1203
46.
1204
1205
colonic mucins. Biosci Biotechnol Biochem 78, 1444–1451 (2014).
47.
1206
1207
Nishiyama, K. et al. Evaluation of bifidobacterial adhesion to acidic sugar chains of porcine
Yasui, K. et al. Improvement of bacterial transformation efficiency using plasmid artificial
modification. Nucleic Acids Res 37, e3–e3 (2009).
48.
Kozakai, T., Shimofusa, Y., Nomura, I. & Suzuki, T. Construction of a reporter system for
1208
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1209
promoters and terminators. Biosci Microbiota Food Health 40, 115–122 (2021).
1210
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1211
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bifidobacteria-infant symbiosis. Sci Adv 5, eaaw7696 (2019).
50.
1213
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Sakanaka, M. et al. Evolutionary adaptation in fucosyllactose uptake systems supports
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Minamisawa, T. & Hirabayashi, J. Fragmentations of isomeric sulfated monosaccharides using
1215
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1217
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1218
1219
Nat Methods 13, 581–583 (2016).
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1221
1224
Segata, N. et al. Metagenomic biomarker discovery and explanation. Genome Biol 12, R60
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54.
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1223
Callahan, B. J. et al. DADA2: High-resolution sample inference from Illumina amplicon data.
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Biol 20, 257 (2019).
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Camacho, C. et al. BLAST+: architecture and applications. BMC Bioinformatics 10, 421
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Benjamini, Y. & Hochberg, Y. Controlling the false discovery rate: a practical and powerful
1226
approach to multiple testing. Journal of the Royal Statistical Society. Series B
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(Methodological) 57, 289–300 (1995).
1228
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candidates for the therapy of osteoarthritis. Chem Biol 8, 701–711 (2001).
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Yamada, C. et al. Molecular insight into evolution of symbiosis between breast-fed infants and
1231
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Sakurama, H. et al. Lacto-N-biosidase encoded by a novel gene of Bifidobacterium longum
1234
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1235
its active expression. J Biol Chem 288, 25194–25206 (2013).
1236
38
100
Non-sulfated O-glycans
Oral gavage
(PBS or B. bifidum 109 cells per day)
Relative intensity (%)
Acclimation 0
(Conv.)
779
5 (Day)
Dissection
PBS (n = 5)
Bb (n = 5)
954
983
50
100
*m/z 1100 (ES)
1158
1332
1228
1590
**
Sulfated O-glycans
**m/z 780 (ES)
50
1246
867 1041
Collect faeces
(Day 0)
Collect faeces
(Day 5)
1951
1764
500
1000
1420
1500
2000
2500
3000
m/z
Estimated amounts of glycans
(m/z) PBS Bb
0.05
1.5
Estimated glycan amounts
(nmol per 0.1 mg faecal extracts)
Non-sulfated
Sulfated
0.10
(nmol per 0.1 mg faecal extracts)
534
779
954
1025
1142
1199
1257
1332
1386
1433
1519
1590
1678
1748
1781
1835
2039
2196
1025
1071
1317
1491
1852
Bb
p = 0.97
0.5
40
p = 0.39
20
Fuc
Gal
GalNAc
−20
−40
PBS
p = 0.044
NeuAc
GlcNAc
GlcNAc-6S
Bb
p = 0.44
PBS
10
p = 0.025
Faeces
p = 0.68
Oligosaccharide length
GlcNAc-6S
(pmol per mg dry faeces)
p = 0.19
Relative abundance (%)
Relative abundance (%)
p = 0.055
p = 0.97
ρ = −0.74
p = 4.0 × 10−9
20
Sulfated
O-glycan
Caecum contents
60
40
Non-sulfated
O-glycan
Sulfated
O-glycans
Non-sulfated
O-glycans
PBS
p = 0.95
% to PBS control
Bb
Day
Day
GlcNAc-6S
0.15
ρ = 0.38
q = 0.017
p = 0.021
100
Infants
ρ = 0.40
p = 0.020
10−1
10−2
Adults
ρ = 0.24
p = 0.35
10−3
10−4
Flintibacter sp.
