臺灣博碩士論文加值系統

出芽酵母菌染色質重塑複合物(RSC complex)具有利用水解三磷酸腺苷(ATP)產生能量進而改變染色質結構的能力。從個別單元突變株的研究中，已知染色質重塑複合物參與多項細胞功能，包括轉錄作用、姐妹染色體連結、去氧核醣核酸損壞修補和細胞壁完整性維持。本論文中探討的HTL1基因是一個染色質重塑複合物的非必須單元，之前的研究發現，HTL1基因剔除會造成單倍體酵母菌染色數增加，以及在致死溫度生長時出現細胞生長週期停滯在G2/M時期。為了進一步研究染色質重塑複合物的細胞功能，我們探討了HTL1基因剔除株在上述部份細胞功能以及染色質重塑複合物結構的影響。結果分為三部份，在第一部份，我們探討HTL1基因剔除株中細胞壁完整性的缺陷。結果顯示HTL1基因剔除株具有細胞壁完整性缺陷的特徵，而且此細胞壁完整性缺陷可以被等壓濃度的山梨醇(sorbitol)拯救，然而染色質重塑複合物結構上受到HTL1基因剔除後的影響並不會因此恢復。第二部份我們探討了HTL1基因剔除株有絲分裂期停滯的機轉，結果顯示HTL1基因剔除株的有絲分裂期停滯受紡錘絲檢查點(spindle checkpoint)影響，另外，細胞壁完整性缺陷也影響HTL1基因剔除株有絲分裂的進行。第三部份我們探討CHA1基因在HTL1基因剔除株的表現。CHA1是一月安基酸代謝相關基因，在絲月安酸(serine)或蘇月安酸(therine)存在時，CHA基因會大量表現，平時則受抑制，然而在染色質重塑複合物單元RSC8基因表現受抑制後，CHA1的表現抑制作用也消失。我們的實驗結果顯示，雖然結構上，HTL1基因剔除株會造成Rsc8蛋白質單元容易與染色質重塑複合物分離，但CHA1的抑制作用在HTL1基因剔除株中並沒有消失，甚至有強化的現象。以上結果顯示，HTL1基因在細胞壁完整性維持與有絲分裂週期的功能上與其他單元不同，而且HTL1基因剔除對染色質重塑複合物結構與功能的影響也似乎不一致。

The Saccharomyces cerevisiae RSC (remodel the structure of chromatin) complex is a chromatin remodeler that hydrolyzes ATP as energy to mediate the chromatin structure. RSC plays a role in the cellular function of transcriptional regulation, sister chromatids cohesion, DNA damage repair and cell wall integrity. The HTL1 (high temperature lethal) gene encodes a nonessential subunit of the RSC complex. Deletion of HTL1 causes increased ploidy at the permissive temperature, and the ∆htl1 mutant accumulates in the cell cycle at the G2/M phase at restrictive temperatures. In this thesis, we investigated the cellular function of the HTL1 gene. The results are described in three chapters. In chapter 1, we explored the defect in cell wall integrity in the ∆htl1 mutant. The HTL1 gene has been shown to genetically interact with the PKC1 pathway, the signaling pathway important for the biogenesis and stress response of yeast cell wall; however, a characteristic defect in cell wall integrity has not been explored in the ∆htl1 single mutants. We show that the ∆htl1 mutant is sensitive to agents that cause cell wall stress; in addition, various phenotypes of htl1 cells, including a defect in cell wall integrity and the mitotic arrest are rescued by the presence of osmotic stabilizer such as sorbitol. We also show that the aberrant structure of the RSC complex in the ∆htl1 mutant is not consistent with the increasing severity of the phenotypes of ∆htl1 cells as the temperature increases. In chapter 2, we explored the mechanism of mitotic arrest in the ∆htl1 mutant. The results show that the mitotic arrest in ∆htl1 cells is dependent on the spindle assembly checkpoint; in addition, the mitotic arrest in the ∆htl1 mutant is partially caused by the defect in the cell wall integrity. In chapter 3, we explored the regulation of CHA1 gene, which encodes the catabolic L-serine deaminase. A previous study showed that the transcriptional expression of CHA1 gene was de-repressed when Rsc8p and Sth1p, two major RSC components, had been depleted. However, our results show that CHA1 is persistently repressed in the ∆htl1 mutant during prolonged incubation under the repressive condition. Furthermore, the RSC complex associates with the CHA1 locus whether serine is present or not, and the association between the RSC complex and the CHA1 locus is enhanced in ∆htl1 cells. Taken together, our present findings suggest that HTL1 may play a role different from that of other RSC components in terms of cell wall integrity and the G2/M transition. The results also suggest that the defects in cell wall integrity and the G2/M transition of the ∆htl1 mutant are interconnected. Moreover, the functional and structural effects of the ∆htl1 mutant on the RSC complex do not seem to be coordinated.

