1. p. 2. p. 3. p. 4. p. 5. p. 6. p. 1 Sd Skt Gt /+ Skt Gt 2 Sd Skt Gt /+ Skt Gt 3 4 Skt 5) 6) 7. p. 1 Skt Gt Sd 2 [Sd Skt Gt /+ Skt Gt ] 3 Skt 4 1
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- ひろき あさぶき
- 5 years ago
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1 熊本大学学術リポジトリ Kumamoto University Repositor Title 直腸肛門奇形に関する発生学的解析 Author(s) 須田, 博子 Citation Issue date Type URL Thesis or Dissertation Right
2 1. p. 2. p. 3. p. 4. p. 5. p. 6. p. 1 Sd Skt Gt /+ Skt Gt 2 Sd Skt Gt /+ Skt Gt 3 4 Skt 5) 6) 7. p. 1 Skt Gt Sd 2 [Sd Skt Gt /+ Skt Gt ] 3 Skt 4 1
3 8. p. 9. p. 10. p. 2
4 1. 100% Danforth s short tail(sd) Sickle tail (Skt) (Skt Gt ) (Sd Skt Gt /+ Skt Gt ) 100% (cloaca) Skt (Sd Skt Gt /+ Skt Gt ) Skt 9.5 (E9.5) E13.5 HE (Skt Gt ) Skt Xgal E10.5 RNA In situ hybridization (ISH) E10.5 cloaca plate E11.5 E13.5 Skt E9.5 cloaca plate E11.5 cloaca plate E13.5 cloaca plate Cdx 5 Hox Hoxa13 Hoxd13 Cdx2 cloacal plate Skt cloaca plate cloaca plate Skt cloaca plate 3
5 ISH Hoxa13 Hoxd13 Cdx2 S kt cloaca plate Hoxa13 Hoxd13 Cdx2 4
6 2. (1) Hiroko Suda Kwang-Jong Lee Kei Semba Fumie Kyushima Takashi Ando Masatake Araki Kimi Araki Yukihiro Inomata Ken-ichi Yamamura The Skt gene, required for anorectal development, is a candidate for a molecular marker of the cloacal plate Pediatr Surg Int. 2011;27(3): (2) Takashi Ando, Kei Semba, Hiroko Suda, Akira Sei, Hiroshi Mizuta, Masatake Araki, Kuniya Abe, Kenji Imai, Naomi Nakagata, Kimi Araki, Ken-ichi Yamamura The floor plate is sufficient for development of the sclerotome and spine without the notochord. Mech Dev. 2011;128(1-2): ,, ;40:
7 3. 6
8 4. ABC : avidin-biotin complex bhlh basic helix-loop-helix β-gal β-galactosidase β-geo β-galactosidase neomycin BMP bone morphogenic protein BSA bovine serum albumin, CAG : cytomegarovirus enhancer, chicken beta actin promoter, rabbit beta-globin polya cm centimorgan, Cdx : caudal type homeobox DAPI : 4',6-diamidino-2-phenylindole DNA Deoxyribo nucleic acid, E Embryonic day, Fgf : fibroblast growth factor GTP : guanosine triphosphate, GDP : Guanosine diphosphate, Gapdh : Glyceraldehyde 3-phosphate dehydrogenase HE Hematoxylin-Eosin HH : Hamburger and Hamilton Hox : homeobox gene IRES : Internal ribosome entry site. n.s. not significant PBS phosphate buffered saline 7
9 PBT phosphate buffered saline containing 0.1% Tween-20 PCR polymerase chain reaction, PFA Paraformaldehyde RT room temperature, SD Standard Deviation, Sd Danforth s short tail Skt Sickle tail Skt Gt B6;CB-Skt GtAyu8021IMEG Shh Sonic hedgehog TBE Tris-borate-EDTA 8
10 5. A. 5, Amussat Pena posterior sagittal anorectoplasty [1] [2, 3] B Ladd & Gross I IV Stephen Wingspread [1] 9
11 X Wingspread Classification (1984) 1. a) b) a) b) C. 50% 10
12 [4] [5] VACTERL (Vertebral anomalies, anal atresia, cardiac malformations, tracheoesophageal fistula, renal anomalies, and limb anomalies) MURCS (Mullerian duct aplasia, renal aplasia, and cervicothoracic somite dysplasia) OEIS (Omphalocele, exstrophy, imperforate anus, and spinal defects) Axial mesodermal dysplasia Klippel-Feil syndrome Sirenomelia-caudal regression Trisomy 21 Trisomy 13 Trisomy 18 Pallister-Killian syndrome Cat-eye syndrome Parental unidisomy 16 Deletion 22q11 syndrome (del22q11.2) Currarino syndrome Pallister-Hall syndrome Townes-Brock syndrome Ulnar-mammary syndrome Okihiro syndrome Rieger syndrome Thanatophoric dwarfism Hirschsprung disease Feingold syndrome Kabuki syndrome Optitz BBB/G syndrome Johanson-Blizzard syndrome Spondylocostal dysostosis Short rib polydactyly syndrome Baller-Gerold syndrome Ciliopathies Fraser syndrome Lowe syndrome Heterotaxia FG syndrome X-linked mental retardation MIDAS syndrome Christian syndrome 11
13 D [6] Townes-Brocks syndrome [7, 8] Currarino's syndrome [9, 10] Pallister-Hall syndrome [11, 12] 21 [2, 5, 6] [13] [14] [15] [16] [15] [16] E cloacal plate cloacal plate [17, 18] cloacal plate [19, 20] cloacal plate [21, 22] cloacal plate 12
14 F. Hamburger and Hamilton HH stage 8 HH stage 13 HH stage Sonic hedgehog (Shh) Shh Bmp4 Hox [23, 24] Shh Shh -/ - Gli2 Gli3 Gli2 Gli3 [25, 26] Shh Hox Hox Hox ' Hox Hoxb-8 Hoxb-9 Hoxc-9 Hoxa-13 Hoxd-13 Hoxa-11 Hoxd-11 Hoxd-12 Hoxa-13 Hoxd-13 Hoxd-12 Hoxa-13 Hoxd-13 [27] Hoxd-12 -/ - Hoxd-13 -/ - [28] Hoxa13 +/ - ;Hoxd13 -/ - [29] Hox 13
15 Hox ParaHox Cdx Cdx2 8.5 [30] Cdx2 Cdx2 [31] Cdx Hox Cdx [32] 9.5 Cdx2 [33] A A [21, 34] Cyp26a1 Cyp26a1 [35-37] Fgf Fgf10 -/ cloacal plate cloacal plate [38][39] Wnt Wnt5a 10.5 Sd [21] Wnt5a -/ - [40] Wnt Dact1 14
16 Dact1 Wnt Wnt/βcatenin PCP [41] Wnt Sall 4 Sall Sall1 Towens-Brocks syndrome Sall4 Okihiro sundrome Sall1 -/ - [42] Sall4 +/ - Sall1 +/ - Sall4 +/ - Sall1 +/ - Sall4 +/ - truncated Sall1 Sall4 [43] VACTER Pcsk5 Pcsk5 TGFβ Gdf1 Hox Caudal regression syndrome Mnx1 Gdf1 -/ - [44] 100% 15