KGMB00164 (%)
Free GlcNAc-6S
(nmol per mg dry faeces)
101
0.10
0.05
0.00
10−5
102
103
104
105
106
107
108
109
bbhII
(Copy number per mg dry faeces)
0.000
0.001
bbhII homologs (%)
Katoh et al., Figure 1
5 (Day)
PBS
WT
B. bifidum
(109 cells per mouse)
Dissection
bbhII
Detection limit
PBS
GalNAc
NeuAc GlcNAc-6S
0.0
5.0
534
Detection
limit
0.0
0.5
1.0
534
Sulfated
1025
1158
534
895
(m/z)
(>0.20)
0.20
PBS-control (GF)
PBS
0.15
WT
1024
779
1228
895
1473
WT-mono-colonized
0.10
0.05
bbhII
534
100
1.5
Concentration
[nmol per caecal contents (g)]
* 1228 1473
Estimated amounts
(nmol per 0.1 mg faecal extracts)
bbhII-mono-colonized
500
1000
1500
2000
2500
3000
m/z
Sulfated O-glycans
867
100
**780 (ES)
1071
PBS-control (GF)
**
Non-sulfated
983
779 895
Sulfated
m/z 867
p = 0.036
p = 2.3 × 10−5
p = 0.0056
p = 0.015
15
p = 1.7 × 10−6
2.0
1.5
p = 5.7 × 10−5
p = 0.0009
1.5
10
1.0
1.0
0.5
0.5
0.0
0.0
PBS WT bbhII
PBS WT bbhII
PBS WT bbhII
100
MS/MS at m/z 867
**
WT-mono-colonized
100
867
**
1071
bbhII-mono-colonized
500
1000
1500
2000
m/z
2500
3000
Relative intensity (%)
Relative intensity (%)
(nmol per 0.1 mg
faecal extracts)
Relative intensity (%)
779
Estimated amounts of glycans
*1100 (ES)
1024 1402 1473
1228
709 895
15.0
Non-sulfated
983
100
10.0
Non-sulfated O-glycans
100
1607
GlcNAc
Detection
limit
Concentration
[nmol per caecal contents (g)]
Gal
GlcNAc-6S
Log [B. bifidum cell number per
caecal contents (g)]
p=
0.0036
1764
p=
0.0032
1403
p=
0.0002
p = 0.0018
1504
p = 0.027 p = 0.0032
ρ = 0.89
p = 1.0 × 10−8
1229
p = 0.0022
p=
0.0046
p=
0.0001
p=
0.0001
1332
p = 0.046
10
Fuc
p = 0.046
PBS
WT
bbhII
15
NeuAc
Log [B. bifidum cell number per
caecal contents (g)]
Concentrations
[nmol per caecal content (g)]
Monosaccharides in caecal contents
bbhII
WT
1607
1317
1474
1041
1923
Acclimation
(2 weeks)
2618
Germ-free mice
Log
[B. bifidum cell number per
caecal contents (g)]
Oral gavage
520
100
747
370
Glycan symbol
520
631
Gal
PI
867
50
370
650747
ol
GalNAc
631
GlcNAc
650
Sulfate
50
300
550
800
m/z
Katoh et al., Figure 2
GH20 catalytic domain
Ca2+
CBM32
N-domain
(46-197)
C-domain
(751-861)
W651
Q640
W607
V649
Y637
P639
W588
GlcNAc-6S
H688
W685
E553
(acid/base)
D552
(stabilizer)
E687
R358
Q640/
M396
W651/408
Y637/393
V649/
L406
W607/361
P639/
D395
W588/344
H688/
T455
E687/444
W685/442
D553/314
D552/313
R358/162
H467/250
Katoh et al., Figure 3
CBM32 N-domain
N126
K85
N89
Ca2+
R95
PUGNAc-6S
(Ki = 15.4 nM)
NAG-thiazoline-6S
(NAGT-6S)
(Ki = 52.3 nM)
W183
E62
PGM O-glycans
Q640
0 mM
NAGT-6S
w/o cultivation
W651
Y637
779
100
Non-sulfated
1041
100
708
1402
953
534
**
1420
867 1246 1491
E553
D552
R358
GH20 catalytic domain
1 mM
NAGT-6S
24 h cultivation
E687
Relative intensity (%)
W607
0 mM
NAGT-6S
24 h cultivation
W685
Sulfated
100 534
100
1402
