production by Propionibacterium freudenreichii. Int. J. Food Microbiol. Kantanen, K., Chamlagain, B., Succi, V., Sepp¨ anen, S., Edelmann, M., Siitonen, A., 444, 111463. https://doi.org/10.1016/j.ijfoodmicro.2025.111463. Cong, S., Zhang, X., Ji, J., Liu, X., Hu, N., 2024. Isolation and identification of blueberry2024. Fermentation of faba bean flour as a pre-treatment in high-moisture extrusion derived lactic acid bacteria and their probiotic, antioxidant, and fermentation properties. Food Biosci. 62, 104497. https://doi.org/10.1016/j.fbio.2024.104497. Cramer, H., Kessler, C.S., Sundberg, T., Leach, M.J., Schumann, D., Adams, J., Lauche, R., 10.3390/10.1016/j.fufo.2024.100518. 2017. Characteristics of americans choosing vegetarian and vegan diets for health reasons. J. Nutr. Educ. Behav. 49 (7), 561-567. https://doi.org/10.1016/j. jneb.2017.04.011. Dank, A., van Mastrigt, O., Boeren, S., Lillevang, S.K., Abee, T., Smid, E.J., 2021. Propionibacterium freudenreichii thrives in microaerobic conditions by complete oxidation of lactate to CO₂. Environ. Microbiol. 23 (6), 3116-3129. https://doi.org/ enzymatic determination of Dand L-lactic acid in selected foods and beverages: 10.1111/1462-2920.15532. De Vuyst, L., Neysens, P., 2005. The sourdough microflora: biodiversity and metabolic interactions. Trends Food Sci. Technol. 16, 43-56. https://doi.org/10.1016/j. tifs.2004.02.012. Deptula, P., Kylli, P., Chamlagain, B., Edelmann, M., Kariluoto, S., Piironen, V., Varmanen, P., 2015. BluB/CobT2 fusion enzyme activity reveals mechanisms responsible for production of active form of vitamin B12 by Propionibacterium freudenreichii. Microb. Cell Factories 14, 186. https://doi.org/10.1186/s12934-015- 835.1974. 0363-9. Deptula, P., Chamlagain, B., Edelmann, M., Sangsuwan, P., Nyman, T.A., Savijoki, K., Piironen, V., Varmanen, P., 2017. Food-like growth conditions support production of (3), 37. https://doi.org/10.3390/microorganisms5030037. active vitamin B12 by Propionibacterium freudenreichii 2067 without DMBI, the lower Litwinek, D., Boreczek, J., Gambu´s, H., Buksa, K., Berski, W., Kowalczyk, M., 2022. ligand base, or cobalt supplementation. Front. Microbiol. 8, 368. https://doi.org/ Developing lactic acid bacteria starter cultures for wholemeal rye flour bread with 10.3389/fmicb.2017.00368. Echegaray, N., Yilmaz, B., Sharma, H., Kumar, M., Pateiro, M., Ozogul, F., Lorenzo, J.M., 17, e0261677. https://doi.org/10.1371/journal.pone.0261677. 2023. A novel approach to Lactiplantibacillus plantarum: from probiotic properties to Loivamaa, I., Greis, M., Nikander, V., Edelmann, M., P¨ oys¨ the omics insights. Microbiol. Res. 268, 127289. https://doi.org/10.1016/j. micres.2022.127289. Edelmann, M., Chamlagain, B., Santin, M., Kariluoto, S., Piironen, V., 2016. Stability of 10.1016/j.fbio.2024.105807. added and in situ-produced vitamin B12 in breadmaking. Food Chem. 204, 21-28. https://doi.org/10.1016/j.foodchem.2016.02.071. EFSA ANS Panel, 2014. Scientific opinion on the re-evaluation of propionic acid (E 280), 10.1139/m68-199. sodium propionate (E 281), calcium propionate (E 282) and potassium propionate (E 283) as food additives. EFSA J. 12 (7), 3779. https://doi.org/10.2903/j. efsa.2014.3779. Falentin, H., Deutsch, S.M., Jan, G., Loux, V., Thierry, A., Parayre, S., Maillard, M.B., Moore, J.F., DuVivier, R., Johanningsmeier, S.D., 2022. Changes in the free amino acid Dherbécourt, J., Cousin, F.J., Jardin, J., Siguier, P., Couloux, A., Barbe, V., Vacherie, B., Wincker, P., Gibrat, J.F., Gaillardin, C., Lortal, S., 2010. The complete 599-611. https://doi.org/10.1111/1750-3841.15990. genome of Propionibacterium freudenreichii CIRM-BIA1T, a hardy actinobacterium with food and probiotic applications. PLoS One 5 (7), e11748. https://doi.org/ 10.1371/journal.pone.0011748. Fernández-Cruz, M.L., Mansilla, M.L., Tadeo, J.L., 2010. Mycotoxins in fruits and their nrgastro.2012.76. processed products: analysis, occurrence and health implications. J. Adv. Res. 1 (2), Niklewicz, A., Smith, A.D., Smith, A., Holzer, A., Klein, A., McCaddon, A., Molloy, A.M., 113-122. https://doi.org/10.1016/j.jare.2010.03.002. Gagnaire, V., Jardin, J., Rabah, H., Briard-Bion, V., Jan, G., 2015. Emmental cheese environment enhances Propionibacterium freudenreichii stress tolerance. PLoS One 10, Owen, P.J., 2023. The importance of vitamin B12 for individuals choosing plant e0135780. https://doi.org/10.1371/journal.pone.0135780. Gientka, I., Synowiec, A., Roszko, M., Nguyen, C.N.K., Pobiega, K., Kot, A.M., 2024. Comparison of physicochemical characteristics and microbial quality between commercially available organic and conventional japanese soy sauces. Appl. Sci. 14 S., 2013. Antimicrobial activity and composition profile of grape (Vitis vinifera) (9), 3784. https://doi.org/10.3390/app14093784. Głowacka-Rutkowska, A., Ulatowska, M., Empel, J., Kowalczyk, M., Boreczek, J., Łobocka, M., 2020. A kayvirus distant homolog of staphylococcal virulence determinants and VISA biomarker is a phage lytic enzyme. Viruses 12 (3), 292. https://doi.org/10.3390/v12030292. Green, R., Allen, L.H., Bjørke-Monsen, A.-L., Brito, A., Guéant, J.-L., Miller, J.W., Molloy, A.M., Nexo, E., Stabler, S., Toh, B.-H., Ueland, P.M., Yajnik, C., 2017. Vitamin B12 deficiency. Nat. Rev. Dis. Primers 3, 17040. https://doi.org/10.1038/ nrdp.2017.40. Guéant, J.L., Guéant-Rodriguez, R.M., Kosgei, V.J., Coelho, D., 2021. Causes and consequences of impaired methionine synthase activity in acquired and inherited disorders of vitamin B12 metabolism. Crit. Rev. Biochem. Mol. Biol. 57 (2), 133-155. Patra, A., Abdullah, S., Pradhan, R.C., 2022. Review on the extraction of bioactive https://doi.org/10.1080/10409238.2021.1979459. Hoteit, M., Khadra, R., Fadlallah, Z., Mourad, Y., Chahine, M., Skaiki, F., Al Manasfi, E., Bioprocess. 