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    ์• ๊ธฐ์žฅ๋Œ€์—์„œ DEMETER์™€ ์ƒํ˜ธ์ž‘์šฉํ•˜๋Š” ์ธ์ž์˜ ์„ ๋ณ„

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    ํ•™์œ„๋…ผ๋ฌธ(์„์‚ฌ)--์„œ์šธ๋Œ€ํ•™๊ต ๋Œ€ํ•™์› :์ž์—ฐ๊ณผํ•™๋Œ€ํ•™ ์ƒ๋ช…๊ณผํ•™๋ถ€,2019. 8. ์ตœ์—ฐํฌ.DNA methylation and demethylation plays significant roles in regulating gene expression on variable stress reactions and TEs silencing. A DEMETER (DME), an active DNA demethylase in plants, is expressed in the central cell of the female gametophyte in Arabidopsis required for seed development and has glycosylase activity that can actively remove 5-mC that is replaced by cytosine via base excision pathway. DME induces maternal allele expression of the imprinted MEDEA (MEA) polycomb gene and the DNA glycosylase activity of DME leads to the DNA demethylation on its targets. In plants, there is no massive methylation reprogramming in embryo like mammal but instead, the companion cells whose DNA contents are not inherited to the next generation go through global demethylation and the hypomethylated state is maintained in late endosperm stage in which DME is no longer expressed. Despite all these distinct significances, little is known about the interactors that associate with DME. Here, to identify the interacting partners of DME, I used Bimolecular Fluorescence Complementation (BiFC). From 83 genes that have been confirmed by the Yeast Two-Hybrid system that interact with DME, I primarily chose 18 candidates to test interaction. While examining these 18 candidates by BiFC, AT5G37930, AT5G60980, AT1G70620, AT5G23090 and the C terminal part of AT1G20960 showed fluorescent signals when it was co-transfected with DME in Arabidopsis Col-0 protoplasts. These interactors contain either E3-ubiquitin ligase activity, RNA binding motif, homologous feature of transcription factor which is related to histone acetylation, or are related to RNA splicing. To further understand its relation with DME in plants one candidate, At1g20960 that is related to RNA splicing, was chosen and its mutant was crossed with dme mutant for phenotyping analyses. dme mutant allele transmission, segregation and seed abortion ratio shown in dme single mutant were not changed in double mutants. Therefore, by doing further experiments, using different candidates, this study would give some specific and clear perspectives and contribute to widen the knowledge by identifying a novel protein involved in the DME demethylation pathway in plants.DNA์˜ ๋ฉ”ํ‹ธํ™”์™€ ๋””๋ฉ”ํ‹ธํ™”๋Š” ์œ ์ „์ž์˜ ๋ฐœํ˜„์„ ์กฐ์ ˆํ•˜์—ฌ ์ƒ๋ฌผ์ฒด๊ฐ€ ์ง๋ฉดํ•˜๋Š” ๋ณ€ํ™”ํ•˜๋Š” ์ฃผ๋ณ€ ํ™˜๊ฒฝ๊ณผ ๋‹ค์–‘ํ•œ ์ŠคํŠธ๋ ˆ์Šค ์ƒํ™ฉ์— ๋Œ€ํ•œ ์ ์ ˆํ•œ ๋ฐ˜์‘์„ ๊ฐ€๋Šฅํ•˜๊ฒŒ ํ•˜๋ฉฐ ์ „์ด ์ธ์ž์˜ ๋ฐœํ˜„์„ ์–ต์ œํ•˜๋Š” ๋“ฑ์˜ ์—ญํ• ์„ ํ•˜๋ฏ€๋กœ ๋งค์šฐ ์ค‘์š”ํ•˜๋‹ค. DEMETER(DME)๋Š” ์‹๋ฌผ์—์„œ ์ž‘์šฉํ•˜๋Š” DNA ๊ธ€๋ฆฌ์ฝ”์‹คํ™” ํšจ์†Œ๋กœ ์• ๊ธฐ์žฅ๋Œ€ ์•”๋ฐฐ์šฐ์ฒด์˜ ์ค‘์‹ฌ์„ธํฌ์—์„œ ๋ฐœํ˜„ํ•˜๋ฉฐ ์ข…์ž์˜ ๋ฐœ๋‹ฌ์— ํ•„์ˆ˜์ ์ด๋‹ค. DME๋Š” ์—ผ๊ธฐ์ ˆ์ œํšŒ๋ณต๋ฐฉ์‹์„ ํ†ตํ•ด 5-mC๋ฅผ ์ง์ ‘ ์ œ๊ฑฐํ•˜๊ณ  ์‹œํ† ์‹ ์œผ๋กœ ์น˜ํ™˜ํ•˜๋Š” ๋ฐฉ์‹์˜ ๋””๋ฉ”ํ‹ธํ™”๋ฅผ ๋งค๊ฐœํ•œ๋‹ค. ์‹๋ฌผ์˜ ๋ฐฐ์—์„œ๋Š” ํฌ์œ ๋™๋ฌผ๊ณผ ๊ฐ™์€ ๋Œ€๊ทœ๋ชจ์˜ ๋ฉ”ํ‹ธ๋ ˆ์ด์…˜ ๋ฆฌํ”„๋กœ๊ทธ๋ž˜๋ฐ์ด ์ผ์–ด๋‚˜์ง€ ์•Š๋Š” ๋Œ€์‹  ๋‹ค์Œ ์„ธ๋Œ€๋กœ ์œ ์ „ ์ •๋ณด๊ฐ€ ์ „๋‹ฌ๋˜์ง€ ์•Š๋Š” ์ƒ์‹์„ธํฌ๋‚˜ ๋ฐฐ์˜ ์ฃผ๋ณ€ ์„ธํฌ์—์„œ ์ „๋ฐ˜์ ์ธ ๋””๋ฉ”ํ‹ธ๋ ˆ์ด์…˜์ด ์ผ์–ด๋‚˜๊ณ  ์ด๋Ÿฌํ•œ ํ•˜์ดํฌ๋ฉ”ํ‹ธ๋ ˆ์ด์…˜ ์ƒํƒœ๊ฐ€ DME๊ฐ€ ๋” ์ด์ƒ ๋ฐœํ˜„ํ•˜์ง€ ์•Š๋Š” ํ›„๊ธฐ ๋ฐฐ์ฃผ๊นŒ์ง€ ์œ ์ง€๋œ๋‹ค. ์ด๋Ÿฌํ•œ ์ค‘์š”์„ฑ์—๋„ ๋ถˆ๊ตฌํ•˜๊ณ  DME์˜ ๋””๋ฉ”ํ‹ธ๋ ˆ์ด์…˜ ๊ณผ์ •์—์„œ DME์™€ ์ƒํ˜ธ์ž‘์šฉํ•˜๋Š” ์ธ์ž๋“ค์— ๋Œ€ํ•œ ์—ฐ๊ตฌ๋Š” ๋งŽ์ด ์•Œ๋ ค์ง„ ๋ฐ”๊ฐ€ ์—†๋‹ค. ๋ณธ ์—ฐ๊ตฌ์—์„œ๋Š” DME์™€ ์ƒํ˜ธ์ž‘์šฉํ•˜๋Š” ์ธ์ž๋“ค์„ ๋ฐํžˆ๊ธฐ ์œ„ํ•˜์—ฌ ์ด๋ถ„์žํ˜•๊ด‘์ƒ๋ณด๊ธฐ๋ฒ•(Bimolecular Fluorescence Complementation)์„ ์ด์šฉํ•˜์˜€๋‹ค. ํšจ๋ชจ์ด์ค‘์žก์ข…ํ™”(Yeast Two-Hybrid)๋ฅผ ์ด์šฉํ•œ ์„ ํ–‰์—ฐ๊ตฌ๋ฅผ ํ†ตํ•ด ๋ฐํ˜€์ง„ DME์™€ ์ƒํ˜ธ์ž‘์šฉํ•˜๋Š” 83๊ฐœ์˜ ์œ ์ „์ž ์ค‘ 18๊ฐœ์˜ ํ›„๋ณด ์œ ์ „์ž๋ฅผ ์šฐ์„ ์ ์œผ๋กœ ์„ ์ •ํ•˜์˜€๊ณ  ๊ทธ ์ค‘ AT5G37930, AT5G60980, AT1G70620, AT5G23090, ๊ทธ๋ฆฌ๊ณ  AT1G20960์˜ C ๋ง๋‹จ๋ถ€๊ฐ€ ์• ๊ธฐ์žฅ๋Œ€ ์•ผ์ƒ์ข…(Col-0)์˜ ์›ํ˜•์งˆ์ฒด์— DME์™€ ํ•จ๊ป˜ ํŠธ๋žœ์ŠคํŽ™์…˜๋˜์—ˆ์„ ๋•Œ ํ˜•๊ด‘ ์‹ ํ˜ธ๋ฅผ ๋ณด์ด๋Š” ๊ฒƒ์„ ํ™•์ธํ•  ์ˆ˜ ์žˆ์—ˆ๋‹ค. ์ด ์œ ์ „์ž๋“ค์€ E3 ์œ ๋น„ํ€ดํ‹ด ๋ผ์ด๊ฒŒ์ด์ฆˆ ํ™œ์„ฑ์„ ๊ฐ€์ง€๊ฑฐ๋‚˜ RNA ๊ฒฐํ•ฉ ๋ชจํ‹ฐ๋ธŒ๋ฅผ ๊ฐ€์ง€๋ฉฐ ํžˆ์Šคํ†ค ์•„์„ธํ‹ธ๋ ˆ์ด์…˜๊ณผ ๊ด€๊ณ„๋œ ์ „์‚ฌ์ธ์ž์˜ ์ƒ๋™ ์œ ์ „์ž๋ฅผ ํฌํ•จํ•œ๋‹ค. ์ถ”๊ฐ€์ ์œผ๋กœ ์ด๋ถ„์žํ˜•๊ด‘์ƒ๋ณด๊ธฐ๋ฒ•์„ ํ†ตํ•ด ๋ฐํ˜€์ง„ ์œ ์ „์ž๋“ค๊ณผ DME์˜ ๊ด€๊ณ„์— ๋Œ€ํ•œ ์ดํ•ด๋ฅผ ์œ„ํ•ด RNA ์Šคํ”Œ๋ผ์ด์‹ฑ์— ๊ด€์—ฌํ•˜๋Š” Brr2a๋ฅผ ์•”ํ˜ธํ™”ํ•˜๋Š” ์œ ์ „์ž์ธ At1g20960์„ ์ด์šฉํ•˜์—ฌ ๋Œ์—ฐ๋ณ€์ด ์—ฐ๊ตฌ๋ฅผ ์ง„ํ–‰ํ•˜์˜€๋‹ค. At1g20960์˜ T-DNA ์‚ฝ์ž… ๋Œ์—ฐ๋ณ€์ด์ฒด์ธ emb1507-1 ์‹๋ฌผ์ฒด๋ฅผ ยฌdme-2 ๋Œ์—ฐ๋ณ€์ด์ฒด์™€ ๊ต๋ฐฐํ•˜์—ฌ ์–ป์€ ์ด์ค‘์ดํ˜•์ ‘ํ•ฉ ๋Œ์—ฐ๋ณ€์ด๋ฅผ ์ด์šฉํ•˜์—ฌ ์ข…์ž์˜ ๋ฐœ๋‹ฌ๋‹จ๊ณ„ ๋ฐ ์ข…์ž์œ ์‚ฐ ์—ฌ๋ถ€์™€ ๋Œ์—ฐ๋ณ€์ด ๋Œ€๋ฆฝ์œ ์ „์ž์˜ ๋ถ„๋ฆฌ๋น„, ํŠธ๋žœ์Šค๋ฏธ์…˜์˜ ๋ณ€ํ™”๋ฅผ ์ค‘์‹ฌ์œผ๋กœ ํ‘œํ˜„ํ˜•์„ ๋ถ„์„ํ•œ ๊ฒฐ๊ณผ dme-2 ๋‹จ์ผ ์ดํ˜•์ ‘ํ•ฉ๋Œ์—ฐ๋ณ€์ด์ฒด์™€ ๋น„๊ตํ–ˆ์„ ๋•Œ ํ•ด๋‹น ํ‘œํ˜„ํ˜•๋“ค์—์„œ์˜ ์ฃผ๋ชฉํ• ๋งŒํ•œ ๋ณ€ํ™”๋ฅผ ๊ด€์ฐฐํ•  ์ˆ˜๋Š” ์—†์—ˆ๋‹ค. ๋•Œ๋ฌธ์— ์‹๋ฌผ์—์„œ DME์˜ ๋””๋ฉ”ํ‹ธ๋ ˆ์ด์…˜ ๊ณผ์ •์— ๊ด€์—ฌํ•˜๋Š” ์ƒˆ๋กœ์šด ์ธ์ž๋ฅผ ๋ฐํ˜€๋ƒ„์œผ๋กœ์จ ์ด์— ๋Œ€ํ•œ ๊ตฌ์ฒด์ ์ธ ์ดํ•ด๋ฅผ ๋”ํ•˜๊ธฐ ์œ„ํ•ด์„œ๋Š” ์ถ”๊ฐ€์ ์ธ ์‹คํ—˜์„ ํ†ตํ•ด ๋ถ„์ž์ƒ๋ฌผํ•™์  ๊ด€์ ์—์„œ์˜ ํ‘œํ˜„ํ˜• ๋ถ„์„์ด ํ•„์š”ํ•˜๋ฉฐ At1g20960 ์™ธ์— ๋‚˜๋จธ์ง€ ํ›„๋ณด ์œ ์ „์ž๋“ค์— ๋Œ€ํ•œ ์—ฐ๊ตฌ ์—ญ์‹œ ํ•„์š”ํ•˜๋‹ค.ABSTRACT โ…ฐ TABLES OF CONTENTS โ…ฒ LIST OF FIGURES โ…ด LIST OF TABLES โ…ต BACKGROUNDS 1 1. DNA Methylation in Arabidopsis 1 2. DNA demethylation and DEMETER 4 3. Purpose of this Study 7 CHAPTER ONE: Interaction Partner Screening for DEMETER using Bimolecular Fluorescence Complementation(BiFC) Assay 8 1. Introduction 9 2. Materials and methods 12 2.1. Plant material and growth conditions 12 2.2. Previous yeast two-hybrid data 12 2.3. List-up for BiFC assay 15 2.4. Cloning for BiFC assay 15 2.5. BiFC assay 21 3. Results and discussion 23 3.1. Cloning for BiFC assay 23 3.2. 5 genes were identified as interactors of DME through BiFC 25 CHAPTER TWO: Phenotypic Study for the DME interactors 31 1. Introduction 32 2. Materials and methods 35 2.1. Plant material and growth conditions 35 2.2. Seed-set analysis and whole-mount clearing 35 3. Results and discussion 38 3.1. EMB/emb1507-1 seed phenotype 38 3.2. EMB/emb1507-1;DME/dme-2 double heterozygote mutant seed phenotype 38 3.3. Transmission of emb and dme alleles in EMB/emb1507-1;DME/dme-2 mutant plants 41 CONCLUSION 43 REFERENCE 46 ABSTRACT IN KOREAN 52Maste

    ๋น„์™ธ์ƒ์„ฑ ๋‘๊ฐœ ๋‚ด ๊ฒฝ๋ง‰ํ•˜ํ˜ˆ์ข… ํ™˜์ž์—์„œ ์ „์‚ฐํ™”๋‹จ์ธต์ดฌ์˜-์ฒ™์ˆ˜๊ฐ•์กฐ์˜์ˆ ์—์„œ์˜ ๋‡Œ์ฒ™์ˆ˜์•ก ๋ˆ„์ถœ ๋นˆ๋„์— ๋Œ€ํ•œ ์—ฐ๊ตฌ

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    ํ•™์œ„๋…ผ๋ฌธ(๋ฐ•์‚ฌ) -- ์„œ์šธ๋Œ€ํ•™๊ต๋Œ€ํ•™์› : ์˜๊ณผ๋Œ€ํ•™ ์ž„์ƒ์˜๊ณผํ•™๊ณผ, 2022. 8. ์ด์ค€์šฐ.In our tertiary medical center, we observed a higher rate of cerebrospinal fluid (CSF) leak on computed tomography (CT) myelography in patients with nontraumatic intracranial subdural hematoma (SDH) than that reported in previous literature. Therefore, we designed the present study to demonstrate which screening criteria were responsible for the difference of the prevalence, and determine potential clinical and imaging parameters favoring CSF leak in patients with nontraumatic intracranial SDH. Patients diagnosed with nontraumatic intracranial SDH who underwent CT myelography (60 patients) and who did not undergo CT myelography (50 patients) between January 2012 and March 2018 were selected. Patients with CT myelography were divided into CSF leak-positive and CSF leak-negative groups according to CT myelography reports. Clinical and magnetic resonance imaging findings were statistically compared among the three groups. Patients with the following findings significantly tended to be selected as CT myelography candidates: lower mean age, no aspirin intake, no clopidogrel administration, no unilateral weakness, no gait disturbance, normal Glasgow coma scale score, orthostatic headache, nonpostural headache, bilateral SDH, thinner maximal hematoma thickness, dural thickening/enhancement, cisternal obliteration, transverse sinus convexity, decreased pontomesencephalic angle and mamillopontine distance. In total 48 of the 60 (80%) patients exhibited CSF leak on CT myelography. Between the