Dietary-fat-induced taurocholic acid promotes pathobiont expansion and colitis in Il10-/- mice

Nature. 2012 Jul 5;487(7405):104-8. doi: 10.1038/nature11225.

Abstract

The composite human microbiome of Western populations has probably changed over the past century, brought on by new environmental triggers that often have a negative impact on human health. Here we show that consumption of a diet high in saturated (milk-derived) fat, but not polyunsaturated (safflower oil) fat, changes the conditions for microbial assemblage and promotes the expansion of a low-abundance, sulphite-reducing pathobiont, Bilophila wadsworthia. This was associated with a pro-inflammatory T helper type 1 (T(H)1) immune response and increased incidence of colitis in genetically susceptible Il10(−/−), but not wild-type mice. These effects are mediated by milk-derived-fat-promoted taurine conjugation of hepatic bile acids, which increases the availability of organic sulphur used by sulphite-reducing microorganisms like B. wadsworthia. When mice were fed a low-fat diet supplemented with taurocholic acid, but not with glycocholic acid, for example, a bloom of B. wadsworthia and development of colitis were observed in Il10(−/−) mice. Together these data show that dietary fats, by promoting changes in host bile acid composition, can markedly alter conditions for gut microbial assemblage, resulting in dysbiosis that can perturb immune homeostasis. The data provide a plausible mechanistic basis by which Western-type diets high in certain saturated fats might increase the prevalence of complex immune-mediated diseases like inflammatory bowel disease in genetically susceptible hosts.

Publication types

  • Research Support, N.I.H., Extramural
  • Research Support, Non-U.S. Gov't

MeSH terms

  • Animals
  • Bile Acids and Salts / metabolism
  • Bilophila / drug effects*
  • Bilophila / growth & development
  • Colitis / chemically induced*
  • Colitis / immunology
  • Colitis / microbiology*
  • Colitis / pathology
  • Diet, Fat-Restricted
  • Dietary Fats / pharmacology*
  • Inflammation / chemically induced
  • Inflammation / immunology
  • Inflammation / microbiology
  • Inflammatory Bowel Diseases / chemically induced
  • Inflammatory Bowel Diseases / microbiology
  • Inflammatory Bowel Diseases / pathology
  • Interleukin-10 / deficiency*
  • Interleukin-10 / genetics
  • Metagenome / drug effects*
  • Mice
  • Mice, Inbred C57BL
  • Milk / chemistry
  • Molecular Sequence Data
  • Safflower Oil / pharmacology
  • Sulfites / metabolism
  • Taurine / metabolism
  • Taurocholic Acid / metabolism*
  • Taurocholic Acid / pharmacology
  • Th1 Cells / drug effects
  • Th1 Cells / immunology

Substances

  • Bile Acids and Salts
  • Dietary Fats
  • Sulfites
  • Interleukin-10
  • Taurine
  • Taurocholic Acid
  • Safflower Oil

Associated data

  • GENBANK/JQ890637
  • GENBANK/JQ890638
  • GENBANK/JQ890639
  • GENBANK/JQ890640
  • GENBANK/JQ890641
  • GENBANK/JQ890642