中文摘要 iv
Abstract v
Introduction 1
Chapter 1. Cell wall integrity in the ∆htl1 mutant 6
Introduction 6
Results 8
1. The characteristics of cell wall defect that occur in the ∆htl1 mutant 8
2. The characteristic defect in cell wall integrity of the ∆htl1 mutant is temperature-dependent 9
3. Htl1p interacts with the Rsc1 and Rsc2 sub-complexes 9
4. Rsc8p and Rsc58p partially disassemble from the RSC complex in Δhtl1 yeast cells examined by the sucrose gradient analysis 10
5. Sorbitol does not restore the disassociation of Rsc8p from RSC complexes in the ∆htl1 mutant 11
6. Temperature and CFW sensitivities of the ∆htl1 mutant are not suppressed by over-expression of the PKC1 gene 12
7. Expression of MAP kinase SLT2 is up-regulated in ∆htl1 cells 13
8. Transcriptional change in the ∆htl1 and GAL1-HTL1 cells 14
Discussion 16
Chapter 2. Ploidy shift and cell cycle progression in the ∆htl1 mutant 19
Introduction 19
Results 21
1. Ploidy of ∆htl1 yeast haploid cells increases to 2N DNA content 21
2. Ploidy increase of the ∆htl1 mutant is temperature-dependent and is suppressed by 1 M sorbitol 21
3. The accumulation of 2N DNA content in the Htl1p-depleted cell at restrictive temperature remains prominent in the ∆mad1 background 22
4. Accumulation of phosphorylated Scc1p in ∆htl1 cells is moderated by the deletion of MAD1 24
5. Sister chromatids cohesion in ∆htl1 yeast cells 25
6. Structure of CEN3 element in the ∆htl1 mutant 27
7. Structure of CEN3 element in the GAL1-HTL1 cell 28
8. G2/M arrest of the htl1 mutant is relieved by the ∆slt2 28
Discussion 30
Chapter 3. Expression of CHA1 gene in the ∆htl1 mutant 32
Introduction 32
Results 34
1. The nucleosome pattern of CHA1 gene tends to remain in a repressed state in the ∆htl1 mutant after prolonged incubation in the presence of serine 34
2. The extent of association between the RSC complex and CHA1 gene is enhanced in the ∆htl1 mutant 35
3. The extent of chromatin-association of the RSC complex is enhanced in the ∆htl1 cell 37
Discussion 39
Materials and methods 43
References 57
List of figures…………………………………………………………………..........64
1. The osmotic stabilizer sorbitol suppresses the CFW sensitivity of the ∆htl1 mutant……………………………………………...………………………..64
2. CFW sensitivity of the ∆htl1 and ∆rsc7 mutants is temperature-dependent…………………………………………………….65
3. Htl1p is associated with both the RSC1 and RSC2 subcomplexes………66
4. Integrity of the RSC complex in the ∆htl1 and ∆rsc7 mutants…………..67
5. Integrity of the RSC complex in the ∆htl1 mutant………………………..68
6. Rsc8p is disassembled from RSC complexes in the ∆htl1 and ∆rsc7 mutants……………………………………………………………………...69
7. 1 M Sorbitol does not restore the disassociation of Rsc8p from RSC complexes in the ∆htl1 mutant……………………………………………..70
8. The CFW and temperature sensitivity of the ∆htl1 mutant are not rescued by over-expression of PKC1……………………………………………...…71
9. Expression of MAP kinase Slt2p is up-regulated in the ∆htl1 mutant…..72
10. Ploidy of the ∆htl1 mutant in the background of S288C and W303 is increased………………………………………………………………….…73
11. DNA content of different yeast strains……………………………...……74
12. Ploidy increase of the ∆htl1 mutant is temperature-dependent and is suppressed by 1 M sorbitol…………………………………………………75
13. Changes of DNA content in the ∆htl1 mutant at restrictive temperature………………………………………………………………....76
14. Changes of cell cycle in the ∆htl1 mutant at restrictive temperature….77
15. Depletion of RSC subunits as examined by Western blotting………….78
16. Changes in the DNA content on the depletion of Htl1p in ∆mad1 or ∆bub2 backgrounds…………………………………………………………80
17. Changes in the DNA content on the depletion of Htl1p in the ∆mad1 background………………………………………………………………….81
18. Changes in the DNA content on the depletion of Sth1p in ∆mad1 or ∆bub2 backgrounds…………………………………………………………82
19. Changes in the DNA content on the depletion of Rsc8p in ∆mad1 or ∆bub2 backgrounds…………………………………………………………83
20. Accumulation of hyperphosphorylated Scc1p in htl1 cells is relieved in a ∆mad1 background…………………………………………………………84
21. Cohesion of sister chromatids is prematurely separated in the ∆htl1 mutant…………………………………………………………………….....85
22. Separation of sister chromatids is delayed in the ∆htl1 mutant………..86
23. Nucleosome pattern analysis of CEN3 franking region………………...87
24. Nucleosome pattern analysis of CEN3 franking region………………...89
25. Nucleosome pattern analysis of CEN3 franking region………………...90
26. Restriction enzyme accessibility of CEN3 element……………………...91
27. Accumulation of phosphorylated Scc1p of the ∆htl1 mutant is rescued by 1M sorbitol………………………………………………………………….92
28. Mitotic arrest of the ∆htl1 mutant is rescued by the deletion of SLT2…93
29. The nucleosome pattern of CHA1 gene tends to remain in a repressed state in the ∆htl1 mutant after prolonged incubation in the presence of serine…………………………………………………………………….…...94
30. Interaction between RSC complex and CHA1 gene examined by chromatin immunoprecipitation (ChIP)……………………………….….95
31. Extent of chromatin association of the RSC complex is increased in the ∆htl1 mutant………………………………………………………………....96
32. Construction of pRS306-CHA1-HA……………………………………...97
33. Expression of Cha1p in the serine present or not……………………......98
List of tables…………………………………………………………………………99
I. List of ORFs whose transcripts are induced at least 4-fold on the depletion of Htl1p…………………………………………………………...99
II. List of ORFs whose transcripts are reduced at least 1.17-fold on the depletion of Htl1p………………………………………………………….100
III. List of ORFs whose transcripts are induced at least 2-fold in the ∆htl1………………………………………………………………………...102
IV. List of ORFs whose transcripts are reduced at least 0.75-fold in the ∆htl1………………………………………………………………………...103
V. Chromosome gain analysis of the ∆htl1 mutant………………………...104
Appendix…………………………………………...………………………………105
Figure A1. Components of the RSC complex and genetic interaction between HTL1 and RSC components……………………………………….105

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