leak-positive and negative groups, an age of < 69 years was significantly associated with the presence of CSF leak. However, patients aged โ‰ฅ 69 years also had tendency to exhibit spontaneous intracranial hypotension (SIH)-induced SDH (14/23; 60.87%). There were no other variables, except age, that showed statistical differences between the leak-positive and negative groups. If variable clinical and imaging factors are taken into consideration when selecting patients who needed CT myelography, more cases with CSF leak can be diagnosed in patients with nontraumatic intracranial SDH. Importantly, clinicians need to broaden the patientsโ€™ age-related indication to a higher age group for CT myelography to detect CSF leak. Also, the absence of postural headache or well-known typical SIH-related MRI features in patients does not mean that SIH can be completely excluded in clinical practice.๋ณธ์›์—์„œ ๋น„์™ธ์ƒ์„ฑ ๋‘๊ฐœ๋‚ด ๊ฒฝ๋ง‰ํ•˜ํ˜ˆ์ข… ํ™˜์ž๋ฅผ ๋Œ€์ƒ์œผ๋กœ CT ์ฒ™์ˆ˜๊ฐ•์กฐ์˜์ˆ ์„ ์‹œํ–‰ํ•˜๋Š” ๊ฒฝ์šฐ ๊ธฐ์กด ๋ฌธํ—Œ์— ๋ณด๊ณ ๋œ ๊ฒƒ๋ณด๋‹ค ๋†’์€ ๋นˆ๋„๋กœ ๋‡Œ์ฒ™์ˆ˜์•ก ๋ˆ„์ˆ˜๊ฐ€ ํ™•์ธ๋˜์–ด, ์ด์— ๋ณธ์› CT ์ฒ™์ˆ˜๊ฐ•์กฐ์˜์ˆ  ์ ์šฉ ๊ธฐ์ค€์„ ์‚ดํŽด๋ณด๊ณ  ๋น„์™ธ์ƒ์„ฑ ๋‘๊ฐœ๋‚ด ๊ฒฝ๋ง‰ํ•˜ํ˜ˆ์ข… ํ™˜์ž์—์„œ CT ์ฒ™์ˆ˜๊ฐ•์กฐ์˜์ˆ ์„ ํ•„์š”๋กœ ํ•˜๋Š” ์œ ์˜๋ฏธํ•œ ์ž„์ƒ ๋ฐ ์˜์ƒ ์†Œ๊ฒฌ์„ ๋ฐํ˜€๋‚ด๊ณ ์ž ํ•˜์˜€๋‹ค. 2012๋…„ 1์›”๋ถ€ํ„ฐ 2018๋…„ 3์›” ์‚ฌ์ด ๋น„์™ธ์ƒ์„ฑ ๋‘๊ฐœ๋‚ด ๊ฒฝ๋ง‰ํ•˜ํ˜ˆ์ข…์œผ๋กœ ์ง„๋‹จ๋ฐ›๊ณ  CT ์ฒ™์ˆ˜๊ฐ•์กฐ์˜์ˆ ์„ ์‹œํ–‰ ๋ฐ›์€ ํ™˜์ž(60๋ช…)์™€ ๋น„์™ธ์ƒ์„ฑ ๋‘๊ฐœ๋‚ด ๊ฒฝ๋ง‰ํ•˜ํ˜ˆ์ข…์œผ๋กœ ์ง„๋‹จ๋ฐ›์•˜์œผ๋‚˜ CT ์ฒ™์ˆ˜๊ฐ• ์กฐ์˜์ˆ ์„ ์‹œํ–‰ ๋ฐ›์ง€ ์•Š์€ ํ™˜์ž(50๋ช…)๋ฅผ ๋Œ€์ƒ์œผ๋กœ ํ•˜์˜€๋‹ค. ์ „์ž์˜ ํ™˜์ž๋Š” CT ์ฒ™์ˆ˜๊ฐ•์กฐ์˜์ˆ  ๊ฒฐ๊ณผ์— ๋”ฐ๋ผ ๋‡Œ์ฒ™์ˆ˜์•ก ๋ˆ„์ˆ˜๊ฐ€ ์กด์žฌํ•˜๋Š” ๊ตฐ๊ณผ ๊ทธ๋ ‡์ง€ ์•Š์€ ๊ตฐ์œผ๋กœ ๋ถ„๋ฅ˜๋˜์—ˆ๋‹ค. ์—ฐ๊ตฌ์ง„์€ ์ด ์„ธ ๊ตฐ ์‚ฌ์ด์˜ ์ž„์ƒ์  ์†Œ๊ฒฌ ๋ฐ ์ž๊ธฐ๊ณต๋ช… ์˜์ƒ ์†Œ๊ฒฌ์„ ํ†ต๊ณ„ํ•™์ ์œผ๋กœ ๋ถ„์„ํ•˜์˜€์œผ๋ฉฐ ๋น„์™ธ์ƒ์„ฑ ๋‘๊ฐœ๋‚ด ๊ฒฝ๋ง‰ํ•˜ํ˜ˆ์ข… ํ™˜์ž๋Š” ๋‹ค์Œ๊ณผ ๊ฐ™์€ ๊ฒฝ์šฐ์— ์œ ์˜๋ฏธํ•˜๊ฒŒ CT myelography๋ฅผ ์‹œํ–‰ ๋ฐ›๋Š” ๊ฒฝํ–ฅ์ด ์žˆ์—ˆ๋‹ค : ๋‚ฎ์€ ์—ฐ๋ น์ธ ๊ฒฝ์šฐ, ์•„์Šคํ”ผ๋ฆฐ์ด๋‚˜ ํด๋กœํ”ผ๋„๊ทธ๋ ์„ ๋ณต์šฉํ•˜์ง€ ์•Š๋Š” ๊ฒฝ์šฐ, ํŽธ์ธก์„ฑ ์œ„์•ฝ๊ฐ์„ ๋ณด์ด์ง€ ์•Š๋Š” ๊ฒฝ์šฐ, ๋ณดํ–‰์žฅ์• ๋ฅผ ๋ณด์ด์ง€ ์•Š๋Š” ๊ฒฝ์šฐ, Glasgow coma scale ์ ์ˆ˜๊ฐ€ ์ •์ƒ์ธ ๊ฒฝ์šฐ, ๊ธฐ๋ฆฝ์„ฑ ๋‘ํ†ต์ด ์žˆ๋Š” ๊ฒฝ์šฐ, ์ž์„ธ์™€ ๋ฌด๊ด€ํ•œ ๋‘ํ†ต์ด ์žˆ๋Š” ๊ฒฝ์šฐ, ์ž๊ธฐ๊ณต๋ช…์˜์ƒ์—์„œ ์–‘์ธก์„ฑ ๊ฒฝ๋ง‰ํ•˜ ํ˜ˆ์ข…, ๋น„๊ต์  ์–‡์€ ํ˜ˆ์ข…๋‘๊ป˜, ๊ฒฝ์งˆ๋ง‰ ๋น„ํ›„/์กฐ์˜์ฆ๊ฐ•, ์ˆ˜์กฐํ์‡„, ํšก๋‹จ์ •๋งฅ์˜ ํŒฝ์ฐฝ, ๊ต๋‡Œ-์ค‘๋‡Œ ์‚ฌ์ด๊ฐ ๊ฐ์†Œ, ์œ ๋‘์ฒด-๊ต๋‡Œ ๊ฐ„๊ฒฉ ๊ฐ์†Œ๋ฅผ ๋ณด์ด๋Š” ๊ฒฝ์šฐ. ์ด๋ ‡๊ฒŒ CT myelography ์‹œํ–‰ ๋ฐ›์€ ๋ก€์˜ 80%์—์„œ ๋‡Œ์ฒ™์ˆ˜์•ก ๋ˆ„์ˆ˜๋ฅผ ๋ณด์˜€์œผ๋ฉฐ, ๋ˆ„์ˆ˜๋ฅผ ๋ณด์ด๋Š” ๊ตฐ์€ ๋ˆ„์ˆ˜๋ฅผ ๋ณด์ด์ง€ ์•Š๋Š” ๊ตฐ์— ๋น„ํ•ด ํ†ต๊ณ„์ ์œผ๋กœ ์œ ์˜๋ฏธํ•˜๊ฒŒ ๋‚ฎ์€ ์—ฐ๋ น(69์„ธ ๋ฏธ๋งŒ)์ด์—ˆ์œผ๋‚˜ 69์„ธ ์ด์ƒ์˜ ํ™˜์ž์—์„œ๋„ ๋‡Œ์ฒ™์ˆ˜์•ก ๋ˆ„์ˆ˜๋ฅผ ๋ณด์ด๋Š” ๊ฒฝํ–ฅ์ด ์žˆ์—ˆ๋‹ค(14/23; 60.87%). ๋น„์™ธ์ƒ์„ฑ ๋‘๊ฐœ๋‚ด ๊ฒฝ๋ง‰ํ•˜ํ˜ˆ์ข… ํ™˜์ž์—์„œ ์ƒ๊ธฐ ๊ธฐ์ˆ ๋œ ๋‹ค์–‘ํ•œ ์ž„์ƒ์ , ์˜์ƒ์  ์†Œ๊ฒฌ์„ ๊ณ ๋ คํ•˜์—ฌ ๋‡Œ์ฒ™์ˆ˜์•ก ๋ˆ„์ˆ˜ ํ™•์ธ ๊ฒ€์‚ฌ๊ฐ€ ํ•„์š”ํ•œ ํ™˜์ž๊ตฐ์„ ์„ ํƒํ•œ๋‹ค๋ฉด ๊ธฐ์กด ๋ฌธํ—Œ์— ๋ณด๊ณ ๋œ ๊ฒƒ๋ณด๋‹ค ๋†’์€ ๋น„์œจ๋กœ ๋‡Œ์ฒ™์ˆ˜์•ก ๋ˆ„์ˆ˜๊ฐ€ ์žˆ๋Š” ์ž๋ฐœ์„ฑ ๋‘๊ฐœ๋‚ด ์ €ํ˜ˆ์•• ํ™˜์ž๋ฅผ ์ง„๋‹จํ•  ์ˆ˜ ์žˆ์–ด ์‹œ๊ธฐ์ ์ ˆํ•œ ์น˜๋ฃŒ๊ฐ€ ๊ฐ€๋Šฅํ•˜๋‹ค. ๊ธฐ์กด ๋ฌธํ—Œ์— ๋ณด๊ณ ๋œ ๊ฒƒ์— ๋น„ํ•ด ๊ณ ์—ฐ๋ น์ธ ๊ฒฝ์šฐ์—๋„ ์ž๋ฐœ์„ฑ ๋‘๊ฐœ๋‚ด ์ €ํ˜ˆ์•• ํ™˜์ž๊ฐ€ ์žˆ์„ ์ˆ˜ ์žˆ๋‹ค๋Š” ์ ์— ์ฃผ์˜ํ•˜์—ฌ์•ผ ํ•œ๋‹ค.Contents 1. Introduction 1 2. Materials and methods 2 3. Results 7 4. Discussion 15 5. Conclusion 17 6. Disclosure 17 References 19 ๊ตญ๋ฌธ์ดˆ๋ก 23 Tables [Table 1] 7 [Table 2] 8 [Table 3] 11 [Table 4] 12 Figures [Figure 1] 13๋ฐ•

    SHORTROOT์™€ NAC-REGULATED SEED MORPHOLOGY 1์— ์˜ํ•œ ์• ๊ธฐ์žฅ๋Œ€ ๋ฟŒ๋ฆฌ ์ฒด๊ด€ ๋ฐœ๋‹ฌ์˜ ์‹œ๊ณต๊ฐ„์  ์กฐ์ ˆ์— ๊ด€ํ•œ ์—ฐ๊ตฌ