17 Shh -/ - [25, 26] Gli-2 -/ - [25] Gli-3 -/ - [25] Gli-2 -/ -; Gli-3 +/ - [25] Gli-2 +/ -; Gli-3 -/ - [25] Gli-2 -/ -; Gli-3 -/ - [25] Hoxd13 -/ - [28] Hoxa13 +/ -; Hoxd13 -/ - [29] Cdx2 -/ - (conditional) [33] Cyp26a1 -/ - sirenomelia [36] Fgf10 -/ - [39] Wnt5a -/ - 25%, 75%) [40] Dact1 -/ - [41] Sall4 +/ - [43] Sall1 +/ - Sall4 +/ - [43] Pcsk5 [44] Gdf1 -/ - [44] G. Danforth s short tail (Sd) Sickle tail (Skt) Sd 1930 Danforth [45, 46] Sd Sd Sd Sd Sd Sd Sd Sd Sd Sd 16
18 Sd [47-49] Cre-loxP Cre loxp Cre-loxP IRES lacz neomycin β-geo polya [50] pu-8 ES Skt Gt [51] Skt (Sickle tail) Sd Sd Skt Gt Sd 0.95cM Skt [51] 2 cis -configuration (Sd Skt Gt /+ +) Skt locus lacz Sd genotyping Skt Skt Gt Skt [51] Skt Gt Skt Gt Sd [Sd +/+ +] [Sd Skt Gt /+ + ] [Sd Skt Gt /+ Skt Gt ] Skt [52] [Sd Skt Gt /+ Skt Gt ] 100% 100% 100% Sd [Sd Skt Gt /+ Skt Gt ] Sd 17
![100% Skt [Sd Skt Gt /+ Skt Gt ] Skt 1.](/docs-images/92/108238164/images/19-0.jpg "Sd Skt (a) Sd Skt Sd Sd (b) Skt Skt 18")
19 100% Skt [Sd Skt Gt /+ Skt Gt ] Skt 1. Sd Skt (a) Sd Skt Sd Sd (b) Skt Skt 18
20 2. Sd locus Skt locus Skt Gt a b (a) Sd, Skt locus Physical / Genetic Map (b) Sd locus Skt Gt cM (b) Skt Gt Skt exon trap pu-8 black box: exon 4. ( [52] 19
21 6. 1) Sd Skt G /+ Skt Gt pu-8 [50] ES ICR (Charles River) C57BL/6 (CLEA) F1 Danforth s short tail (Sd) (Bar Harbor, ME, USA) Sd Skt G /+ Skt Gt Sd +/+ + (C57BL/6 genetic background) C57BL/6 genetic background Skt Gt/Gt Sd Skt Gt [Sd Skt Gt /+ +; cis configuration] Skt Gt/Gt genome PCR Skt wildtype allele Skt 14 GTS (5 - CCACCCCTACATGTGTCTTT -3 ) GTA (5 - CGAGTAAGTAACATCCCTCC -3 ) 339bp PCR [+ +/+ Skt Gt ] 2) Sd Skt G /+ Skt Gt Sd Skt G /+ Skt Gt % formaldehyde/pbs HISTOS5 (Leica Microsystems) 4μm Hematoxylin-Eosin (HE) 3) % formaldehyde/pbs HISTOS5 (Leica 20
22 Microsystems) 4μm Hematoxylin-Eosin (HE) 4) Skt Skt Skt Sd Skt Skt Skt [51] [52] β Whole-mount X-gal Allen [53] 1% formaldehyde, 0.2% glutaraldehyde, and 0.02% NP-40 in phosphate-buffered saline (PBS) 0.5 PBS X-gal (5 mm potassium ferricyanide, 5 mm potassium ferrocyanide, 2 mm MgCl2, 0.5% X-gal in PBS) 30 PBS 4% paraformaldehyde Whole body (25% 50% 70% 100% and 100%, 1 ) benzylalcohol/benzylbenzoate (1:2) X-gal 8μm Nuclear Fast red 5) Sd Sd Skt [Sd Skt Gt /+ Skt Gt ] 21
