阿拉伯芥自噬作用相關蛋白ATG8f與ATG8h的功能分析

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研究生：	林莉晏 Lin, Li-Yen
論文名稱：	阿拉伯芥自噬作用相關蛋白ATG8f與ATG8h的功能分析 Functional analysis of the Arabidopsis autophagy-related proteins ATG8f and ATG8h
指導教授：	劉姿吟 Liu, Tzu-Yin
口試委員:	汪宏達 Wang, Horng-Dar 邱子珍 Chiou, Tzyy-Jen
學位類別：	碩士 Master
系所名稱：	生命科學暨醫學院 - 生物資訊與結構生物研究所 Institute of Bioinformatics and Structural Biology
論文出版年：	2021
畢業學年度：	109
語文別：	英文
論文頁數：	67
中文關鍵詞：	阿拉伯芥、自噬作用、自噬作用相關蛋白、缺磷反應、根部發育
外文關鍵詞：	autophagy-related proteins, ATG8f, ATG8h, phosphate starvation response, root development
相關次數：	點閱：3 下載：0
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植物自噬作用是一個透過液泡進行細胞質分解的過程並且維持細胞穩定狀態。自噬作用相關蛋白ATG8在自噬小體(autophagosome)的形成過程中扮演中心角色，自噬小體包裹細胞質中的物質並且送入液泡中分解。阿拉伯芥(Arabidopsis thaliana)的ATG8基因家族有九個蛋白異構體(isoform) (AtATG8a–AtATG8i)。對於缺乏磷酸時如何提升植物自噬作用以及AtATG8蛋白如何召集特定物質到液泡分解尚未清楚。我們發現在缺磷時，ATG8f與ATG8h在地上部與根部的表現會被明顯地調升。這個磷酸缺乏時的調升作用在phosphate starvation response 1 (phr1)的突變株中會部分地被抑制，PHR1負責編碼轉錄活化子(transcription activator)以增加缺磷反應基因(Pi starvation responsive genes)的轉錄。我們假設ATG8f與ATG8h是PHR1調控磷酸平衡路徑的下游目標。在此研究中我們發現PHR1並未直接提升ATG8f與ATG8h的基因表現。PHR1活化ATG8f與ATG8h的基因表現可能需要仰賴其他蛋白一同參與，或是藉由PHR1引導第二波的轉錄活化(transcriptional activation)。此外，我們得到atg8f, atg8h, atg8f/atg8h T-DNA插入突變株。在缺磷時，atg8f/atg8h根部組織的自噬通量(autophagic flux)比野生型(wild-type)來得少，意味著ATG8f與ATG8h會調控在缺磷下的自噬作用活性。我們也觀察到無論是否缺磷，atg8f/atg8h的側根數量比野生型少。ATG8f與ATG8h大量表現在主根的維管束組織，尤其是側根即將長出的地方。這些結果提示著ATG8f與ATG8h有潛在調節側根發育的功能。

Plant autophagy is a vacuolar-mediated process of cytoplasmic degradation and maintains cellular homeostasis. Autophagy-related 8 (ATG8) protein plays a central role in the formation of autophagosomes which enclose and deliver the cytoplasmic components into vacuoles for degradation. The Arabidopsis thaliana ATG8 gene family comprises nine isoforms (AtATG8a–AtATG8i). It remains unclear how phosphate (Pi) limitation regulates the induction of plant autophagy and how AtATG8s proteins recruit specific cargos for vacuolar degradation under Pi starvation. We revealed that AtATG8f and AtATG8h are upregulated in the shoot and root by Pi starvation. Such upregulation by Pi starvation can be partially suppressed in the shoot and root of phosphate starvation response 1 (phr1) mutant encoding a transcriptional activator of Pi starvation responsive genes. We hypothesized that AtATG8f and AtATG8h may be the downstream targets of the PHR1-dependent regulatory pathway of Pi homeostasis. In this study, we showed that PHR1 seemed not to directly transactivate the expression of AtATG8f and AtATG8h. PHR1-dependent Pi starvation-induction of AtATG8f and AtATG8h may rely on the cooperation of PHR1 and other components or a second wave of transcriptional activation triggered by PHR1. In addition, we obtained atg8f, atg8h, atg8f/atg8h T-DNA mutants and demonstrated that the autophagic flux was reduced in the Pi-starved root of atg8f/atg8h compared to that of wild-type (WT), indicating that AtATG8f and AtATG8h play a role in regulating autophagic activities under Pi limitation. Moreover, the atg8f/atg8h mutant has fewer lateral root number than WT does regardless of Pi supply. Spatial expression patterns of AtATG8f and AtATG8h are expressed in the vascular tissues of primary roots and strongly at the sites where lateral roots initiate to emerge. These results suggested that AtATG8f and AtATG8h have a potential role in modulating lateral root development.