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    ํ•™์œ„๋…ผ๋ฌธ(๋ฐ•์‚ฌ) -- ์„œ์šธ๋Œ€ํ•™๊ต๋Œ€ํ•™์› : ์ž์—ฐ๊ณผํ•™๋Œ€ํ•™ ์ƒ๋ช…๊ณผํ•™๋ถ€, 2021.8. ๊น€ํšจ์ง„.์‹๋ฌผ์˜ ๋ฟŒ๋ฆฌ๋Š” ์ •๋‹จ๋ถ„์—ด์กฐ์ง์—์„œ ์ผ์–ด๋‚˜๋Š” ์„ธํฌ ๋ถ„์—ด์„ ํ†ตํ•˜์—ฌ ๊ธธ์ด ์„ฑ์žฅ์„ ํ•œ๋‹ค. ์ง€์†์ ์ธ ๊ธธ์ด ์„ฑ์žฅ์„ ์œ„ํ•ด์„œ๋Š” ๋ถ„์—ด๋Šฅ์ด ์™•์„ฑํ•œ ์„ธํฌ๋ฅผ ๊ณต๊ธ‰ํ•˜๋Š” ๋ฏธ๋ถ„ํ™” ์ƒํƒœ์˜ ์ค„๊ธฐ์„ธํฌ(stem cell)๋“ค์ด ์ •๋‹จ๋ถ„์—ด์กฐ์ง์—์„œ ์œ ์ง€ํ•˜๊ณ  ์žˆ๋‹ค. ํŠนํžˆ, ์• ๊ธฐ์žฅ๋Œ€ ๋ฟŒ๋ฆฌ๋ถ„์—ด์กฐ์ง์—์„œ๋Š” ๋ถ„์—ด์ง€์—ฐ์ค‘์‹ฌ๋ถ€๋ฅผ ์—์›Œ์‹ผ ์ค„๊ธฐ์„ธํฌ์˜ ์ •๋ ฌ๋œ ๋น„๋Œ€์นญ๋ถ„์—ด์„ ํ†ตํ•˜์—ฌ ๋ฟŒ๋ฆฌ ์ค‘์‹ฌ์ฃผ์— ๋ฌด๊ธฐ์–‘๋ถ„์„ ์ˆ˜์†กํ•˜๋Š” ๋ฌผ๊ด€๋ถ€๊ฐ€ ์ผ๋ ฌ๋กœ ํ˜•์„ฑ๋˜๊ณ , ์–‘๋ถ„๊ณผ ํฌ๋„๋‹น ๋ฐ ๋‹ค์–‘ํ•œ ํ˜•ํƒœ์˜ ์‹ ํ˜ธ์ „๋‹ฌ ๋ฌผ์งˆ์„ ์ˆ˜์†กํ•˜๋Š” ๋‘ ๊ฐœ ์ถ•์˜ ์ฒด๊ด€๋ถ€๊ฐ€ ๋ฌผ๊ด€๋ถ€์˜ ์ˆ˜์ง ๋ฐฉํ–ฅ์œผ๋กœ ํ˜•์„ฑ๋œ๋‹ค. ์ด ๋•Œ, ์ฒด๊ด€์„ธํฌ๋Š” ์‹๋ฌผ์ด ์„ฑ์žฅํ•˜๋Š” ๋ฐ ํ•„์ˆ˜ ์˜์–‘์†Œ์™€ ์‹ ํ˜ธ ์ „๋‹ฌ ๋ถ„์ž์˜ ์šด๋ฐ˜์„ ์ค‘์žฌํ•˜๋Š” ์ค‘์š”ํ•œ ์—ญํ• ์„ ํ•œ๋‹ค. ์ด๋Ÿฌํ•œ ๊ธฐ๋Šฅ์„ ๊ฐ€์ง€๋Š” ์ฒด๊ด€์„ธํฌ๋Š” ๋‘ ๊ฐ€์ง€ ์ฃผ์š” ๋น„๋Œ€์นญ ์„ธํฌ ๋ถ„์—ด์„ ํ†ตํ•ด ๋ฐ˜์„ธํฌ(Companion cells, CCs)์™€ phloem sieve elements (phloem SEs)๋กœ ๋ฐœ๋‹ฌ๋œ๋‹ค. ์• ๊ธฐ์žฅ๋Œ€ ๋ฟŒ๋ฆฌ ๋ถ„์—ด์กฐ์ง์—์„œ๋Š” ๋ฐ˜์„ธํฌ๋Š” phloem SEs๋ฅผ ๋งˆ์ฃผํ•˜๋Š” procambial ์„ธํฌ์™€ ๋‚ดํ”ผ(pericycle)์˜ ๋น„๋Œ€์นญ๋ถ„์—ด์„ ํ†ตํ•ด ์ƒ์„ฑ๋˜๋ฉฐ, ์ด์™€ ๋‹ค๋ฅด๊ฒŒ proto phloem SE์™€ metaphloem SE๋กœ ๊ตฌ์„ฑ๋œ ์ฒด๊ด€ SE๋Š” ๋‘ ๋ฒˆ์˜ ์—ฐ์†์ ์ธ ์ฒด๊ด€ ์ „๊ตฌ์„ธํฌ์˜ ๋น„๋Œ€์นญ๋ถ„์—ด๋กœ ๋ฐœ๋‹ฌํ•˜๊ฒŒ ๋œ๋‹ค. ์‹คํ—˜์‹ค์˜ ์ด์ „ ์—ฐ๊ตฌ์—์„œ GRAS ๊ณ„์—ด ์ „์‚ฌ ์ธ์ž์ธ SHORTROOT (SHR)๊ฐ€ ๋ฟŒ๋ฆฌ์˜ ์ฒด๊ด€ ๋ฐœ๋‹ฌ๊ณผ์ •์—์„œ sieve element(SE)์™€ ๋ฐ˜์„ธํฌ๋ฅผ ํ˜•์„ฑํ•˜๊ธฐ ์œ„ํ•œ ๋น„๋Œ€์นญ๋ถ„์—ด์— ์ฃผ์š”ํ•œ ์—ญํ• ์„ ํ•œ๋‹ค๋Š” ๊ฒƒ์„ ๋ฐํ˜”๋‹ค. SHR์˜ ์ „์‚ฌ์ฒด๋Š” ๋ฟŒ๋ฆฌ์—์„œ ์ฒด๊ด€์„ธํฌ๋ฅผ ์ œ์™ธํ•œ ๋ฌผ๊ด€, procambium ์„ธํฌ์ธต์—์„œ ๋ฐœํ˜„๋˜์ง€๋งŒ, SHR์˜ ๋‹จ๋ฐฑ์งˆ์€ ์ฒด๊ด€๋ถ€๋กœ ์ด๋™ํ•˜์—ฌ ์ฒด๊ด€ SE ํ˜•์„ฑ์— ํ•„์š”ํ•œ ๋น„๋Œ€์นญ๋ถ„์—ด์„ ์œ ๋„ํ•œ๋‹ค. ์ด๋ฏธ endodermis ๋ฐ ๋ถ„์—ด์ง€์—ฐ์ค‘์‹ฌ๋ถ€ (Quiescent center, QC)์—์„œ SHR์€ SCRACOW (SCR)์˜ ๋ฐœํ˜„์„ ํ™œ์„ฑํ™”ํ•˜๊ณ  SCR๊ณผ ์ƒํ˜ธ ์ž‘์šฉํ•œ๋‹ค๋Š” ๊ฒƒ์€ ์•Œ๋ ค์ ธ ์žˆ๋‹ค. ์ด ๋‘˜์˜ ๋‹จ๋ฐฑ์งˆ ๋ณตํ•ฉ์ฒด๋Š” microRNA165/6 (miR165/6)์˜ ๋ฐœํ˜„์„ ์œ ๋ฐœ์‹œํ‚ค๊ณ  ์ด๋“ค์€ ๋ฟŒ๋ฆฌ ์ค‘์‹ฌ๋ถ€๋กœ ์ด๋™ํ•˜์—ฌ Class III homeodomain leucine zipper (HD-ZIPIII) mRNA๋ฅผ ๋ถ„ํ•ดํ•˜์—ฌ HD-ZIPIII์˜ ๋†๋„ ๊ตฌ๋ฐฐ๋ฅผ ํ˜•์„ฑํ•œ๋‹ค. ์ด๋Ÿฌํ•œ ๋†๋„ ๊ตฌ๋ฐฐ๋Š” protoxylem๊ณผ metaxylem์„ ํ˜•์„ฑํ•˜์—ฌ ๋ฌผ๊ด€๋ถ€๋ฅผ ์กฐ์งํ™”ํ•œ๋‹ค. ์ด์ „ ์ฝ”๋„ฌ๋Œ€ํ•™๊ต์—์„œ ์ด์ง€์˜ ๊ต์ˆ˜๋‹˜๊ป˜ ์ง€๋„๋ฅผ ๋ฐ›์•˜๋˜ ๋ฐ•์‚ฌ๊ณผ์ • ํ•™์ƒ, Dr. Jing Zhou๋Š” ์ฒด๊ด€๋ถ€๋กœ์˜ SHR ์ด๋™์ด ์ฒด๊ด€์„ธํฌ ๋น„๋Œ€์นญ ์„ธํฌ ๋ถ„์—ด์„ ์ด‰์ง„ํ•˜์—ฌ ์ฒด๊ด€์„ธํฌ ๋ฐœ๋‹ฌ์— ์ฃผ์š”ํ•œ ์—ญํ• ์„ ํ•œ๋‹ค๋Š” ๊ฒƒ์„ ๊ด€์ฐฐํ•˜์˜€๋‹ค. ๊ทธ๋Ÿฌ๋‚˜, ๋ฟŒ๋ฆฌ์ค‘์‹ฌ๋ถ€์˜ SHR์˜ ์ด๋™์ด phloem SEs ๋ฐ CCs ํ˜•์„ฑ์„ ์œ„ํ•œ ๋น„๋Œ€์นญ๋ถ„์—ด์„ ์–ด๋–ป๊ฒŒ ์กฐ์ ˆํ•˜๋Š”์ง€์— ๊ด€ํ•œ ๊ตฌ์ฒด์ ์ธ ๊ณผ์ •์€ ์—ฌ์ „ํžˆ ๋ถˆ๋ถ„๋ช…ํ–ˆ๋‹ค. ๋”ฐ๋ผ์„œ ๋ณธ ์—ฐ๊ตฌ๋Š” SHR์™€ ๊ทธ ํ•˜์œ„ ์œ ์ „์ž๋“ค์ด ์–ด๋–ป๊ฒŒ ์ฒด๊ด€์„ธํฌ ๋ฐœ๋‹ฌ์— ๊ธฐ์—ฌํ•˜๋Š”์ง€ ์•Œ์•„๋‚ด๋Š” ๊ฒƒ์„ ๋ชฉํ‘œ๋กœ ์‚ผ์•˜๋‹ค. ์ด๋ฅผ ๋ฐํžˆ๊ธฐ ์œ„ํ•ด, SHR ํ•˜์œ„ ์œ ์ „์ž๋กœ์„œ์˜ NARS1, SND2๊ฐ€ ์ฒด๊ด€์„ธํฌ ํ˜•์„ฑ์„ ์œ„ํ•ด ์–ด๋– ํ•œ ๊ธฐ๋Šฅ์„ ํ•˜๋Š”์ง€ ๋ถ„์„ํ•˜์˜€๊ณ  ์ด๋“ค์˜ ๊ด€๊ณ„๋ฅผ ๋ฐํžˆ๊ณ ์ž ํ•˜์˜€๋‹ค. ๋ณธ ์—ฐ๊ตฌ๋ฅผ ํ†ตํ•ด SHR์™€ ๊ทธ์˜ ์ง์ ‘์ ์ธ ํ•˜์œ„ ์ „์‚ฌ ์ธ์ž์ธ NAC-REGULATED SEED MORPHOLOGY 1 (NARS1)๊ณผ SECONDARY WALL-ASSOCIATED NAC DOMAIN PROTEIN 2 (SND2)๊ฐ€ positive feedforward loop๋ฅผ ํ˜•์„ฑํ•˜๋ฉฐ ์ฒด๊ด€์„ธํฌ ๋ฐœ๋‹ฌ์— ๊ด€์—ฌํ•œ๋‹ค๋Š” ์‚ฌ์‹ค์„ ๋ฐํž ์ˆ˜ ์žˆ์—ˆ๋‹ค. ํŠนํžˆ, NARS1์ด โ€˜Topโ€“down long distance signalโ€™์„ ํ†ตํ•œ ์ „์‚ฌ ์กฐ์ ˆ์ธ์ž๋กœ ์ฒด๊ด€ SE ์ „๊ตฌ(precursor)์„ธํฌ์˜ ๋น„๋Œ€์นญ ๋ถ„์—ด์„ ์กฐ์ ˆํ•  ๊ฒƒ์ด๋ผ๋Š” ์ž ์žฌ์  ๊ฐ€๋Šฅ์„ฑ์„ ์ œ์‹œํ•˜์˜€๋‹ค. ํฅ๋ฏธ๋กญ๊ฒŒ๋„, NARS1์˜ ์ „์‚ฌ์ฒด๋Š” ๋ฟŒ๋ฆฌ์˜ ๋ถ„์—ด์กฐ์ง(meristem)์ด ์•„๋‹Œ ์„ธํฌ๋“ค์˜ ๋น„๋Œ€์นญ๋ถ„์—ด์ด ๋๋‚˜๊ณ  ์„ฑ์žฅํ•˜๋Š” ๋ถ„ํ™”๋Œ€(differentiation zone)์˜ ๋ฐ˜์„ธํฌ์—์„œ ๋ฐœํ˜„์ด ๋œ๋‹ค. ๊ทธ๋Ÿฌ๋‚˜ nars1์˜ ๋Œ์—ฐ๋ณ€์ด์ฒด๋Š” ์ฒด๊ด€ ์ „๊ตฌ์„ธํฌ์˜ ๋น„๋Œ€์นญ๋ถ„์—ด์ด ๋น„์ •์ƒ์ ์œผ๋กœ ์ผ์–ด๋‚œ๋‹ค. ์ด๋Š” NARS1์˜ ์œ„์ชฝ ๋ฐ˜์„ธํฌ์—์„œ์˜ ๋ฐœํ˜„์ด ์•„๋ž˜ ๋ถ„์—ด์กฐ์ง์—์„œ์˜ ๋น„๋Œ€์นญ๋ถ„์—ด์„ ์กฐ์ ˆํ•œ๋‹ค๋Š” โ€˜Long distance top-down signalingโ€™์˜ ๊ฐ€๋Šฅ์„ฑ์„ ์ œ์‹œํ•œ๋‹ค. ๊ฒŒ๋‹ค๊ฐ€ SND2๋Š” NARS1์„ ํ†ตํ•ด ๋ฟŒ๋ฆฌ์˜ ๋ถ„์—ด์กฐ์ง์ธ protophloem SE์—์„œ ๋ฐœํ˜„๋˜๋ฉฐ NARS1์˜ ๋ฐœํ˜„์„ ํ™œ์„ฑํ™”ํ•˜์—ฌ positive feedback loop๋ฅผ ํ˜•์„ฑํ•˜๋Š” ๋ฐ ๋’ท๋ฐ›์นจ๋œ๋‹ค. ์ด๋Ÿฌํ•œ ๊ฒฐ๊ณผ๋“ค์„ ์ข…ํ•ฉํ•ด๋ณผ ๋•Œ, ๋ณธ ์—ฐ๊ตฌ๋Š” NARS1์ด phloem SE์˜ ๋น„๋Œ€์นญ๋ถ„์—ด์„ ์œ„ํ•ด โ€˜Long distance top-down signalingโ€™์„ ํ†ตํ•ด ์กฐ์ ˆํ•˜๋ฉฐ, ์ด๋Š” ๋ถ„ํ™”๋œ phloem ์„ธํฌ๋“ค์ด ์•„์ง ๋ถ„ํ™”๋˜์ง€ ์•Š์€ ์ „๊ตฌ์„ธํฌ๋ฅผ ๊ทœ์ œํ•  ์ˆ˜ ์žˆ๊ฒŒ ํ•œ๋‹ค๋Š” ๊ฐ€๋Šฅ์„ฑ์„ ์ œ์‹œํ•œ๋‹ค. ์ด๋Ÿฌํ•œ ์—ฐ๊ตฌ๋ฅผ ํ†ตํ•ด ๋‹ค์ด๋‚˜๋ฏนํ•œ ์‹๋ฌผ ์„ฑ์žฅ ๊ณผ์ • ์ค‘ ๊ด€๋‹ค๋ฐœ ์กฐ์ง ์‹œ์Šคํ…œ์˜ ์„ธํฌ ์œ ํ˜• ํŒจํ„ด ํ˜•์„ฑ๊ณผ ์„ธํฌ ๋ถ„ํ™”์— ๋Œ€ํ•œ ์ƒˆ๋กœ์šด ํ†ต์ฐฐ๋ ฅ์„ ์ œ๊ณตํ•  ๊ฒƒ์ด๋ผ ๊ธฐ๋Œ€ํ•œ๋‹ค.In the plant vascular system, phloem plays an important role in mediating the transportation of essential nutrients and signaling molecules. Two major phloem cell types are generated by asymmetric cell divisions (ACDs); companion cells (CCs) and phloem sieve elements (SEs). In the root apical meristem of Arabidopsis thaliana, CCs are generated by the ACDs of procambial cells facing the phloem SEs and pericycle cells. On the other hand, phloem SEs which are composed of the protophloem and metaphloem SEs, are originated from two sequential ACDs of a phloem SE-procambium precursor and a SE precursor. A previous study in the lab suggested that SHORTROOT (SHR), a GRAS family transcription factor, regulates the ACDs for CCs from the endodermis and for SEs from the phloem. SHR mRNA is transcribed in the stele (the xylem, procambium, pericycle cells) except for the phloem pole. But, SHR protein moves into the phloem pole, endodermis, and quiescent center (QC). In the endodermis and QC, SHR activates the expression of SCARECROW (SCR), and then interacts with SCR. This protein complex promotes the expression of microRNA 165/6 (miR165/6). miR165/6 move into the stele and establish the Class III homeodomain leucine zipper transcription factors (HD-ZIP III) gradient by degrading HD-ZIP III mRNAs. This process specifies the identities of proto- and metaxylem. Dr. Jing Zhou, the former Ph.D. student, observed that SHR moving into the phloem pole promotes ACDs for phloem development. However, it was still unclear how movement of SHR into the phloem pole regulates ACDs for SE and CC formation. Therefore, my dissertation research was aimed to find how SHR and its downstream genes serve in phloem development. To address this goal, I performed the functional analysis of NARS1 and SND2, SHR downstream target genes for phloem formation. In this dissertation, I present that SHORTROOT (SHR) and its direct downstream genes encoding two transcription factors, NAC-REGULATED SEED MORPHOLOGY 1 (NARS1) and SECONDARY WALL-ASSOCIATED NAC DOMAIN PROTEIN 2 (SND2), are involved in the phloem development via long-distance top-down signaling while forming a positive feedforward loop. NARS1 is a pivotal regulator of ACDs of phloem SE precursor in the meristem, although it is transcribed in the CCs of the root differentiation zone. Thus, NARS1 might be involved in the long-distance top-down signaling pathway. In addition, SND2 is expressed in the protophloem SEs via NARS1 and activates NARS1 expression, thereby setting up a positive feedback loop. Together, this study indicates that ACDs for the development of phloem SE occur via long-distance top-down signaling, which enable differentiated phloem cells to regulate the undifferentiated precursor cells. Findings from this study provide new insights into the cell type patterning and differentiation of vascular system during indeterminate plant growth.1.Introduction 1 1.1 Phloem development in the root apical meristem of Arabidopsis thaliana 1 1.2 Recent progress in molecular mechanisms of phloem sieve element development 6 1.3. Asymmetric cell division is essential to the development of multicellular organisms 8 1.4. SHORTROOT (SHR) is a master regulator in the vascular patterning 12 1.5. SHR regulates development of both phloem SEs and CCs 16 2. Material and Methods 21 2.1. Plant Materials 21 2.2. Plasmid Construction for this study 21 2.3. Microscopic observation 22 2.4. Cross-section 23 2.5. Immunostaining SE-ENOD 24 2.6. CFDA dye test for tracing symplastic loading pathway 25 2.7. ClearSee for confocal imaging . .25 2.8. Callose deposition assay with Aniline blue staining 26 2.9. RealTime-quantitative Polymerase Chain Reaction: RT-qPCR 27 2.10. Statistical Analysis 27 3. Results 30 3.1. SHORTROOT(SHR) is responsible for the phloem development 30 3.1.1. SHR moving into the phloem pole controls ACD of phloem SE precursor 30 3.1.2. Quantification and classification of phloem SE development and cell counting.. 