  • GENBANK/JQ890643
  • GENBANK/JQ890644
  • GENBANK/JQ890645
  • GENBANK/JQ890646
  • GENBANK/JQ890647
  • GENBANK/JQ890648
  • GENBANK/JQ890649
  • GENBANK/JQ890650
  • GENBANK/JQ890651
  • GENBANK/JQ890652
  • GENBANK/JQ890653
  • GENBANK/JQ890654
  • GENBANK/JQ890655
  • GENBANK/JQ890656
  • GENBANK/JQ890657
  • GENBANK/JQ890658
  • GENBANK/JQ890659
  • GENBANK/JQ890660
  • GENBANK/JQ890661
  • GENBANK/JQ890662
  • GENBANK/JQ890663
  • GENBANK/JQ890664
  • GENBANK/JQ890665
  • GENBANK/JQ890666
  • GENBANK/JQ890667
  • GENBANK/JQ890668
  • GENBANK/JQ890669
  • GENBANK/JQ890670
  • GENBANK/JQ890671
  • GENBANK/JQ890672
  • GENBANK/JQ890673
  • GENBANK/JQ890674
  • GENBANK/JQ890675
  • GENBANK/JQ890676
  • GENBANK/JQ890677
  • GENBANK/JQ890678
  • GENBANK/JQ890679
  • GENBANK/JQ890680
  • GENBANK/JQ890681
  • GENBANK/JQ890682
  • GENBANK/JQ890683
  • GENBANK/JQ890684
  • GENBANK/JQ890685
  • GENBANK/JQ890686
  • GENBANK/JQ890687
  • GENBANK/JQ890688
  • GENBANK/JQ890689
  • GENBANK/JQ890690
  • GENBANK/JQ890691
  • GENBANK/JQ890692
  • GENBANK/JQ890693
  • GENBANK/JQ890694
  • GENBANK/JQ890695
  • GENBANK/JQ890696
  • GENBANK/JQ890697
  • GENBANK/JQ890698
  • GENBANK/JQ890699
  • GENBANK/JQ890700
  • GENBANK/JQ890701
  • GENBANK/JQ890702
  • GENBANK/JQ890703
  • GENBANK/JQ890704
  • GENBANK/JQ890705
  • GENBANK/JQ890706
  • GENBANK/JQ890707
  • GENBANK/JQ890708
  • GENBANK/JQ890709
  • GENBANK/JQ890710
  • GENBANK/JQ890711
  • GENBANK/JQ890712
  • GENBANK/JQ890713
  • GENBANK/JQ890714
  • GENBANK/JQ890715
  • GENBANK/JQ890716
  • GENBANK/JQ890717
  • GENBANK/JQ890718
  • GENBANK/JQ890719
  • GENBANK/JQ890720
  • GENBANK/JQ890721
  • GENBANK/JQ890722
  • GENBANK/JQ890723
  • GENBANK/JQ890724
  • GENBANK/JQ890725
  • GENBANK/JQ890726
  • GENBANK/JQ890727
  • GENBANK/JQ890728
  • GENBANK/JQ890729
  • GENBANK/JQ890730
  • GENBANK/JQ890731
  • GENBANK/JQ890732
  • GENBANK/JQ890733
  • GENBANK/JQ890734
  • GENBANK/JQ890735
  • GENBANK/JQ890736
  • GENBANK/JQ890737
  • GENBANK/JQ890738
  • GENBANK/JQ890739
  • GENBANK/JQ890740
  • GENBANK/JQ890741
  • GENBANK/JQ890742
  • GENBANK/JQ890743
  • GENBANK/JQ890744
  • GENBANK/JQ890745
  • GENBANK/JQ890746
  • GENBANK/JQ890747
  • GENBANK/JQ890748
  • GENBANK/JQ890749
  • GENBANK/JQ890750
  • GENBANK/JQ890751
  • GENBANK/JQ890752
  • GENBANK/JQ890753
  • GENBANK/JQ890754
  • GENBANK/JQ890755
  • GENBANK/JQ890756
  • GENBANK/JQ890757
  • GENBANK/JQ890758
  • GENBANK/JQ890759
  • GENBANK/JQ890760
  • GENBANK/JQ890761
  • GENBANK/JQ890762
  • GENBANK/JQ890763
  • GENBANK/JQ890764
  • GENBANK/JQ890765
  • GENBANK/JQ890766
  • GENBANK/JQ890767
  • GENBANK/JQ890768
  • GENBANK/JQ890769
  • GENBANK/JQ890770
  • GENBANK/JQ890771
  • GENBANK/JQ890772
  • GENBANK/JQ890773
  • GENBANK/JQ890774
  • GENBANK/JQ890775
  • GENBANK/JQ890776
  • GENBANK/JQ890777
  • GENBANK/JQ890778
  • GENBANK/JQ890779
  • GENBANK/JQ890780
  • GENBANK/JQ890781
  • GENBANK/JQ890782
  • GENBANK/JQ890783
  • GENBANK/JQ890784