953
**
1025
779
867
1041
1316
1491
708
100
534
867
100
779
1041
**
9531025
1402
708
400
1316
1491
1400
2400
400
1400
m/z
2400
m/z
Non-sulfated O-glycans
Sulfated O-glycans
2.0
50
m/z 1025
m/z 534
m/z 1332
m/z 953
m/z 1474
30
20
m/z 1402
m/z 708
10
m/z 779
0.1
24
NAGT-6S (mM)
Incubation time (h)
Glycan symbol
L-Fuc
Others
m/z 1316
m/z 867
m/z 1246
m/z 1287
m/z 1491
1.5
1.0
m/z 1420
Gal
GalNAc
0.5
m/z 1041
GlcNAc
Sulfate
0.1
24
Estimated glycan amounts
(nmol per 100 μg mucin protein)
Others
Estimated glycan amounts
(nmol per 100 μg mucin protein)
40
NAGT-6S (mM)
Incubation time (h)
Katoh et al., Figure 4
GlcNAc-6S
GlcNAc-6S
(GlcN)3
GlcNAc-Man
K85
N89
Ca2+
Ca2+
E62
W183
R95
N126
S97
T127
BbhII CBM32
Ca2+
Paenibacillus DD2
p < 0.0001
60
p < 0.0001
0.05
80
40
20
0.00
10
15
p < 0.0001
Binding (%)
A450
100
p < 0.0001
120
0.10
NagH CBM32-2
p = 0.00077
S184
20
Mucin (μg/ml)
PGM with CBM
PCM w/o CBM
PCM with CBM
pNP-β-GlcNAc-6S
Purified enzymes
PGM
Purified enzymes
Recombinant cells
p = 0.0006
20
10
0.008
0.006
0.004
0.002
0.000
p = 0.0184
2.0
Activity (s−1)
30
Activity (unit/OD600)
Activity (s−1)
40
p = 0.455
p < 0.0001
p < 0.0001
p = 0.935
50
Recombinant cells
1.5
1.0
0.5
0.0
p = 0.0001
Activity (unit/OD600)
PGM w/o CBM
0.00010
p < 0.0001
0.00008
0.00006
0.00004
0.00002
0.00000
Katoh et al., Figure 5
Number of mucCBM in muc-GH
30
20
abc
bc
abc
ab
10
bcd
cd
CGC64_00690
muc-GH with muc-CBM
per total muc-GH (%)
GH16_3
50
40
bc
30
20
bc
bc
19
(+)
10
(+)
12
(+)
(─)
10
Genome exam.
GH16_3
11
(─)
37
(─)
(+)
92
(+)
(+)
B. bifidum GH16
(ORF fragmented)
2.0
B. bifidum
GH
CBM
Crossfeeding
Goblet
cell
Selffeeding
Glycan symbol
CBM-dependent
breakdown
L-Fuc
Gal
GalNAc
GlcNAc
Mono-, disaccharides
Intestinal lumen
Mucus layer
Epithelial
cell
Sulfate
Katoh et al., Figure 6
Relative abundance
Relative abundance at family level of taxonomy
B. bifidumadministered
PBS control
Day 0
PBS control
B. bifidumadministered
Day 5
Linear discriminant analysis Effect Size (LEfSe)
f__Desulfovibrionaceae.g__
f__Desulfovibrionaceae.g__.s__
f__Rikenellaceae
f__Rikenellaceae.g__.s__
f__Rikenellaceae.g__
Bifidobacterium longum
g__Bacteroides.s__
p__Proteobacteria
o__Desulfovibrionales
f__Desulfovibrionaceae
c__Deltaproteobacteria
f__Bifidobacteriaceae
g__Bifidobacterium
c__Actinobacteria
o__Bifidobacteriales
f__S24_7.g__
f__S24_7.g__.s__
f__S24_7
o__Clostridiales
o__Clostridiales
o__Clostridiales
g__Parabacteroides.s__
g__Parabacteroides
f__Porphyromonadaceae
Linear Discriminant Analysis (LDA) Score (log 10)
Katoh et al., Extended Data Fig. 1
Relative abundance
Relative abundance at the family level of taxonomy
Post-cultivation
Pre-cultivation
GlcNAc-6Sadded
None-added
β-Diversity
Linear discriminant analysis Effect Size (LEfSe)
UniFrac (weighted)
