Thesis LJ Westerman
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Spirochaetosis
L.J. Westerman
L.J. Westerm
anH
uman Intestinal Spirochaetosis
2013
Uitnodigingvoor het bijwonen van de
openbare verdediging van mijn
op dinsdag 10 december 2013
om 10:30
in het Academiegebouw van de
Universiteit Utrecht
Domplein 29 Utrecht
harte uitgenodigd voor
Kanunnikenzaal van de Faculty
Club (achter de Senaatszaal)
Laurens Westerman
ParanimfenMarcus Westerman
06-46238506
Lysanne Ng-A-Foe
3605697890399
ISBN 978-90-393-6056-9
-
Chapter 1 7
Chapter 2 39
Chapter 3 61
Chapter 4 77
Chapter 5 89
Chapter 6 125
Chapter 7 145
Chapter 8 161
Chapter 9 181
Addendum 197
203
Table of Contents
-
8
-
9
A brief history of the Brachyspiraceae
.
.
a .
-
10
0.02 (~24bp)
Figure 1:
-
11
.
Figure 2:
-
12
Chapter 1
4
.
-
13
.
.
Figure 3:
-
14
Chapter 1
Chapter 9
The various Brachyspira species
Brachyspira hyodysenteriae (hog: pig; dysentery: severe diarrhoea)
-
15
Tabl
e 1:
56
Spec
ies
Indo
leHi
ppur
ate
Type
of r
efer
ence
stra
in*
++
++
+
++
++
+ +
++ +
*
44+ +
++ +
++
*
+
-
16
0 . 03
' B . i b a r a k i ' H I S 2 4 / 1 1 / 9 9
B.innocen
s B256
'B.suanana' AN
4859/03
'B.m
uris '
A10
/06
B . a a l b o r g i 5 1 3 A
B.a
l vi n
i pu
l li
C1
'B .hampsoni i ' c l
uster I NSH-16
B.hyodysenter iae B
78
B. intermedia PW
S/A
'B.murid
arum'
A58/06
B.murdoch i i 56-150
'B.hominis' adhucomuHca09
B. p
i lo s
i co l
i P 4
3 /6 /
7 8
'B.r
aus
' A
249/
07
' B .hampson i i ' c lus ter I I NSH-24
'B.corv i ' AN
968/2/04
' B. p
u ll i '
AN
304/
04
'B.ca
nis' D
og A2
T .p r imi
a ZAS - 2
A
0 . 0 4
B.murd
ochi i 5
6-150
B.canis Dog A2
B. p
u ll i
AN3
04/0
4
B. a
l vi n
i pu
l li
C1
B.suana na AN 4859/03
B.intermedia PWS/A
B.ham
pson i i c lus ter I I NSH
-24B
.ham
pson
i i
c lus
ter
I N
SH-1
6
B.in
noce
ns B
256
B.hyodysenter iae B78
B.aa lborg i 513A
B .corv i AN968/2/04B . pi l o s
i c ol i P
4 3 /6 / 7
8
T . p r i m i a Z A S - 2
B
Figure 4:
-
17
Brachyspira innocens (innocens: innocent)
.
Brachyspira murdochii (Murdoch: Australian university where much research into
.
Brachyspira pilosicoli (pilo: hairy; colon: colon)
chapter 9
-
18
Chapter 1
Brachyspira intermedia (intemediate: referring to the fact that the phenotype is between B. hyodystenteriae and B. innocens)
..
In
Brachyspira alvinipulli
-
19
Brachyspira aalborgi
Brachyspira canis (canine: dog)
p In
Brachyspira corvi (corvus: crows)
-
20
Chapter 1
44
Brachyspira pulli
Brachyspira muridarum and Brachyspira muris
.
.
.
(sus: pig ( Anas platyrhynchos))
-
21
Brachyspira hampsonii (Prof. Hampson: prominent Brachyspira
Brachyspira ibaraki
-
22
Tabl
e 2:
Spec
ies
B. a
albo
rgi
B. a
lvin
ipul
liB.
hyo
dyse
nter
iae
B. in
noce
nsB.
inte
rmed
iaB.
mur
doch
iiB.
pilo
sico
liB
. can
is
B. c
orvi
-
23
Spec
ies
B. h
omin
is
B. m
urid
arum
B
. mur
is
B. p
ulli
B. h
amps
onii
B
. ham
pson
ii
-
24
Chapter 1
Brachyspira hominis (hominis: human)
Zoonosis
-
25
Epidemiological evidence
-
26
Chapter 1
.
