Lactobacillus exopolysaccharides: New perspectives on engineering strategies, physiochemical functions, and immunomodulatory effects on host health

Trends in Food Science & Technology - Tập 103 - Trang 36-48 - 2020
Muhammad Shahid Riaz Rajoka1,2, Yiguang Wu1, Hafiza Mahreen Mehwish2, Manisha Bansal2, Liqing Zhao1
1College of Chemistry and Environmental Engineering, Shenzhen University, Shenzhen, Guangdong 518060, China
2School of Pharmaceutical Science, Guangdong Key Laboratory for Genome Stability & Human Disease Prevention, Shenzhen Key Laboratory of Novel Natural Health Care Products, Engineering Laboratory of Shenzhen Natural Small Molecule Innovative Drugs, Health Science Center, Shenzhen University, Shenzhen 518060, PR China

Tài liệu tham khảo

Altmann, 2016, Genome analysis and characterisation of the exopolysaccharide produced by Bifidobacterium longum subsp. longum 35624™, PloS One, 11, 10.1371/journal.pone.0162983 Amiri, 2019, Exopolysaccharides production by Lactobacillus acidophilus LA5 and Bifidobacterium animalis subsp. lactis BB12: Optimization of fermentation variables and characterization of structure and bioactivities, International Journal of Biological Macromolecule, 123, 752, 10.1016/j.ijbiomac.2018.11.084 Anwar, 2010, Inulin and levan synthesis by probiotic Lactobacillus gasseri strains: Characterization of three novel fructansucrase enzymes and their fructan products, Microbiology, 156, 1264, 10.1099/mic.0.036616-0 Aryantini, 2017, Complete genome sequence of Lactobacillus fermentum MTCC 25067 (Formerly TDS030603), a viscous exopolysaccharide producing strain isolated from indian fermented milk, Genome Announcements, 5, 10.1128/genomeA.00091-17 Ates, 2015, Systems biology of microbial exopolysaccharides production, Frontiers in Bioengineering and Biotechnology, 3, 10.3389/fbioe.2015.00200 Badel, 2011, New perspectives for Lactobacilli exopolysaccharides, Biotechnology Advances, 29, 54, 10.1016/j.biotechadv.2010.08.011 Balzaretti, 2017, A novel rhamnose-rich hetero-exopolysaccharide isolated from Lactobacillus paracasei DG activates THP-1 human monocytic cells, Applied and Environmental Microbiology, 83, 10.1128/AEM.02702-16 Bernal, 2012, Promising biotechnological applications of antibiofilm exopolysaccharides, Microbial Biotechnology, 5, 670, 10.1111/j.1751-7915.2012.00359.x Bhat, 2019, Hypocholesterolemic potential and bioactivity spectrum of an exopolysaccharide from a probiotic isolate Lactobacillus paracasei M7, Bioactive Carbohydrate and Dietary Fibre, 19, 100191, 10.1016/j.bcdf.2019.100191 Bleau, 2010, Intermediate chains of exopolysaccharides from Lactobacillus rhamnosus RW-9595M increase IL-10 production by macrophages, Journal of Applied Microbiology, 108, 666, 10.1111/j.1365-2672.2009.04450.x Boels, 2001, Sugar catabolism and its impact on the biosynthesis and engineering of exopolysaccharide production in lactic acid bacteria, International Dairy Journal, 11, 723, 10.1016/S0958-6946(01)00116-9 Boels, 2001, Functional analysis of the Lactococcus lactis galU and galE genes and their impact on sugar nucleotide and exopolysaccharide biosynthesis, Applied and Environmental Microbiology, 67, 3033, 10.1128/AEM.67.7.3033-3040.2001 Bouazzaoui, 2006, Use of antisense RNA to modulate glycosyltransferase gene expression and exopolysaccharide molecular mass in Lactobacillus rhamnosus, Journal of Microbiological Methods, 65, 216, 10.1016/j.mimet.2005.07.011 Burns, 2011, Technological characterization and survival of the exopolysaccharide-producing strain Lactobacillus delbrueckii subsp. lactis 193 and its bile-resistant derivative 193+ in simulated gastric and intestinal juices, Journal of Dairy Reserach, 78, 357, 10.1017/S0022029911000355 Caggianiello, 2016, Exopolysaccharides produced by lactic acid bacteria: From health-promoting benefits to stress tolerance mechanisms, Applied Microbiology and Biotechnology, 100, 3877, 10.1007/s00253-016-7471-2 Charchoghlyan, 2017, Rheological properties and volatile composition of fermented milk prepared by exopolysaccharide-producing Lactobacillus acidophilus n.v. Er2 317/402 strain Narine, Biotechnology and Bioprocess Engineering, 22, 327, 10.1007/s12257-017-0065-8 Chawla, 