TABLE 1.
Prevalence of virulence factors in ExPEC according to the source (Korhonen et al., 1985; Guyer et al., 2000, 2000; Russo et al., 2001; Bingen-Bidois et al., 2002; Johnson et al., 2002, 2004, 2008; Marrs et al., 2002; Bonacorsi et al., 2003; Russo, 2003; Srinivasan et al., 2003; Buckles et al., 2004; Bonacorsi and Bingen, 2005; Parham et al., 2005a; Rendón et al., 2007; Restieri et al., 2007; Ananias and Yano, 2008; Henderson et al., 2009; Schubert et al., 2010; Ejrnæs et al., 2011; Mahjoub-Messai et al., 2011; Vejborg et al., 2011; Lehti et al., 2012; Qin et al., 2013; Tarchouna et al., 2013; Firoozeh et al., 2014; Basmaci et al., 2015; Nagarjuna et al., 2015; Wijetunge et al., 2015; Beck et al., 2016; Cordeiro et al., 2016; Lee and Lee, 2018; Daga et al., 2019; Nojoomi and Ghasemian, 2019; Raimondi et al., 2019; Sarowska et al., 2019).
| DESIGNATION | ADHESIN | FUNCTION AND ASSOCIATED INFECTIONS | Feces (%) | Cystitis (%) | APN (%) | Blood (%) | LCS (%) | PATHOTYPE |
| Type 1 fimbriae | fimH | Colonization factor and biofilm formation in EIs, especially cystitis and meningitidis, by binding receptor D-Mannose | 68 to 99 | 62 to 100 | 88 to 99 | 90 to 98 | 92 | UPEC, SEPEC, MNEC, APEC |
| ECP fimbriae | Mat | Meningitis associated and temperature regulated fimbriae | 48 | 91 to 98 | 92 | 100 | 96 | NMEC |
| Curli fimbriae | csgAB | Attachment and invasion of host cells, interaction with host proteins, activation of the immune system, bacterial aggregation and biofilm formation | 34 to 36 | 44 to 100 | 24 | 53 to 92 | NF | UPEC, SEPEC, APEC |
| P fimbriae | papGII | Proinflammatory activity and colonization factor in EIs especially pyelonephritis by binding GbO4 and GbO3 | 17 to 23 | 13 | 64 to 67 | 48 to 68 | 20 to 50 | UPEC, SEPEC, MNEC, APEC |
| papGIII | Proinflammatory activity and colonization factor in EIs especially cystitis by binding receptors GbO5 and GloboA | 8 to 9 | 22 | 19 to 35 | 14 to 27 | 1 to 6 | UPEC, SEPEC, APEC | |
| S fimbriae | sfaS | Adhesion to intestinal and urinary tract cells and facilitate penetration into the tissues by binding receptors NeuNAc (α2-3)Gal in UTIs, meningitidis | 7 to 32 | 13 to 34 | 88 to 99 | 39 to 97 | 28 to 59 | UPEC, SEPEC, NMEC |
| F1C fimbriae | focG | Adhesion to epithelial and endothelial cells in the bladder and kidney | 3 to 6 | 14 to 39 | 35 | 21 | 1 to 4 | UPEC, SEPEC |
| G fimbriae | gafD | Binding GlcNAc receptor | 5 | 0 to 4 | 0 to 18 | 0 | 3 | |
| Auf fimbriae | auf | Auf fimbriae | 21 to 27 | 67 | NF | NF | 62 | UPEC, NMEC |
| RTX protein TosA | tosA | Adhesion to host cells derived from the upper urinary tract, enhances survival in disseminated infections and lethality during sepsis | 11 | 16 | 29 | UPEC | ||
| Dr fimbriae | dra | Binding to the receptors DAF of epithelial cells, type IV collagen and CEACAMs. Promotes host cells invasion | 6 | 0 to 4 | 6 to 12 | 5 | NF | UPEC |