.
.
-
27
-
28
Chapter 1
.
-
29
Figure 5:
-
30
Chapter 1
Table 3:
Country Colon biopsies Appendices Faeces*
Concluding
-
31
Aim of this thesischapter 2
chapter 4 (chapters 2 8
In chapter 3 Chapter 5
chapter 8. Chapter 6chapter 7
chapter 5
-
32
Chapter 1
45
16
41
17
41
86 10
33
36
11
189
11
10
1
31
1
50
49
3
88
27
12
37
85
70
82
22
34
57
31
40
33
125
63
Int J 46
References
-
33
160
Int J 29
Int J 47
48
35
36
28
9
14
2
24
7
23
39
53
136
178
8
67
59
106
53
130
67
Occurrence
153
15
52
-
34
Chapter 1
146
158
124
137
37
102
134
46
23
47
111
23
119
53
59
146
12
52
121
99
21
22
28
53
35
91
63
32
105
15
121
36
-
35
41
68
12
13
61
42
59
39
2
59
J 22
21
J 113
113
26
93
146
34
153
16
158
53
21
33
64
65
J 31
141
48
138
143
-
36
Chapter 1
7
103
31
134
179
150
38
38
224
187
13
74
134
37
Int J 18
4
45
49
7
30
49
35
30
59
101
21
90
39
41
23
-
37
-
38
-
4
4
-
40
Chapter 2
Abstract
Methods
Results
p
Conclusions
-
41
4
Jensen
-
42
Chapter 2
.
Aims of this study
-
43
MethodsEthics statement
-
44
Chapter 2
Processing of FFPE biopsy samples
-
45
-
46
Table 1:
Primer set 53 Total length C Time Purpose
PCR volume
PCR consumables
PCR protocol
-
47
m
Sequencing
Figure 1:
StepA (212bp)60 271
StepB (253bp)203 445
StepC (258bp)366 623
Bspp (82bp)135 216
5 31000 200 300 400 500 600 700
AmpliconPrimer
-
48
Chapter 2
-
49
Results
A
PCR results
p
p p
-
50
-2
0
2
4
6
8
10
12
66 68 70 72 74 76 78 80 82 84 86 88 90 92 94 96
B. aalborgi)
B. pilosicoli)
Figure 2:
Table 2
Sequence result
-
51
B. pilosicoli
B. burgdorferi
B. murdochii B. innocens
B. hyodysenteriae B. intermedia
B. alvinipulli
B. aalborgi
0.01
B. pilosicoli
Brachyspira
B. hominis
Brachyspira aalborgi
Figure 3:
-
52
Chapter 2
(p
Table 3:
p
# + +
-
53
Discussion
.
-
54
Chapter 2
m
-
55
-
56
Chapter 2
-
57
48
35
160
2
Int J 46
40
67
63
106
62
41
16
12
53
39
References
23
53
42
16
31
49
7
41
115
12
2
39
59
116
-
58
Chapter 2
197
25 Suppl 3
37
27
12
132
83
95
32
23
40
67
136
178
35
-
59
Supplementary Table S1:
-
60
-
62
Chapter 3
Abstract
Methods
Results
Conclusion
-
63
Appendiceal Spirochaetosis in Children
.
.
-
64
Chapter 3
.
-
65
Appendiceal Spirochaetosis in Children
Materials and methodsEthics statement
Sample processing
-
66
Chapter 3
-
67
Table 1:
53 Amplicon
PCR volume
PCR protocol
-
68
Chapter 3
Histopathology
-
69
Appendiceal Spirochaetosis in Children
Results
Table 2:
Immunohistochemistry
PCR result + ++ +++
p
(p
-
70
Chapter 3
Discussion
Table 3:
Total appendices Control group Other diagnoses
4
-
71
Appendiceal Spirochaetosis in Children
(p
-
72
Chapter 3
-
73
Appendiceal Spirochaetosis in Children
24
17
12
27
88
90
23
8
3
16
7
160
67
35
2
142
2
59
39
53
39
23
154
52
J 38
53
4
31
Am J 132
55
25 Suppl 3
51
7
References
-
74
Chapter 3
37
33
65
-
75
-
76
-
44
-
78
Chapter 4
Abstract
(p
-
79
As a
-
80
Chapter 4
Materials and methods
Table 1: a
a
-
81
Results
p
p
-
82
Chapter 4
Discussion
p
-
83
-
84
Chapter 4
-
85
16
3
7
53
23
59
2
J 22
45
82
4
53
J 38
42
16
31
7
48
85
101
21
21
39
70
82
22
53
132
31
References
-
86
Supplementary table S1:
Sample ID
BacteriaParasites
S. entericaC. jejuni
S. dysenteriaeSTEC/EPEC
G. lam
bliaD. fragilis
C. parvum
+++
*+
+*
*+
*+
*
+*
*
+#
+++
-
87
-
88
-
A
4
4
-
90
Chapter 5
Abstract
-
91
Brachyspira species
.