None-added
GlcNAc-6S-added
0.3
f__Bacteroidaceae
g__Bacteroides
g__Clostridium
Clostridium hathewayi
PCo2 (24.5%)
0.2
0.1
Post-cultivation
GlcNAc-6S-added
0.0
g__Achromobacter.s__
g__Achromobacter
f__Enterobacteriaceae
f__Enterobacteriaceae
Linear Discriminant Analysis (LDA) Score (log 10)
Post-cultivation
None-added
-0.1
Pre-cultivation
-0.2
-0.3
-0.2
-0.1
0.0
0.1
0.2
PCo1 (52.4%)
Katoh et al., Extended Data Fig. 2
Sgl clade
BbhII clade
Substrate specificity
6-SO3-β-N-Acetylglucosaminidase
β-N-Acetylhexosaminidase (Broad substrate specificity)
Chitinolytic β-N-acetylglucosaminidase
N-Glycan processing β-N-acetyglucosaminidase
Disperisin B (β-1,6-specific N-acetylglucosaminidase)
β-1,3-N-Acetylglucosaminidase acting on lacto-N-triose II and mucin core 3
Lacto-N-biosidase
Taxonomic classification
Eukaryotes
Mammal
Plant
Insect/prawn
Fungi
Nematode (C. elegans)
Others
Bacterial phyla
Actinomycetota
Bacillota
Bacteroidota
Chlamydiae
Pseudomonadota
Verrucomicrobiota
Katoh et al., Extended Data Fig. 3
Relative abundances of species-specific reads vs. bbhII homologspecific reads in the metagenomic dataset obtained for mothers
Relative abundances of species-specific reads vs. sgl homologspecific reads in the metagenomic dataset obtained for infants
Species
q value
(FDR correction,
Q = 5 %)
Species
q value
(FDR correction,
Q = 5 %)
Flintibacter sp. KGMB00164
0.0167
0.378
Bacteroides thetaiotaomicron
Bacteroides caccae
Bacteroides sp. CBA7301
Butyricimonas faecalis
Bacteroides sp. A1C1
Bacteroides sp. HF 162
Bacteroides sp. CACC 737
Paraprevotella xylaniphila
Barnesiella viscericola
Bacteroides uniformis
Bacteroides intestinalis
Phocaeicola dorei
Bacteroides sp. M10
Odoribacter splanchnicus
Bacteroides sp. HF 5287
Bacteroides helcogenes
Bacteroides caecimuris
Bacteroides cellulosilyticus
Bacteroides sp. HF 5141
Parabacteroides sp. CT06
Bacteroides ovatus
Parabacteroides distasonis
Phocaeicola salanitronis
Alistipes megaguti
Collinsella aerofaciens
Clostridium perfringens
Bacteroides xylanisolvens
Phocaeicola vulgatus
Sutterella faecalis
Parolsenella catena
Alistipes finegoldii
Alistipes shahii
Alistipes communis
2.28E-14
3.43E-13
3.84E-08
1.57E-07
4.23E-07
4.23E-07
4.23E-07
2.21E-06
1.29E-05
1.29E-05
1.78E-05
1.78E-05
2.45E-05
2.93E-05
3.14E-05
3.95E-05
0.0000500
7.80E-05
7.80E-05
0.000213
0.000262
0.000416
0.000461
0.00112
0.00132
0.00207
0.00337
0.00349
0.00425
0.00754
0.00923
0.00983
0.0156
0.764
0.737
0.617
0.595
0.573
0.572
0.572
0.545
0.513
0.511
0.504
0.502
0.495
0.490
0.487
0.481
0.476
0.465
0.464
0.442
0.436
0.424
0.421
0.397
0.392
-0.379
0.365
0.363
0.356
0.338
0.330
0.327
0.311
Relative abundances of species-specific reads vs. sgl homologspecific reads in the metagenomic dataset obtained for mothers
Species
