桿狀病毒早期基因對內源及外源啟動子表現之影響

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研究生：	吳佩穎 Wu, Pei-Yin
論文名稱：	桿狀病毒早期基因對內源及外源啟動子表現之影響 Functional analyses of baculovirus early genes on the expression of exogenous and endogenous promoters
指導教授：	趙裕展 Chao, Yu-Chan 林俊成 Lin, Chun-Cheng
口試委員:
學位類別：	博士 Doctor
系所名稱：	理學院 - 化學系 Department of Chemistry
論文出版年：	2010
畢業學年度：	98
語文別：	英文
論文頁數：	127
中文關鍵詞：	桿狀病毒、polyhedrin 上游序列、啟動子活化、增強子、早期基因-1 、晚期表現因子-2
外文關鍵詞：	Autographa californica polyhedrovirus, polyhedrin upstream sequence, enhancer, promoter stimulation, immediate-early 1, late expression factor 2
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Baculovirus viral DNA replication and transcriptional regulation are essential components to the life cycle of the virus and also important issues for expression of the engineered protein using this useful viral tool. The polyhedrin upstream sequence (pu) has been previously shown to function as an enhancer in an infection dependent manner. It is also the second known enhancer identified in baculovirus in addition to the homologous regions (hrs). Differing from hrs, the enhancer’s function of pu requires the presence of viral infection, suggesting that virus-encoded factors are required for pu’s enhancer function. To identify these factor(s), a cosmid library of Autographa californica multiple nucleopolyhedrovirus (AcMNPV) genomic DNA was constructed. Sequential fragmentation/analysis reveals 3 virus-encoded factors affecting pu’s enhancer function. These factors include IE1, IE2, and PE38 with IE1 being essential and IE2/PE38 being stimulatory. Northern blot analysis result further showed that pu and IE1 function together to stimulate the transcription activity of target promoter, and pu is therefore defined as a transcriptional enhancer.
Of the three orfs that constitute pu, orf6 encodes the late expression factor 2 (lef-2) of AcMNPV and is the best studied one. It is one of the 20 late expression factors which are essential for baculoviral late gene expression. It’s been implicated to involve in both origin-dependent DNA replication and late gene expression. To better study the role of lef-2 in the life cycle of the virus, a lef-2 deletion bacmid was constructed and analyzed. The results show that lef-2 is indeed required for efficient viral DNA replication and late/very late gene expression. However, low-level of viral DNA replication could be detected in the absence of lef-2, suggesting that the function of this gene may be stimulatory rather than essential, as previously suggested, for viral DNA replication. The temporal/spatial distribution of LEF-2 in the infected cells was also analyzed. In addition, it was found that LEF-2 was associated with the nucleocapsids of both budded viruses and occluded-derived viruses, suggesting that its function may be required immediately after viral entry into host cells.
In addition to arthropods, baculovirus can also enter a variety of mammalian cells with varied efficiencies and has therefore been proposed as an alternative gene delivery tool. To optimize baculovirus transduced gene expression in target mammalian cells, we found that the known baculovirus trans-activator, IE1, could stimulate the expression of cytomegalovirus immediate early (CMVie) promoter. This IE1-mediated stimulation of CMVie promoter activity can be further enhanced by the presence of hr, the known baculovirus enhancer for better gene expression.

