SH2B1 藉由影響染色質狀態及MyoD-DNA 結合能力來促進肌肉基因的表現

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研究生：	陳冠維 Chen, Kuan-Wei
論文名稱：	SH2B1 藉由影響染色質狀態及MyoD-DNA 結合能力來促進肌肉基因的表現 SH2B1 modulates chromatin state and MyoD occupancy to enhance expressions of myogenic genes
指導教授：	陳令儀 Chen, Linyi
口試委員:	林玉俊 Lin, Yu-Chun 高承福 Kao, Cheng-Fu 陳盛良 Chen, Shen-Liang 鄭世進 Cheng, Shih-Chin
學位類別：	博士 Doctor
系所名稱：	生命科學暨醫學院 - 分子醫學研究所 Institute of Molecular Medicine
論文出版年：	2017
畢業學年度：	105
語文別：	英文
論文頁數：	201
中文關鍵詞：	染色質、分化、組蛋白修飾、肌肉生成、轉錄調控
外文關鍵詞：	Chromatin, Differentiation, Histone modifications, Myogenesis, Transcriptional regulation
相關次數：	點閱：3 下載：0
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肌肉生成對於胚胎發育及肌肉再生是必要的。當中胚層的細胞決定走向肌肉生成時，一連串的訊息蛋白包括類胰島素生長因子將啟動許多的訊號傳遞路徑，最後調控染色質的重組及肌肉基因的轉錄。然而在這過程中，染色質的重組，轉錄因子與DNA的結合，連接組蛋白(linker histone)及組蛋白(histone)之修飾的交互作用並不清楚。因此，我們證明一個在肌肉生成中可以調控泛染色質聚集和肌肉管生成的新蛋白−SH2B1。SH2B1和連接組蛋白H1形成複合體並可以幫助組蛋白H1從轉錄起始點(transcription start site)離開，進一步促進肌肉基因Igf2和Myog的表現。這暗示SH2B1是一個有潛力的連接組蛋白H1的伴護蛋白(histone H1 chaperone)。再者，SH2B1對於在Igf2及Myog附近的啟動子(promoter)和增強子(enhancer)上的轉錄因子MyoD-DNA之結合，MyoD活性及組蛋白H3第四離胺酸三個甲基化(H3K4me3)的上升和第九離胺酸三個甲基化(H3K9me3)的下降都是必要的。總結以上的實驗可以指出SH2B1是一個具有潛力的連接組蛋白H1之伴護蛋白，可以精確調控染色質的狀態及肌肉基因的表現，最後促進肌肉生成。肌肉細胞之間的融合是形成肌肉組織的一個關鍵步驟。我們在肌肉分化早期發現肌肉細胞(C2C12 cells)的細胞膜有泡狀結構。這泡狀結構僅出現在分化的肌肉細胞而與細胞凋亡無關。這樣的結果暗示泡狀結構出現在肌肉細胞融合之前。除此之外，對於細胞融合關鍵的蛋白質−肌動蛋白(actin)，肌球蛋白重鏈(myosin heavy chain)，肌肉黏著分子(M-cadherin)及β-catenin−也出現在泡狀結構裡面。這些實驗暗示著泡狀結構與肌肉細胞融合是有關的。在這篇論文中，我們展示在肌肉生成時，SH2B1可以透過表關遺傳學的方式影響肌肉基因的表現以及一個在肌肉細胞融合前的特殊細胞形態。

Myogenesis is essential during embryogenesis as well as muscle regeneration. As mesoderm-derived cell lineage commits to myogenesis, a spectrum of signaling molecules, including insulin-like growth factor (IGF), activate signaling pathways and ultimately instruct chromatin remodeling and the transcription of myogenic genes. Nonetheless, the interplay among chromatin remodeling, transcription factor binding, linker histone and histone modifications during myogenesis remains to be clarified. To this end, we have identified and characterized a novel myogenic regulator, SH2B1, which is required for global chromatin condensation during myogenesis and the formation of myotubes. SH2B1 interacts with histone H1 and is required for the removal of histone H1 from active transcription sites, allowing for the expressions of myogenic genes, Igf2 and Myog, indicating SH2B1 is a potential histone H1 chaperone. Furthermore, SH2B1 is required for the transcriptional activity of MyoD and MyoD occupancy, the induction of histone H3 lysine 4 trimethylation as well as the reduction of histone H3 lysine 9 trimethylation at the enhancer/promoter regions of Igf2 and Myog during myogenesis. Together, these findings demonstrate that SH2B1, as a putative histone H1 chaperone, fine-tunes global-local chromatin states, expressions of myogenic genes and ultimately promotes myogenesis.
Fusion between myoblasts is a key step for forming muscle tissue. During early stage of myogenesis, bubble-like structures protruding from plasma membrane are observed. These bubble-like structures are formed only on differentiating C2C12 cells and not correlated with apoptosis. Our findings suggest that bubble-like structures appear before the fusion of the two myoblasts. Furthermore, actin, myosin heavy chain, M-cadherin as well as β-catenin that are crucial for fusion localize within the bubble-like structures. These data imply that the formation of bubble-like structures may be required for the fusion. Taken together, this thesis demonstrates an epigenetic role of SH2B1 on myogenic genes and describes a differentiation specific morphological change before fusion of myoblasts.