  • GENBANK/JQ890785
  • GENBANK/JQ890786
  • GENBANK/JQ890787
  • GENBANK/JQ890788
  • GENBANK/JQ890789
  • GENBANK/JQ890790
  • GENBANK/JQ890791
  • GENBANK/JQ890792
  • GENBANK/JQ890793
  • GENBANK/JQ890794
  • GENBANK/JQ890795
  • GENBANK/JQ890796
  • GENBANK/JQ890797
  • GENBANK/JQ890798
  • GENBANK/JQ890799
  • GENBANK/JQ890800
  • GENBANK/JQ890801
  • GENBANK/JQ890802
  • GENBANK/JQ890803
  • GENBANK/JQ890804
  • GENBANK/JQ890805
  • GENBANK/JQ890806
  • GENBANK/JQ890807
  • GENBANK/JQ890808
  • GENBANK/JQ890809
  • GENBANK/JQ890810
  • GENBANK/JQ890811
  • GENBANK/JQ890812
  • GENBANK/JQ890813
  • GENBANK/JQ890814
  • GENBANK/JQ890815
  • GENBANK/JQ890816
  • GENBANK/JQ890817
  • GENBANK/JQ890818
  • GENBANK/JQ890819
  • GENBANK/JQ890820
  • GENBANK/JQ890821
  • GENBANK/JQ890822
  • GENBANK/JQ890823
  • GENBANK/JQ890824
  • GENBANK/JQ890825
  • GENBANK/JQ890826
  • GENBANK/JQ890827
  • GENBANK/JQ890828
  • GENBANK/JQ890829
  • GENBANK/JQ890830
  • GENBANK/JQ890831
  • GENBANK/JQ890832
  • GENBANK/JQ890833
  • GENBANK/JQ890834
  • GENBANK/JQ890835
  • GENBANK/JQ890836
  • GENBANK/JQ890837
  • GENBANK/JQ890838
  • GENBANK/JQ890839
  • GENBANK/JQ890840
  • GENBANK/JQ890841
  • GENBANK/JQ890842
  • GENBANK/JQ890843
  • GENBANK/JQ890844
  • GENBANK/JQ890845
  • GENBANK/JQ890846
  • GENBANK/JQ890847
  • GENBANK/JQ890848
  • GENBANK/JQ890849
  • GENBANK/JQ890850
  • GENBANK/JQ890851
  • GENBANK/JQ890852
  • GENBANK/JQ890853
  • GENBANK/JQ890854
  • GENBANK/JQ890855
  • GENBANK/JQ890856
  • GENBANK/JQ890857
  • GENBANK/JQ890858
  • GENBANK/JQ890859
  • GENBANK/JQ890860
  • GENBANK/JQ890861
  • GENBANK/JQ890862
  • GENBANK/JQ890863
  • GENBANK/JQ890864
  • GENBANK/JQ890865
  • GENBANK/JQ890866
  • GENBANK/JQ890867
  • GENBANK/JQ890868
  • GENBANK/JQ890869
  • GENBANK/JQ890870
  • GENBANK/JQ890871
  • GENBANK/JQ890872
  • GENBANK/JQ890873
  • GENBANK/JQ890874
  • GENBANK/JQ890875
  • GENBANK/JQ890876
  • GENBANK/JQ890877
  • GENBANK/JQ890878
  • GENBANK/JQ890879
  • GENBANK/JQ890880
  • GENBANK/JQ890881
  • GENBANK/JQ890882
  • GENBANK/JQ890883
  • GENBANK/JQ890884
  • GENBANK/JQ890885
  • GENBANK/JQ890886
  • GENBANK/JQ890887
  • GENBANK/JQ890888
  • GENBANK/JQ890889
  • GENBANK/JQ890890
  • GENBANK/JQ890891
  • GENBANK/JQ890892
  • GENBANK/JQ890893
  • GENBANK/JQ890894
  • GENBANK/JQ890895
  • GENBANK/JQ890896
  • GENBANK/JQ890897
  • GENBANK/JQ890898
  • GENBANK/JQ890899
  • GENBANK/JQ890900
  • GENBANK/JQ890901
  • GENBANK/JQ890902
  • GENBANK/JQ890903
  • GENBANK/JQ890904
  • GENBANK/JQ890905
  • GENBANK/JQ890906
  • GENBANK/JQ890907
  • GENBANK/JQ890908
  • GENBANK/JQ890909
  • GENBANK/JQ890910
  • GENBANK/JQ890911
  • GENBANK/JQ890912
  • GENBANK/JQ890913
  • GENBANK/JQ890914
  • GENBANK/JQ890915
  • GENBANK/JQ890916
  • GENBANK/JQ890917
  • GENBANK/JQ890918
  • GENBANK/JQ890919
  • GENBANK/JQ890920
  • GENBANK/JQ890921
  • GENBANK/JQ890922
  • GENBANK/JQ890923
  • GENBANK/JQ890924
  • GENBANK/JQ890925
  • GENBANK/JQ890926
