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| DC 欄位 | 值 | 語言 |
|---|---|---|
| dc.contributor.advisor | 吳益群 | |
| dc.contributor.author | Tsung-Yuan Hsu | en |
| dc.contributor.author | 徐綜遠 | zh_TW |
| dc.date.accessioned | 2021-06-08T04:19:26Z | - |
| dc.date.copyright | 2010-07-27 | |
| dc.date.issued | 2010 | |
| dc.date.submitted | 2010-07-22 | |
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(2001a). The C. elegans PH domain protein CED-12 regulates cytoskeletal reorganization via a Rho/Rac GTPase signaling pathway. Dev Cell 1, 477-89. Zhou, Z., Hartwieg, E. and Horvitz, H. R. (2001b). CED-1 is a transmembrane receptor that mediates cell corpse engulfment in C. elegans. Cell 104, 43-56. Zhou, Z., Mangahas, P. M. and Yu, X. (2004). The genetics of hiding the corpse: engulfment and degradation of apoptotic cells in C. elegans and D. melanogaster. Curr Top Dev Biol 63, 91-143. | |
| dc.identifier.uri | http://tdr.lib.ntu.edu.tw/jspui/handle/123456789/22506 | - |
| dc.description.abstract | 計劃性細胞死亡對多細胞生物體的發育與維持其細胞總體數目的恆定相當重要，而死細胞最終必須被吞噬細胞適時的清除以防止其釋出有害物質或引起發炎反應。吞噬細胞藉由細胞膜上的吞噬受器辨認死細胞，進而將其吞噬、分解。先前線蟲的研究顯示，吞噬受器PSR-1會傳遞訊息CED-2/CrkII-CED-5/Dock180-CED-12/ELMO複合體，而活化CED-10/Rac GTPase，促使細胞膜延展並包圍死細胞。然而在遺傳分析實驗中，psr-1突變株的吞噬作用缺失遠低於下游基因突變所造成的影響，顯示可能有其他受器作用於此吞噬路徑。藉由遺傳分析實驗，我們發現插入素 INA-1-PAT-3 與PAT-2-PAT-3以及受器型酪胺酸激酶RTK-X是吞噬受器，且INA-1與RTK-X皆經由CED-2/CrkII傳遞訊號，而與PAT-2則是作用在不同的訊息傳遞路徑。在活體研究發現，INA-1的胞外區域具有辨認死細胞的功能，可直接或間接辨識死細胞上外翻的吞噬訊號phosphatidylserine (PS)，其胞內區域在體外的研究中可與非受器型酪胺酸磷酸酶SRC-1結合，而CED-2/CrkII也可與SRC-1結合。由進一步的遺傳分析我們證實INA-1作為吞噬受器，接收並傳遞吞噬訊息至SRC-1，調控下游的CED-2/CrkII-CED-5/Dock180-CED-12 /ELMO複合體，進而活化CED-10/Rac GTPase，促進細胞骨架移動吞噬死細胞。我們除了確認插入素在活體生物中可以扮演吞噬受器外，我們也提出插入素可以經由與FAK無關的訊息傳遞路徑來活化CED-2/CrkII-CED-5/Dock180-CED-12 /ELMO複合體而這個非傳統的路徑在演化上是具有保守性的。 | zh_TW |
| dc.description.abstract | Engulfment of apoptotic cells is important for cellular homeostasis and the development of multicellular organisms. Engulfment receptor recognizes and promotes the phagocytosis of apoptotic cells by engulfing cells. Previous studies have shown that more than one engulfment receptors act upstream of the conserved signaling module CED-2/CrkII-CED-5/Dock180-CED-12/ELMO for cell corpse removal in C. elegans, but little is known about their identities, except for PSR-1. We show that, in C. elegans, integrins INA-1-PAT-3 and PAT-2- PAT-3 and receptor tyrosine kinase RTK-X function as engulfment receptors. Genetic studies show that ina-1 and rtk-x act through ced-2, whereas pat-2 likely defines a novel engulfment pathway, distinct from the previously identified ones. The INA-1 extracellular domain is shown to bind to the surface of apoptotic cells in vivo. This binding requires phospholipid scramblase SCRM-1, which promotes the exposure of phosphatidylserine, a key “eat-me” signal in apoptotic cells. Furthermore, we identified an essential role of non-receptor tyrosine kinase SRC-1 in INA-1-mediated cell corpse removal. Finally, our genetic and biochemical data suggest that SRC-1 relays the scrm-1-dependent engulfment signal from INA-1 to the conserved motility-promoting signaling complex CED-2/CrkII-CED-5/Dock180- CED-12 /ELMO for CED-10/Rac activation, probably by interactions with CED-2 and the INA-1 cytoplasmic domain, leading to the internalization of apoptotic cells. Our findings provide evidence that integrin functions as an engulfment receptor at the whole organism level and reveal a non-conventional signaling pathway, in which SRC provides a FAK-independent linkage between integrin α and the common motility–promoting signaling module CED-2/CrkII-CED-5/Dock180- CED-12/ELMO to promote the internalization of apoptotic cells. | en |
| dc.description.provenance | Made available in DSpace on 2021-06-08T04:19:26Z (GMT). No. of bitstreams: 1 ntu-99-D92B43002-1.pdf: 2783973 bytes, checksum: 78df413e475be3cfa7733106051a761b (MD5) Previous issue date: 2010 | en |