Bacteroides thetaiotaomicron
Bacteroides caccae
Parabacteroides sp. CT06
Bacteroides sp. HF 162
Bacteroides caecimuris
Bacteroides sp. HF 5141
Bacteroides sp. CBA7301
Bacteroides fragilis
Bacteroides sp. CACC 737
Bacteroides ovatus
Bacteroides uniformis
Bacteroides xylanisolvens
Parabacteroides distasonis
Bacteroides sp. A1C1
Lacrimispora saccharolytica
Bifidobacterium angulatum
Bacteroides sp. M10
q value
(FDR correction,
Q = 5 %)
0.0000437
0.00313
0.00313
0.00313
0.00313
0.00313
0.00313
0.00393
0.00647
0.0127
0.0152
0.0152
0.0232
0.0294
0.0300
0.0483
0.798
0.5228
0.423
0.418
0.416
0.411
0.405
0.405
0.396
0.378
0.356
0.348
0.346
0.331
-0.321
-0.319
0.301
Katoh et al., Extended Data Fig. 4
qPCR
target region
Pr-MS955
B. bifidum
B. bifidum
pNP-GlcNAc6S-hydrolyzing activity
(nmol/min/mg protein)
bbhIIrt-P2-F
Wild-type
(kDa)
Pr-MS956
bbhIIrt-P2-R
250 180 -
Homologous region (501 bp)
130 -
Suicide plasmid pHT33
(CmR)
CmR
95 72 55 43 -
qPCR
target region
Single crossover
recombination
34 26 -
bbhIIrt-P2-F
17 11 -
250 180 130 95 72 55 43 34 26 -
80
60
40
20
WT
bbhII
Cell-free extracts
of B. bifidum
17 11 -
Disruptant
Antibodies: BbhII
GLBP
bbhIIrt-P2-R
BbhII-His6 (39-861aa)
CBM32
(kDa)
(kDa)
116 -
66.4 55.6 42.7 34.6 -
97.2 -
27.0 -
66.4 55.6 -
B. longum JCM 31944
expressing BbhII variants
(kDa)
190 135 100 -
20.0 -
80 -
14.3 -
58 -
BbhII
42.7 58 -
GLBP
46 -
BbhII-His6
Antibodies:
(kDa)
116 97.2 66.4 55.6 42.7 -
Katoh et al., Extended Data Fig. 5
Fuc
GlcNAc
GalNAc
Gal
GlcNAc-6S
NeuAc
Std.
PGM
HS
KS
CS
B. bifidum
HA
Katoh et al., Extended Data Fig. 6
Non-sulfated O-glycans of PCM
779
100
Sulfated O-glycans of PCM
100
BbhII (−)
BbhII (−)
**
1041
708
50
50
1491
1420
1286
1403
1246
Abundance (%)
953
534
1648
1736
2115
779
100
**
100
BbhII (+)
708
BbhII (+)
1403
953
50
50
1246
1648
534
1491
400
800
1041
1200
1600
2000
2400
400
800
−log10(q-value)
1736
1286
1200
m/z
1420
2115
1600
2000
2400
m/z
1041
Non-sulfated O-glycans
1112
1286
Sulfated O-glycans
1025
−4
−2
log2 (fold-change of
estimated glycan amount)
MS/MS at m/z 1041
100
Relative intensity (%)
370
631
694
921
Glycan symbol
L-Fuc
694
50
Molecular ion
[M+2Na−H]+
1041
853
370 433
ol
Gal
853
GalNAc
631
GlcNAc
921
100
600
1100
Sulfate
433
m/z
Katoh et al., Extended Data Fig. 7
MS/MS at m/z 1246 obtained from non-treated PCM
520
50
520
m/z 1246
ol
Molecular ion
[M+2Na−H]+
1246
(Resistant)
Mixture
Relative intensity (%)
748
748
100
970
100
600
1100
m/z
m/z 1246
(Resistant)
ol
970
748
520
MS/MS at m/z 1491 obtained from non-treated PCM
Relative intensity (%)
370
993
100
50
853
678 734
660
748 882
520
765
631
433
370
200
433
Molecular ion
[M+2Na−H]+
1491
1009
m/z 1491
(possibly
susceptible)
853
660
1200
1081
748
433
MS/MS at m/z 1491 obtained from BbhII-treated PCM
993
100
631