  • GENBANK/JQ890927
  • GENBANK/JQ890928
  • GENBANK/JQ890929
  • GENBANK/JQ890930
  • GENBANK/JQ890931
  • GENBANK/JQ890932
  • GENBANK/JQ890933
  • GENBANK/JQ890934
  • GENBANK/JQ890935
  • GENBANK/JQ890936
  • GENBANK/JQ890937
  • GENBANK/JQ890938
  • GENBANK/JQ890939
  • GENBANK/JQ890940
  • GENBANK/JQ890941
  • GENBANK/JQ890942
  • GENBANK/JQ890943
  • GENBANK/JQ890944
  • GENBANK/JQ890945
  • GENBANK/JQ890946
  • GENBANK/JQ890947
  • GENBANK/JQ890948
  • GENBANK/JQ890949
  • GENBANK/JQ890950
  • GENBANK/JQ890951
  • GENBANK/JQ890952
  • GENBANK/JQ890953
  • GENBANK/JQ890954
  • GENBANK/JQ890955
  • GENBANK/JQ890956
  • GENBANK/JQ890957
  • GENBANK/JQ890958
  • GENBANK/JQ890959
  • GENBANK/JQ890960
  • GENBANK/JQ890961
  • GENBANK/JQ890962
  • GENBANK/JQ890963
  • GENBANK/JQ890964
  • GENBANK/JQ890965
  • GENBANK/JQ890966
  • GENBANK/JQ890967
  • GENBANK/JQ890968
  • GENBANK/JQ890969
  • GENBANK/JQ890970
  • GENBANK/JQ890971
  • GENBANK/JQ890972
  • GENBANK/JQ890973
  • GENBANK/JQ890974
  • GENBANK/JQ890975
  • GENBANK/JQ890976
  • GENBANK/JQ890977
  • GENBANK/JQ890978
  • GENBANK/JQ890979
  • GENBANK/JQ890980
  • GENBANK/JQ890981
  • GENBANK/JQ890982
  • GENBANK/JQ890983
  • GENBANK/JQ890984
  • GENBANK/JQ890985
  • GENBANK/JQ890986
  • GENBANK/JQ890987
  • GENBANK/JQ890988
  • GENBANK/JQ890989
  • GENBANK/JQ890990
  • GENBANK/JQ890991
  • GENBANK/JQ890992
  • GENBANK/JQ890993
  • GENBANK/JQ890994
  • GENBANK/JQ890995
  • GENBANK/JQ890996
  • GENBANK/JQ890997
  • GENBANK/JQ890998
  • GENBANK/JQ890999
  • GENBANK/JQ891000
  • GENBANK/JQ891001
  • GENBANK/JQ891002
  • GENBANK/JQ891003
  • GENBANK/JQ891004
  • GENBANK/JQ891005
  • GENBANK/JQ891006
  • GENBANK/JQ891007
  • GENBANK/JQ891008
  • GENBANK/JQ891009
  • GENBANK/JQ891010
  • GENBANK/JQ891011
  • GENBANK/JQ891012
  • GENBANK/JQ891013
  • GENBANK/JQ891014
  • GENBANK/JQ891015
  • GENBANK/JQ891016
  • GENBANK/JQ891017
  • GENBANK/JQ891018
  • GENBANK/JQ891019
  • GENBANK/JQ891020
  • GENBANK/JQ891021
  • GENBANK/JQ891022
  • GENBANK/JQ891023
  • GENBANK/JQ891024
  • GENBANK/JQ891025
  • GENBANK/JQ891026
  • GENBANK/JQ891027
  • GENBANK/JQ891028
  • GENBANK/JQ891029
  • GENBANK/JQ891030
  • GENBANK/JQ891031
  • GENBANK/JQ891032
  • GENBANK/JQ891033
  • GENBANK/JQ891034
  • GENBANK/JQ891035
  • GENBANK/JQ891036
  • GENBANK/JQ891037
  • GENBANK/JQ891038
  • GENBANK/JQ891039
  • GENBANK/JQ891040
  • GENBANK/JQ891041
  • GENBANK/JQ891042
  • GENBANK/JQ891043
  • GENBANK/JQ891044
  • GENBANK/JQ891045
  • GENBANK/JQ891046
  • GENBANK/JQ891047
  • GENBANK/JQ891048
  • GENBANK/JQ891049
  • GENBANK/JQ891050
  • GENBANK/JQ891051
  • GENBANK/JQ891052
  • GENBANK/JQ891053
  • GENBANK/JQ891054
  • GENBANK/JQ891055
  • GENBANK/JQ891056
  • GENBANK/JQ891057
  • GENBANK/JQ891058
  • GENBANK/JQ891059
  • GENBANK/JQ891060
  • GENBANK/JQ891061
  • GENBANK/JQ891062
  • GENBANK/JQ891063
  • GENBANK/JQ891064
  • GENBANK/JQ891065
  • GENBANK/JQ891066
  • GENBANK/JQ891067
  • GENBANK/JQ891068