2009, Microbial cellulose: Fermentative production and applications, Food Technology and Biotechnology, 47, 107 Chen, 2016, Structure-function relationships of bacterial and enzymatically produced reuterans and dextran in sourdough bread baking application, International Journal of Food Microbiology, 239, 95, 10.1016/j.ijfoodmicro.2016.06.010 Chen, 2014, Exopolysaccharides synthesized by Lactobacillus reuteri protect against enterotoxigenic Escherichia coli in piglets, Applied and Environmental Microbiology, 80, 5752, 10.1128/AEM.01782-14 Chen, 2019, A role of exopolysaccharide produced by Streptococcus thermophilus in the intestinal inflammation and mucosal barrier in Caco-2 monolayer and dextran sulphate sodium-induced experimental murine colitis, Molecules, 24, 513, 10.3390/molecules24030513 Ciszek-Lenda, 2011, Biological functions of exopolysaccharides from probiotic bacteria, Central European Journal of Immunology, 36, 51 Ciszek-Lenda, 2011, Immunoregulatory potential of exopolysaccharide from Lactobacillus rhamnosus KL37: Effects on the production of inflammatory mediators by mouse macrophages, International Journal of Experimental Pathology, 92, 382, 10.1111/j.1365-2613.2011.00788.x Ciszek-Lenda, 2013, Further studies on immunomodulatory effects of exopolysaccharide isolated from Lactobacillus rhamnosus KL37C, Central European Journal of Immunology, 38, 289, 10.5114/ceji.2013.37743 Côté, 2012, Cloning, expression, and characterization of an insoluble glucan-producing glucansucrase from Leuconostoc mesenteroides NRRL B-1118, Applied Microbiology and Biotechnology, 93, 2387, 10.1007/s00253-011-3562-2 Cui, 2017, New advances in exopolysaccharides production of Streptococcus thermophilus, Archives of Microbiology, 199, 799, 10.1007/s00203-017-1366-1 Cuthbertson, 2009, Pivotal roles of the outer membrane polysaccharide export and polysaccharide copolymerase protein families in export of extracellular polysaccharides in gram-negative bacteria, Microbiology and Molecular Biology Reviews, 73, 155, 10.1128/MMBR.00024-08 De Vuyst, 2001, Recent developments in the biosynthesis and applications of heteropolysaccharides from lactic acid bacteria, Interenational Dairy Journal, 11, 687, 10.1016/S0958-6946(01)00114-5 De Vuyst, 1999, Heteropolysaccharides from lactic acid bacteria, FEMS Microbiology Reviews, 23, 153, 10.1016/S0168-6445(98)00042-4 De Vuyst, 2003, Exopolysaccharide-producing Streptococcus thermophilus strains as functional starter cultures in the production of fermented milks, International Dairy Journal, 13, 707, 10.1016/S0958-6946(03)00105-5 Delgado, 2019, Fermentation of commercial soy beverages with lactobacilli and bifidobacteria strains featuring high β-glucosidase activity, Innovative Food Science & Emerging Technologies, 51, 148, 10.1016/j.ifset.2018.03.018 Dertli, 2018, Structural analysis of the α-d-glucan produced by the sourdough isolate Lactobacillus brevis E25, Food Chemistry, 242, 45, 10.1016/j.foodchem.2017.09.017 Di, 2018, Research methods for structural analysis of lactic acid bacteria induced exopolysaccharides, Chinese Journal of Analytical Chemistry, 46, 875, 10.1016/S1872-2040(18)61091-6 Dols, 1997, Dextransucrase production by Leuconostoc mesenteroides NRRL B-1299. comparison with L. mesenteroides NRRL B-512F, Enzyme and Microbial Technology, 20, 523, 10.1016/S0141-0229(96)00189-5 Domingos-Lopes, 2017, Immunomodulatory activity of exopolysaccharide producing Leuconostoc citreum strain isolated from Pico cheese, Journal of Functional Foods, 33, 235, 10.1016/j.jff.2017.03.054 Du, 2017, Molecular weight and helix conformation determine intestinal anti-inflammatory effects of exopolysaccharide from Schizophyllum commune, Carbohydrate Polymers, 172, 68, 10.1016/j.carbpol.2017.05.032 Du, 2017, Optimization, purification and structural characterization of a dextran produced by L. mesenteroides isolated from Chinese sauerkraut, Carbohydrate Polymers, 174, 409, 10.1016/j.carbpol.2017.06.084 Dueñas-Chasco, 1998, Structural analysis of the exopolysaccharides produced by Lactobacillus spp. G-77, Carbohydrate Research, 307, 125, 10.1016/S0008-6215(98)00034-2 Faber, 2001, Structure of the extracellular polysaccharide produced by