23 cloacal plate 10.5 n=7 RNAeasy mini kit QIAGEN total RNA Filgen CodeLink TM Mouse Whole Genome Bioarray [+ Skt Gt /+ +] [Sd Skt Gt /+ Skt Gt ] [Sd Skt Gt /+ Skt Gt ] / [+ Skt Gt /+ +] Microarray Data Analysis tool WikiPathways NCBI BioSystems Skt Skt Skt CAG Skt BMT10 (Nucleofector, Lonza) Skt BMT10 PFA PBT BSA Skt Calnexine DAPI Phalloidin Tubulin Calnexin rabbit anti-mouse EGFR antibody (Cell Signaling Technology, Inc. Beverly, MA), 1:1,000. Phalloidin Green Fluorescent Conjugate, Acti-stain (Cytoskeleton, Inc. ), 100nM. Monoclonal Anti-β-Tubulin FITC antibody produced in mouse (Sigma Aldrich Corp.), 1:100. Purified Mouse Anti-Calnexin anti body (BD Transduction Laboratories), 1:50. RhoGTP ase 22
24 Rho Rho G G GTP GDP 10.5 n= n=5 [+ Skt Gt /+ +] [Sd Skt Gt /+ Skt Gt ] Rho RhoA Cdc42 GTP RhoA GTP Cdc42 G-LIZA Small G-Activation Kit Cytoskeleton, Inc Lysis Buffer lysate GTP incubate RhoA Cdc42 [Sd Skt Gt /+ Skt Gt ] 20 Hoxa-13 Hoxd-13 Cdx2 Real time PCR In situ hybridization Real time PCR total RNA cdna Taqman Hoxa13 ; Mm _m1 Hoxd13 ; Mm _m1 Cdx2 ; Mm _m1 Gapdh ; Mm _g1 TaqMan Gene Expression Assays Gapdh Whole mount in situ hybridization DIG labeled RNA probe Cdx2 Hoxa13 Hoxd13 23
25 Cdx2 Forward primer 5 -aca gca gca gca gca aca ac-3 Reverse primer 5 -tga ctc gaa cag cag caa ac- 3 Hoxa13 Forward primer 5 - ctt ccc atg gaa agc tat c -3 Reverse primer 5 - ttt ctc ttt gac cct cct g -3 Hoxd13 Forward primer 5 - tgg gct atg gct acc act tc -3 Reverse primer 5 - gat gaa gac tca gtg gag ac -3 Shh ShhcDNA PCR pgem -T Easy Vector Promega QIAGEN Plasmid Maxi Kit(QIAGEN) DIG-labelled RNA probe cdna 5 MCS (37 ) RNA Polymerase (T3, T7 or Sp6 )(Roche Diagnostics) DIG RNA Labeling Mix(Roche Diagnostics) 37 2 incubate in vitro transcription RNA 30 Tris RNase free (BioRad) 95 3 denature 1 TBE 100V 15 In situ hybridization DIG labeled RNA probe Rosen Kokubu Whole mount in situ hybridization [54, 55] 4% PFA (4 ) PBT (25% 50% 75% 100% 100% 30 ) -80 Hybridization 75% 50% 25% /PBS 10 incubate 6% /PBS 60 ( ) PBT ( ) RIPA (0.05% SDS, 0.15M NaCl, 1% NP40, 0.5%, 1mM EDTA, 5mM Tris-HCl ph8.0) 10 incubate ( ) PBT ( ) 4 PFA/0.2% /PBT 20 hybe-buffer (50% formamide, 5 SSC, 500µg heparin, 0.02% Tween20) + trna (100µg/ml, invitrogen) 68 3 incubate 24
26 prehybridization RNA 80, 3 denature hybe-buffer + trna (100µg/ml) 1:100 (250ng/ml) 68 hybridization DIG hybe-buffer 65 RNase solution (0.5M NaCl, 10mM Tris-HCl ph7.5, 0.1% Tween20) RNase A(100µg/ml) incubate SSC/FA/Tween20 (50% formamide, 2 SSC, 0.1% Tween20) 5 /2 10 /3 30 /5 (65 ) TBST (RT) MABT(100mM, 150mM NaCl, ph7.5, 0.1% Tween-20) (RT) blocking buffer (MABT, 1.5% blocking reagent [Roche