  • GENBANK/JQ891069
  • GENBANK/JQ891070
  • GENBANK/JQ891071
  • GENBANK/JQ891072
  • GENBANK/JQ891073
  • GENBANK/JQ891074
  • GENBANK/JQ891075
  • GENBANK/JQ891076
  • GENBANK/JQ891077
  • GENBANK/JQ891078
  • GENBANK/JQ891079
  • GENBANK/JQ891080
  • GENBANK/JQ891081
  • GENBANK/JQ891082
  • GENBANK/JQ891083
  • GENBANK/JQ891084
  • GENBANK/JQ891085
  • GENBANK/JQ891086
  • GENBANK/JQ891087
  • GENBANK/JQ891088
  • GENBANK/JQ891089
  • GENBANK/JQ891090
  • GENBANK/JQ891091
  • GENBANK/JQ891092
  • GENBANK/JQ891093
  • GENBANK/JQ891094
  • GENBANK/JQ891095
  • GENBANK/JQ891096
  • GENBANK/JQ891097
  • GENBANK/JQ891098
  • GENBANK/JQ891099
  • GENBANK/JQ891100
  • GENBANK/JQ891101
  • GENBANK/JQ891102
  • GENBANK/JQ891103
  • GENBANK/JQ891104
  • GENBANK/JQ891105
  • GENBANK/JQ891106
  • GENBANK/JQ891107
  • GENBANK/JQ891108
  • GENBANK/JQ891109
  • GENBANK/JQ891110
  • GENBANK/JQ891111
  • GENBANK/JQ891112
  • GENBANK/JQ891113
  • GENBANK/JQ891114
  • GENBANK/JQ891115
  • GENBANK/JQ891116
  • GENBANK/JQ891117
  • GENBANK/JQ891118
  • GENBANK/JQ891119
  • GENBANK/JQ891120
  • GENBANK/JQ891121
  • GENBANK/JQ891122
  • GENBANK/JQ891123
  • GENBANK/JQ891124
  • GENBANK/JQ891125
  • GENBANK/JQ891126
  • GENBANK/JQ891127
  • GENBANK/JQ891128
  • GENBANK/JQ891129
  • GENBANK/JQ891130
  • GENBANK/JQ891131
  • GENBANK/JQ891132
  • GENBANK/JQ891133
  • GENBANK/JQ891134
  • GENBANK/JQ891135
  • GENBANK/JQ891136
  • GENBANK/JQ891137
  • GENBANK/JQ891138
  • GENBANK/JQ891139
  • GENBANK/JQ891140
  • GENBANK/JQ891141
  • GENBANK/JQ891142
  • GENBANK/JQ891143
  • GENBANK/JQ891144
  • GENBANK/JQ891145
  • GENBANK/JQ891146
  • GENBANK/JQ891147
  • GENBANK/JQ891148
  • GENBANK/JQ891149
  • GENBANK/JQ891150
  • GENBANK/JQ891151
  • GENBANK/JQ891152
  • GENBANK/JQ891153
  • GENBANK/JQ891154
  • GENBANK/JQ891155
  • GENBANK/JQ891156
  • GENBANK/JQ891157
  • GENBANK/JQ891158
  • GENBANK/JQ891159
  • GENBANK/JQ891160
  • GENBANK/JQ891161
  • GENBANK/JQ891162
  • GENBANK/JQ891163
  • GENBANK/JQ891164
  • GENBANK/JQ891165
  • GENBANK/JQ891166
  • GENBANK/JQ891167
  • GENBANK/JQ891168
  • GENBANK/JQ891169
  • GENBANK/JQ891170
  • GENBANK/JQ891171
  • GENBANK/JQ891172
  • GENBANK/JQ891173
  • GENBANK/JQ891174
  • GENBANK/JQ891175
  • GENBANK/JQ891176
  • GENBANK/JQ891177
  • GENBANK/JQ891178
  • GENBANK/JQ891179
  • GENBANK/JQ891180
  • GENBANK/JQ891181
  • GENBANK/JQ891182
  • GENBANK/JQ891183
  • GENBANK/JQ891184
  • GENBANK/JQ891185
  • GENBANK/JQ891186
  • GENBANK/JQ891187
  • GENBANK/JQ891188
  • GENBANK/JQ891189
  • GENBANK/JQ891190
  • GENBANK/JQ891191
  • GENBANK/JQ891192
  • GENBANK/JQ891193
  • GENBANK/JQ891194
  • GENBANK/JQ891195
  • GENBANK/JQ891196
  • GENBANK/JQ891197
  • GENBANK/JQ891198
  • GENBANK/JQ891199
  • GENBANK/JQ891200
  • GENBANK/JQ891201
  • GENBANK/JQ891202
  • GENBANK/JQ891203
  • GENBANK/JQ891204
  • GENBANK/JQ891205
  • GENBANK/JQ891206
  • GENBANK/JQ891207
  • GENBANK/JQ891208
  • GENBANK/JQ891209
  • GENBANK/JQ891210