TABLE CONTENT    V
FIGURE CONTENT    VI
TABLE OF ABBREVIATION    VIII
ABSTRACT    IX
中文摘要    XI
CHAPTER 1: INTRODUCTION    1
CHAPTER 2: LITERATURE REVIEW    3
SECTION 1: Baculovirus Classification    3
SECTION 2: Morphology and Structures    3
SECTION 3: Infection cycle of baculovirus    4
SECTION 4: Baculovirus genome replication    5
SECTION 5: Baculovirus late and very late gene expression    6
SECTION 6: Baculovirus vector expression system    7
SECTION 7: Generation of recombinant baculovirus by Bac-to-BacTM   system    8
SECTION 8: Baculovirus as an alternative gene delivery tool in mammalian cells    9
CHAPTER 3: IDENTIFICATION OF ACMNPV-ENCODED FACTORS                                     REQUIRED FOR THE ENHANCER FUNCTION OF POLYHEDRIN UPSTREAM (PU) SEQUENCE    10
SECTION 1: ABSTRACT    10
SECTION 2: INTRODUCTION    11
SECTION 3: MATERIALS AND METHODS    13
3.1 Cell culture and virus    13
3.2 Preparation of viral DNA    13
3.3 Construction of phr5, pLP, and reporter plasmids    14
3.4 Replication assay    14
3.5 Construction of AcMNPV Cosmid library    15
3.6 Cloning of sub-fragments of Cosmid 8    15
3.7 Construction of frame-shift mutation clones    16
3.8 Transient transfection assay    16
3.9 Luciferase activity assay    17
3.10 Northern blotting    17
SECTION 4: RESULTS    19
4.1 pu-dependent transcriptional activation is not associated with DNA replication    19
4.2 Identification of AcMNPV sub-genomic region(s) that support pu-dependent promoter activation    19
4.3 ie1, ie2, and pe38 are required for the pu-dependent activation of CMVm promoter    21
4.4 IE1 acts through pu to stimulate transcriptional activity of CMVm promoter    23
4.5 pu sequence per se is responsible for pu-mediated promoter activation    24
4.6 pu enhances baculovirus p143 promoter activity in the presence of IE1    24
4.7 hr stimulates pu-mediated activation of CMVm promoter    25
SECTION 5: DISCUSSION    27
CHAPTER 4: BACULOVIRUS LATE EXPRESSION FACTOR 2 (LEF-2) IS ESSENTIAL FOR PRODUCTIVE VIRAL DNA REPLICATION    46
SECTION 1: ABSTRACT    46
SECTION 2:  INTRODUCTION    47
SECTION 3: MATERIALS AND METHODS    50
3.1 Cell Culture and viruses    50
3.2 Construction of lef-2 knockout and repair bacmids    50
3.3 Construction of p143 knockout bacmid    52
3.4 Bacmid DNA purification and transfection    53
3.5 Growth curve assay    53
3.6 Dpn-I replication assay    54
3.7 Northern Blot Analysis    54
3.8 Cellular localization of LEF-2 in nucleus and cytoplasm    55
3.9 Immunofluorescence Microscopy    55
3.10 Purification of virus particles from BV and ODV    57
3.11 Separation of nucleocapsid and envelope proteins of BV and ODV    58
3.12 Western blot analysis    58
SECTION 4: RESULTS    60
4.1 Generation of lef-2 knockout and repair AcMNPV bacmids    60
4.2 Analysis of wt, lef-2KO, and repair bacmids in Sf-21 cells    60
4.3 lef-2 deletion abolished budded virus production    61
4.4 DpnI-sensitivity DNA replication assay    63
4.5 Analysis of baculovirus early, late, and very late gene expressions in lef-2 knockout virus    64
4.6 Spatial Distribution of LEF-2    65
4.7 Localization of LEF-2 at the replication foci    66
4.8 Association of LEF-2 with nucleocapsids of both BVs and ODVs    67
SECTION 5: DISCUSSION    69
CHAPTER 5: BACULOVIRUS TRANSACTIVATOR IE1 AND ENHANCER HR COOPERATIVELY STIMULATE HUMAN CMV PROMOTER IN MAMMALIAN CELLS    86
SECTION 1: ABSTRACT    86
SECTION 2: INTRODUCTION    87
SECTION 3: MATERIAL AND METHODS    89
3.1 Plasmid Construction    89
3.2 Cell Culture    90
3.3 Transfection    90
3.4 Generation of recombinant virus    91
3.5 Mammalian Cell Culture and Virus Transduction Experiments    91
3.6 Luciferase Activity Assay    91
SECTION 4: RESULTS    93
4.1 Baculovirus IE1 can trans-activate CMV promoter in mammalian cells    93
4.2 Cis-linked baculovirus hr1 enhanced CMV promoter activation mediated by IE1    93
4.3 Baculovirus homologous region could augment IE1-mediated transactivation in trans    94
4.4 CMV enhancer region could be functionally substituted by hr1 for IE1 trans-activation in mammalian cells    95
4.5 Activation of CMV promoter by IE1 can be achieved through baculovirus transduction    97
SECTION 5: DISCUSSION    99
CHAPTER 6    110
CONCLUSIONS    110
REFERENCES    112
APPENDIX    126

                                

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