Diagnostics]) 1 blocking RT DIG (Anti-Digoxigenin-AP, Fab fragments [Roche Diagnostics]) blocking buffer (1 2000) 4, 1 preabsorption blocking buffer DIG blocking buffer (4 ) TBST 1 8 (RT) TBST 4 NTMT[100mM NaCl, 100mM Tris-HCl (ph 9.5), 50mM MgCl2, 0.1% Tween20] incubate (RT) NBT/BCIP Stock Solution (Roche Diagnostics)/ NTMT ( 4 ) 4% PFA 30 (RT) Whole mount (25% 50% 75% 100% 100% 30 ) benzylalcohol/benzylbenzoate (1:2) 8μm Nuclear Fast red 4%PFA 4μm 0.3% / PBS ABC Hematoxylin Cdx2 ( cell signaling #3977) 1/100 25
27 (5) ± (SE) VARA t p <
28 7. Skt Gt Sd 19.5 [+ Skt Gt /+ Skt Gt ] (n=7) [Sd +/+ +] 50% (n=12) [Sd Skt Gt /+ + ] 80% (n=10) [Sd Skt Gt /+ Skt Gt ] 100% (n=8) [Sd +/+ +] 33% [Sd Skt Gt /+ + ] 75% [Sd Skt Gt /+ Skt Gt ] 75% ( 1 ) [Sd +/+ +] 33%(n=12) [Sd Skt Gt /+ + ] 25% (n=10) [Sd Skt Gt /+ Skt Gt ] 13% (n=8) (data not shown) [Sd Skt Gt /+ Skt Gt ] E9.5 E10.5 E11.5 E12.5 E13.5 [+ +/+ Skt Gt ] 9.5 cloacal plate 10.5 cloacal plate cloacal plate 13.5 cloacal plate cloacal plate [Sd Skt Gt /+ Skt Gt ] cloacal plate cloacal plate
29 cloacal plateを欠如した部分で間葉組織が増殖し 尿直腸中隔のcloacal plateへの接着が妨げ られていた 胎生13.5日になっても尿直腸中隔の下降が適切に起こっていないため 尿生 殖洞と肛門直腸洞との分離は不完全となり 両者を交通する瘻孔が形成されていた 図 3 図 3. 直腸肛門発生過程の比較 胎生9.5日から胎生13.5日総排腔発生過程を正中矢状断で示した コントロールマウス [+ SktGt /+ +] では cloacal plateは増殖し尾根部まで伸長している 尿直腸中隔が下降し 腹側の尿生殖洞と背側の直腸肛門洞に 分離される ( a-e ) 胎生13.5日に肛門膜が消失し肛門が開口する 直腸肛門奇形モデル [Sd SktGt/+ SktGt] では clocal plateの尾側へ伸長していない 尿直腸中隔は下降しているがcloacal plateと接触することなく 尿生殖 洞と直腸肛門洞は完全に分離されていない ( f-j ) 黒矢 瘻孔 白矢 増殖している間葉組織 括弧 cloacal plate. Bars=100µm, HG: hind gut, TG: tail gut, URS: urorectal septum, UGS: urogenital sinus, GT: genital tubercle. 28
30 Skt Skt X-gal [+ Skt Gt /+ Skt Gt ] [+ Skt Gt /+ +] [+ Skt Gt /+ +] [+ Skt Gt /+ Skt Gt ] [+ Skt Gt /+ +] [+ Skt Gt /+ Skt Gt ] [Sd Skt Gt /+ Skt Gt ] 3 [+ Skt Gt /+ +] cloacal plate β 10.5 [+ Skt Gt /+ Skt Gt ] 9.5 cloacal plate β [+ Skt Gt /+ +] cloacal plate cloacal plate cloacal plate [Sd Skt Gt /+ Skt Gt ] β cloacal plate cloacal plate 4 In situ hybridization cloacal plate (data not shown) Skt cloacal plate cloacal plate Skt Xgal 29
![図 4. 総排腔周囲のXgal染色 胎生9.5日から胎生12.5日の[+ SktGt /+ +] ( a-d ) [+ SktGt/+ SktGt] ( e-h ) [Sd SktGt/+ SktGt] ( i-l )のXgal染色を 示す βガラクトシダーゼ活性は [+ SktGt /+ +] では胎生10.5日から12.5日までcloacal plateで確認され 胎生 11.](/docs-images/92/108238164/images/31-0.jpg "5日からはcloacal plateだけでなくcloacal plate周囲の間葉組織でも確認されるようになる [+ SktGt/+ SktGt] では[+ SktGt /+ +]より強い発現が得られ 胎生9.")