Lactobacillus delbrueckii subsp. bulgaricus 291, Carbohydrate Research, 331, 183, 10.1016/S0008-6215(01)00012-X Faber, 2002, Modeling of the structure in aqueous solution of the exopolysaccharide produced by Lactobacillus helveticus 766, Biopolymers, 63, 66, 10.1002/bip.1063 Fanning, 2012, Bifidobacterial surface-exopolysaccharide facilitates commensal-host interaction through immune modulation and pathogen protection, Proceeding of the National Academy of Science of the Unitated States of America, 109, 2108, 10.1073/pnas.1115621109 Fearon, 1996, The instructive role of innate immunity in the acquired immune response, Science, 272, 50, 10.1126/science.272.5258.50 Franken, 2016, Macrophages: Sentinels and regulators of the immune system, Cellular Microbiology, 18, 475, 10.1111/cmi.12580 Fraunhofer, 2018, Characterization of β-glucan formation by Lactobacillus brevis TMW 1.2112 isolated from slimy spoiled beer, International Journal of Biological Macromolecules, 107, 874, 10.1016/j.ijbiomac.2017.09.063 Fraunhofer, 2018, Influence of different sugars and initial pH on β-glucan formation by Lactobacillus brevis TMW 1.2112, Current Microbiology, 75, 794, 10.1007/s00284-018-1450-z Garai-Ibabe, 2010, Naturally occurring 2-substituted (1,3)-β-d-glucan producing Lactobacillus suebicus and Pediococcus parvulus strains with potential utility in the production of functional foods, Bioresource Technology, 101, 9254, 10.1016/j.biortech.2010.07.050 Gentès, 2011, Gel formation and rheological properties of fermented milk with in situ exopolysaccharide production by lactic acid bacteria, Dairy Science & Technology, 91, 645, 10.1007/s13594-011-0039-0 Gentès, 2011, Gel formation and rheological properties of fermented milk with in situ exopolysaccharide production by lactic acid bacteria, Dairy Science & Technology, 91, 645, 10.1007/s13594-011-0039-0 Gerwig, 2013, Structure determination of the exopolysaccharide of Lactobacillus fermentum TDS030603—a revision, Carbohydrate Research, 378, 84, 10.1016/j.carres.2013.04.026 Girard, 2007, Gelation of skim milk containing anionic exopolysaccharides and recovery of texture after shearing, Food Hydrocolloids, 21, 1031, 10.1016/j.foodhyd.2006.07.012 Gontijo, 2020, Phylogenetic distribution of the bacteriocin repertoire of lactic acid bacteria species associated with artisanal cheese, Food Research International, 128, 108783, 10.1016/j.foodres.2019.108783 Górska-Frączek, 2013, The structure and immunoreactivity of exopolysaccharide isolated from Lactobacillus johnsonii strain 151, Carbohydrate Research, 378, 148, 10.1016/j.carres.2013.05.012 Górska-Frączek, 2011, Structural studies of the exopolysaccharide consisting of a monosaccharide repeating unit isolated from Lactobacillus rhamnosus KL37B, Carbohydrate Research, 346, 2926, 10.1016/j.carres.2011.10.024 Górska, 2016, Chemical characterization and immunomodulatory properties of polysaccharides isolated from probiotic Lactobacillus casei LOCK 0919, Glycobiology, 26, 1014, 10.1093/glycob/cww047 Gotoh, 2017, Effect of orally administered exopolysaccharides produced by Lactococcus lactis subsp. cremoris FC on a mouse model of dermatitis induced by repeated exposure to 2,4,6-trinitro-1-chlorobenzene, Journal of Functional Foods, 35, 43, 10.1016/j.jff.2017.04.045 Guo, 2013, Antioxidant and immunomodulatory activity of selenium exopolysaccharide produced by Lactococcus lactis subsp. lactis, Food Chemistry, 138, 84, 10.1016/j.foodchem.2012.10.029 Han, 2016, Improvement of the texture of yogurt by use of exopolysaccharide producing lactic acid bacteria, BioMed Research International, 10.1155/2016/7945675 Hidalgo-Cantabrana, 2014, Exopolysaccharide-producing Bifidobacterium animalis subsp. lactis strains and their polymers elicit different responses on immune cells from blood and gut associated lymphoid tissue, Anaerobe, 26, 24, 10.1016/j.anaerobe.2014.01.003 Hosono, 1997, Characterization of a water-soluble polysaccharide fraction with immunopotentiating activity from Bifidobacterium adolescentis M101-4, Bioscience Biotechnology and Biochemistry, 61, 312, 10.1271/bbb.61.312 Ibrahim, 2015, Effects of exopolysaccharide-producing starter cultures on physicochemical, rheological