9, 14. https://doi.org/10.1186/s40643-022-00498-3. Chahine, A., Poh, O.B.J., Tzenios, N., 2024. Prevalence and time trends of low serum Pieszka, M., Gogol, P., Pietras, M., Pieszka, M., 2015. Valuable components of dried B12 levels and inadequate B12 dietary intake in Lebanese adults amidst the food insecurity situation: findings from a nationally representative cross-sectional study. natural antioxidants and nutraceuticals in the animal diet. Ann. Anim. Sci. 15 (2), Nutrients 16 (2), 226. https://doi.org/10.3390/nu16020226. Iqbal, A., Schulz, P., Rizvi, S.S.H., 2021. Valorization of bioactive compounds in fruit Piwowarek, K., Lipi´ nska, E., Ha´c-Szyma´ nczuk, E., Kot, A.M., Kieliszek, M., Bonin, S., pomace from agro-fruit industries: present insights and future challenges. Food Biosci. 44 (Part A), 101384. https://doi.org/10.1016/j.fbio.2021.101384. Jin, Q., O'Hair, J., Stewart, A.C., O'Keefe, S.F., Neilson, A.P., Kim, Y.T., McGuire, M., wastewater. Molecules 26 (13), 3965. https://doi.org/10.3390/molecules26133965. Lee, A., Wilder, G., Huang, H., 2019. Compositional characterization of different industrial white and red grape pomaces in Virginia and the potential valorization of obtaining valuable bacterial metabolites. Appl. Microbiol. Biotechnol. 107, the major components. Foods 8 (12), 667. https://doi.org/10.3390/foods8120667. Juarez del Valle, M., Lai˜ no, J.E., Savoy de Giori, G., LeBlanc, J.G., 2014. Riboflavin Piwowarek, K., Ruzik, L., Pobiega, K., Kalisz, S., 2025. Valorization of fruit by-products producing lactic acid bacteria as a biotechnological strategy to obtain bio-enriched with Propionibacterium freudenreichii: fortification of vitamin B12 in different types of soymilk. Food Res. Int. 62, 1015-1019. https://doi.org/10.1016/j. foodres.2014.05.029. Kahala, M., Blasco, L., Bragge, R., Porcellato, D., Østlie, H.M., Rundberget, T., Baz- Lomba, J.A., Pihlava, J.-M., Hellström, J., Gullberg Jørgensen, E., Joutsjoki, V., biological significance. Annu. Rev. Microbiol. 50, 137-181. https://doi.org/ Devold, T.G., Pihlanto, A., 2024. Lactic and propionic acid bacteria starter cultures 10.1146/annurev.micro.50.1.137. for improved nutritional properties of pea, faba bean and lentil. LWT-Food Sci. Kemppinen, L., Kemppinen, A., Kangas, M., Varmanen, P., Piironen, V., Jouppila, K., for simultaneous fortification of vitamin B12 and reduction of raffinose oligosaccharides in meat analogues. Future Foods. 10, 100518. https://doi.org/ Kruk, M., Varmanen, P., Edelmann, M., Chamlagain, B., Trząskowska, M., 2024. Food by product valorisation in nutrients through spent brewer's yeast bioprocessing with Propionibacterium freudenreichii. J. Clean. Prod. 434, 140102. https://doi.org/ 10.1016/j.jclepro.2023.140102. Lacorn, M., Hektor, T., 2025. Validation of Enzytec™ liquid D− /L-lactic acid for official method 2024.08 first action. J. AOAC Int. 108 (5), 692-707. https://doi.org/ 10.1093/jaoacint/qsaf046. Lau, K.Q., Sabran, M.R., Shafie, S.R., 2021. Utilization of vegetable and fruit by-products as functional ingredient and food. Front. Nutr. 8, 661693. https://doi.org/10.3389/ fnut.2021.661693. Lee, I.H., Fredrickson, A., Tsuchiya, H., 1974. Diauxic growth of Propionibacterium shermanii. Appl. Microbiol. 