(
-
92
Chapter 5
. In
(alp
.
-
93
Material and methodsMLST design
Table 1:
Criterion Database
-
94
Chapter 5
atpG
-
95
atpG
-
96
Table 2:
Locus TargetAmplicon (bp)
Primers**Total Useful Min
atpG
PCR volume
PCR consumables
PCR protocol
-
0.02 (~24bp)
Colour-plate A:
-
Colour-plate B:
-
Colour-plate C:
-
-2
0
2
4
6
8
10
12
65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96
513A (B. aalborgi)
WesB (B. pilosicoli)
Colour-plate D:
-
-2
0
2
4
6
8
10
12
65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96
513A (B. aalborgi)
WesB (B. pilosicoli)
Colour-plate E:
-
B. aalborgi
B. pilosicoli
B. hominis
B. hyodysenteriae
B. murdochii
B. alvinipulli
B. innocens
B. intermedia
Colour-plate F:
-
B. innocens
B. intermedia
B. aalborgi
B. pilosicoli
B. hominis
B. hyodysenteriae
B. murdochii
B. alvinipulli
B. innocens
B. intermedia
Colour-plate G:
-
Colour-plate H:
-
Colour-plate I:
-
Colour-plate J:
0.05
WW25
200900828
H4-2
200909930
Q95.3281.0
WF1
201124725
Q97.2224.3.1
201221374
200513182a
Q95.0635.6
SC1
Q98.0026.12
VTN021f
513A
HRM_4BHRM_2B
Q95.2078.2
Q97.2110.4.11
3295.90BW015/C138
Q1588.5
EM1
OF_11
199800660
Q97.000.6.8
N1
QF1
HRM7A
7072i
201102362
Q98.0033.72
201204927
P43/6/78
Q98.0072.08
200831665
201009156
K1
B256
B-78
Q98.0037.6
T.primitia_ZAS-2
HRM7
201101966
T1
9803.1
PWS/A
f199600014
201105386
KP5
201013692
H21
200410620a
M1
V1_H141
L72200300944
WesB
201125230
WIA8
C1
D9201243
V1_H11
201009139
201200678VTN022f
RV1
Q1404.6A
200410620b
H38-2
H60-2
Q91.1223.3
200805431
200517505
Q98.0028.3
J_F2
200709033
WA1
95/1000
200232081
200513182b
Q94.0354.0.6
56-150
Q98.0072.31
Human
Pig
Chicken
Dog
-
0.05
9803.1
T1
HRM7A
D9201243
Q91.1223.3
HRM_4B200909930
HRM_2B
WW25
Q95.3281.0
SC1
Q98.0072.31
200805431
201009156
W015/C138
JF2
B-78
Q94.0354.0.6
201200678
N1
OF_11
200709033
M1
C1
V1_H141
200300944
201221374
WA1
56-150
200513182a
200517505
H21
7072i
201013692
Q98.0028.3
201101966
VTN022f
Q95.2078.2
201125230
WesB
95/1000
Q97.2224.3.1
Q1404.6A
Q97.2110.4.11Q95.0635.6
VTN021f
WIA8
200513182b
200831665
201009139
201102362
P43/6/78
QF1
200410620b
Q97.000.6.8
513A
201105386
Q98.0026.12
200900828
200232081
201204927
WF1
Q1588.5
3295.90B
200410620a
PWS/A
Q98.0072.08
HRM7
EM1
H38-2
K1
KP5
H60-2
T.primitia_ZAS-2
201124725
B256
RV1
V1_H11
H4-2
A
B
C
Human
Pig
Chicken
Dog
Colour-plate K:
-
Human PigChicken Dog
Colour-plate L:
The Netherlands
Australia
Italy
Papua New Guinea
United Kingdom
United States of America
-
The Netherlands
Australia
Italy
Papua New Guinea
