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DOI 10.1007/s00253-015-6946-x
MINI-REVIEW
Received: 14 April 2015 / Revised: 17 August 2015 / Accepted: 20 August 2015 / Published online: 8 September 2015
# Springer-Verlag Berlin Heidelberg 2015
* Guoqiang Zhu
yzgqzhu@yzu.edu.cn
Mingxu Zhou
zhoumingxu@outlook.com
Yang Yang
yy@yzu.edu.cn
Panlin Chen
chen_panlin@outlook.com
Huijie Hu
hhj0417@163.com
Philip R. Hardwidge
hardwidg@gmail.com
1
Introduction
Escherichia coli (E. coli) is a Gram-negative, rod-shaped, facultatively anaerobic bacterium that resides in the gastrointestinal tract of warm-blooded animals (Donnenberg 2013). Most
E. coli are motile, typically with peritrichous flagella. The
length of the typical flagellum of E. coli is about 10 m and
the diameter is 20 nm. Flagellin, also known as H-antigen
(FliC protein, encoded by fliC gene in E. coli), is a major
surface antigen for E. coli serotyping (Nataro and Kaper
1998). Fifty-three H-antigens are recognized while a designation of NM (H) indicates the absence of flagellin (Orskov
and Orskov 1984). About 20,000 flagellin monomers polymerize to form a filament (OBrien and Bennett 1972). A
bacterial flagellum also consists of a basal body that acts as
a rotary motor and a hook that connects the motor and propeller (Terashima et al. 2008). Over 50 genes participate in flagellar biosynthesis (Macnab 2003). The genetic regulation of
flagellar assembly has been reviewed recently (Deane et al.
2010; Kazmierczak and Hendrixson 2013) and will not be
discussed in detail here. The flagellum is primarily a locomotive organelle, allowing bacteria to move from nutrient-poor
environments toward nutrient-rich ones (Wadhams and
Armitage 2004). In the past few years, studies have revealed
that flagella also affect bacterial virulence by delivering virulence factors and mediating biofilm formation, bacterial adhesion, and invasion. This review describes the different flagellin variants in E. coli and highlights the role of flagella in
bacterial functions other than motility.
E. coli flagellin variants
Based on considerable variability of E. coli flagellin in ultrastructure and 53 different H-antigens recognized, Lawn et al.
grouped 50 different flagella H serotypes into six
8884
8885
H serotype
Morphotypea
H1
CP007799
1788
H2
H3
AY249138
AB128916
1494
1590
C
U
H4
H5
AY249989
AY249990
1050
1311
A
B
H6
AY249991
1647
H7
H8
AY249992
AJ865465
AJ884569
1758
1479
E
D
AY249994
AY249995
2013
1263
U
C
-a
-b
H9
H10
H11
FN649414
1464
H12
AY249997
1788
H14
H15
AY249998
AY249999
1653
1689
F
F
H16
H17
AB128919
NC017663
1575
1524
U
A
H18
H19
H20
AY250001
AY250002
AY250003
1665
1842
1731
F
F
F
H21
H23
H24
H25
H26
H27
DQ862122
AY250005
AY250006
LM996317
AY250008
AF345848
1476
1767
1479c
1332
1674
1464
D
E
C
B
F
D
H28
H29
H30
H31
H32
AY250010
AY250012
AY250011
AY250013
AY250014
1740
1332
1713
1668
1713
F
C
F
F
F
H33
H34
H35
H36
H37
H38
H39
H40
1287
1638
1509
1671
1686
1344
1299
1479
B
E
U
U
F
B
B
D
H41
H42
H43
H44
H45
H46
H47
AY250015
AY250016
EF392692
EF392693
AY250017
AY250018
AY250019
AJ884568
AJ865464
AY250020
AY250021
AY250022
AB269770
AY250023
AY250024
EF392694
1680
1281
1506
1725
1707
1719
1107
F
B
C
F
E
F
B
H48
H49
AY250025
AY250026
1497
1695
C
E
-a
-b
Descriptionb
flkA
fliC; 2 genotypes
flnA
flkA
flkA
fliC; 2 genotypes
fllA
flkA
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Table 1 (continued)
H serotype
Morphotypea
Descriptionb
H51
AY250027
1818
H52
H53
AY250028
AB128917
1344
1272
B
C
flkA
H54
AB128918
1551
ND
flmA
H55
H56
AB269771
AY250029
1869
1344
ND
ND
fllA
Morphotypes: A thin, long pitch helix; B thin, subunit pattern; C rough subunit pattern; D polar subunit pattern; E short pitch loop pattern; F long pitch
loop pattern. Serotypes labeled BU^ have a unique surface structure not readily grouped with those of any other serotype. The morphotype of three
serotypes (H54, H55, and H56) are not determined (ND)
Except H3, H17, H35, H36, H44, H47, H53, H54, and H55 serotypes, the other 44 H-antigens are all encoded by fliC genes
As lacking of N- and C-terminal regions sequence for the H24-antigen, the ORF length here is based on the conserved N- and C-terminal length of other
serotypes
Homologous proteins of the E. coli fT3SS and T3SSs from different bacteria species
Shigella ssp.
Description
FliH
FliI
FliJ
FlhA
FlhB
FliO
FliP
FliQ
FliR
FlgN
FliS
FliT
OrgB
InvC
InvI
InvA
SpaS
N/A
SpaP
SpaQ
SpaR
N/A
SicA
PipC
MxiN
Spa47
Spa13
MxiA
Spa40
N/A
Spa24
Spa9
Spa29
N/A
IpgC
IpgF
ATPase regulator
ATPase
Chaperone
Major export apparatus protein, interacts with soluble components
Export switch protein, interacts with soluble components
Minor export apparatus protein, maintains stability of FliP
Minor export apparatus protein, cleaved signal sequence
Minor export apparatus protein
Minor export apparatus protein
Chaperone
Chaperone
Chaperone
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coordinate bacterial functions that promote survival in infected (Kansal et al. 2013). The myriad relationships between
flagella and bacterial virulence remain an active area of
investigation.
Conclusions
The flagellum controls many important functions other than
motility, including protein secretion, adhesion, biofilm formation, and pro-inflammatory host responses as well as apoptosis. Flagellar master genes also coregulate other virulence factors in pathogenic E. coli (Lane et al. 2007; Lehti et al. 2012;
Simms and Mobley 2008; Zhou et al. 2013), which may help
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