31 図 4. 総排腔周囲のXgal染色 胎生9.5日から胎生12.5日の[+ SktGt /+ +] ( a-d ) [+ SktGt/+ SktGt] ( e-h ) [Sd SktGt/+ SktGt] ( i-l )のXgal染色を 示す βガラクトシダーゼ活性は [+ SktGt /+ +] では胎生10.5日から12.5日までcloacal plateで確認され 胎生 11.5日からはcloacal plateだけでなくcloacal plate周囲の間葉組織でも確認されるようになる [+ SktGt/+ SktGt] では[+ SktGt /+ +]より強い発現が得られ 胎生9.5日のcloacal plateとなる内胚葉 外胚葉の部分でβガラクトシ ダーゼ活性が確認できる [Sd SktGt/+ SktGt]のβガラクトシダーゼ活性のある組織は [+ SktGt /+ +] ) [+ SktGt/+ SktGt]とほぼ同じである 胎生12.5日では 後腸の代わりに増殖している間葉組織でのβガラクトシダーゼ活 性が認められる 黒矢 βガラクトシダーゼ活性 白矢 増殖している間葉組織 括弧 cloacal plate Bars=100µm, HG: hind gut, TG: tail gut, URS: urorectal septum, UGS: urogenital sinus. 30
![４ 脊索と直腸肛門奇形 直腸肛門奇形モデルの脊索の形態について総排泄腔レベルの脊索に注目して解析した まず 全体像を把握するためWhole mount で観察を行った [+ SktGt/+ +] 胚では 神経管腹 側に頭尾則に発達した脊索が観察される [Sd SktGt/+ SktGt] 胚では 胎生9.5日の尾側の脊 索は正常に発達しているが 胸椎レベルでは脊索の分断化が認められた 胎生10.](/docs-images/92/108238164/images/32-0.jpg "5日にな ると総排腔背側のレベルでも脊索形成が障害されており 分断化している部分に加え 腹 側に枝状に伸長する脊索が観察された この異常組織を組織切片にて正中矢状断で観察す ると背側大動脈が一部欠損し その欠損部を通り前方へ枝状に伸びる脊索組織が観察され た 図5 脊索で発現する遺伝子としてよく知られてるShhの発現の有無を確認したとこ ろ 異常をきたした脊索においてもShhの発現は保たれていた")
32 ４ 脊索と直腸肛門奇形 直腸肛門奇形モデルの脊索の形態について総排泄腔レベルの脊索に注目して解析した まず 全体像を把握するためWhole mount で観察を行った [+ SktGt/+ +] 胚では 神経管腹 側に頭尾則に発達した脊索が観察される [Sd SktGt/+ SktGt] 胚では 胎生9.5日の尾側の脊 索は正常に発達しているが 胸椎レベルでは脊索の分断化が認められた 胎生10.5日にな ると総排腔背側のレベルでも脊索形成が障害されており 分断化している部分に加え 腹 側に枝状に伸長する脊索が観察された この異常組織を組織切片にて正中矢状断で観察す ると背側大動脈が一部欠損し その欠損部を通り前方へ枝状に伸びる脊索組織が観察され た 図5 脊索で発現する遺伝子としてよく知られてるShhの発現の有無を確認したとこ ろ 異常をきたした脊索においてもShhの発現は保たれていた また 横断像で同部位を 観察したところ 血管の欠損部は背側大動脈の融合が障害された正中部分であった 図 5 図5. 総排腔周囲の脊索形成 31
![Xgal 9. 5 12.5 whole mount [+ Skt Gt /+ +] ( a-c ) [Sd Skt Gt /+ Skt Gt ] ( d-f ) 10.](/docs-images/92/108238164/images/33-0.jpg "5 [+ Skt Gt /+ +] ( i ) [Sd Skt Gt / + Skt Gt ] ( j, k ) β [+ Skt Gt /+ +] 10.5 12.5 [Sd Skt Gt /+ Skt Gt ] 9.5 [+ Skt Gt /+ +] 10.")
