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Gutierrez et al. Microbiome Res Rep 2023;2:36 https://dx.doi.org/10.20517/mrr.2023.37 Page 7 of 23

B. bifidum TMC 3116
B. bifidum TMC 3117
B. breve TMC 3207
B. breve TMC 3217
B. breve TMC 3218
B. breve TMC 3219
B. infantis ATCC 15697 Glycan array [86]

Table 2. Literature review of mucus and cell binding adhesins in Bifidobacterium species and strains
Bifidobacterium species Adhesin Ref.
B. bifidum PRL2010 Sortase-dependent pili [76]
B. bifidum ATCC 15696 Extracellular sialidase [77]
B. bifidum A8 Extracellular transaldolase [79]
B. longum NCC2705 Extracellular transaldolase [78]
B. longum BBMN68 Putative adhesion proteins [75]
B. longum BBMN68 FimM [75]
B. bifidum 85B FimM homologs [75]
B. gallinarum CACC 514 FimM homologs [75]
B. longum NCC2705 Type 2 glycoprotein-binding fimbriae homolog [80]
B. longum NCC2705 EF-Tu [81]
B. longum NCC2705 Enolase [81]
B. animalis subsp. lactis Bl07 Enolase [82]
B. animalis subsp. lactis KLDS 2.0603 GroEL [83]
B. longum VMKB44 Blap-1 [84]
B. longum JCM1217 Endo-α-N-acetylgalactosaminidase [85]
B. longum subsp. infantis ATCC 15697 Family 1 of solute binding proteins (F1SBPs) [86,87]
B. breve UCC2003 Type IVb pilus-type proteins [82,83,88]
B. longum NCC2705 Extracellular vesicles [89]

Figure 1. Schematic outlining examples of the various mechanisms by which Bifidobacterium adhere to mucus. (A) Extracellular vesicles
released by Bifidobacterium can bind to mucus, and in turn, this binding can inhibit pathogen colonization; (B) Bifidobacterium possess a
wide array of pili and fimbriae, including FimM and its homologs, type 2 fimbriae, and type IVb pili, which bind to mucus; (C) Other
proteins such as F1SBPs (family-1 binding proteins), BL0155 (a type of ABC transport transmembrane protein), GroEL (a heat shock
protein), and EF-Tu (Elongation Factor Tu) are involved in mucus binding; (D) Endo-α-N-acetylgalactosaminidase, transaldolase,
sialidase, and enolase are enzymes that facilitate mucus adhesion.