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研究生:桑達
研究生(外文):Kalaipandian Sundaravelpandian
論文名稱:錳及活性氧對阿拉伯芥根毛分化之影響
論文名稱(外文):A study on the effects of manganese and reactive oxygen species on root hair differentiation in Arabidopsis
指導教授:施臥虎
指導教授(外文):Wolfgang Schmidt
口試委員:陳榮芳麥卓琍鄭萬興邱子珍
口試委員(外文):Long-Fang ChenMarjori MatzkeWan-Hsing ChengTzyy-Jen Chiou
口試日期:2012-12-28
學位類別:博士
校院名稱:國立中興大學
系所名稱:生物科技學研究所
學門:生命科學學門
學類:生物科技學類
論文種類:學術論文
論文出版年:2012
畢業學年度:101
語文別:英文
論文頁數:96
中文關鍵詞:根毛活性氧分子轉錄因子
外文關鍵詞:root hairsmanganesereactive oxygen speciestranscriptional Mediators
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Root hairs, long tubular-shaped outgrowths from root epidermal cells, are an excellent model to study cell specification and differentiation. Their wide distribution among all major groups of vascular plants gives evidence for a long evolutionary history and their importance to the success of diverse plant groups in the adaption to changing environmental conditions. Root hairs increase the root surface area to aid plants in nutrient acquisition, anchorage, and interactions with microbes. Understanding of root hair differentiation might help to uncover the nutrient transport activities in the root. In order to gain an insight into root hair differentiation mechanism, we aimed to study the effects of manganese (Mn) and reactive oxygen species (ROS) on root hair formation.
Manganese is the second most prevalent transition metal in the Earth’s crust but its availability is often limited due to rapid oxidation and low mobility of the oxidized forms. Acclimation to low Mn availability was studied in Arabidopsis seedlings subjected to Mn deficiency. As reported here, Mn deficiency caused a thorough change in the arrangement and characteristics of the root epidermal cells. A proportion of the extra hairs formed upon Mn deficiency were located in atrichoblast positions, indicative of a post-embryonic reprogramming of the cell fate acquired during embryogenesis. When plants were grown under a light intensity of >50 μmol m-2 s-1 in the presence of manganese root hair elongation was substantially inhibited, whereas Mn-deficient seedlings displayed stimulated root hair development. GeneChip analysis revealed several candidate genes with potential roles in the reprogramming of rhizodermal cells. None of the genes that function in epidermal cell fate specification were affected by Mn deficiency, indicating that the patterning mechanism which controls the differentiation of rhizodermal cells during embryogenesis have been bypassed under Mn-deficient conditions. This assumption is supported by the partial rescue of the hairless cpc mutant by Mn deficiency. Inductively coupled plasma-optical emission spectroscopy (ICP-OES) analysis revealed that, besides the anticipated reduction in Mn concentration, Mn deficiency caused an increase in iron concentration. This increase was associated with a decreased transcript level of the iron transporter IRT1, indicative of a more efficient transport of iron in the absence of Mn. To identify genes involved in Mn deficiency altered root hair differentiation, we performed both forward and reverse genetic screening strategies. Two mutants were identified through forward genetic screening namely mai1 and mai2 (manganese deficiency root hair defective). The mai1 mutant showed Mn specific root hair defective phenotype and a T-DNA insertion found at the MED6, a subunit of transcriptional Mediator complex in the mai1 genome. However, transgenic plant studies suggest that the MED6 gene was not responsible for the mai1 mutant phenotype. As the mai2 mutant plants had several T-DNA insertions, it was backcrossed for two generations with wild-type (Col-0). After backcross, mai2 mutant plants retained the root hair defective phenotype without any T-DNA insertions. Hence, we employed a map-based cloning and identified a mutation in the upper arm of chromosome 1 which was flanking between CER450618 and CER461064 InDel markers about ~450 Kb.
Although we did not provide evidence to show that the MED6 gene was responsible for the mai1 mutant phenotype, we acquired an importance of transcriptional Mediator complex in growth and development of eukaryotes. To elucidate a possible role of transcriptional Mediators in root hair formation, we analyzed the root hair phenotype of T-DNA insertion lines that harbor homozygous mutations in genes encoding Mediator subunits. Three mutant lines of two Mediator subunits, MED8 (SALK_092406) and MED25 (also called PFT1, SALK_129555 and SALK_059316) showed root hair defective phenotype. Genetic evidence indicates that the Mediator subunits PFT1/MED25 and MED8 are critical for root hair differentiation, but act via separate mechanisms. Transcriptional profiling of pft1 roots revealed that PFT1 activates a subset of H2O2-producing class III peroxidases. pft1 mutants showed perturbed hydrogen peroxide and superoxide distribution, suggesting that PFT1 is essential to maintain redox homeostasis in the root. Chemical treatments rescued the pft1mutant phenotype, indicating that correct ROS distribution is an essential prerequisite for root hair differentiation. In addition, PFT1 positively regulates cell wall remodeling genes that are essential for root hair formation. Our results demonstrate that PFT1 maintains ROS distribution that, in turn, controls root hair differentiation. Thus, our findings revealed a novel mechanism in which the Mediator controls ROS homeostasis by regulating the transcriptional machinery.
Abstract…………………………………………………………………………………i
List of figures………………………………………………………………………iv
List of tables…………………………………………………………………………vi
List of appendices………………………………………………………………vii
Part 1. Manganese deficiency alters the patterning and development of root hairs in Arabidopsis………1
Abstract…………………………………………………………………………………2
Introduction…………………………………………………………………………3
Materials and methods………………………………………………………6
Results…………………………………………………………………………………10
Discussion……………………………………………………………………………16
Part 2. PFT1, a transcriptional Mediator complex subunit, controls root hair differentiation through ROS distribution in Arabidopsis…………………………………………………41
Abstract……………………………………………………………………………42
Introduction……………………………………………………………………………43
Materials and methods……………………………………………………46
Results…………………………………………………………………………………49
Discussion…………………………………………………………………………53
Part 3. Conclusion and future prospects…………………………………………75
References…………………………………………………………………………78
Appendices…………………………………………………………………………87
Curriculum vitae…………………………………………………………………95
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