| Afimbrial adhesin | afa | Binding to receptor DAF on the epithelial cells and hemagglutination capacity | 2 to 17 | 19 to 81 | 16 | 49 | 9 | UPEC |
| IrgA homolog adhesin | iha | Iron-regulated-gene-homolog adhesin | 14 to 24 | 22 | 41 | 33 to 40 | 19 | UPEC, SEPEC |
| DESIGNATION | INVASIN | FUCNTION AND ASSOCIATED INFECTIONS | Feces (%) | Cystitis (%) | APN (%) | Blood (%) | LCS (%) | PATHOTYPE |
| hemagglutinin | hek/hra | Promote autoaggregation, hemagglutination, epithelial cell invasion and is associated with bacteremia in newborns | 14 to 28 | 28 | 33 to 66 | 45 | 11 | UPEC, SEPEC, NMEC, APEC |
| Endothelial brain invasion | ibeA | Cell invasion into the host tissues such as endothelial cell by binding Vimentine and Caspr1 to promote penetration of the blood–brain barrier | 1 to 15 | 13 to 23 | 29 | 4 to 11 | 32 to 38 | UPEC, NMEC, SEPEC, APEC |
| Arylsulfatase | aslA | Contributes to brain microvascular endothelial cell invasion | 46 | NF | NF | NF | 87 to 89 | NMEC |
| DESIGNATION | TOXINS | FUCNTION AND ASSOCIATED INFECTIONS | Feces (%) | Cystitis (%) | APN (%) | Blood (%) | LCS (%) | PATHOTYPE |
| α-hemolysin | hly | Pore-forming RTX toxin responsible for kidney injury and inflammation | 4 to 15 | 15 to 43 | 38 to 47 | 34 to 77 | 9 to 30 | UPEC, NMEC |
| Cytotoxic necrotizing factor 1 | cnf1 | activation of Rho GTPases, engaging in cell necrosis and promoting resistance to phagocytosis, inflammation, recurrent UTIs and blood-brain barrier penetration | 3 to 13 | 13 to 39 | 28 to 47 | 17 to 49 | 6 to 27 | UPEC, SEPEC, NMEC |
| Cytolethal distending toxin | cdt | DNAse inducing apoptosis and cellular senescence resulting in cell distention | 0 to 1 | 9 to 17 | 12 | 0 to10 | 46 | SEPEC |
| Colibactin | clb | PK-NRP compounds inducing DNA damage resulting preferentially in apoptosis in immune cells and premature cellular senescence in epithelial cells. | 11 to 32 | 33 | 44 | 18 to 58 | 75 | UPEC, SEPEC, NMEC, APEC |
| DESIGNATION | ATs | FUCNTION AND ASSOCIATED INFECTIONS | Feces (%) | Cystitis (%) | APN (%) | Blood (%) | LCS (%) | PATHOTYPE |
| Antigen43 | agn43 | Autotransporter involved in auto-aggregation, adhesion to host cells, biofilm development and persistence | 41 | 29 to 94 | 65 | 69 | NF | UPEC |
| Sap | sap | Autotransporter adhesin | NF | 1,5 | NF | NF | NF | |
| Secreted autotransporter toxin | Sat | Autotransporter having serine protease activity inducing cytotoxic effect and kidney injury | 14 to 24 | 23 to 56 | 55 to 68 | 26 to 39 | 49 | UPEC, NMEC |
| Vacuolating autotransporter toxin | Vat | Autotransporter having serine protease activity affecting mucins, facilitating epithelium colonization and inducing host cell vacuolization | 0 to 49 | 42 to 68 | 44 | 12 to 71 | 51 to 76 | UPEC, APEC, NMEC |
| Temperature-sensitive hemagglutinin | tsh | Autotransporter having temperature-sensitive hemagglutinin activity | 1 to 5 | 4 to 8 | 18 | 20 | 11 | APEC, NMEC |