| dc.description.tableofcontents | Table of contents
致謝…………………………………………1 中文摘要...........................................2 Abstract……………………4 Part I Introduction……………………………..6 Materials and Methods……………………………………………21 Strains and Alleles........21 RNAi experiments.............21 Plasmid construction………………22 Transgenic and Rescue .....………...24 Yeast Two Hybrid system…………………………………………..……...………..25 Kinase assay……………………………….................26 Microscopy………………………………........................26 Results…………………………………………..28 1. ina-1 is important for the efficiency of cell-corpse engulfment during early and mid embryogenesis…………………………………..28 2. Ina-1 acts in a genetic pathway with ced-2, ced-5, and ced-12 for clearance of apoptotic cells……………………30 3. pat-2 may define a novel engulfment pathway………………31 4. Ina-1 acts genetically upstream of ced-2, ced-5 and ced-12 to promote engulfment of apoptotic cells. …………31 5. ina-1 functions in engulfing cells, but not dying cells, to promote cell-corpse removal……………….32 6. ina-1 was important for the initiation and extension of pseudopods around apoptotic cells during the engulfment process …………………………………33 7. Mutations in ina-1 affect the internalization of apoptotic cells………………35 8. ina-1 recognizes the surface of apoptotic cells……………………..………….35 9. Screen for intracellular molecules that may act downstream of ina-1 for cell engulfment ……………………………………………………...............36 10. src-1 likely acts downstream of ina-1 and upstream of ced-2 in the genetic pathway that controls cell corpse engulfment……..……………………….…37 11. SRC-1 autophosphorylates and interacts with INA-1 and CED-2 in vitro........................................38 12. SRC-1 colocalizes with INA-1 to the pseudopods extending around apoptotic cells………………………………39 Part II Results……………………………….42 1. Screen for receptor tyrosine kinases required for the cell-corpse engulfment during embryogenesis in C.elegans……………………………………………..42 2. RTK-X is important for cell corpse engulfment during embryogenesis……..43 3. rtk-x increases apoptotic cell numbers in ina-1 and ced-1 mutants, but not ced-2 , ced-5 and ced-12 mutants………………………………………...…43 4. RTK-X is widely expressed but functions in engulfing cells for cell-corpse removal.………………………………...44 5. RTK-X is able to autophosphorylate itself and physically interacts with CED-2 in the yeast two-hybrid assay……………………………………………………45 6. RTK-X is important for DTC path-finding and is transcriptionally up-regulated in L4.……………………………………………………………….46 Discussion…………………………………………….47 References…………………………………………51 Figure and tables………………………………………62 Figure 1. The ina-1(gm39) animals have a normal but generate division pattern extra cell surviving cells……………………………………..…………...62 Figure 2. ina-1 affects cell corpse engulfment during embryonic stage.……....63 Figure 3. INA-1 is widely expresses in many cells including dying and engulfing cells………………………….64 Figure 4. The Pegl-1 promoter expresse ina-1 in dying cells………………………66 Figure 5. INA-1::GFP is enriched in the pseudopods extending around apoptotic cells..…………………………….67 Figure 6. Recruitment of HGRS-1::mRFP to the apoptotic cell-containing phagosomes…….…………………………………68 Figure 7. ina-1 is important for the initiation and/or extension of membrane processes around apoptotic cells for phagocytic cup formation during the engulfment process…………………………………………………69 Figure 8. ina-1 mutations reduce the efficiency of cell corpse internalization in engulfing cells……………………………………………………………..70 Figure 9. Internalization of an apoptotic cell in an engulfing cell monitored using the transgene Pced-1myri::gfp…………………………….