882
m/z
1302
765
50
1009
734
520
678
660
748
765
853
1215
1231
Molecular ion
[M+2Na−H]+
1491
ol
1371
1302
m/z 1491
(Resistant)
Mixture
Relative intensity (%)
ol
1215
1231
1081
1371
1144 1302
700
1144
1081
1009
835
200
700
678
1200
m/z
853
660
1231
520
o o
1302
993
ol
m/z 1491
(Resistant)
734
835
678
Glycan symbol
853
660
L-Fuc
1231
Gal
GalNAc
GlcNAc
Sulfate
1302
ol
m/z 1491
(Resistant)
1215
993
520
734
Katoh et al., Extended Data Fig. 8
Inhibition of BbhII by PUGNAc-6S
Inhibitor: PUGNAc-6S
Inhibitor: PUGNAc-6S
35
35
[I]
30
30
20
20
40
15
15
1/Turnover (sec)
20
-1
v (s-1)
0.20
0.20
10
25
25
Turnover (s )
0.25
0.25
(nM)
v-1 (s)
100
0.15
0.15
0.10
0.10
10
10
0.05
0.05
55
00
0.0
0.2
0.2
0.4
0.4
0.6
0.6
0.8
0.8
1.0
1.0
-10
-10
1.2
1.2
-8
-8
-6
-6
[S]0 (mM)
-4
-4
-2
-2
22
44
66
-1
1/[pNP- -GlcNAc-6S]
-1) )
[S-1] (mM(mM
[pNP- -GlcNAc-6S] (mM)
Inhibition of BbhII by NAGT-6S
Inhibitor: Thiazoline-GlcNAc-6S
Inhibitor: Thiazoline-GlcNAc-6S
[I]
(nM)
0.14
0.14
35
35
30
30
50
20
20
250
15
15
1/Turnover (sec)
v-1 (s)
0.10
0.10
100
-1
v (s-1)
25
25
Turnover (s )
0.12
0.12
25
0.08
0.08
0.06
0.06
10
10
0.04
0.04
55
0.02
0.02
00
0.0
0.2
0.2
0.4
0.4
0.6
0.6
0.8
0.8
1.0
1.0
-10
-10
1.2
1.2
-8
-8
-4
-4
-2
-2
Mode
Km (μM)
kcat (s-1)
Ki (nM)
PUGNAc-6S
Competitive
115 ± 14
32.3 ± 0.7
15.4 ± 1.9
NAGT-6S
Competitive
122 ± 16
32.6 ± 0.8
52.3 ± 7.2
Inhibitor
ITC analysis of CBM-His6 (WT)
pNP-β-GlcNAc-6S
pNP-β-GlcNAc
Ligand: 1 mM
Protein: 0.1 mM
c-value: 40.2
pNP-β-GlcNAcc-6S
00
44
66
[S-1]0 (mM-1)
[S]0 (mM)
-6
-6
1/[pNP- -GlcNAc-6S] (mM)
[pNP- -GlcNAc-6S] (mM)
Equation
kcat [E]0 [S]
v=
Km (1 + [ I ] / Ki ) + [S]
ITC analysis of CBM-His6 (W183A)
pNP-β-GlcNAc-6S
Ligand: 1 mM
Protein: 0.1 mM
pNP-β-GlcNAc
Ligand: 1 mM
Protein: 0.1 mM
Ligand: 1 mM
Protein: 0.1 mM
Ka
(103 M−1)
Kd
(μM)
ΔG0
(kJ mol−1)
ΔH
(kJ mol−1)
−TΔS0
(kJ mol−1)
40.2 ± 3.4
24.9 ± 2.1
−26.7
−49.2± 1.5
22.5
0.924 ± 0.018
Katoh et al., Extended Data Fig. 9
VECTORS
NMDS1
muc-GH
R2
NMDS2
P (>r)
GH16_3
0.01789
-0.99984
0.5380
0.0001
GH109
0.08254
-0.99659
0.5344
0.0001
GH112
-0.08254
0.99659
0.5344
0.0001
GH101
-0.02202
0.99976
0.4907
0.0001
GH42
0.02708
0.99963
0.4381
0.0001
GH110
-0.03276
-0.99946
0.4358
0.0001
GH35
-0.17121
-0.98523
0.4095
0.0001
GH16
0.89625
-0.44354
0.3438
0.0001
GH98
-0.01299
0.99992
0.2307
0.0001
GH20
-0.09459
-0.99552
0.2009
0.0001
GH123
-0.92867
-0.37091
0.1303
0.0001
GH36
-0.96505
0.26208
0.1253
0.0003
GH89
-0.55027
-0.83499
0.1076
0.0001
GH95
-0.66414
0.74761
0.0793
0.0002
GH136
0.97722
-0.21225
0.0612
0.0028
GH31
-0.98287
0.1843
0.0514
0.0092
GH129
0.98287
-0.1843
0.0514
0.0092
GH84
0.71886
-0.69515
0.0322
0.0397
GH2
GH29
Permutation: free
Number of permutations: 9999
Katoh et al., Extended Data Fig. 10
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