United Kingdom
United States of AmericaColour-plate M:
-
OverijsselSIR biopsies: 0.94 (0.93 - 0.95)
SIR HIS: 0.96 (0.72 - 1.24)Pigs 483.2/km2
Poultry 2,943.3/km2
HIV-centres 2/30
GelderlandSIR biopsies: 1.08 (1.08 - 1.09)
SIR HIS: 0.48 (0.36 - 0.64)Pigs 442.9/km2
Poultry 3,439.3/km2
HIV-centres 2/30
DrentheSIR biopsies: 0.92 (0.90 - 0.93)
SIR HIS: 0.54 (0.29 - 0.90)Pigs 107,5/km2
Poultry 2,363.8/km2
HIV-centres 0/30
FrieslandSIR biopsies: 1.04 (1.02 - 1.05)
SIR HIS: 0.21 (0.08 - 0.42)Pigs 31.8/km2
Poultry 2,187.2/km2
HIV-centres 1/30
GroningenSIR biopsies: 0.89 (0.88 - 0.90)
SIR HIS: 0.29 (0.13 - 0.55)Pigs 63.7/km2
Poultry 2.376.6/km2
HIV-centres 1/30
FlevolandSIR biopsies: 1.20 (1.18 - 1.22)
SIR HIS: 2.24 (1.62 - 3.02)Pigs 40.8/km2
Poultry 1,461.9/km2
HIV-centres 2/30
UtrechtSIR biopsies: 0.97 (0.96 - 0.98)
SIR HIS: 0.93 (0.71 - 1.20)Pigs 219.1/km2
Poultry 1,227.9/km2
HIV-centres 2/30
Noord-HollandSIR biopsies: 1.01 (1.01 - 1.02)
SIR HIS: 1.84 (1.63 - 2.08)Pigs 9.7/km2
Poultry 493.4/km2
HIV-centres 9/30
Zuid-HollandSIR biopsies: 0.99 (0.98 - 0.99)
SIR HIS: 0.46 (0.37 - 0.57)Pigs 56.5/km2
Poultry 390.0/km2
HIV-centres 7/30
Noord-BrabantSIR biopsies: 0.82 (0.81 - 0.82)
SIR HIS: 0.58 (0.46 - 0.73)Pigs 1,078.6/km2
Poultry 5,330.5/km2
HIV-centres 2/30
LimburgSIR biopsies: 1.27 (1.26 - 1.28)
SIR HIS: 2.92 (2.51 - 3.38)Pigs 790.6/km2
Poultry 6,880.0/km2
HIV-centres 1/30
ZeelandSIR biopsies: 0.86 (0.85 - 0.88)
SIR HIS: 1.03 (0.64 - 1.58)Pigs 43.6/km2
Poultry 1,015.2/km2
HIV-centres 1/30
Colour-plate N:
-
OverijsselSIR biopsies: 0.94 (0.93 - 0.95)
SIR HIS: 0.96 (0.72 - 1.24)Pigs 483.2/km2
Poultry 2,943.3/km2
HIV-centres 2/30
GelderlandSIR biopsies: 1.08 (1.08 - 1.09)
SIR HIS: 0.48 (0.36 - 0.64)Pigs 442.9/km2
Poultry 3,439.3/km2
HIV-centres 2/30
DrentheSIR biopsies: 0.92 (0.90 - 0.93)
SIR HIS: 0.54 (0.29 - 0.90)Pigs 107,5/km2
Poultry 2,363.8/km2
HIV-centres 0/30
FrieslandSIR biopsies: 1.04 (1.02 - 1.05)
SIR HIS: 0.21 (0.08 - 0.42)Pigs 31.8/km2
Poultry 2,187.2/km2
HIV-centres 1/30
GroningenSIR biopsies: 0.89 (0.88 - 0.90)
SIR HIS: 0.29 (0.13 - 0.55)Pigs 63.7/km2
Poultry 2.376.6/km2
HIV-centres 1/30
FlevolandSIR biopsies: 1.20 (1.18 - 1.22)
SIR HIS: 2.24 (1.62 - 3.02)Pigs 40.8/km2
Poultry 1,461.9/km2
HIV-centres 2/30
UtrechtSIR biopsies: 0.97 (0.96 - 0.98)
SIR HIS: 0.93 (0.71 - 1.20)Pigs 219.1/km2
Poultry 1,227.9/km2
HIV-centres 2/30
Noord-HollandSIR biopsies: 1.01 (1.01 - 1.02)
SIR HIS: 1.84 (1.63 - 2.08)Pigs 9.7/km2
Poultry 493.4/km2
HIV-centres 9/30
Zuid-HollandSIR biopsies: 0.99 (0.98 - 0.99)
SIR HIS: 0.46 (0.37 - 0.57)Pigs 56.5/km2
Poultry 390.0/km2
HIV-centres 7/30
Noord-BrabantSIR biopsies: 0.82 (0.81 - 0.82)
SIR HIS: 0.58 (0.46 - 0.73)Pigs 1,078.6/km2
Poultry 5,330.5/km2
HIV-centres 2/30
LimburgSIR biopsies: 1.27 (1.26 - 1.28)
SIR HIS: 2.92 (2.51 - 3.38)Pigs 790.6/km2
Poultry 6,880.0/km2
HIV-centres 1/30
ZeelandSIR biopsies: 0.86 (0.85 - 0.88)
SIR HIS: 1.03 (0.64 - 1.58)Pigs 43.6/km2
Poultry 1,015.2/km2
HIV-centres 1/30
1 Appendix2 Coecum3 Ascending colon4 Transversing colon5 Descending colon6 Sigmoid7 Rectum
12
3
4
5
67
A
12
3
4
5
67
B
12
3
4
5
67
C
12
3
4
5
67
D
Colour-plate O:
-
Colour-plate P:
A
B
-
97
ResultsSelected MSLT genes
( atpG
atpG
-
98
0 . 05
201122
867_B.
aalborg
i
201022967g_B.aalborgi
95 /1000_GenBank_B.p i los ico l i
201024300_B.aalborgi
ilocisolip.B
_669101102
201105351b_B.aalborgi
B256_B.innocens
200704900_B.aalborgi
200216407_B.aalborgi
ilocis
olip.B
_931
9001
02200410620a_B .p i l os i co l i
201129345_B.aalborgi
PWS/A_B
.intermedia
sinimoh.B_b295300102
200920244_B.aalborgi
ilocis
olip.
B_44
9003
002
200917097_B.aalborgi
201050465_B.aalborgi
20111037
4b_B.aalb
orgi
ilocisolip.B_
032521102
200232081_B .p i l o s i co l i
igroblaa.B_e153501102
200902757_B.
aalborgi
201003592a_B
.hom
inis
200913478_
B.aalborgi
201022967d_B.aalborgi
200430368_B.aalborgi
2011
1160
3_B.aalbo
rgi
iloc
isoli
p.B_
6519
0010
2
ilocisolip
.B_26320
1102
201104608_B.aalbo
rgi
200813138f_B.aa lborg i
200813138a_B.aalborgi
200813138g_B.aalborgi
201022967b_B.aalborgi
201122669_B.aalborgi
201120
460_B.
aalbor
gi
iloc
isoli
p.B_
7294
0210
2
200805639_B.aalborgi
200813138e_B.aalborgi
200813340_B.a
alborg
i
200922746_B.aalborgi
200925995_B.hominis
201003
249_B.
aalborgi
201021258_B.hominis
200414761_B.aalborgi
200811646_B.aalborgi
201024348_B.hominis
200608126*_B.hom
inis
sinimoh.
B_343806002
200813138d_B.aalborgi
201013692_B .p i l o s i co l i
2011
2389
8_B.aalbo
rgi
200703075_B.a
alborg
i
201108603_B.hom
inis
201024610_B.aalborgi
igro
blaa
.B_
c153
5011
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ominis
200813138e_B.aa
lborgi
B256_B
.innocens
200813138h_B.aalborgi
200811646_
B.aalborgi
201022967b_B
.aalborgi
ilocisol
ip.B_87
600210
2
200410620a_B .p i l os i co l i
200702703_B.a
alborg
i
2011
2463
9_B.aalbo
rgi
201024610_B.aalborgi
201111
603_B.a
alborgi
200900639_B.aalborgi
201105351f_B.aa lborgi
201004
769_B.
aalbor
gi
200918072_B.aalborgi
ilocis
olip.
B_50
2306
991
200813138c_B.aalborgi
ilocis
olip.