and sensory properties of fermented camel's milk, Emirates Journal of Food and Agriculture, 27, 374, 10.9755/ejfa.v27i4.19911 Inturri, 2017, Chemical and biological properties of the novel exopolysaccharide produced by a probiotic strain of Bifidobacterium longum, Carbohydrate Polymers, 174, 1172, 10.1016/j.carbpol.2017.07.039 Islam, 2014, Synthesis of bacterial polysaccharides via the Wzx/Wzy-dependent pathway, Canadian Journal of Microbiology, 60, 697, 10.1139/cjm-2014-0595 Ismail, 2010, Production, purification and structural characterization of an exopolysaccharide produced by a probiotic Lactobacillus plantarum MTCC 9510, Archives of Microbiology, 192, 1049, 10.1007/s00203-010-0636-y Ismail, 2013, Exposition of antitumour activity of a chemically characterized exopolysaccharide from a probiotic Lactobacillus plantarum MTCC 9510, Biologia, 68, 1041, 10.2478/s11756-013-0275-2 Ismail, 2014, Molecular characterization of an exopolysaccharide from a probiotic Lactobacillus plantarum MTCC 9510 and its efficacy to improve the texture of starchy food, Journal of Food Science & Technology, 51, 4012, 10.1007/s13197-013-0928-8 Jiménez-Guzmán, 2009, Use of an exopolysaccharide-producing strain of Streptococcus thermophilus in the manufacture of Mexican Panela cheese, Lebensmittel-Wissenschaft und -Technologie- Food Science and Technology, 42, 1508, 10.1016/j.lwt.2009.04.009 Jin, 2010, Preparation and biological activities of an exopolysaccharide produced by Enterobacter cloacae Z0206, Carbohydrate Polymers, 81, 607, 10.1016/j.carbpol.2010.03.020 Kanamarlapudi, 2017, Characterization of exopolysaccharide produced by Streptococcus thermophilus CC30, BioMed Research International, 10.1155/2017/4201809 Kanmani, 2018, Genomic characterization of Lactobacillus delbrueckii TUA4408L and evaluation of the antiviral activities of its extracellular polysaccharides in porcine intestinal epithelial cells, Frontiers in Immunology, 9, 10.3389/fimmu.2018.02178 Kanmani, 2018, Exopolysaccharides from Lactobacillus delbrueckii OLL1073R-1 modulate innate antiviral immune response in porcine intestinal epithelial cells, Molecular Immunology, 93, 253, 10.1016/j.molimm.2017.07.009 Kavitake, 2016, Characterization of a novel galactan produced by Weissella confusa KR780676 from an acidic fermented food, International Journal of Biological Macromolecules, 86, 681, 10.1016/j.ijbiomac.2016.01.099 Kotzamanidis, 2010, Evaluation of adhesion capacity, cell surface traits and immunomodulatory activity of presumptive probiotic Lactobacillus strains, International Journal of Food Microbiology, 140, 154, 10.1016/j.ijfoodmicro.2010.04.004 Kralj, 2004, Glucan synthesis in the genus Lactobacillus: Isolation and characterization of glucansucrase genes, enzymes and glucan products from six different strains, Microbiology, 150, 3681, 10.1099/mic.0.27321-0 Kristo, 2011, The role of exopolysaccharide produced by Lactococcus lactis subsp. cremoris in structure formation and recovery of acid milk gels, International Dairy Journal, 21, 656, 10.1016/j.idairyj.2011.02.002 Kubo, 2017, Cross talk of innate and acquired immunity in allergy, Arerugi, 66, 179 Laws, 2001, Biosynthesis, characterisation, and design of bacterial exopolysaccharides from lactic acid bacteria, Biotechnology Advances, 19, 597, 10.1016/S0734-9750(01)00084-2 Leathers, 2004, Transformation of alternan-producing strains of Leuconostoc by electroporation, Biotechnology Letters, 26, 1119, 10.1023/B:BILE.0000035482.41491.7e Lee, 2016, Strain-specific features of extracellular polysaccharides and their impact on Lactobacillus plantarum host interactions, Applied and Environmental Microbiology, 82, 3959, 10.1128/AEM.00306-16 Levander, 2002, Enhanced exopolysaccharide production by metabolic engineering of Streptococcus thermophilus, Applied and Environmental Microbiology, 68, 784, 10.1128/AEM.68.2.784-790.2002 Li, 2010, Strain improvement and metabolic flux modeling of wild-type and mutant Alcaligenes sp. NX-3 for synthesis of exopolysaccharide welan gum, Biotechnology and Bioprocess Engineering, 15, 777, 10.1007/s12257-010-0021-3 Li, 2015, Two UDP-glucuronic acid decarboxylases involved in the biosynthesis of a bacterial exopolysaccharide