33 Xgal whole mount [+ Skt Gt /+ +] ( a-c ) [Sd Skt Gt /+ Skt Gt ] ( d-f ) 10.5 [+ Skt Gt /+ +] ( i ) [Sd Skt Gt / + Skt Gt ] ( j, k ) β [+ Skt Gt /+ +] [Sd Skt Gt /+ Skt Gt ] 9.5 [+ Skt Gt /+ +] 10.5 ( j ) Bars=100µm, NC: notochord, TG: tail gut, CL:cloaca, NT: neural tube, DA: dorsal aorta 32
34 図6. 脊索でのShhの発現 In situ hybridization法により 胎生10.5日胚脊索のShhの発現を横断面にて解析した (a), (b) は [+ SktGt /+ +] (c), (d)は [Sd SktGt/+ SktGt] を示す (a), (c) は(b), (d) のそれぞれ拡大像を示す 黒四角 [Sd SktGt/+ SktGt] の 異常な形態を示す脊索でもShhは発現していた 神経底板が誘導されておらず 背側大動脈の融合が障害され ていた Bars=100µm, NC: notochord, CL:cloaca, NT: neural tube, DA: dorsal aorta, FP: floor plate ５ 遺伝子発現プロファイリング解析 ①シグナルパスウェイ解析 マイクロアレイで得られた全マウス36227遺伝子の解析結果より 発現強度比 [Sd SktGt/+ SktGt] / [+ SktGt/+ +]が2.0以上の遺伝子もしくは 0.5以下であった遺伝子を抽出したところ 1291遺伝子が抽出された これらの遺伝子を用いてパスウェイ解析を行ったところ９つの シグナルパスウェイで有意差が得られた 表５ いずれも過去の報告で直腸肛門発生と の関連は指摘されていないシグナル経路であり 各々の変動していた遺伝子についてもこ れまで直腸肛門発生との関連は示唆されていない 我々は以前 ヒトSkt遺伝子の解析で S k t 蛋白が細胞骨格の一つである微小管と共局在するという結果を得ている そこで 33
35 WikiPathways (Regulation of actin cytoskeleton) BioSystems (G13 signaling pathway) Skt Rho G (a) Skt Skt BMT10 Skt Skt Skt Tubulin Skt 7 Phalloidin Calnexin (b) Rho G 10.5 n= n=5 Rho G RhoA Cdc RhoA Cdc RhoA Cdc42 8. Changed Total Z score P-value Pathway name genes genes ( >0) ( 0.05) Oocyte meiosis (BioSystems) Regulation of actin cytoskeleton (WikiPathways) Ovarian infertility genes (WikiPathways) G13 signaling pathway (WikiPathways) Beta oxidation meta pathway (WikiPathways) Axon guidance (BioSystems) B cell receptor signaling pathway (WikiPathways) T cell receptor signaling pathway (BioSystems) FcγR-mediated phagocytosis (BioSystems)
36 7. Skt Skt BMT10 (a, d, g) Skt (b, e, h) DAPI Skt 35
37 8. RhoGTPase 10.5 (n=5) 11.5 (n=5) Cdc42 RhoA Cdc42 RhoA Hoxa-13 / ; Hoxd-13 / ; Cdx2 / ; PCR 10.5 Hoxd-13 Hoxa-13 Cdx2 36
38 In situ hybrydization 9.5, 10.5 Hoxa-13 Hoxd-13 Cdx2 Hoxa-13 Hoxd-13 Whole mount in situ hybridization Hoxd-13 whole body whole body Cdx2 whole body 9.5 cloacal plate
39 . Gene Normalized intensity {Sd Skt Gt /+ Skt Gt } ( n=7) Normalized intensity {++/+ Skt Gt } ( n=7) Normalized intensity Ratio / Shh Gli Hoxa Hoxd Cdx Cyp26a Rara Rarb Fgf Wnt5a Dact Sall Sall Noggin Efnb EphB EphB PCR 38