  • GENBANK/JQ891211
  • GENBANK/JQ891212
  • GENBANK/JQ891213
  • GENBANK/JQ891214
  • GENBANK/JQ891215
  • GENBANK/JQ891216
  • GENBANK/JQ891217
  • GENBANK/JQ891218
  • GENBANK/JQ891219
  • GENBANK/JQ891220
  • GENBANK/JQ891221
  • GENBANK/JQ891222
  • GENBANK/JQ891223
  • GENBANK/JQ891224
  • GENBANK/JQ891225
  • GENBANK/JQ891226
  • GENBANK/JQ891227
  • GENBANK/JQ891228
  • GENBANK/JQ891229
  • GENBANK/JQ891230
  • GENBANK/JQ891231
  • GENBANK/JQ891232
  • GENBANK/JQ891233
  • GENBANK/JQ891234
  • GENBANK/JQ891235
  • GENBANK/JQ891236
  • GENBANK/JQ891237
  • GENBANK/JQ891238
  • GENBANK/JQ891239
  • GENBANK/JQ891240
  • GENBANK/JQ891241
  • GENBANK/JQ891242
  • GENBANK/JQ891243
  • GENBANK/JQ891244
  • GENBANK/JQ891245
  • GENBANK/JQ891246
  • GENBANK/JQ891247
  • GENBANK/JQ891248
  • GENBANK/JQ891249
  • GENBANK/JQ891250
  • GENBANK/JQ891251
  • GENBANK/JQ891252
  • GENBANK/JQ891253
  • GENBANK/JQ891254
  • GENBANK/JQ891255
  • GENBANK/JQ891256
  • GENBANK/JQ891257
  • GENBANK/JQ891258
  • GENBANK/JQ891259
  • GENBANK/JQ891260
  • GENBANK/JQ891261
  • GENBANK/JQ891262
  • GENBANK/JQ891263
  • GENBANK/JQ891264
  • GENBANK/JQ891265
  • GENBANK/JQ891266
  • GENBANK/JQ891267
  • GENBANK/JQ891268
  • GENBANK/JQ891269
  • GENBANK/JQ891270
  • GENBANK/JQ891271
  • GENBANK/JQ891272
  • GENBANK/JQ891273
  • GENBANK/JQ891274
  • GENBANK/JQ891275
  • GENBANK/JQ891276
  • GENBANK/JQ891277
  • GENBANK/JQ891278
  • GENBANK/JQ891279
  • GENBANK/JQ891280
  • GENBANK/JQ891281
  • GENBANK/JQ891282
  • GENBANK/JQ891283
  • GENBANK/JQ891284
  • GENBANK/JQ891285
  • GENBANK/JQ891286
  • GENBANK/JQ891287
  • GENBANK/JQ891288
  • GENBANK/JQ891289
  • GENBANK/JQ891290
  • GENBANK/JQ891291
  • GENBANK/JQ891292
  • GENBANK/JQ891293
  • GENBANK/JQ891294
  • GENBANK/JQ891295
  • GENBANK/JQ891296
  • GENBANK/JQ891297
  • GENBANK/JQ891298
  • GENBANK/JQ891299
  • GENBANK/JQ891300
  • GENBANK/JQ891301
  • GENBANK/JQ891302
  • GENBANK/JQ891303
  • GENBANK/JQ891304
  • GENBANK/JQ891305
  • GENBANK/JQ891306
  • GENBANK/JQ891307
  • GENBANK/JQ891308
  • GENBANK/JQ891309
  • GENBANK/JQ891310
  • GENBANK/JQ891311
  • GENBANK/JQ891312
  • GENBANK/JQ891313
  • GENBANK/JQ891314
  • GENBANK/JQ891315
  • GENBANK/JQ891316
  • GENBANK/JQ891317
  • GENBANK/JQ891318
  • GENBANK/JQ891319
  • GENBANK/JQ891320
  • GENBANK/JQ891321
  • GENBANK/JQ891322
  • GENBANK/JQ891323
  • GENBANK/JQ891324
  • GENBANK/JQ891325
  • GENBANK/JQ891326
  • GENBANK/JQ891327
  • GENBANK/JQ891328
  • GENBANK/JQ891329
  • GENBANK/JQ891330
  • GENBANK/JQ891331
  • GENBANK/JQ891332
  • GENBANK/JQ891333
  • GENBANK/JQ891334
  • GENBANK/JQ891335
  • GENBANK/JQ891336
  • GENBANK/JQ891337
  • GENBANK/JQ891338
  • GENBANK/JQ891339
  • GENBANK/JQ891340
  • GENBANK/JQ891341
  • GENBANK/JQ891342
  • GENBANK/JQ891343
  • GENBANK/JQ891344
  • GENBANK/JQ891345
  • GENBANK/JQ891346
  • GENBANK/JQ891347
  • GENBANK/JQ891348
  • GENBANK/JQ891349
  • GENBANK/JQ891350
  • GENBANK/JQ891351
  • GENBANK/JQ891352
