細胞核中的DNA上有許多基因，而基因儲存著遺傳訊息，這些訊息像是龐大藍圖刻劃出特定蛋白才能夠進行特定機制。我們先前對於第II型AAA-ATPase (ATPase associated with diverse cellular activities)其中的TER94進行研究，發現顯性抑制(dominant negative)突變TER94K2A的表達引起果蠅感光細胞（R細胞）核增大，透過FK2免疫染色發現核內有許多泛素化蛋白(ubiquitinated proteins)累積，此外我們發現異常的核中有較多的γH2Av累積，代表DNA損傷反應（DDR） 受到阻礙而無法順利進行。這些數據共同為TER94缺陷引起的核變大與DDR損傷建立了基礎。為了進一步探索TER94調控核大小的潛在機制，我們透過基因篩選(genetic screening)發現SWM1，NTH1，DNA2，REV3L和HELQ RNAi抑制了TER94K2A誘導的核擴大， 由此可知，DDR參與了TER94調控的核大小控制。由於蛋白酶體與自噬失調可能導致泛素化蛋白積累，因此我利用RT-PCR觀察表現TER94K2A是否改變RNA的量在各個不同自噬基因，結果顯示表達TER94K2A 的時候自噬體RNA的上升。為了進一步證實這些自噬蛋白可能抑制TER94K2A所造成核變大的重要性，我發現knockdown atg4，atg7，atg8，atg10，atg12，atg14，atg18和atg101可以顯著抑制表現TER94K2A所造成的核增大。重要的是，這些自噬蛋白大多數是atg8活化所必需，atg8活化與自噬體的延伸有關。因此發現將自噬作用失調與核變大有密切的關係。我們的研究中顯示細胞核型態變化與DDR進行有相當的關聯性，而TER94功能受損將導致DDR無法進行，進一步阻礙自噬作用。這些研究讓我們更深入了解核變大的各種原因，但對於新增的核膜來源我們還不清楚，且DDR和autophagy之間的關係也尚未明瞭。

The nucleus houses genetic information to control virtually every aspect of eukaryotic cell activity. In our previous study of type II AAA-ATPase, TER94, we found that expression of a dominant-negative mutant TER94K2A causes nuclear enlargement of fly photoreceptor cells (R cell) with the accumulation of ubiquitinated protein as revealed by FK2. Furthermore, the enlarged nucleus displayed a high level of γH2Av, an incomplete DNA damage response (DDR). Together, these data established a foundation that TER94-deficit induced nuclear size expansion links to the impairment of DDR. To further explore the underlying mechanism of TER94 in regulating the nuclear size, I conducted a genetic screen and identified that SWM1, NTH1, DNA2, REV3L, and HELQ RNAi suppressed TER94K2A-induced nuclear enlargement, supporting DDR is indeed involved in TER94-regulated nuclear size control. As ubiquitinated protein accumulation may result from proteasome and/or autophagy misregulations, I performed RT-PCR to test whether the expression of specific autophagic genes was altered by TER94K2A induction. The result showed some atg genes, especially those associated with autophagosome formation, were upregulated. To further confirm the significance of those autophagic components in regulating TER94K2A-induced nuclear size change, I used the RNA interference approach to knockdown atg genes in the whole genome. Indeed, I found the knockdown of atg4, atg7, atg8, atg10, atg12, atg14, atg18, and atg101, could significantly suppress TER94K2A-induced nuclear size enlargement. Importantly, the majority of these candidates are required for atg8 activation, which associates with autophagosome elongation. This finding thus links the aberrant nuclear expansion to the misregulation of autophagy. Together, our work suggests TER94 AAA ATPase is essential to maintain nuclear integrity by sustain DDR process, and the impairment of TER94 function would lead to DDR deficit by which to impede autophagic process. This study sheds light on autophagy in regulating nuclear size.

Abstract 2
中文摘要 4
Introduction 9
Materials and Methods 13
Drosophila stocks and genetics 13
Immunohistochemistry 13
RT-PCR 14
Results 15
TER94 loss of function causes an aberrant nuclear enlargement and DDR impairment in photoreceptor cells 15
Loss of TER94 Function affects DNA damage repair 15
Knockdown other DDR-related genes reduces the enlarged nuclear caused by expressing TER94K2A 18
Knockdown p53 or mu2 inhibits autophagy regulation 20
Expressing TER94K2A or p53 could alter the expression of a subset autophagic genes 20
The knockdown of a subset ATG genes could partially suppress the nuclear enlargement caused by TER94K2A 21
Discussion 24
Figure 27
Tables 44
Reference 51

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