40 Cdx2 Hoxa13 Hoxd13のE9.5 (n=7) E10.5 (n=4) における発現量について定量PCRで評価した 縦軸には Gapdhとの相対比を示す E10.5のHoxd13のARMでの発現上昇は有意差を認めた 図10. Hoxa-13 Hoxd-13発現パターン (Whole mount In situ hybridization法) 胎生9.5日 胎生10.5日の Hoxa-13 Hoxd-13の発現パターン をWhole mount In situ hybridization法にて示し た 上段にコントロール [+ +/+ SktGt] 下段に直腸肛門奇形モデル [Sd SktGt/+ SktGt] を示す Hoxa-13, Hoxd-13の発現パターンは類似しているが Hoxd-13の方が発現が強い 胎生9.5日ではコントロール モデル で明らかな違いはない 胎生10.5日になるとコントロールでは認められなかった尾腹側に二重線のシグナル が得られた 矢印 39
41 図11. Hoxd-13発現パターン (In situ hybridization) 胎生9.5日を正中矢状断 胎生10.5日は冠状断で示した 胎生9.5日 コントロールマウスで は 後腸内胚葉 腹側でのみシグナルが確認できるが(a) 直腸肛門奇形モデルでは後腸内胚葉背側 矢印 尾芽と尾根部の 間葉組織でもシグナルが検出された (c) 胎生10.5日ではコントロールマウスでは生殖結節での発現が 認められる (b) 直腸肛門奇形モデルでは生殖結節の発現強度は上昇しており 矢頭 また 尾腸内胚葉 その周囲の間葉組織での発現 を認めた (d) HG: hind gut, TG: tail gut, CL: cloaca, NT: neural tube, NC: notochord, GT: genital tubercle, TB: tail bud, WD: wolffian duct. 40
42 図12. Cdx2発現パターン 胎生9.5日と胎生10.5日のCdx2発現パターンをWhole mount in situ hybrydization (a, c, e, g)と正中矢状断像の免 疫染色像を示した (b, d, f, h) Whole bodyでは胎生9.5日の尾芽 尾側の神経管と腸管での発現を 胎生10.5日 の尾先端での発現をコントロール モデルともに認めている (a, c, e, g) 組織切片にて詳細に観察すると 直 腸肛門奇形モデルで 胎生9.5日にcloacal plate周辺組織の発現低下が観察された 矢印 (b, f) HG: hind gut, TG: tail gut, CL: cloaca, NT: neural tube, TB: tail bud. 41
43 8. Sd Skt [Sd Skt Gt /+ Skt Gt ] 100% Skt cloacal plate cloacal plate Skt cloacal plate Skt Sd Cdx2 5 Hox Sd Sd Sd Skt Gt 100% Sd Skt Sd [52] Skt cloacal plate cloacal plate cloacal plate cloacal plate urethral plate [56-58] Skt cloacal plate cloacal plate Sd 42
44 [Sd Skt Gt /+ Skt Gt ] cloacal plate Skt Gt cloacal plate Skt β Skt Skt 1, [51] PSOAT N proline-rich ( a.a.) coiled-coil ( a.a.) Skt 14 Skt coiled-coil proline-rich β-geo Skt Skt coiled-coil [59] proline-rich SH3 WW [60] [61] in Vitro Skt [62] Skt Skt Sd Skt cloacal plate 43
45 [51] [52] [63-65] 6.5 (node) definitive endoderm 8.0 definitive endoderm definitive endoderm definitive endoderm [66][67] 1998 Hebrok Shh HH stage 11 Shh HH stage 15 Shh [68] Shh Pdx1 [68] Bmp Noggin Noggin Bmp [63-65] Shh Bmp Shh Shh 44
46 VEGF [69] cloacal plate Sd Skt Skt Skt Skt [70] Rho G G GTP GDP Rho RhoA Rac1 Cdc Rho Rac1 Cdc42 Rho [71, 72] Rho Cdc42 Rac1 Cdc42/Rac1 RhoA [73] Rho G 45
47 [74] Skt N proline-rich N proline-rich G [75] RhoA Cdc42 clocal plate Rho Hoxa-13, Hoxd-13 Cdx2 Sd PCR 10.5 Hoxd-13 Hoxd-13 Hoxa-13 Cdx2 PCR Hoxa-13, Hoxd-13 Cdx2 cloacal plate Young Cdx2 Hox-13 Sd [76] Young Hox Hox-13 Hox Young 10.5 Hox Cdx Hox-13 Cdx Hox-13 Sd Cdx Wnt Fgf [77][78] Sd 46
48 Sd in Vivo 12. Sd Skt 76 Sd critical Sd 47
49 Cdx2 Hox13 Skt Sd 48
50 9. Skt cloaca plate Hoxa13 Hoxd13 Cdx2 49
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