  • GENBANK/JQ891353
  • GENBANK/JQ891354
  • GENBANK/JQ891355
  • GENBANK/JQ891356
  • GENBANK/JQ891357
  • GENBANK/JQ891358
  • GENBANK/JQ891359
  • GENBANK/JQ891360
  • GENBANK/JQ891361
  • GENBANK/JQ891362
  • GENBANK/JQ891363
  • GENBANK/JQ891364
  • GENBANK/JQ891365
  • GENBANK/JQ891366
  • GENBANK/JQ891367
  • GENBANK/JQ891368
  • GENBANK/JQ891369
  • GENBANK/JQ891370
  • GENBANK/JQ891371
  • GENBANK/JQ891372
  • GENBANK/JQ891373
  • GENBANK/JQ891374
  • GENBANK/JQ891375
  • GENBANK/JQ891376
  • GENBANK/JQ891377
  • GENBANK/JQ891378
  • GENBANK/JQ891379
  • GENBANK/JQ891380
  • GENBANK/JQ891381
  • GENBANK/JQ891382
  • GENBANK/JQ891383
  • GENBANK/JQ891384
  • GENBANK/JQ891385
  • GENBANK/JQ891386
  • GENBANK/JQ891387
  • GENBANK/JQ891388
  • GENBANK/JQ891389
  • GENBANK/JQ891390
  • GENBANK/JQ891391
  • GENBANK/JQ891392
  • GENBANK/JQ891393
  • GENBANK/JQ891394
  • GENBANK/JQ891395
  • GENBANK/JQ891396
  • GENBANK/JQ891397
  • GENBANK/JQ891398
  • GENBANK/JQ891399
  • GENBANK/JQ891400
  • GENBANK/JQ891401
  • GENBANK/JQ891402
  • GENBANK/JQ891403
  • GENBANK/JQ891404
  • GENBANK/JQ891405
  • GENBANK/JQ891406
  • GENBANK/JQ891407
  • GENBANK/JQ891408
  • GENBANK/JQ891409
  • GENBANK/JQ891410
  • GENBANK/JQ891411
  • GENBANK/JQ891412
  • GENBANK/JQ891413
  • GENBANK/JQ891414
  • GENBANK/JQ891415
  • GENBANK/JQ891416
  • GENBANK/JQ891417
  • GENBANK/JQ891418
  • GENBANK/JQ891419
  • GENBANK/JQ891420
  • GENBANK/JQ891421
  • GENBANK/JQ891422
  • GENBANK/JQ891423
  • GENBANK/JQ891424
  • GENBANK/JQ891425
  • GENBANK/JQ891426
  • GENBANK/JQ891427
  • GENBANK/JQ891428
  • GENBANK/JQ891429
  • GENBANK/JQ891430
  • GENBANK/JQ891431
  • GENBANK/JQ891432
  • GENBANK/JQ891433
  • GENBANK/JQ891434
  • GENBANK/JQ891435
  • GENBANK/JQ891436
  • GENBANK/JQ891437
  • GENBANK/JQ891438
  • GENBANK/JQ891439
  • GENBANK/JQ891440
  • GENBANK/JQ891441
  • GENBANK/JQ891442
  • GENBANK/JQ891443
  • GENBANK/JQ891444
  • GENBANK/JQ891445
  • GENBANK/JQ891446
  • GENBANK/JQ891447
  • GENBANK/JQ891448
  • GENBANK/JQ891449
  • GENBANK/JQ891450
  • GENBANK/JQ891451
  • GENBANK/JQ891452
  • GENBANK/JQ891453
  • GENBANK/JQ891454
  • GENBANK/JQ891455
  • GENBANK/JQ891456
  • GENBANK/JQ891457
  • GENBANK/JQ891458
  • GENBANK/JQ891459
  • GENBANK/JQ891460
  • GENBANK/JQ891461
  • GENBANK/JQ891462
  • GENBANK/JQ891463
  • GENBANK/JQ891464
  • GENBANK/JQ891465
  • GENBANK/JQ891466
  • GENBANK/JQ891467
  • GENBANK/JQ891468
  • GENBANK/JQ891469
  • GENBANK/JQ891470
  • GENBANK/JQ891471
  • GENBANK/JQ891472
  • GENBANK/JQ891473
  • GENBANK/JQ891474
  • GENBANK/JQ891475
  • GENBANK/JQ891476
  • GENBANK/JQ891477
  • GENBANK/JQ891478
  • GENBANK/JQ891479
  • GENBANK/JQ891480
  • GENBANK/JQ891481
  • GENBANK/JQ891482
  • GENBANK/JQ891483
  • GENBANK/JQ891484
  • GENBANK/JQ891485
  • GENBANK/JQ891486
  • GENBANK/JQ891487
  • GENBANK/JQ891488
  • GENBANK/JQ891489
  • GENBANK/JQ891490
  • GENBANK/JQ891491
  • GENBANK/JQ891492
  • GENBANK/JQ891493
  • GENBANK/JQ891494