28, 831-835. https://doi.org/10.1128/am.28.5.831- Leyva Salas, M., Mounier, J., Valence, F., Coton, M., Thierry, A., Coton, E., 2017. Antifungal microbial agents for food biopreservation-a review. Microorganisms 5 improved functionality, nutritional value, taste, appearance and safety. PLoS One a, M., Varmanen, P., Saris, P.E. J., 2025. Two-step fermentation to produce vitamin B12 containing beer using Propionibacterium freudenreichii and yeast. Food Biosci. 63, 105807. https://doi.org/ Malik, A.C., Reinbold, G.W., Vedamuthu, E.R., 1968. An evaluation of the taxonomy of Propionibacterium. Can. J. Microbiol. 14 (11), 1185-1191. https://doi.org/ Martens, J.H., Barg, H., Warren, M.A., Jahn, D., 2002. Microbial production of vitamin B12. Appl. Microbiol. Biotechnol. 58, 275-285. https://doi.org/10.1007/s00253- 001-0902-7.profile of pickling cucumber during lactic acid fermentation. J. Food Sci. 87 (2), Nielsen, M.J., Rasmussen, M.R., Andersen, C.B., Nexø, E., Moestrup, S.K., 2012. Vitamin B12 transport from food to the body's cells—a sophisticated, multistep pathway. Nat. Rev. Gastroenterol. Hepatol. 9, 345-354. https://doi.org/10.1038/ Wolffenbuttel, B.H.R., Nexo, E., McNulty, H., Refsum, H., Gueant, J.L., Dib, M.J., Ward, M., Murphy, M., Green, R., Ahmadi, K.R., Hannibal, L., Warren, M.J., based diets. Eur. J. Nutr. 62 (3), 1551-1559. https://doi.org/10.1007/s00394-022- 03025-4. Oliveira, D.A., Salvador, A.A., Smˆ ania, A., Smˆ ania, E.F.A., Maraschin, M., Ferreira, S.R. pomace extracts obtained by supercritical fluids. J. Biotechnol. 164 (3), 423-432. https://doi.org/10.1016/j.jbiotec.2012.09.014. Oszmiański, J., Wojdyło, A., Lachowicz, S., 2016. Effect of dried powder preparation process on polyphenolic content and antioxidant activity of blue honeysuckle berries (Lonicera caerulea L. var. kamtschatica) . LWT-Food Sci. Technol. 67, 214-222. https://doi.org/10.1016/j.lwt.2015.11.051. Palíková, I., Heinrich, J., Bednáˇr, P., Marhol, P., Kˇren, V., Cvak, L., Valentová, K., Růˇziˇcka, F., Hol´ a, V., Koláˇr, M., Sim´ ˇ anek, V., Ulrichov´ a, J., 2008. Constituents and antimicrobial properties of blue honeysuckle: a novel source for phenolic antioxidants. J. Agric. Food Chem. 56 (24), 11883-11889. https://doi.org/10.1021/ jf8026233.compounds and characterization of fruit industry by-products. Bioresour. pomaces of chokeberry, black currant, strawberry, apple and carrot as a source of 475-491. https://doi.org/10.2478/aoas-2014-0072. 2021. Use of Propionibacterium freudenreichii T82 strain for effective biosynthesis of propionic acid and trehalose in a medium with apple pomace extract and potato Piwowarek, K., Lipi´ nska, E., Kieliszek, M., 2023. Reprocessing of side-streams towards 2169-2208. https://doi.org/10.1007/s00253-023-12458-8. fruit pomace. LWT-Food Sci. Technol. 237, 118785. https://doi.org/10.1016/j. lwt.2025.118785. Roth, J.R., Lawrence, J.G., Bobik, T.A., 1996. Cobalamin (Coenzyme B12): synthesis and Sirohi, R., Tarafdar, A., Singh, S., Negi, T., Gaur, V.K., Gnansounou, E., Bharathiraja, B., trends and opportunities for sustainable biorefinery. Bioresour. Technol. 314, 123771. https://doi.org/10.1016/j.biortech.2020.123771. Smid, E.J., Lacroix, C., 2013. Microbe-microbe interactions in mixed culture food fermentations. Curr. Opin. Biotechnol. 