in Paenibacillus elgii, Applied Microbiology and Biotechnology, 99, 3127, 10.1007/s00253-014-6362-7 Li, 2016, Characterization, anti-inflammatory and antiproliferative activities of natural and sulfonated exo-polysaccharides from Streptococcus thermophilus ASCC 1275, Journal of Food Science, 81, M1167, 10.1111/1750-3841.13276 Li, 2015, Characterization of a novel polysaccharide with anti-colon cancer activity from Lactobacillus helveticus MB2-1, Carbohydrate Research, 411, 6, 10.1016/j.carres.2014.12.014 Limoli, 2015, Bacterial extracellular polysaccharides in biofilm formation and function, Microbiology Spectrum, 3, 1, 10.1128/microbiolspec.MB-0011-2014 Liu, 2017, Increasing the bioflocculant production and identifying the effect of overexpressing epsB on the synthesis of polysaccharide and γ-PGA in Bacillus licheniformis, Microbial Cell Factories, 16, 163, 10.1186/s12934-017-0775-9 Lynch, 2018, Exopolysaccharide producing lactic acid bacteria: Their techno-functional role and potential application in gluten-free bread products, Food Research International, 110, 52, 10.1016/j.foodres.2017.03.012 Lynch, 2018, Exopolysaccharide producing lactic acid bacteria: Their techno-functional role and potential application in gluten-free bread products, Food Research International, 110, 52, 10.1016/j.foodres.2017.03.012 Lynch, 2018, Lactic acid bacteria exopolysaccharides in foods and beverages: Isolation, properties, characterization, and health benefits, Annual Reviews of Food Science and Technology, 9, 155, 10.1146/annurev-food-030117-012537 Makino, 2006, Immunomodulatory effects of polysaccharides produced by Lactobacillus delbrueckii ssp. bulgaricus OLL1073R-1, Journal of Dairy Science, 89, 2873, 10.3168/jds.S0022-0302(06)72560-7 Makino, 2016, Enhanced natural killer cell activation by exopolysaccharides derived from yogurt fermented with Lactobacillus delbrueckii ssp. bulgaricus OLL1073R-1, Journal of Dairy Science, 99, 915, 10.3168/jds.2015-10376 Matsuzaki, 2015, Exopolysaccharides produced by Leuconostoc mesenteroides strain NTM048 as an immunostimulant to enhance the mucosal barrier and influence the systemic immune response, Journal of Agricultural and Food Chemistry, 63, 7009, 10.1021/acs.jafc.5b01960 Matsuzaki, 2014, Immunomodulating activity of exopolysaccharide-producing Leuconostoc mesenteroides strain NTM048 from green peas, Journal of Applied Microbiology, 116, 980, 10.1111/jam.12411 Mazhar, 2020, The Prospects for the therapeutic implications of genetically engineered probiotics, Journal of Food Quality, 10.1155/2020/9676452 Mende, 2012, Exopolysaccharide production by three different strains of Streptococcus thermophilus and its effect on physical properties of acidified milk, Engineering in Life Sciences, 12, 466, 10.1002/elsc.201100114 Mende, 2013, Addition of purified exopolysaccharide isolates from Streptococcus thermophilus to milk and their impact on the rheology of acid gels, Food Hydrocolloids, 32, 178, 10.1016/j.foodhyd.2012.12.011 Mileti, 2009, Comparison of the immunomodulatory properties of three probiotic strains of lactobacilli using complex culture systems: Prediction for in vivo efficacy, PloS One, 4, 10.1371/journal.pone.0007056 Navarini, 2001, Isolation and characterization of the exopolysaccharide produced by Streptococcus thermophilus SFi201, International Journal of Biological Macromolecules, 28, 219, 10.1016/S0141-8130(01)00118-0 Nederby, 2018, Quantification of NK cell activity using whole blood: Methodological aspects of a new test, Journal of Immunological Methods, 458, 21, 10.1016/j.jim.2018.04.002 Newbrun, 1968, Physico-chemical characteristics of the levan produced by Streptococcus salivarius, Carbohydrate Research, 6, 165, 10.1016/S0008-6215(00)81506-2 Ni, 2018, Biosynthesis of levan from sucrose using a thermostable levansucrase from Lactobacillus reuteri LTH5448, International Journal of Biological Macromolecules, 113, 29, 10.1016/j.ijbiomac.2018.01.187 Nikolic, 2012, Characterisation of the exopolysaccharide (EPS)-producing Lactobacillus paraplantarum BGCG11 and its non-EPS producing derivative strains as potential probiotics, International Journal of Food Microbiology, 158, 155, 