……………….71 Figure 10. INA-1(N) binds to the surface of apoptotic cells in a scrm-1-dependent manner…………………………………72 Figure 11. Secreted GFP does not bind to apoptotic cells………………………73 Figure 12. SRC-1 interacts with CED-2 and the intracellular region of INA-1...............................74 Figure 13. SRC-1 has an in vitro protein kinase activity………………………....75 Figure 14. INA-1 and SRC-1 are co-localized around cell corpse……………….76 Figure 15. INA-1 and SRC-1 are co-localized on the phagocytic cup around cell corpse…………………..77 Figure 16. SRC-1 surrounds dying cell during early phagocytic cup formation.....................78 Figure 17.rtk-x is important for cell corpse engulfment during embryogenesis………………………………………………….....79 Figure 18. RTK-X is expressed in many cells including dying cells.………...…...80 Figure 19. RTK-X::mCherry is localized around apoptotic cell prior to the recruitment of endosomes in the engulfing cell……………………...81 Figure 20. RTK-X is able to autophosphorylate …….……………………………82 Figure 21. RTK-X physically interacts with CED-2 in the Yeast two-Hybrid assay……………………………………...83 Figure 22. RTK-X is expressed on neuron, muscle and intestine cells during larva to adult stage…………………………………………………………..84 Figure 23. DTC migration is defective in rtk-x strain…………………….......…..86 Figure 24. rtk-x is transcriptionally up-regulated at the L4 stage……………….87 Figure 25. The signaling model of rtk-x, pat-2 and ina-1 mediated engulfment pathway………………………………………………………………..88 Figure 26. The intracellular region of INA-1 has specific function in cell-corpse engulfment in C.elegans………………………………………………89 Figure 27. The binding of an apoptotic cell by INA-1(N)::GFP is independent on CD36-like family…………………………………………………..90 Figure 28. SRC-2 expresses on dying cells……………………………………….91 Table 1. ina-1(gm39) mutation results in the survival of cells that normally die in the ventral cord……………………………………………………………92 Table 2. The extra cell corpses of ina-1 mutants are generated by programmed cell death but not anoikis……………………………….93 Table 3. Numbers of persistent cell corpses in ina-1(RNAi) and other ced mutant backgrounds……………………………………………………94 Table 4. pat-2 defines a novel cell-corpse engulfment pathway...........................96 Table 5. ina-1 genetically acts upstream of ced-2,ced-5 and ced-12…………….97 Table 6. Expression of ina-1 cDNA in engulfing cells rescued the ina-1 engulfment defect………………………………………………………...98 Table 7. Detection of corpse GFP circle around cell corpse using Pced-1myr::gfp ………………………………………………99 Table 8. INA-1(N)::GFP binds to apoptotic cells in a scrm-1 dependent manner…………………………………….100 Table 9. Cell corpse counts of kin-32 and other mutants…………………….101 Table 10. src-1 acts downstream of ina-1 and upstream of ced-2 during cell corpse engulfment…………………………………………………....102 Table 11. Number of apoptotic cells in rtk-x, and other engulfment-defective mutants…………………103 Appendix……………………………...….104 | |
| dc.language.iso | en | |
| dc.subject | 細胞凋亡 | zh_TW |
| dc.subject | 線蟲 | zh_TW |
| dc.subject | 插入素 | zh_TW |
| dc.subject | 受器型酪胺酸激酶 | zh_TW |
| dc.subject | 吞噬 | zh_TW |
| dc.subject | engulfment | en |
| dc.subject | C.elegans | en |
| dc.subject | integrin | en |
| dc.subject | receptor tyrosine kinase | en |
| dc.subject | apoptosis | en |
| dc.title | 線蟲插入素與受器型酪胺酸激酶促進清除死細胞 | zh_TW |
| dc.title | Engulfment of apoptotic cells in C. elegans is mediated by integrin and receptor tyrosine kinase | en |
| dc.type | Thesis | |
| dc.date.schoolyear | 98-2 | |
| dc.description.degree | 博士 | |
| dc.contributor.oralexamcommittee | 陳瑞華,廖秀娟,汪宏達,陳昌熙 | |
| dc.subject.keyword | 線蟲,插入素,受器型酪胺酸激酶,吞噬,細胞凋亡, | zh_TW |
| dc.subject.keyword | integrin,receptor tyrosine kinase,apoptosis,engulfment,C.elegans, | en |
| dc.relation.page | 104 | |
| dc.rights.note | 未授權 | |
| dc.date.accepted | 2010-07-22 | |
| dc.contributor.author-college | 生命科學院 | zh_TW |
| dc.contributor.author-dept | 分子與細胞生物學研究所 | zh_TW |
| 顯示於系所單位： | 分子與細胞生物學研究所 | |
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