24, 148-154. https://doi.org/10.1016/j. copbio.2012.11.007. Soares, M.G., Bevilaqua, G.C., Tassi, E.M.M., ´ Schmidt, V.C.R., 2023. Fermented foods 312-317. https://doi.org/10.1016/j.indcrop.2013.04.039. and beverages: a potential in situ vitamin B12 biofortification - a literature review. Watanabe, F., 2007. Vitamin B12 sources and bioavailability. Exp. Biol. Med. (Maywood) Int. J. Food Sci. 74 (6), 655-667. https://doi.org/10.1080/ 09637486.2023.2248422. SOP M 3535, 2024. Bestimmung von Vitamin B12 in Lebensmittel. Futtermittel und Nahrungserg¨ anzungsmittel mittels LC-MS/MS (ang. Determination of vitamin B12 in food, feed and dietary supplements using LC-MS/MS). Tangyu, M., Fritz, M., Ye, L., Arag˜ ao Börner, R., Morin-Rivron, D., Campos-Gim´ Bolten, C.J., Bogicevic, B., Wittmann, C., 2022. Co-cultures of Propionibacterium freudenreichii and Bacillus amyloliquefaciens cooperatively upgrade sunflower seed milk to high levels of vitamin B12 and multiple co-benefits. Microb. Cell Factories fortification. LWT-Food Sci. Technol. 137, 110431. https://doi.org/10.1016/j. 21, 48. https://doi.org/10.1186/s12934-022-01773-w. Thierry, A., Deutsch, S.-M., Falentin, H., Dalmasso, M., Cousin, F.J., Jan, G., 2011. New Ye, K., Shijo, M., Jin, S., Shimizu, K., 1996. Efficient production of vitamin B 12 from insights into physiology and metabolism of Propionibacterium freudenreichii. Int. J. propionic acid bacteria under periodic variation of dissolved oxygen concentration. Food Microbiol. 149 (1), 19-27. https://doi.org/10.1016/j. ijfoodmicro.2011.04.026. Tumbas Saponjac, ˇ V., Gironés-Vilaplana, A., Djilas, S., Mena, P., Cetkovi´ A., Canadanovi´ ˇ c-Brunet, J., Vuli´c, J., Stajˇci´c, S., Vinˇci´c, M., 2015. Chemical bioconversion, and bioactivities of vitamin B12 for vegetarian diet. Food Chem. 463, composition and potential bioactivity of strawberry pomace. RSC Adv. 5, Vargas-Luna, C., Godoy, L., Benavides, S., Ceppi de Lecco, C., Urtubia, A., Franco, W., 2025. Screening and selection of native lactic acid bacteria isolated from Chilean grapes. Foods 14 (1), 143. https://doi.org/10.3390/foods14010143. Wang, L., Yang, X., Qin, P., Shan, F., Ren, G., 2013. Flavonoid composition, antibacterial and antioxidant properties of tartary buckwheat bran extract. Ind. Crop. Prod. 49, 232 (10), 1266-1274. https://doi.org/10.3181/0703-MR-67. Xie, C., Coda, R., Chamlagain, B., Varmanen, P., Piironen, V., Katina, K., 2019. Co fermentation of Propionibacterium freudenreichii and Lactobacillus brevis in wheat bran for in situ production of vitamin B12. Front. Microbiol. 10, 1541. https://doi.org/ 10.3389/fmicb.2019.01541. enez, E., Xie, C., Coda, R., Chamlagain, B., Edelmann, M., Varmanen, P., Piironen, V., Katina, K., 2021. Fermentation of cereal, pseudo-cereal and legume materials with Propionibacterium freudenreichii and Levilactobacillus brevis for vitamin B12 lwt.2020.110431. J. Ferment. Bioeng. 82 (5), 484-491. https://doi.org/10.1016/S0922-338X(97) 86988-7. ´ c, G., Moreno, D. Zhou, Y., He, A., Xu, B., 2025. Natural resources, quantification, microbial 140849. https://doi.org/10.1016/j.foodchem.2024.14084.