Table of the content    I
Abstract    V
中文摘要    VII
Acknowledgements    VIII
Publication List    X
Introduction    1
Myogenesis    1
Signal pathways initiate by insulin like growth factors 1 and 2 (IGF1 and 2) during myogenesis    2
Epigenetic regulation of Myog gene    5
Histone Chaperone    7
SH2B adaptor protein 1 (SH2B1)    10
Rhabdomyosarcoma (RMS)    11
Myoblast fusion    12
Adhesion molecules and actin cytoskeletons    12
Lipids at plasma membrane    14
Materials and Methods    17
Reagents    17
Plasmids    18
Cell culture    18
Isolation and culture of primary myoblasts    19
Knockdown of endogenous SH2B1 and SET via RNA interference    19
Immunoblotting and immunoprecipitation    20
Fractionation    21
Immunofluorescence staining, timelapse imaging and fusion index    21
Proteomic analysis using liquid chromatograph-tandem mass spectrometry    22
Semi-quantitative real-time polymerase chain reaction (qPCR)    22
Luciferase assays    23
Chromatin immunoprecipitation (ChIP) assays    23
Restriction endonuclease digestion    24
Micrococcal nuclease (MNase) digestion assays    25
Duolink in situ proximity ligation assay (PLA)    25
Glutathione S-transferase (GST) pull down assay    26
Statistical analysis    26
Results    27
Differential expression of SH2B family members during myogenesis    27
SH2B1 is required for myotube formation    27
SH2B1 is required for chromatin re-organization and condensation during myogenesis    29
SH2B1 is required for the expression of Igf2, transcriptional activity of MyoD and histone modifications of Igf2    31
SH2B1 regulates Myog expression through histone modifications and MyoD occupancy    35
Morphological changes before myoblasts fusion    39
Actin and MyHC are localized in the bubble-like structures during myogenesis    40
Farnesylated-GFP marks the bubble-like structure    41
M-cadherin and β-catenin are localized on the bubble-like structures    41
Differentiating myoblasts with bubble-like structures are not apoptotic cells    42
Discussion    43
Figures    51
Figure 1. The mRNA expression of SH2B family members during myogenesis.    51
Figure 2. The protein level of SH2B1 during myogenesis.    52
Figure 3. Establishment of C2C12 cells stably knocking down SH2B1.    53
Figure 4. SH2B1 is required for myotube formation.    54
Figure 5. Knockdown of SH2B1 reduces the length of myotube and the expression of MyHC in primary myoblast.    55
Figure 6. Overexpressing SH2B1 increases the level of Myogenin during myogenesis.    56
Figure 7. Overexpression of SH2B1 promotes myotube formation.    57
Figure 8. Overexpressing SH2B1 elevates the length of myotube and the expression of MyHC in primary myoblast.    58
Figure 9. Overexpressing SH2B1 in C2C12-shSH2B1 cells rescues myogenesis.    59
Figure 10. SH2B1 does not affect proliferation and cell cycle exist.    60
Figure 11. SH2B1 is required for chromatin re-organization during myogenesis.    61
Figure 12. Potential SH2B1 interacting proteins.    62
Figure 13. SH2B1 interacts with histone H1 in vitro.    63
Figure 14. SH2B1 interacts with histone H1 in vivo.    65
Figure 15. SH2B1 promotes chromatin condensation during myogenesis.    66
Figure 16. SET is required for myogenesis.    67
Figure 17. SH2B1 is required for the expression of Igf2.    68
Figure 18. SH2B1 did not affect the expression of IGF1R.    69
Figure 19. SH2B1 interacts with IGF1R.    70
Figure 20. Knocking down SH2B1 decreases the accessibility of chromatin at Igf2 enhancer during myogenesis.    71
Figure 21. SH2B1 is required for the removal of histone H1 at the promoter of Igf2.    72
Figure 22. SH2B1 modulates histone modifications at Igf2 promoter and enhancer during myogenesis.    73
Figure 23. Knockdown of SH2B1 decreases the level of pAKT but not pERK1/2.    74
Figure 24. SH2B1 regulates MyoD transcriptional activity and MyoD occupancy at Igf2 enhancer region.    75
Figure 25. SH2B1 promotes the nuclear export of FOXO3a.    77
Figure 26. SH2B1 is required for the expression of Myogenin but not MyoD.    78
Figure 27. Knocking down SH2B1 decreases the accessibility of chromatin at Myog promoter during myogenesis.    79
Figure 28. SH2B1 is required for the removal of histone H1 at the promoter of Myog.    80
Figure 29. SH2B1 modulates histone modifications at Myog promoter region during myogenesis.    81
Figure 30. SH2B1 regulates Myog promoter activity and MyoD occupancy at Myog promoter and enhancer regions.    82
Figure 31. SH2B1 regulates epigenetic modifiers during myogenesis.    84
Figure 32. The occupancy of MyoD increases at the 5’ region of Sh2b1 during myogenesis.    85
Figure 33. SH2B1 enhances differentiation of RD cells.    86
Figure 34. Membrane protrusions were observed during myogenesis.    87
Figure 35. Bubble-like structures only form during myogenesis.    89
Figure 36. Farnesylated-GFP marks the bubble-like structure.    90
Figure 37. M-cadherin and β-catenin are localized at bubble-like structre.    91
Figure 38. C2C12 cells with blebs are not apoptotic cells.    92
Figure 39. MyoD occupancy at mouse Myog gene during myogenesis.    93
Figure 40. Schematic model of how SH2B1 regulates myogenesis.    94
Tables    96
Table 1. List of antibodies dilution and their applications    96
Table 2. List of primers for gene expression, ChIP and restriction endonuclease digestion    97
Table 3. SH2B1 binding partners during myogenesis    100
Reference    105

                                

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