10.1016/j.ijfoodmicro.2012.07.015 Notararigo, 2014, Immunomodulation of human macrophages and myeloid cells by 2-substituted (1-3)-β-D-glucan from P. parvulus 2.6, Carbohydrate Polymers, 112, 109, 10.1016/j.carbpol.2014.05.073 Nourikyan, 2015, Autophosphorylation of the bacterial tyrosine-kinase CpsD connects capsule synthesis with the cell cycle in Streptococcus pneumoniae, PLoS Genetics, 11, 10.1371/journal.pgen.1005518 Nwodo, 2012, Bacterial exopolysaccharides: Functionality and prospects, International Journal of Molecular Science, 13, 14002, 10.3390/ijms131114002 Ołdak, 2017, Comparison of antibacterial activity of Lactobacillus plantarum strains isolated from two different kinds of regional cheeses from Poland: Oscypek and korycinski cheese, BioMed Research International, 10.1155/2017/6820369 Patel, 2012, Potentials of exopolysaccharides from lactic Acid bacteria, Indian Journal of Microbiology, 52, 3, 10.1007/s12088-011-0148-8 Patten, 2014, The structure and immunomodulatory activity on intestinal epithelial cells of the EPSs isolated from Lactobacillus helveticus sp. Rosyjski and Lactobacillus acidophilus sp. 5e2, Carbohydrate Research, 384, 119, 10.1016/j.carres.2013.12.008 Petersone, 2018, T cell/B cell collaboration and autoimmunity: An intimate relationship, Frontiers in Immunology, 9, 10.3389/fimmu.2018.01941 Prechtl, 2018, Cold and salt stress modulate amount, molecular and macromolecular structure of a Lactobacillus sakei dextran, Food Hydrocolloids, 82, 73, 10.1016/j.foodhyd.2018.04.003 Qin, 2015, Comparative transcriptome analysis reveals that lactose acts as an inducer and provides proper carbon sources for enhancing exopolysaccharide yield in the deep-sea bacterium Zunongwangia profunda SM-A87, PloS One, 10 Rahbar Saadat, 2019, A comprehensive review of anticancer, immunomodulatory and health beneficial effects of the lactic acid bacteria exopolysaccharides, Carbohydrate Polymers, 217, 79, 10.1016/j.carbpol.2019.04.025 Rajoka, 2019, Characterization, the antioxidant and antimicrobial activity of exopolysaccharide isolated from poultry origin Lactobacilli, Probiotics and Antimicrobial Proteins, 11, 1132, 10.1007/s12602-018-9494-8 Rajoka, 2019, Characterization and anti-tumor activity of exopolysaccharide produced by Lactobacillus kefiri isolated from Chinese kefir grains, Journal of Functional Foods, 63, 103588, 10.1016/j.jff.2019.103588 Rajoka, 2018, Functional characterization and biotechnological potential of exopolysaccharide produced by Lactobacillus rhamnosus strains isolated from human breast milk, LWT-Food Science and Technology, 89, 638, 10.1016/j.lwt.2017.11.034 Rajoka, 2017, Identification, characterization, and probiotic potential of Lactobacillus rhamnosus isolated from human milk, LWT-Food Science and Technology, 84, 271, 10.1016/j.lwt.2017.05.055 Rajoka, 2017, Interaction between diet composition and gut microbiota and its impact on gastrointestinal tract health, Food Science and Human Wellness, 6, 121, 10.1016/j.fshw.2017.07.003 Rajoka, 2017, Capacity of lactic acid bacteria in immunity enhancement and cancer prevention, Applied Microbiology and Biotechnology, 101, 35, 10.1007/s00253-016-8005-7 Ren, 2016, Bioactive exopolysaccharides from a Streptococcus thermophilus strain: Screening, purification and characterization, International Journal of Biological Macromolecules, 86, 402, 10.1016/j.ijbiomac.2016.01.085 Roobab, 2020, Sources, formulations, advanced delivery and health benefits of probiotics, Current Opinion in Food Science, 32, 17, 10.1016/j.cofs.2020.01.003 Rodríguez, 2010, Therapeutic effect of Streptococcus thermophilus CRL 1190-fermented milk on chronic gastritis, World Journal of Gastroenterology, 16, 1622, 10.3748/wjg.v16.i13.1622 Ruas-Madiedo, 2002, An overview of the functionality of exopolysaccharides produced by lactic acid bacteria, International Dairy Journal, 12, 163, 10.1016/S0958-6946(01)00160-1 Rütering, 2017, Tailor-made exopolysaccharides—CRISPR-Cas9 mediated genome editing in Paenibacillus polymyxa, Synthetic Biology, 2, 1, 10.1093/synbio/ysx007 Sá-Correia, 2002, Gellan gum biosynthesis in Sphingomonas paucimobilis ATCC 31461: Genes, enzymes and exopolysaccharide production engineering, Journal of