B_03
9909
002
201024466_B.aalborgi
200414
761_B.aa
lborgi
201050
465_B.a
alborgi
200917337_B.aalborgi
201104149_B.aalborgi
201202009_B.a
alborgi
200831665_B .p i l o s i co l i
201003592c_B.homini
s
201120460_B.aalborgi
200501138_B.aalborgi
2 00805431_B .p i l os i co l i
200410620b_B .p i l os i co l i
201022967g_B.aalborgi
201105
351g_B
.aalborg
i
200904021_B.aalborgi
201114303_B.aalborgi
Figure 4:
-
102
0 . 06
200813138b_B.aalborgi
201024348_B.hominis
200922746_B.aalborgi
200716125_B.aalborgi
igroblaa.B_646118002
iloc
isol
ip.
B_33
0907
002
201105351d_B.aalborgi
201122669_B.aalborgi
200803874_B.hominis
201116
007_B.
hominis
201024
300_B.a
alborgi
201110374b_B.aalborgi
201111603_B.aalborgi
201014350_B.aalborgi
201003249_B.aa
lborgi
iloc
isoli
p.B_
5057
1500
2
200704900_
B.aalborgi
201124639_B.aalborgi
20041476
1_B.aalbor
gi
iloc
isoli
p.B_
4490
0300
2
201018213_B.aalborgi
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inis
200501138_B.aalborgi
201114303_B.aalborgi
iloc
isoli
p.B_
7294
0210
2
200808214_B.aalborgi
201111933_B.aalborgi
201021258_B.hominis
201105351b_B.aalborgi
ilocisolip.B_263201102
201104140_B.hominis
200917337_B.aalborgi
201122
867_B.
aalbor
gi
iloc
isoli
p.B_
b026
0140
02
200813138c_B.aalborgi
iloc
isoli
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8760
0210
220
0902
757_
B.aalbo
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200113
332_B.
aalborg
i
201104608_B.aalborgi
201105351c_B.aalborgiilocisolip.
B_566138002
201003592d
_B.hominis
201015580_B.aalborgi
201108681_B.aalborgi
200905511_B.h
ominis
200920244_B.aalborgi
WA1_GenBank_B.hyodysenteriae
20100359
2b_B.hom
inis
iloc
isol
ip.
B_47
3122
102
201050211_B.aalborgi
201104
406_B.aa
lborgi
2011
0305
9_B.aalbo
rgi
201123898_B.aalborgi
iloci
soli
p.B_
2963
1010
2
200900639_B.aalborgi
201007259_B.aalborgi
95/100
0_Gen
Ban
k_B.pi lo
s icol i
200923973_B.aalborgi
200513182b_B.p i l os i co l i
2 01009156_B .p i l o s i co l i
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0325
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2
200608343_B.hominis
P43
/ 6/ 78 _
B. pi lo s
i co li
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138e_B
.aalbo
rgi201
050465
_B.aalb
orgi
200410620a_B.p i l os i co l i
200925995_B.hominis
201024610_B.aalborgi
iloc
isoli
p.B_
6835
0110
2200805639_B.aalborgi
C 1 _ B . a l v i n i p u l l i
ilocis
olip.
B_82
8009
002
200513182a_B.p i l os i co l i
20051470
2_B.homi
nis
201022967e_B.aalborgi
2011
2914
4_B.aalbo
rgi
201022967c_B.aa
lborgi
200813138a_B.aalborgi
200702665_B.hominis
200816817_B.a
alborg
i
200904021_B.aalborgi
200813138d_B.aa
lborgi
B256_B.innocens
201104149_B.aalborgi
200813138g_B.aalborgi
200702703_B.aalborgi
ilocis
olip.B
_180
2320
02
200602985_B.aalborgi
ilocisolip.B_134508002
200918072_B.aalborgi
WesB_B .p i l o s i co l i
201022967d_B.aa
lborgi
B-78_B.hyodysenter iae
200707443_B.aalborgi
200709653_B.aalborgi
200913478_B.aalborgi
iloc
isoli
p.B_
5274
2110
2
201150177_B.aalborgi
201150471_B.aalborgi
513A
_B.aa lbo
rgi
201119045_B.aalborgi
201202009_B.aalborgi
56-150__B.murdochi i
200901135_B.aalborgi
PWS/A_B.intermedia
ilocis
olip.B
_66910
1102
200914805_B.a
alborg
i
200909930_B .p i l o s i co l i
201110374d_B.aalborgi
200917097_B.aalborgi
200624827_B.aalborgi
ilocisolip.B_931900102
T . p r im i t i a_ZAS-2
Figure 5:
-
103
T .p r im i t i a_ZAS-2
atpG
Table 3:
Locus MLST Length (bp)variable sites
AATLength
variable sites
atpG
**
-
104
Figure 6:
B. aalborgi
B. hominis
B. pilosicoli
B. hyodysenteriae
B. murdochii
B. alvinipulli B. intermedia
B. innocens
-
105
B. intermedia
B. aalborgi
B. hominis
B. pilosicoli
B. hyodysenteriae
B. murdochii
B. alvinipulli B. intermedia
B. innocens
Figure 7:
-
106
Chapter 5
-
107
Figure 8:
-
108
Chapter 5
Discussion
-
109
As
atpG
.