  • GENBANK/JQ891495
  • GENBANK/JQ891496
  • GENBANK/JQ891497
  • GENBANK/JQ891498
  • GENBANK/JQ891499
  • GENBANK/JQ891500
  • GENBANK/JQ891501
  • GENBANK/JQ891502
  • GENBANK/JQ891503
  • GENBANK/JQ891504
  • GENBANK/JQ891505
  • GENBANK/JQ891506
  • GENBANK/JQ891507
  • GENBANK/JQ891508
  • GENBANK/JQ891509
  • GENBANK/JQ891510
  • GENBANK/JQ891511
  • GENBANK/JQ891512
  • GENBANK/JQ891513
  • GENBANK/JQ891514
  • GENBANK/JQ891515
  • GENBANK/JQ891516
  • GENBANK/JQ891517
  • GENBANK/JQ891518
  • GENBANK/JQ891519
  • GENBANK/JQ891520
  • GENBANK/JQ891521
  • GENBANK/JQ891522
  • GENBANK/JQ891523
  • GENBANK/JQ891524
  • GENBANK/JQ891525
  • GENBANK/JQ891526
  • GENBANK/JQ891527
  • GENBANK/JQ891528
  • GENBANK/JQ891529
  • GENBANK/JQ891530
  • GENBANK/JQ891531
  • GENBANK/JQ891532
  • GENBANK/JQ891533
  • GENBANK/JQ891534
  • GENBANK/JQ891535
  • GENBANK/JQ891536
  • GENBANK/JQ891537
  • GENBANK/JQ891538
  • GENBANK/JQ891539
  • GENBANK/JQ891540
  • GENBANK/JQ891541
  • GENBANK/JQ891542
  • GENBANK/JQ891543
  • GENBANK/JQ891544
  • GENBANK/JQ891545
  • GENBANK/JQ891546
  • GENBANK/JQ891547
  • GENBANK/JQ891548
  • GENBANK/JQ891549
  • GENBANK/JQ891550
  • GENBANK/JQ891551
  • GENBANK/JQ891552
  • GENBANK/JQ891553
  • GENBANK/JQ891554
  • GENBANK/JQ891555
  • GENBANK/JQ891556
  • GENBANK/JQ891557
  • GENBANK/JQ891558
  • GENBANK/JQ891559
  • GENBANK/JQ891560
  • GENBANK/JQ891561
  • GENBANK/JQ891562
  • GENBANK/JQ891563
  • GENBANK/JQ891564
  • GENBANK/JQ891565
  • GENBANK/JQ891566
  • GENBANK/JQ891567
  • GENBANK/JQ891568
  • GENBANK/JQ891569
  • GENBANK/JQ891570
  • GENBANK/JQ891571
  • GENBANK/JQ891572
  • GENBANK/JQ891573
  • GENBANK/JQ891574
  • GENBANK/JQ891575
  • GENBANK/JQ891576
  • GENBANK/JQ891577
  • GENBANK/JQ891578
  • GENBANK/JQ891579
  • GENBANK/JQ891580
  • GENBANK/JQ891581
  • GENBANK/JQ891582
  • GENBANK/JQ891583
  • GENBANK/JQ891584
  • GENBANK/JQ891585
  • GENBANK/JQ891586
  • GENBANK/JQ891587
  • GENBANK/JQ891588
  • GENBANK/JQ891589
  • GENBANK/JQ891590
  • GENBANK/JQ891591
  • GENBANK/JQ891592
  • GENBANK/JQ891593
  • GENBANK/JQ891594
  • GENBANK/JQ891595
  • GENBANK/JQ891596
  • GENBANK/JQ891597
  • GENBANK/JQ891598
  • GENBANK/JQ891599
  • GENBANK/JQ891600
  • GENBANK/JQ891601
  • GENBANK/JQ891602
  • GENBANK/JQ891603
  • GENBANK/JQ891604
  • GENBANK/JQ891605
  • GENBANK/JQ891606
  • GENBANK/JQ891607
  • GENBANK/JQ891608
  • GENBANK/JQ891609
  • GENBANK/JQ891610
  • GENBANK/JQ891611
  • GENBANK/JQ891612
  • GENBANK/JQ891613
  • GENBANK/JQ891614
  • GENBANK/JQ891615
  • GENBANK/JQ891616
  • GENBANK/JQ891617
  • GENBANK/JQ891618
  • GENBANK/JQ891619
  • GENBANK/JQ891620
  • GENBANK/JQ891621
  • GENBANK/JQ891622
  • GENBANK/JQ891623
  • GENBANK/JQ891624
  • GENBANK/JQ891625
  • GENBANK/JQ891626
  • GENBANK/JQ891627
  • GENBANK/JQ891628
  • GENBANK/JQ891629
  • GENBANK/JQ891630
  • GENBANK/JQ891631
  • GENBANK/JQ891632
  • GENBANK/JQ891633
  • GENBANK/JQ891634
  • GENBANK/JQ891635
  • GENBANK/JQ891636