Industrial Microbiology and Biotechnology, 29, 170, 10.1038/sj.jim.7000266 Salazar, 2014, Immune modulating capability of two exopolysaccharide-producing Bifidobacterium strains in a Wistar rat model, BioMed Research International, 10.1155/2014/106290 Sasaki, 2015, Cell-bound exopolysaccharides of Lactobacillus brevis KB290 enhance cytotoxic activity of mouse splenocytes, Journal of Applied Microbiology, 118, 506, 10.1111/jam.12686 Sato, 2004, Dextran from Leuconostoc mesenteroides augments immunostimulatory effects by the introduction of phosphate groups, Journal of Food Protection, 67, 1719, 10.4315/0362-028X-67.8.1719 Satpute, 2010, Biosurfactants, bioemulsifiers and exopolysaccharides from marine microorganisms, Biotechnology Advances, 28, 436, 10.1016/j.biotechadv.2010.02.006 Schiavi, 2016, The surface-associated exopolysaccharide of Bifidobacterium longum 35624 plays an essential role in dampening host proinflammatory responses and repressing local TH17 responses, Applied and Environmental Microbiology, 82, 7185, 10.1128/AEM.02238-16 Schmid, 2015, Bacterial exopolysaccharides: Biosynthesis pathways and engineering strategies, Frontiers in Microbiology, 6, 10.3389/fmicb.2015.00496 Shin, 2016, Exopolysaccharide fraction from Pediococcus pentosaceus KFT18 induces immunostimulatory activity in macrophages and immunosuppressed mice, Journal of Applied Microbiology, 120, 1390, 10.1111/jam.13099 Simms, 1990, The structural analysis of a levan produced by Streptococcus salivarius SS2, Carbohydrate Research, 208, 193, 10.1016/0008-6215(90)80099-O Son, 2018, Antioxidant and immunostimulatory effect of potential probiotic Lactobacillus paraplantarum SC61 isolated from Korean traditional fermented food, jangajji, Microbial Pathogenesis, 125, 486, 10.1016/j.micpath.2018.10.018 Song, 2018, Relationship between putative eps genes and production of exopolysaccharide in Lactobacillus casei LC2W, Frontiers in Microbiology, 9, 10.3389/fmicb.2018.01882 Stingele, 1996, Identification and characterization of the eps (Exopolysaccharide) gene cluster from Streptococcus thermophilus Sfi6, Journal of Bacteriology, 178, 1680, 10.1128/JB.178.6.1680-1690.1996 Stingele, 1999, Introduction of the exopolysaccharide gene cluster from Streptococcus thermophilus Sfi6 into Lactococcus lactis MG1363: Production and characterization of an altered polysaccharide, Molecular Microbiology, 32, 1287, 10.1046/j.1365-2958.1999.01441.x Surayot, 2014, Exopolysaccharides from lactic acid bacteria: Structural analysis, molecular weight effect on immunomodulation, International Journal of Biological Macromolecules, 68, 233, 10.1016/j.ijbiomac.2014.05.005 Tang, 2015, Exopolysaccharide produced by Lactobacillus plantarum induces maturation of dendritic cells in BALB/c mice, PloS One, 10, 10.1371/journal.pone.0143743 Tidona, 2016, Selection of Streptococcus thermophilus strains able to produce exopolysaccharides in milk, International Journal of Dairy Technology, 69, 569, 10.1111/1471-0307.12295 Tuinier, 2001, Effects of structural modifications on some physical characteristics of exopolysaccharides from Lactococcus lactis, Biopolymers, 59, 160, 10.1002/1097-0282(200109)59:3<160::AID-BIP1015>3.0.CO;2-V van Hijum, 2001, Purification of a novel fructosyltransferase from Lactobacillus reuteri strain 121 and characterization of the levan produced, FEMS Microbiology Letters, 205, 323, 10.1016/S0378-1097(01)00490-6 van Hijum, 2002, Characterization of a novel fructosyltransferase from Lactobacillus reuteri that synthesizes high-molecular-weight inulin and inulin oligosaccharides, Applied and Environmental Microbiology, 68, 4390, 10.1128/AEM.68.9.4390-4398.2002 van Kranenburg, 1999, Functional analysis of glycosyltransferase genes from Lactococcus lactis and other gram-positive cocci: Complementation, expression, and diversity, Journal of Bacteriology, 181, 6347, 10.1128/JB.181.20.6347-6353.1999 Vaningelgem, 2004, Biodiversity of exopolysaccharides produced by Streptococcus thermophilus strains is reflected in their production and their molecular and functional characteristics, Applied and Environmental Microbiology, 70, 900, 10.1128/AEM.70.2.900-912.2004 Vastano, 2016, Transcriptional analysis