-
110
Chapter 5
.
-
111
-
112
Chapter 5
32
4
36
36
3
77
153
43
14
43
62
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114
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200813138b
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201102752_B.aalborgi
ilocisolip
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200900639_B.aalborgi
200607003_B.hominis
200902757_B.aalborgi
201024610_B.a
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200716125_B.aalborgi
201111933_B.aalborgi
200624827_B.aalborgi
200608126*_B.hominis
2 00831665_B .p i l o s i co l i
201018213_B.aalborgi
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sini
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8308
002
ilocisolip.B_
651900102
201108603_B.hominis
201119045_B.aalborgi
200922746_B.aalborgi
200513182b_B .p i l os i co l i
201003592b_B.hominis
200113332_B.a
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i
201022967e_B.aalborgi
201014350_B.aa
lborgi
igro
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2
513A
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201129
345_B.a
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201003592c_B.hominis
201004
769_B.
aalbor
gi
201022967g_B.aalborgi
201124639_B
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iloc
isoli
p.B_
9319
0010
2
201003249_B.aalborgi
200813138c_B
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200901135_B.aalborgi
WesB_B .p i l o s i co l i
2011
0535
1d_B
.aalbo
rgi
201024466_B.aalborgi
200410620b_B .p i l os i co l i
200704900_B.aalborgi
200913478_B.aalborgi
iloci
soli
p.B_
8280
0900
2
20070965
3_B.aalb
orgi
200602985_B.aalborgi
200414761_B.aalborgi
200513182a_B .p i l os i co l i
201110374g_B.aalborgi
200410620a_B .p i l os i co l i
igroblaa.B_863034002
201022967h_B.aalborgi
2011
0535
1e_B
.aalbo
rgi
200918072_B.aalborgi
201050211_B.a
alborg
i
2011
5017
7_B.aalbo
rgi
200920
244_B.
aalborg
i
eair
etne
sydo
yh.
B_kn
aBn
eG_
1A
W
ilocisolip.B_729402102
201003592a_B.hominis
20081313
8a_B.aalb
orgi
201110374a_B.aalborgi
igro
blaa
.B_
4128
0800
2
201104406_B.aalborgi
B-78_B
.hyodysenter iae20
1105
351b
_B.aalbo
rgi
aide
mret
ni.B_
A/S
WP
200702665_B.hominis
igro
blaa
.B_
7907
1900
2
201003592d_B.hominis
201024348_B.hominis
201114303_B.aalborgi
200813138g_B.aalborgi
200813340_B.aalborgi
201022967c_B.aalborgi
200703075_B.aalborgi
200816
817_B.
aalbor
gi
201003592e_B.hominis
201122
669_B.
aalborg
i
ilocis
olip.B_
039909
002
201150471_B.aalborgi
201023408_B.aalborgi 20110414
9_B.aalbo
rgi
201110374b_B.aalborgi
200905511_B.hominis
200813138h_B.aalborgi
201015580_B.aalborgi
2009
2397
3_B.aalbo
rgi
ilocis
olip.
B_68
3501
102
201109833_B.aalborgi
201104140_B.hom
inis
iloc
isoli
p.B_
2963
1010
2
200813138d_B.a
alborgi
ilocisolip.B
_180232002
2009
1733
7_B.aalbo
rgi
ilocis
olip.
