[논문]히스톤 3 아세틸화(H3Ac)를 통한 De-Etiolated 1 (DET1)의 애기장대 생체시계 조절

송해룡

doi:10.5352/jls.2012.22.8.999

히스톤 3 아세틸화(H3Ac)를 통한 De-Etiolated 1 (DET1)의 애기장대 생체시계 조절
Regulation of Arabidopsis Circadian Clock by De-Etiolated 1 (DET1) Possibly via Histone 3 Acetylation (H3Ac) 원문보기

생명과학회지 = Journal of life science, v.22 no.8 = no.148, 2012년, pp.999 - 1008

초록
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자기 현가적(self-sustaining) 조절 장치인 생체시계는 24시간 주기의 생체리듬을 조절하며 또한 생물체로 하여금 매일 변화하는 자연환경의 외부 신호를 인지할 수 있도록 해준다. 생체시계 유전자의 발현 조절은 전사/해독의 역환류 기작을 통해 이루어진다. 애기장대 LATE ELONGATED HYPOCOTYL (LHY)와 CIRCADIAN CLOCK-ASSOCIATED 1 (CCA1)는 아침에 최고조로 발현되며 해독된 LHY and CCA1는 저녁에 최고로 발현되는 TIMING OF CAB EXPRESSION1 (TOC1)의 발현을 억제한다. TOC1단백질은 LHY와 CCA1 발현을 촉진시킴으로써 생체시계의 핵심 진자(oscillator)를 형성한다. 동물에서 생체시계의 주요 전사 인자인CLOCK은 아세틸화효소 활성 기능을 가지며, 이는 생체시계의 기능 유지에 아세틸화의 중요함을 의미한다. 하지만 애기장대 생체시계에 아세틸화를 담당하는 인자에 대한 정보는 현재 보고된 바가 없다. 본 연구에서 DET1 (De-Etiolated1)는 암조건하에서 애기장대 생체시계 관련 핵심인자 중 하나인 LHY발현을 억제하는데 필요하며 이의 억제는 H3Ac 조절을 통해 이루어짐을 증명하였다. 하지만 LHY 아세틸화를 담당하는 효소의 발굴 및 이들 효소와 DET1과의 연결을 찾는 문제는 여전히 미재로 남아있다.

Abstract ▼ AI-Helper

The circadian clock is a self-sustaining 24-hour timekeeper that allows organisms to anticipate daily-changing environmental time cues. Circadian clock genes are regulated by a transcriptional-translational feedback loop. In Arabidopsis, LATE ELONGATED HYPOCOTYL (LHY) and CIRCADIAN CLOCK-ASSOCIATED 1 (CCA1) transcripts are highly expressed in the morning. Translated LHY and CCA1 proteins repress the expression of the TIMING OF CAB EXPRESSION 1 (TOC1) transcripts, which peaks in the evening. The TOC1 protein elevates the expression of the LHY and CCA1 transcripts, forming a negative feedback loop that is believed to constitute the oscillatory mechanism of the clock. In mammals, the transcription factor protein CLOCK, which is a central component of the circadian clock, was reported to have an intrinsic histone acetyltransferase (HAT) activity, suggesting that histone acetylation is important for core clock mechanisms. However, little is known about the components necessary for the histone acetylation of the Arabidopsis clock-related genes. Here, I report that DET1 (De-Etiolated1) functions as a negative regulator of a key component of the Arabidopsis circadian clock gene LHY in constant dark phases (DD) and is required for the down-regulation of LHY expression through the acetylation of histone 3 (H3Ac). However, the HATs directly responsible for the acetylation of H3 within LHY chromatin need to be identified, and a link connecting the HATs and DET1 protein is still absent.

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가설 설정

3B). To test whether LHY:luc reflects well the level of LHY mRNAs. WT and det1-1 seedlings were grown in 12L12D, released to DD and then harvested at ZT24, 48, 60 and 72 for RT-PCR experiments.

제안 방법

WT and det1-1 seedlings were grown in 12L12D, and harvested at ZT12 and ZT20 in which the level of LHY transcripts between WT and det1-1 was greatly distinguishable. ChIP assays were performed with antibody specific to H3Ac and the enrichments of LHY, CCA1 and TOC1 loci were measured by using diverse sets of primers (Fig. 2A). H3Ac levels in LP1 and LX1 but not LX4 and L3U regions of LHY were higher at ZT20 than at ZT12 in both WT and det1-1 (Fig.
(B) Expression of CCA1 and TOC1 transcripts in Col and det1-1 seedlings grown in 12L12D. Col and det1-1 seedlings were grown under 12L12D for 7 days, harvested at every 2 hours for one day and used for RT-PCR analyses. UBQ was included as an expression control.
Pre-sprays were repeated every 8 hours over a 24-hour period to remove luciferase activity accumu- lated prior to the first luciferin treatment [25]. Luminescence was imaged for 30 minutes every 2 hours for 3 days using photon-counting cameras (Hamamatsu). The luminescence levels were quantified from the images using Metamorph™ software (Universal Imaging Corp.
Seeds were sown on Murashige and Skoog (MS)-agar medium supplemented with 1% sucrose, and refrigerated for at least 3-4 days before growing at 22℃ under 70-100 μE m^-2 s^-1 cool white fluorescent light. Photoperiod durations (12L12D, LL or DD) were programmed according to the purpose of the experiments.
Col and det1-1 seedlings were grown under 12L12D conditions for 7 days, transferred to DD at ZT 0. Plants were harvested at ZT 24, ZT 48, ZT 60 and ZT 72, respectively, and used for RT-PCR analysis. UBQ was included as an expression control.
Furthermore, other studies suggested that DET1 might recruit HATs through binding to DDB1 and cause chromatin remodeling[3,37]. To test if the rhythmic oscillation of LHY, CCA1, and TOC1 correlates with changes in chromatin structure, ChIP assays were performed. WT and det1-1 seedlings were grown in 12L12D, and harvested at ZT12 and ZT20 in which the level of LHY transcripts between WT and det1-1 was greatly distinguishable.
To test the effect of det1 mutation on LHY expression, the LHY:luc reporter gene was introduced into the det1-1 mutant by genetic crossing. Expression of homozygous LHY:luc was assayed in wild-type (WT) and det1-1 plants under 12L12D (12 hours of light/12 hours of dark phases).
To test whether LHY:luc reflects well the level of LHY mRNAs. WT and det1-1 seedlings were grown in 12L12D, released to DD and then harvested at ZT24, 48, 60 and 72 for RT-PCR experiments. Like LHY:luc results (Fig.

이론/모형

The comparative ΔΔCT method was used to evaluate the relative quantities of each amplified product in the samples.

성능/효과

2B and 2E). Taken together, these results indicate that the transcription activity of all Arabidopsis central clock genes, LHY, CCA1, and TOC1, in 12L12D is positively correlated with the levels of H3Ac.

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