of exopolysaccharides biosynthesis gene clusters in Lactobacillus plantarum, Archives of Microbiology, 198, 295, 10.1007/s00203-015-1169-1 Verkhnyatskaya, 2019, Shaping the infant microbiome with non-digestible carbohydrates, Frontiers in Microbiology, 10, 10.3389/fmicb.2019.00343 Wang, 2019, In vitro immunomodulatory effects of acidic exopolysaccharide produced by Lactobacillus planetarium JLAU103 on RAW264.7 macrophages, International Journal of Biological Macromolecules Wang, 2014, Catalytic synthesis of sulfated polysaccharides. II: Comparative studies of solution conformation and antioxidant activities, Carbohydrate Polymers, 107, 221, 10.1016/j.carbpol.2014.02.074 Wang, 2016, Functional and bioinformatics analysis of an exopolysaccharide-related gene (epsN) from Lactobacillus kefiranofaciens ZW3, Archive of Microbiology, 198, 611, 10.1007/s00203-016-1217-5 Wang, 2018, Characterization and immunomodulatory activity of an exopolysaccharide produced by Lactobacillus plantarum JLK0142 isolated from fermented dairy tofu, International Journal of Biological Macromolecules, 115, 985, 10.1016/j.ijbiomac.2018.04.099 Wang, 2015, Characterization and bioactivities of an exopolysaccharide produced by Lactobacillus plantarum YW32, International Journal of Biological Macromolecules, 74, 119, 10.1016/j.ijbiomac.2014.12.006 Wang, 2015, Chemical modification, characterization and bioactivity of a released exopolysaccharide (r-EPS1) from Lactobacillus plantarum 70810, Glycoconjugate Journal, 32, 17, 10.1007/s10719-014-9567-1 Xu, 2017, Structural analysis and mucosal immune regulation of exopolysaccharide fraction from Bifidobacterium animalis RH, Food and Agricultural Immunology, 28, 1226, 10.1080/09540105.2017.1333578 Yasuda, 2008, Suppressive effect on activation of macrophages by Lactobacillus casei strain Shirota genes determining the synthesis of cell wall-associated polysaccharides, Applied and Environmental Microbiology, 74, 4746, 10.1128/AEM.00412-08 Yatim, 2015, A brief journey through the immune system, Clinical Journal of the American Society of Nephrology, 10, 1274, 10.2215/CJN.10031014 Yegorenkova, 2018, Immunomodulatory activity of exopolysaccharide from the rhizobacterium Paenibacillus polymyxa CCM 1465, Archives of Microbiology, 200, 1471, 10.1007/s00203-018-1564-5 Zannini, 2016, Production, properties, and industrial food application of lactic acid bacteria-derived exopolysaccharides, Applied Microbiology and Biotechnology, 100, 1121, 10.1007/s00253-015-7172-2 Zaporozhets, 2016, Prospects for the therapeutic application of sulfated polysaccharides of brown algae in diseases of the cardiovascular system: Review, Pharmaceutical Biology, 54, 3126, 10.1080/13880209.2016.1185444 Zarour, 2017, Rheology and bioactivity of high molecular weight dextrans synthesised by lactic acid bacteria, Carbohydrate Polymers, 174, 646, 10.1016/j.carbpol.2017.06.113 Zhang, 2016, Physicochemical characteristics and bioactivities of the exopolysaccharide and its sulphated polymer from Streptococcus thermophilus GST-6, Carbohydrate Polymers, 146, 368, 10.1016/j.carbpol.2016.03.063 Zhang, 2016, Characterization and sulfated modification of an exopolysaccharide from Lactobacillus plantarum ZDY2013 and its biological activities, Carbohydrate Polymers, 153, 25, 10.1016/j.carbpol.2016.07.084 Zhou, 2018, Exopolysaccharides from Lactobacillus plantarum NCU116 regulate intestinal barrier function via STAT3 signaling pathway, Journal of Agricultural and Food Chemistry, 66, 9719, 10.1021/acs.jafc.8b03340 Zhu, 2014, Screening and characterization of Sphingomonas sp. mutant for welan gum biosynthesis at an elevated temperature, Bioprocess and Biosystem Engineering, 37, 1849, 10.1007/s00449-014-1159-8 Zhu, 2019, Exopolysaccharides produced by yogurt-texture improving Lactobacillus plantarum RS20D and the immunoregulatory activity, International Journal of Biological Macromolecules, 121, 342, 10.1016/j.ijbiomac.2018.09.201 Zununi Vahed, 2017, Leuconostoc mesenteroides derived anticancer pharmaceuticals hinder inflammation and cell survival in colon cancer cells by modulating NF-κB/AKT/PTEN/MAPK pathways, Biomedicine & Pharmacotherapy, 94, 1094, 10.1016/j.biopha.2017.08.033