摘要    i
Abstract    ii
Acknowledgement    iii
Table of contents    iv
List of abbreviations    viii
Chapter 1. Introduction    1
1.1 An overview of autophagy    1
1.1.1 The role of ATG8 protein in the formation of autophagosome    2
1.1.2 Autophagic flux as determined by ATG8 accumulation    3
1.1.3 Arabidopsis ATG8 gene family    5
1.2 Phosphorus for plant growth    6
1.3 The maintenance of Pi homeostasis in plants    6
1.4 Pi starvation-induced autophagy in the root apical meristem    8
1.5 Aims of the study    9
Chapter 2. Materials and methods    10
2.1 Plant materials and growth conditions    10
2.1.1 Plant material    10
2.1.2 Sterilization of Arabidopsis seeds    10
2.1.3 Plant medium    10
2.1.4 Plant soil matrix    11
2.2 Plasmid construction    11
2.3 Bacterial growth medium    12
2.4 Bacterial host cell strains and plasmid transformation    12
2.4.1 Bacterial host cell strains    12
2.4.2 Plasmid transformation into E.coli    13
2.4.3 Plasmid transformation into agrobacteria    13
2.5 Protoplast isolation and transfection assay    13
2.5.1 Protoplast isolation    13
2.5.2 Protoplast transfection    15
2.6 Transient dual-luciferase assay    15
2.7 Genomic DNA extraction    16
2.8 RNA extraction    16
2.9 Validation of T-DNA insertion mutants    16
2.9.1 PCR genotyping of homozygous T-DNA insertion lines    16
2.9.2 RT-PCR analysis of homozygous T-DNA insertion knockout lines    17
2.10 Real-time quantitative reverse transcription PCR (qRT-PCR) analysis    17
2.11 Pi level measurement    18
2.12 Protein assay    18
2.12.1 Total protein extraction    18
2.12.2 Protein concentration measurement    19
2.12.3 Immunoblotting analysis    20
2.13 Analysis of root phenotypes    20
2.14 Agrobacterium-mediated transformation of Arabidopsis    21
2.15 Transgenic plant selection    21
2.16 Microscopy analysis of GFP-reporter lines    21
2.17 Histochemical staining of β-glucuronidase (GUS)    22
Chapter 3. Results    23
3.1 AtATG8f and AtATG8h promoter activities are not directly activated by PHR1    23
3.2 Characterization of atg8f and atg8h T-DNA insertion knockout mutants    24
3.3 The transcript levels of the other AtATG8 genes are not increased to compensate for the loss of AtATG8f and AtATG8h    25
3.4 The Pi level of atg8f/atg8h is comparable to that of WT    25
3.5 The autophagic flux decreases in the root of Pi-starved atg8f/atg8h    26
3.6 Loss of AtATG8f and AtATG8h suppresses the lateral root formation independent of Pi status    26
3.7 Analysis of AtATG8f and AtATG8h promoter activities under Pi- and N-deficient conditions    27
3.8 Analysis of AtATG8f and AtATG8h promoter activities at the tissue level    29
Chapter 4. Discussion    31
4.1 PHR1 acts upstream of AtATG8f and AtATG8h    31
4.2 Partially functional redundancy of AtATG8f and AtATG8h    32
4.3 Role of AtATG8f and AtATG8h in the maintenance of Pi homeostasis    33
4.4 Role of AtATG8f and AtATG8h in the lateral root development    33
Chapter 5. Figures    35
Figure 1. Transient reporter assay in Arabidopsis protoplast co-expressing PHR1 and AtATG8f/AtATG8h promoter-fused dual-luciferase reporter    35
Figure 2. Characterization of atg8f and atg8h T-DNA insertion mutants    37
Figure 3. Pi levels of atg8f, atg8h, and atg8f/atg8h mutants    38
Figure 4. The ATG8s protein expression of the Pi-starved roots of WT and atg8f/atg8h    39
Figure 5. Phenotypical analysis of root of WT, atg7-3, and atg8f/atg8h    41
Figure 6. Schematic design of AtATG8f and AtATG8h promoters-fused GFP and GUS reporter constructs    42
Figure 7. GFP expression patterns of ProATG8f::GFP transgenic lines    43
Figure 8. GFP expression patterns of ProATG8h::GFP transgenic lines    44
Figure 9. qRT-PCR analysis of GFP and endogenous AtATG8f and AtATG8h expression in ProATG8f::GFP and ProATG8h::GFP transgenic lines    45
Figure 10. Histochemical GUS staining of ProATG8f::GUS transgenic lines    46
Figure 11. Histochemical GUS staining of ProATG8h::GUS transgenic lines    47
Chapter 6. Supplemental data    48
Figure S1. qRT-PCR analysis of AtATGs gene expression in WT seedlings in response to Pi and N starvation    49
Figure S2. PHR1-dependent Pi starvation-induction of ATG8f and ATG8h    50
Figure S3. qRT-PCR analysis of AtATG8s gene expression in the root of WT and atg8f/atg8h    51
Table S1. T-DNA insertion lines for AtATG8f and AtATG8h    52
Table S2. List of primer sequences used for PCR-based gene cloning    53
Table S3. List of constructs used for generating transgenic lines    54
Table S4. Antibiotics used in this study    55
Table S5. List of primer sequences used for PCR genotyping and RT-PCR analyses    56
Table S6. List of primer sequences used for qRT-PCR analysis    57
Table S7. Generation and selection of GUS- and GFP-reporter transgenic lines for ATG8f and ATG8h    58
Table S8. The predicted P1BS element and AtPHR1 binding matrix    59
References    60

                                

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