B_87
6002
102
2011
0535
1c_B
.aalbo
rgi
201122867_B.aalborgi
201111603_B.a
alborg
i
ilocisolip.B_263
201102
iloc
isoli
p.B_
0660
0899
1
201110374d_B.aalborgi
20081164
6_B.aalb
orgi
201001908*_B.aalborgi
201022967d_B.aalborgi
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isoli
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4731
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P 4 3 / 6 / 7 8 _B . p i l o s i c o l i
201050465_B.aalborg
i
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isoli
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2110
2
C1_B
. a l v i n i p u l l i
200904021_B.aalborgi
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isoli
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4490
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2
200813138f_B.aa lborg i
56-150_B.m
urdochi i
igroblaa.B_f153501102
201103059_B.aalborgi
201024300_B.aalborgi
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_50571
5002
199603205_B .p i l o s i co l i
95/1000_GenBank_B.p i los ico l i
2 01124725_B .p i l os i co l i
B256_B
.innocens200101917_B.aalborgi
200901
634_B.aa
lborgi
200914
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lborgi201
108681
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borgi
2012
0200
9_B.aalbo
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01102
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201040
076_B.a
alborgi
200813138e_B.aalbor
gi
201022967f_B.aa lborg i
200501138_B.aalborgi
sini
moh.
B_59
9529
002
200608343_B.hominis
201116007_B.hominis
201007259_
B.aalborgi
201123898_B.aalborgi
201129
144_B.
aalbor
gi
201104608_B.aalborgi
200707443_B.aalborgi
201105351a_B
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200514702_B.hominis
201022967i_B.aa lborg i
0 . 06
T.pr im
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201102
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alborgi
201119045_B.aalborgi
201003592c_B.hominis
20100725
9_B.aalb
orgi
201024348_B.hominis
201021258_B.hominis
201015580_B.aalborgi
200901634_B.aalborgi
201003592b_B.hominis
200918072_B.aalborgi
20110460
8_B.aalbo
rgi
201003592a_B.hominis
200805639_B.
aalborgi
201003592d_B.hominis
201122867_B.aal
borgi
201050465_B.aalborgi
200905511_B.hominis
200602985_
B.aalborgi
-
115
200624827_B.aalborgi
201014350_B.aa
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639_B.aalbor
gi
200414761_B.aalborgi
200920
244_B.
aalborg
i
20110414
9_B.aalbo
rgi2008
11646_B.
aalborgi
201050465_B.aalborg
i
201040
076_B.a
alborgi
201007259_
B.aalborgi
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200917097_B. a
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102
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201015580_B.aalborgi
201104149_B.aalborgi
PWS/A_B.intermedia
200813138b_B
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2010
2340
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201105351d_B.aalborgi
201108681_B.aalborgi
WesB_B .p i l o s i co l i
201111933_B.aalborgi
200901634_B.aalborgi
200917337_B.aalborgi
200914805_B.aalborgi
200808214_B.a
alborg
i
200913478_B.aalborgi
iloc
isoli
p.B_
8760
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2
201004769_B.aalborgi
201003592b_B.hominis
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200707443_B.aalborgi
200920244_B.aalborgi
B256_B.in
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200918072_B.aalborgi
iloc
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102
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201105351c_B.aalborgi
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B_300706002
201022967h
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200702703_B.aalborgi
20110460
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200513182b_B.p i l os i co l i
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201103059_B.aalborgi
201105351g_B.aa
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201003592a_B.hominis
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1900
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200608126*_B.hominis
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201022967g
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200904021_B.aalborgi
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borgi
201024610_B.aalborgi
T .p r im i t i a_ZAS-2
201110374g_B.aalborgi
201114303_B.aalborgi
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201110374d_B.aalborgi
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20102296
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201022967i_B.
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200813
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201129345_B.aalborgi
201123898_B.aalborgi
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200813138a_B.aalborgi
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200905511_B.hominis
201024466_B.aalborgi
200624827_B.aalborgi201109833_B.aalborgi
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200813138c_B.aalborgi
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5661
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201003249_B. a
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201024610_B. aal bor g
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200414761_B. aal bor g
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B.aalborgi
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117
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200703075_B.aalborgi
C1 _
B. al vi ni pu ll i
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95 /1000_GenBank_B.p i los ico l i
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B-78_
B.hyodysenteriae
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1193
3_B.aalbo
rgi
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P43 / 6 / 7 8 _B
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201108681_B.aalborgi
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201105351d_B.aalborgi
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i
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9530
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8c_B.aalb
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1_B.aalbor
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580_B.
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138b_B
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2-S
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mirp
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200813138b_B.aalborgi
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119
ililocis
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iloc
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i l oci s ol i p. B_032521102
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340_B.
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511_B.
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2011
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121
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513A_B.aa lborg i
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2296
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511_B.
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2296
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211_B.
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138_B.a
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