| Serin protease autotransporter | pic | Autotransporter exhibiting serine protease activity affecting mucins, facilitating epithelium colonization and damaging host cell membrane | 0 to 23 | 21 to 34 | 14 to 40 | 0 to 36 | NF | UPEC |
| Secreted and surface associated lipoprotein | sslE/yghJ | Cytotoxic metalloprotease exihiting mucinase and proinflammatory activity | NF | NF | NF | NF | NF | NMEC,IPEC |
| Contact-dependent growth inibition | cdiAB | Induce the growth inhibition of the bacterial target | NF | NF | NF | NF | NF | UPEC |
| DESIGNATION | IRON UPTAKE | FUNCTION AND ASSOCIATED INFECTIONS | Feces (%) | Cystitis (%) | APN (%) | Blood (%) | LCS (%) | PATHOTYPE |
| Enterobactin | ent, fepA | Acquisition of siderophore Enterobactin-Fe3 + | 100 | 100 | * | * | * | UPEC, SEPEC, MNEC, APEC |
| Aerobactin | aer, iuc, iut | Acquisition of siderophore Aerobactin-Fe ions in the host involved in UTI | 16 to 41 | 24 ot 52 | 38 to 77 | 54 to 80 | 61 to 88 | UPEC, APEC, NMEC |
| Salmochelin | iroN | Acquisition of siderophore Salmochelin-Fe ions in the host involved in UTI and meningitidis | 5 to 26 | 33 to 74 | 76 to 78 | 44 to 63 | 38 to 75 | UPEC, NMEC, SEPEC APEC |
| Yersiniabactin | irp, fuyA | Acquisition of siderophore Yersiniabactin-Fe ions in the host involved in UTI and meningitidis | 23 to 70 | 63 to 96 | 97 | 82 | 90 to 99 | NMEC |
| ChuA, Hma | chu, hma | Acquisition of Fe from hemoglobin in the host system | 53 to 59 | 66 to 80 | 66 to 71 | 48 to 73 | 89 | UPEC, SEPEC |
| Sit | sitDCBA | Transportation of Fe and Mn | 25 | 44 | 56 | 96 | 92 | UPEC, APEC |
| IreA | ireA | Iron-regulated homolog of siderophore receptors involved in UTI | 7 to 19 | 13 | 47 | 32 | NF | UPEC |
| Fec | fecARI | Acquisition of citrate-Fe3 + | NF | NF | NF | NF | NF | |
| Fbp | fbpABCD | Fe acquisition | NF | 58 | 59 | 45 | NF | |
| K1 capsule | kpsMI-neuA | Enable resistance to complement activation and phagocytosis, immunological tolerance and intracellular survival promoting both bacteremia and meningitis | 0 to 27 | 35 | 13 to 47 | 8 to 82 | 75 to 90 | SEPEC, NMEC |
| TcpC | tcpC | Impairs the innate immune response by inhibiting Toll-like receptor and MyD88-specific signaling | 8 | 22 | 42 | 32 to 39 | 0 | UPEC, SEPEC |
| D-serine deaminase | dsdCXA | Promotes bacterial growth by using D-serine as carbon, nitrogen, and energy source, and prevent the bacteriostatic activity of D-serine in the urine and brain. | 0 to 4 | 20 to 22 | NF | 6 | 73 to 87 | UPEC, NMEC |
| TrapT | traT | Inhibition of the classical pathway of complement activity and serum survival | 43 to 88 | 36 to 68 | 49 to 81 | 50 to 81 | NF | UPEC, SEPEC |
| Iss | Resistance to complement and serum survival | 4 to 20 | 13 | 35 | 10 to 34 | 25 | UPEC, SEPEC, NMEC | |
| Outer membrane protein T | OmpT | Resistance to protamine and urine survival | 15 to 68 | 70 | 88 to 95 | 21 to 81 | 96 | UPEC, SEPEC, NMEC |
NF, data not found, ∗Present in almost all Escherichia coli genomes.