36 3.2. Identification of NARS1 and SND2 as downstream targets of SHR during phloem development 39 3.2.1. Genetic analysis of NARS1 and SND2 as downstream targets of SHR in terms of phloem d evelopment 39 3.2.2. Characteristics of NAC domain transcription factors 45 3.2.3. Expression domain analysis of NARS1 and SND2 in WT and shr-2 47 3.2.4. Characterization of nars1 and snd2 mutant phenotype. 51 3.2.5. Marker analysis in the nars1 and snd2 mutants. 54 3.2.6. Abnormal formation of phloem SEs in the nars1 mutant. 58 3.3. NARS1 regulates the ACD of the phloem SE as the potential long-distance top-down signal 62 3.3.1. NARS1 is a key regulator of ACD of the phloem SE precursor 62 3.3.2. Analysis of phloem unloading in the root meristem. 68 3.3.3. NARS1 acts as a potential long-distance top-down signal to regulate ACD in phloem SEs. 70 3.3.4. Ectopic expression of NARS1 and SND2 promote the extra phloem SE formation 75 3.3.5. Observation of NARS1 proteins using an estradiol-mediated transactivation system 79 3.3.6. Artificial restriction of phloem unloading using PEP1 treatment as a way to monitor NARS1 85 4. Discussion 89 5. References 94 6. Abstract in Korean 108 7. Appendix 111๋ฐ•

    CO2 Uptake Behavior and Vegetative Growth of Doritaenopsis Mantefon Orchids as Influenced by Light/Dark Cycle Manipulation

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    ํ•™์œ„๋…ผ๋ฌธ (์„์‚ฌ)-- ์„œ์šธ๋Œ€ํ•™๊ต ๋Œ€ํ•™์› ๋†์—…์ƒ๋ช…๊ณผํ•™๋Œ€ํ•™ ์‹๋ฌผ์ƒ์‚ฐ๊ณผํ•™๋ถ€, 2017. 8. ๊น€๊ธฐ์„ .This study was performed to investigate how changing the period of light and dark influences the vegetation growth and the photosynthesis of Doritaenopsis. Clones of Doritaenopsis Queen Beer Mantefon at 4-month-old stage were grown in a closed-plant factory system with four different light/dark cycles06/06 h, 08/08 h, 10/10 h, and 12/12 h. Temperature and relative humidity were set at 28oC and 80%, respectively, with a photosynthetic photon flux density of 160 ยฑ 10 ยตmolยทm-2ยทs-1. Repetitive measurements showed that the leaf length and the leaf width were the longest under 12/12 h closely followed by 10/10 h. The fresh weight and the dry weight of leaves and roots were the heaviest at 10/10 h treated samples. Different CO2 uptake patterns were observed from different light/dark cycles. Under 10/10 h and 12/12 h treatments, the CO2 uptake started at early dark period. When the light/dark cycles were shortened to 06/06 h and 08/08 h, the CO2 uptake started at the middle of dark period. Total CO2 uptake amounts were the highest under 12/12 h treatment followed by 10/10 h, 06/06 h, and 08/08 h treatments. Quantitative measurements showed that the vegetative growths under 10/10 h treatment were comparable with that of 12/12 h treatment. These studies indicated that manipulating light/dark can modify the photosynthesis patterns and vegetative growth of Doritaenopsis Mantefon, resulting in the reduction of the production period.INTRODUCTION 1 LITERATURE REVIEW 5 MATERIALS AND METHODS 8 Plant and Growth Conditions 8 Photoperiod and Temperature Treatments 8 Data Collection and Analysis 10 RESULTS AND DISCUSSION 12 Effects of Light/Dark Cycle Manipulation on CO2 Uptake Patterns 12 Effects of Light/Dark Cycle Manipulation on Vegetative Growth 20 LITERATURE CITED 27 ABSTRACT IN KOREAN 32Maste

    ๋๋‚˜์ง€ ์•Š๋Š” ์ „์Ÿ์„ ๊ธฐ์–ตํ•˜๋‹ค

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    [์„œํ‰] 1. ๋‹ค์นด์‹œ ํ›„์ง€ํƒ€๋‹ˆใƒป์ œํ”„๋ฆฌ ํ™”์ดํŠธใƒป์š”๋„ค์•ผ๋งˆ ๋ฆฌ์‚ฌ ์—ฎ์Œ, ใ€Ž์œ„ํ—˜ํ•œ ๊ธฐ์–ตใ€, 2001 2. ๋„๋ฏธ์•ผ๋งˆ ์ด์น˜๋กœ ์—ฎ์Œ, ใ€Ž๊ธฐ์–ต์ด ๋งํ•˜๊ธฐ ์‹œ์ž‘ํ•œ๋‹คใ€, 2006 3. ๋งˆํฌ ๊ฐˆ๋ฆฌ์น˜์˜ค ์—ฎ์Œ, ใ€Ž์˜ˆ์ธกํ•  ์ˆ˜ ์—†๋Š” ๊ณผ๊ฑฐใ€, 200721์„ธ๊ธฐ์— ์ ‘์–ด๋“  ์ง€ ์–ด๋Š๋ง 10๋…„์ด ์ง€๋‚œ 2010๋…„์—๋„ ์šฐ๋ฆฌ๋ฅผ ๋‘˜๋Ÿฌ์‹ผ ์ •์„ธ๋Š” ํ‰ํ™”์™€๋Š” ๊ฑฐ๋ฆฌ๊ฐ€ ๋ฉ€๋‹ค. 21์„ธ๊ธฐ์˜ ๊ฐœ๋ง‰๊ณผ ํ•จ๊ป˜ ์ „์„ธ๊ณ„๋ฅผ ์ถฉ๊ฒฉ์— ๋น ๋œจ๋ฆฐ 9ยท11์‚ฌ๊ฑด์— ๋’ค์ด์€ ํ…Œ๋Ÿฌ์™€์˜ ์ „์Ÿ, ๋ฏธ๊ตญ์˜ ์ด๋ผํฌ ์นจ๊ณต ๋ฐ ์•„ํ”„๊ฐ€๋‹ˆ์Šคํƒ„ ๋‚ด์ „์€ ์˜ค๋ฐ”๋งˆ ์ •๊ถŒ์ด ์ง‘๊ถŒํ•œ ์ง€๊ธˆ๋„ ์ง€์†๋˜๊ณ  ์žˆ๊ณ , ์ด์Šค๋ผ์—˜-ํŒ”๋ ˆ์Šคํƒ€์ธ, ์ธ๋„-ํŒŒํ‚ค์Šคํƒ„ ๋“ฑ์˜ ์ง€์—ญ๋ถ„์Ÿ์œผ๋กœ ๋ถˆ๋ฆฌ๋Š” ๋ฌด๋ ฅ์ถฉ๋Œ์„ ๋™๋ฐ˜ํ•œ ๊ตญ๊ฐ€ ๊ฐ„์˜ ๊ฐˆ๋“ฑ์€ ์˜คํžˆ๋ ค ๊ทธ ๋ฒ”์œ„๊ฐ€ ์ ์ฐจ ํ™•๋Œ€๋˜๊ณ  ์žˆ๋Š” ์ถ”์„ธ์ด๋‹ค. ๋ฒ ๋ฅผ๋ฆฐ ์žฅ๋ฒฝ์˜ ๋ถ•๊ดด์™€ ์†Œ๋น„์—ํŠธ ์—ฐ๋ฐฉ์˜ ํ•ด์ฒด๋กœ ์‹œ์ž‘ํ•œ 20์„ธ๊ธฐ์˜ ๋งˆ์ง€๋ง‰ 10๋…„, ํ”„๋žœ์‹œ์Šค ํ›„์ฟ ์•ผ๋งˆ๊ฐ€ ์ฃผ์ฐฝํ•œ ์—ญ์‚ฌ์˜ ์ข…๋ง์ด ํ•™๊ณ„๋ฅผ ๋น„๋กฏํ•œ ๊ฐ๊ณ„์—์„œ ์ง„์ง€ํ•˜๊ฒŒ ๋ฐ›์•„๋“ค์—ฌ์กŒ๋˜ ๊ณผ๊ฑฐ๋ฅผ ์ƒ๊ฐํ•ด ๋ณธ๋‹ค๋ฉด, ํ˜„์žฌ์˜ ์ด๋Ÿฐ ์ƒํ™ฉ์€ ๊ฒฉ์„ธ์ง€๊ฐ์„ ๋Š๋ผ๊ฒŒ ํ•œ๋‹ค. ํ˜„์‹ค ์‚ฌํšŒ์ฃผ์˜๊ฐ€ ๋ฌด๋„ˆ์ง„ ๋นˆ ์ž๋ฆฌ๋ฅผ ๋ฏธ๊ตญ์‹์˜ ์ž์œ ๋ฏผ์ฃผ์ฃผ์˜๊ฐ€ ์ฑ„์›€์œผ๋กœ์จ ์ฐพ์•„์˜ฌ ๊ฒƒ์œผ๋กœ ๋ฏฟ์—ˆ๋˜ ์—ญ์‚ฌ์˜ ์ข…์–ธ์€ ๊ฒฐ์ฝ” ์˜ค์ง€ ์•Š์•˜๋‹ค. ๋Œ€์‹  ๊ทธ ์ž๋ฆฌ๋ฅผ ์ฑ„์šด ๊ฒƒ์€ ์ „์ง€๊ตฌํ™”๋ผ๋Š” ํ๋ฆ„ ์†์— ๊ฐ•ํ™”๋œ ์ž๋ณธ์˜ ์ฐฉ์ทจ์™€ ๋‹ค์–‘ํ•œ ์ข…๋ฅ˜์˜ ์ง€์—ญ๊ณผ ๋ฏผ์กฑ ๋ฐ ์ข…๊ต๋ถ„์Ÿ โ€” ์ง€์—ญํ™”๋˜๊ณ  ์„ธ๋ถ„ํ™”๋˜์–ด ์˜คํžˆ๋ ค ๊ทธ ๋ฒ”์œ„๋ฅผ ํ™•๋Œ€ํ•œ ์ „์Ÿ โ€” ์ด์—ˆ๋‹ค

    ํ•œ๊ตญ์–ด ์—ฐ์‡„๋™์‚ฌ ๊ตฌ๋ฌธ์˜ ๋…ผํ•ญ ๊ตฌ์กฐ์™€ ์‚ฌ๊ฑด ๊ตฌ์กฐ ํ•ฉ์„ฑ ์œ ํ˜• -'V-์–ด ๊ฐ€๋‹ค/์˜ค๋‹ค', 'V-์–ด ์ฃผ๋‹ค'๋ฅผ ์ค‘์‹ฌ์œผ๋กœ-

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    ํ•™์œ„๋…ผ๋ฌธ (๋ฐ•์‚ฌ)-- ์„œ์šธ๋Œ€ํ•™๊ต ๋Œ€ํ•™์› ์ธ๋ฌธ๋Œ€ํ•™ ์–ธ์–ดํ•™๊ณผ, 2017. 8. ๋‚จ์Šนํ˜ธ.๋ณธ๊ณ ์—์„œ๋Š” ํ•œ๊ตญ์–ด์˜ V-์–ด ๊ฐ€๋‹ค/์˜ค๋‹ค, V-์–ด ์ฃผ๋‹ค๊ฐ€ ์ƒ์„ฑ ๊ธฐ์ œ๋ฉด์—์„œ ๋ฒ”์–ธ์–ด์ ์œผ๋กœ ๋‚˜ํƒ€๋‚˜๋Š” ์—ฐ์‡„๋™์‚ฌ ๊ตฌ๋ฌธ(Serial Verb Construction)์˜ ํŠน์„ฑ์„ ์ง€๋‹Œ ๊ตฌ๋ฌธ์ž„์„ ๋ฐํžˆ๊ณ  ๊ฐ€๋‹ค/์˜ค๋‹ค, ์ฃผ๋‹ค๊ฐ€ ๋‹ค๋ฅธ ๋™์‚ฌ๋“ค๊ณผ ์—ฐ์‡„๋˜์–ด ๋‚˜ํƒ€๋‚  ๋•Œ ๋…ผํ•ญ ๊ตฌ์กฐ์™€ ์‚ฌ๊ฑด ๊ตฌ์กฐ๊ฐ€ ํ•ฉ์„ฑ๋˜๋Š” ๋ฐฉ์‹์„ ์œ ํ˜•ํ™”ํ•˜์—ฌ ์ œ์‹œํ•œ๋‹ค. ์—ฐ์‡„๋™์‚ฌ ๊ตฌ๋ฌธ์ด ๋‹ค๋ฅธ ๋ณตํ•ฉ ๋™์‚ฌ ๊ตฌ๋ฌธ๊ณผ ๊ตฌ๋ถ„๋˜๋Š” ๊ฐ€์žฅ ํฐ ํŠน์ง•์€ ๋‘ ๋™์‚ฌ๊ฐ€ ๊ฒฐํ•ฉํ•  ๋•Œ ์ ์–ด๋„ ํ•˜๋‚˜ ์ด์ƒ์˜ ๋…ผํ•ญ์„ ๊ณต์œ ํ•˜๋ฉฐ ํ•˜๋‚˜์˜ ๋‹จ์ผ ์‚ฌ๊ฑด์„ ๋‚˜ํƒ€๋‚ธ๋‹ค๋Š” ๊ฒƒ์ด๋‹ค. ์—ฌ๊ธฐ์„œ๋Š” ์‹ฌ์ธต์ ์ธ ํ†ต์‚ฌ ๊ตฌ์กฐ๋ฅผ ์ƒ์ •ํ•˜์ง€๋Š” ์•Š์ง€๋งŒ ๋™์‚ฌ๊ฐ€ ๊ฐ–๋Š” ์˜๋ฏธ ๊ตฌ์กฐ๋ฅผ ๋ฐ˜์˜ํ•˜๋Š” ํ‘œ๋ฉด์  ํ†ต์‚ฌ ๊ตฌ์กฐ๋กœ, ๋…ผํ•ญ ๊ตฌ์กฐ์˜ ํ•ฉ์„ฑ ๋ฐฉ์‹์„ ๋จผ์ € ์ œ์‹œํ•œ๋‹ค. ๋‹ค์Œ์œผ๋กœ ์ˆ ์–ด ๋ถ„ํ•ด(predicate decomposition) ๋ฐฉ์‹์„ ์ด์šฉํ•œ Rappaport Hovav & Levin(19982010)์‹์˜ ์‚ฌ๊ฑด ๊ตฌ์กฐ๋ฅผ ํ†ตํ•ด ๋™์‚ฌ๊ฐ€ ๊ฐ–๋Š” ์˜๋ฏธ ๊ตฌ์กฐ๋ฅผ ํ‘œ์ƒํ•˜์—ฌ ๊ฐ€๋‹ค/์˜ค๋‹ค, ์ฃผ๋‹ค๊ฐ€ ๋‹ค๋ฅธ ๋™์‚ฌ๋“ค๊ณผ ์—ฐ์‡„๋™์‚ฌ ๊ตฌ๋ฌธ์„ ์ด๋ฃฐ ๋•Œ, ๋‘ ๋™์‚ฌ์˜ ํ•˜์œ„ ์‚ฌ๊ฑด์ด ์–ด๋–ป๊ฒŒ ํ•˜๋‚˜์˜ ์‚ฌ๊ฑด์œผ๋กœ ํ•ฉ์„ฑ๋˜๋Š”์ง€๋ฅผ ๋ฐํžŒ๋‹ค. ๊ฐ€๋‹ค/์˜ค๋‹ค, ์ฃผ๋‹ค๋Š” ์ฐฉ์  ๋…ผํ•ญ์„ ํ•„์ˆ˜ ์„ฑ๋ถ„์œผ๋กœ ์ทจํ•˜๋ฉฐ ์ฒ˜์†Œ ๋ณ€ํ™”๋ผ๋Š” ๊ฒฐ๊ณผ ์ƒํƒœ๋ฅผ ํ•˜์œ„ ์‚ฌ๊ฑด์œผ๋กœ ๊ฐ–๋Š” ์ฒ˜์†Œ ์ด๋™ ์‚ฌ๊ฑด์„ ๋‚˜ํƒ€๋‚ธ๋‹ค. ์ด๋Ÿฌํ•œ ๊ฐ€๋‹ค/์˜ค๋‹ค, ์ฃผ๋‹ค๊ฐ€ ๋‹ค๋ฅธ ๋™์‚ฌ๋“ค๊ณผ ๊ฒฐํ•ฉํ•˜์—ฌ ํ•˜๋‚˜์˜ ๋ฌธ์žฅ์„ ๊ตฌ์„ฑํ•  ๋•Œ, ๊ฐ€๋‹ค/์˜ค๋‹ค๋Š” ๊ธฐ๋ณธ์ ์œผ๋กœ ์ฃผ์–ด(ํ–‰๋™์ฃผ)๋ฅผ, ์ฃผ๋‹ค๋Š” ์ฃผ์–ด(ํ–‰๋™์ฃผ)์™€ ๋ชฉ์ ์–ด(๋Œ€์ƒ)๋ฅผ ๊ณต์œ ํ•œ๋‹ค. ์—ฌ๊ธฐ์„œ๋Š” ๊ฐ€๋‹ค/์˜ค๋‹ค, ์ฃผ๋‹ค๊ฐ€ ์ทจํ•˜๋Š” ์ฐฉ์  ๋…ผํ•ญ์ด ํ•ฉ์„ฑ๋˜๋Š” ๋ฐฉ์‹์— ์ฃผ๋ชฉํ•œ๋‹ค. ๊ทธ๋Ÿฐ๋ฐ ๊ฐ€๋‹ค/์˜ค๋‹ค๋Š” ์ฒ˜์†Œ ๋…ผํ•ญ์˜ [ยฑ์ฒ˜์†Œ ๋ณ€ํ™” ]์˜ ์†์„ฑ์— ๋”ฐ๋ผ ๋‹ค๋ฅธ ๋™์‚ฌ์™€์˜ ํ•ฉ์„ฑ ์—ฌ๋ถ€๊ฐ€ ๊ฒฐ์ •๋˜๊ฑฐ๋‚˜ ํ•ฉ์„ฑ ์–‘์ƒ์ด ๋‹ฌ๋ผ์ง„๋‹ค. ์ฃผ๋‹ค๋Š” V1์˜ ์ฐฉ์  ๋…ผํ•ญ ์—ฌ๋ถ€, V1์˜ ๋Œ€์ƒ ๋…ผํ•ญ์ด ๊ตฌ์ฒด๋ฌผ์ธ๊ฐ€ ๋˜๋Š” ์ถ”์ƒ๋ฌผ์ธ๊ฐ€, ๊ทธ๋Ÿฌํ•œ ๋Œ€์ƒ ๋…ผํ•ญ์ด ์ด๋™์ด๋‚˜ ์†Œ์œ  ๋ณ€ํ™”์˜ ๊ฐ€๋Šฅ์„ฑ์ด ์žˆ๋Š” ๊ฒƒ์ธ๊ฐ€์— ๋”ฐ๋ผ ์ฐฉ์  -์—๊ฒŒ ๋…ผํ•ญ์˜ ์‹คํ˜„ ์—ฌ๋ถ€์™€ ํ•ฉ์„ฑ ์œ ํ˜•์ด ๋‹ฌ๋ผ์ง„๋‹ค. ์—ฌ๊ธฐ์„œ๋Š” ๊ฐ€๋‹ค/์˜ค๋‹ค, ์ฃผ๋‹ค๊ฐ€ ๋‹ค๋ฅธ ๋™์‚ฌ๋“ค๊ณผ ๊ฒฐํ•ฉํ•˜์—ฌ ์—ฐ์‡„๋™์‚ฌ ๊ตฌ๋ฌธ์„ ์ด๋ฃฐ ๋•Œ ๋…ผํ•ญ ๊ตฌ์กฐ์™€ ์‚ฌ๊ฑด ๊ตฌ์กฐ๊ฐ€ ํ•ฉ์„ฑ๋˜๋Š” ๋ฐฉ์‹์„ ํฌ๊ฒŒ (1) ๋ณ‘๋ ฌ์  ํ•ฉ์„ฑ ์œ ํ˜•(Summation Type), (2) ํ†ตํ•ฉ์  ํ•ฉ์„ฑ ์œ ํ˜•(Unification Type), (3) ํ˜ผํ•ฉ์  ํ•ฉ์„ฑ ์œ ํ˜•(Mixed Composition Type)์œผ๋กœ ๊ตฌ๋ถ„ํ•œ๋‹ค. ์‚ฌ๊ฑด ๊ตฌ์กฐ์—์„œ ๋ณ‘๋ ฌ์  ํ•ฉ์„ฑ ์œ ํ˜•์€ (1-1) ๋ถ€๊ฐ€์  ๋ณ‘๋ ฌ ํ•ฉ์„ฑ๊ณผ (1-2) ๋ถ€๋ถ„์  ๋ณ‘๋ ฌ ํ•ฉ์„ฑ์œผ๋กœ, ํ†ตํ•ฉ์  ํ•ฉ์„ฑ ์œ ํ˜•์€ (2-1) ์™„์ „ ํ†ตํ•ฉ, (2-2) ๋‚ดํฌ ํ†ตํ•ฉ, (2-3) ๋ถ€๋ถ„ ํก์ˆ˜ ํ†ตํ•ฉ ์œ ํ˜•์œผ๋กœ ์„ธ๋ถ„ํ™”๋œ๋‹ค. ๋ณ‘๋ ฌ์  ํ•ฉ์„ฑ ์œ ํ˜•์€ V1์ด V2์™€ ๊ฒฐํ•ฉํ•˜๋ฉด์„œ ๋™์ผํ•œ ๋…ผํ•ญ์€ ํ•˜๋‚˜๋กœ ํ•ฉ์น˜๋˜ V2์— ์˜ํ•ด ์ฐฉ์  ๋…ผํ•ญ๊ณผ ์‚ฌ๊ฑด ๊ตฌ์กฐ ๋‚ด์— ๋ผ๋Š” ๊ฒฐ๊ณผ ์ƒํƒœ๊ฐ€ ํฌํ•จ๋œ ํ•˜์œ„ ์‚ฌ๊ฑด์„ ์ƒˆ๋กœ์ด ๋„์ž…ํ•˜๊ฒŒ ๋œ๋‹ค. ์ด๋Ÿฌํ•œ ์œ ํ˜•์€ V2์— ์˜ํ•ด ์—ฐ์‡„๋™์‚ฌ ๊ตฌ๋ฌธ์˜ ๋…ผํ•ญ ๊ตฌ์กฐ์™€ ์‚ฌ๊ฑด ๊ตฌ์กฐ๊ฐ€ ์ฃผ๋„์ ์œผ๋กœ ๊ฒฐ์ •๋˜๋Š” ๊ฒฝ์šฐ๋ผ ํ•  ์ˆ˜ ์žˆ๋‹ค. ํ†ตํ•ฉ์  ํ•ฉ์„ฑ ์œ ํ˜•์€ V1๊ณผ V2๊ฐ€ ์ทจํ•˜๋Š” ๋™์ผํ•œ ์†์„ฑ์˜ ์ฒ˜์†Œ ๋…ผํ•ญ๊ณผ ๋™์ผํ•˜๊ฑฐ๋‚˜ ํ•จ์˜ ๊ด€๊ณ„์— ์žˆ๋Š” ์‚ฌ๊ฑด ํ•จ์ˆ˜(event function) ๋ฐ ๊ฒฐ๊ณผ ์ƒํƒœ๋ฅผ ๋‚ดํฌํ•œ ํ•˜์œ„ ์‚ฌ๊ฑด์„ ํ•˜๋‚˜๋กœ ํ†ตํ•ฉํ•˜์—ฌ ์—ฐ์‡„๋™์‚ฌ ๊ตฌ๋ฌธ์„ ์ด๋ฃจ๋Š” ๊ฒฝ์šฐ์ด๋‹ค. ํ‘œ๋ฉด์ ์œผ๋กœ๋Š” V1์˜ ๋…ผํ•ญ ๊ตฌ์กฐ๊ฐ€ V2์™€ ๊ฒฐํ•ฉํ•œ ํ›„์—๋„ ๋™์ผํ•˜๊ฒŒ ์œ ์ง€๋˜๋Š” ๊ฒƒ์œผ๋กœ ๋ณด์—ฌ V2๋ณด๋‹ค๋Š” V1์ด ์ฃผ๋„์ ์ธ ์—ญํ• ์„ ํ•˜๋Š” ์œ ํ˜•์ด๋ผ๊ณ  ํ•  ์ˆ˜ ์žˆ๋‹ค. ํ˜ผํ•ฉ์  ํ•ฉ์„ฑ ์œ ํ˜•์— ์†ํ•˜๋Š” ๋ถ€๋ฅ˜๋“ค์€ V1๊ณผ V2์˜ ์ฒ˜์†Œ ๋…ผํ•ญ์˜ ์†์„ฑ์ด ์ƒ์ถฉ๋˜์–ด ํ•˜๋‚˜๋กœ ํ†ตํ•ฉ๋˜์ง€ ์•Š๊ฑฐ๋‚˜ V2๊ฐ€ ๊ฒฐํ•ฉ๋œ ํ˜•ํƒœ๋กœ๋งŒ ์“ฐ์ด๋Š” ๋“ฑ ์‚ฌ๊ฑด ๊ตฌ์กฐ์ƒ์—์„œ ํ†ตํ•ฉ๋˜์ง€ ์•Š๋Š” ๋…ผํ•ญ๋“ค์€ ๋…ผํ•ญ ๊ตฌ์กฐ์ƒ์—์„œ๋„ ํ—ˆ์šฉ๋˜์ง€ ์•Š๋Š” ๊ฒฝ์šฐ๋“ค์ด๋‹ค. ๋ช‡๋ช‡ ๋…ผํ•ญ ๊ตฌ์กฐ์™€ ์‚ฌ๊ฑด ๊ตฌ์กฐ์˜ ํ•ฉ์„ฑ ๋ฐฉ์‹์ด ๋ฐ˜๋“œ์‹œ ์ผ์น˜ํ•˜์ง€ ์•Š๋Š” ๊ฒฝ์šฐ๋„ ๋‚˜ํƒ€๋‚˜์ง€๋งŒ ๋…ผํ•ญ ๊ตฌ์กฐ์— ์ƒˆ๋กœ์ด ๋„์ž…๋œ ๋…ผํ•ญ์€ ์‚ฌ๊ฑด ๊ตฌ์กฐ ๋‚ด์—์„œ ๋™์ผํ•˜๊ฒŒ ํ•˜๋‚˜์˜ ํ•˜์œ„ ์‚ฌ๊ฑด์œผ๋กœ ๋„์ž…๋˜๋ฉฐ ๋ฐ˜๋Œ€๋กœ ๋…ผํ•ญ ๊ตฌ์กฐ์— ๋ณ€ํ™”๊ฐ€ ์—†์œผ๋ฉด ์‚ฌ๊ฑด ๊ตฌ์กฐ์—์„œ๋„ ์˜ํ–ฅ์„ ๋ฏธ์น˜์ง€ ๋ชปํ•˜๋Š” ๋“ฑ ๋‘ ๋™์‚ฌ๊ฐ€ ๊ฒฐํ•ฉํ•˜์—ฌ ์—ฐ์‡„๋™์‚ฌ ๊ตฌ๋ฌธ์„ ์ด๋ฃจ๋Š” ๊ฒฝ์šฐ์—๋„ ๋…ผํ•ญ ๊ตฌ์กฐ์™€ ์‚ฌ๊ฑด ๊ตฌ์กฐ ์‚ฌ์ด์— ์ผ์ •ํ•œ ๋Œ€์‘ ๊ด€๊ณ„๊ฐ€ ์„ฑ๋ฆฝ๋จ์„ ํ™•์ธํ•  ์ˆ˜ ์žˆ๋‹ค.๊ตญ๋ฌธ ์ดˆ๋ก i 1. ์„œ ๋ก  1 1.1. ์—ฐ๊ตฌ ๋ชฉ์  ๋ฐ ํ•„์š”์„ฑ 1 1.2. ์—ฐ๊ตฌ ๋Œ€์ƒ๊ณผ ๋ฐฉ๋ฒ• 2 1.3. ๋…ผ์˜์˜ ๊ตฌ์„ฑ 4 2. ์—ฐ์‡„๋™์‚ฌ ๊ตฌ๋ฌธ์˜ ์œ ํ˜•๋ก ๊ณผ ํ•œ๊ตญ์–ด์˜ [V1+์–ด+V2] ๊ตฌ๋ฌธ 6 2.1. ์—ฐ์‡„๋™์‚ฌ ๊ตฌ๋ฌธ 6 2.2. ํ•œ๊ตญ์–ด [V1+์–ด+V2] ๊ตฌ๋ฌธ์˜ ํ˜•์„ฑ 17 2.2.1. ์ ‘์†์— ์˜ํ•œ ํ†ต์‚ฌ์  ๊ตฌ์„ฑ ๋˜๋Š” ํ•ฉ์„ฑ๋™์‚ฌ 18 2.2.2. ๋ถ€๋™์‚ฌ ๊ตฌ๋ฌธ 23 2.2.3. ํ•œ๊ตญ์–ด์˜ ์—ฐ์‡„๋™์‚ฌ ๊ตฌ๋ฌธ 30 2.2.3.1. ํ˜•ํƒœ์  ํŠน์„ฑ 31 2.2.3.2. ํ†ต์‚ฌ์  ํŠน์„ฑ 36 2.2.3.2.1. ๋…ผํ•ญ ๊ณต์œ  37 2.2.3.2.2. ๋‹จ์ผ์ ˆ ์†์„ฑ 42 2.2.3.2.3. ์—ฐ์‡„๋™์‚ฌ ๊ตฌ๋ฌธ๊ณผ ํ•ฉ์„ฑ๋™์‚ฌ์˜ ํ†ต์‚ฌ์  ํŠน์„ฑ ๋น„๊ต 54 2.2.3.3. ์˜๋ฏธ์  ํŠน์„ฑ 62 3. ๋™์‚ฌ์˜ ๋…ผํ•ญ ๊ตฌ์กฐ์™€ ์‚ฌ๊ฑด ๊ตฌ์กฐ 68 3.1. ๋…ผํ•ญ ๊ตฌ์กฐ์™€ ์‚ฌ๊ฑด ๊ตฌ์กฐ 68 3.2. ๊ฐ€๋‹ค/์˜ค๋‹ค์˜ ๋…ผํ•ญ ๊ตฌ์กฐ์™€ ์‚ฌ๊ฑด ๊ตฌ์กฐ 73 3.3. ์ฃผ๋‹ค์˜ ๋…ผํ•ญ ๊ตฌ์กฐ์™€ ์‚ฌ๊ฑด ๊ตฌ์กฐ 79 3.3.1. ์˜์–ด give์˜ ๋…ผํ•ญ ๊ตฌ์กฐ์™€ ์‚ฌ๊ฑด ๊ตฌ์กฐ 79 3.3.2. ํ•œ๊ตญ์–ด ์ฃผ๋‹ค์˜ ๋…ผํ•ญ ๊ตฌ์กฐ์™€ ์‚ฌ๊ฑด ๊ตฌ์กฐ 81 3.4. [V1+์–ด+V2{๊ฐ€๋‹ค/์˜ค๋‹ค, ์ฃผ๋‹ค}] ์—ฐ์‡„๋™์‚ฌ ๊ตฌ๋ฌธ์˜ ํ˜•์„ฑ 86 4. ํ•œ๊ตญ์–ด ์—ฐ์‡„๋™์‚ฌ ๊ตฌ๋ฌธ์˜ ๋…ผํ•ญ ๊ตฌ์กฐ ํ•ฉ์„ฑ ์œ ํ˜• 94 4.1. ์—ฐ์‡„๋™์‚ฌ ๊ตฌ๋ฌธ์˜ ๋…ผํ•ญ ๊ตฌ์กฐ ํ•ฉ์„ฑ ๋ฐฉ์‹๊ณผ ์œ ํ˜• 94 4.1.1. ๋…ผํ•ญ ๊ตฌ์กฐ์™€ ์˜๋ฏธ์—ญ ์œ„๊ณ„ ๊ตฌ์กฐ 94 4.1.2. ๊ฐ€๋‹ค/์˜ค๋‹ค, ์ฃผ๋‹ค ์—ฐ์‡„๋™์‚ฌ ๊ตฌ๋ฌธ์˜ ๋…ผํ•ญ ๊ตฌ์กฐ ํ•ฉ์„ฑ ๊ทœ์น™ 101 4.2. [V1+์–ด+V2{๊ฐ€๋‹ค/์˜ค๋‹ค}] ์—ฐ์‡„๋™์‚ฌ ๊ตฌ๋ฌธ์˜ ๋…ผํ•ญ ๊ตฌ์กฐ ํ•ฉ์„ฑ ์œ ํ˜• 102 4.2.1. ๋ณ‘๋ ฌ์  ํ•ฉ์„ฑ ์œ ํ˜• 102 4.2.2. ํ†ตํ•ฉ์  ํ•ฉ์„ฑ ์œ ํ˜• 109 4.2.2.1. ์ฃผ์–ด ๋…ผํ•ญ๊ณผ ์ฒ˜์†Œ ๋…ผํ•ญ ๊ณต์œ  110 4.2.2.2. V1์˜ ๋ชฉ์ ์–ด(๋Œ€์ƒ)-V2์˜ ์ฃผ์–ด(๋Œ€์ƒ) ๋…ผํ•ญ๊ณผ ์ฒ˜์†Œ ๋…ผํ•ญ ๊ณต์œ  120 4.2.3. ํ˜ผํ•ฉ์  ํ•ฉ์„ฑ ์œ ํ˜• 123 4.2.3.1. [V1] - -๋ฅผ: [ํ–‰๋กœ] 123 4.2.3.2. [V1+V2] - -๋ฅผ: [ํ–‰๋กœ] 124 4.2.4. ์ •๋ฆฌ: [V1+์–ด+V2{๊ฐ€๋‹ค/์˜ค๋‹ค}]์˜ ๋…ผํ•ญ ๊ตฌ์กฐ ํ•ฉ์„ฑ ์œ ํ˜• 126 4.3. [V1+์–ด+V2{์ฃผ๋‹ค}] ์—ฐ์‡„๋™์‚ฌ ๊ตฌ๋ฌธ์˜ ๋…ผํ•ญ ๊ตฌ์กฐ ํ•ฉ์„ฑ ์œ ํ˜• 130 4.3.1. ๋ณ‘๋ ฌ์  ํ•ฉ์„ฑ ์œ ํ˜• 130 4.3.2. ํ†ตํ•ฉ์  ํ•ฉ์„ฑ ์œ ํ˜• 141 4.3.2.1. ์™„์ „ ํ†ตํ•ฉ 141 4.3.2.2. ๋‚ดํฌ ํ†ตํ•ฉ 145 4.3.3. ํ˜ผํ•ฉ์  ํ•ฉ์„ฑ ์œ ํ˜• 146 4.3.3.1. V1์˜ ๊ตฌ๋ฌธ ์œ ์ง€: ์ˆ˜ํ˜œ ๊ตฌ๋ฌธ 146 4.3.3.2. -(์–ด)์„œ ๊ตฌ๋ฌธ 152 4.3.3.3. V2์— ์˜ํ•ด ๋ณ„๋„์˜ ์ˆ˜ํ˜œ์ž๋ฅผ ์š”๊ตฌํ•˜๋Š” ๊ตฌ๋ฌธ 157 4.3.3.4. V1์˜ ํ–‰๋™์ฃผ๊ฐ€ V2์˜ ์ฐฉ์ ์ด ๋˜๋Š” ๊ฒฝ์šฐ 158 4.3.3.5. ์–ดํœ˜์  ํ•ฉ์„ฑ์–ด 159 4.3.4. ์ •๋ฆฌ: [V1+์–ด+V2{์ฃผ๋‹ค}]์˜ ๋…ผํ•ญ ๊ตฌ์กฐ ํ•ฉ์„ฑ ์œ ํ˜• 162 5. ํ•œ๊ตญ์–ด ์—ฐ์‡„๋™์‚ฌ ๊ตฌ๋ฌธ์˜ ์‚ฌ๊ฑด ๊ตฌ์กฐ ํ•ฉ์„ฑ ์œ ํ˜• 166 5.1. ๊ฐ€๋‹ค/์˜ค๋‹ค, ์ฃผ๋‹ค ์—ฐ์‡„๋™์‚ฌ ๊ตฌ๋ฌธ์˜ ์‚ฌ๊ฑด ๊ตฌ์กฐ ํ•ฉ์„ฑ 166 5.1.1. ์‚ฌ๊ฑด ๊ตฌ์กฐ์˜ ํ•ฉ์„ฑ ๊ทœ์น™๊ณผ ํ•ฉ์„ฑ ์œ ํ˜• 168 5.1.2. ์‚ฌ๊ฑด ํ•จ์ˆ˜์™€ ์—ฐ์‚ฐ์ž ๋ชฉ๋ก 177 5.1.3. ์‚ฌ๊ฑด ๊ตฌ์กฐ์˜ ๊ฒฐํ•ฉ ์œ ํ˜• ๊ตฌ๋ถ„: -์–ด์™€ -(์–ด)์„œ์˜ ๊ต์ฒด 178 5.2. [V1+์–ด+V2{๊ฐ€๋‹ค/์˜ค๋‹ค}] ์—ฐ์‡„๋™์‚ฌ ๊ตฌ๋ฌธ์˜ ์‚ฌ๊ฑด ๊ตฌ์กฐ ํ•ฉ์„ฑ ์œ ํ˜• 180 5.2.1. ํ†ตํ•ฉ์  ํ•ฉ์„ฑ ์œ ํ˜• 180 5.2.1.1. ์™„์ „ ํ†ตํ•ฉ 180 5.2.1.2. ๋‚ดํฌ ํ†ตํ•ฉ 183 5.2.2. ๋ณ‘๋ ฌ์  ํ•ฉ์„ฑ ์œ ํ˜• 187 5.2.2.1. ๋ถ€๋ถ„์  ๋ณ‘๋ ฌ ํ•ฉ์„ฑ 188 5.2.2.2. ๋ถ€๊ฐ€์  ๋ณ‘๋ ฌ ํ•ฉ์„ฑ 196 5.2.3. ํ˜ผํ•ฉ์  ํ•ฉ์„ฑ ์œ ํ˜• 203 5.2.4. ์ •๋ฆฌ: [V1+์–ด+V2{๊ฐ€๋‹ค/์˜ค๋‹ค}]์˜ ์‚ฌ๊ฑด ๊ตฌ์กฐ ํ•ฉ์„ฑ ์œ ํ˜• 208 5.3. [V1+์–ด+V2{์ฃผ๋‹ค}] ์—ฐ์‡„๋™์‚ฌ ๊ตฌ๋ฌธ์˜ ์‚ฌ๊ฑด ๊ตฌ์กฐ ํ•ฉ์„ฑ ์œ ํ˜• 209 5.3.1. ํ†ตํ•ฉ์  ํ•ฉ์„ฑ ์œ ํ˜• 209 5.3.1.1. ์™„์ „ ํ†ตํ•ฉ 209 5.3.1.2. ๋‚ดํฌ ํ†ตํ•ฉ 212 5.3.1.3. ๋ถ€๋ถ„ ํก์ˆ˜ ํ†ตํ•ฉ 213 5.3.2. ๋ณ‘๋ ฌ์  ํ•ฉ์„ฑ ์œ ํ˜•: ๋ถ€๋ถ„์  ๋ณ‘๋ ฌ ํ•ฉ์„ฑ 217 5.3.3. ํ˜ผํ•ฉ์  ํ•ฉ์„ฑ ์œ ํ˜• 221 5.3.3.1. V1์˜ ์‚ฌ๊ฑด ๊ตฌ์กฐ ์œ ์ง€, V2 ์ฃผ๋‹ค์˜ ์ˆ˜ํ˜œ ์‚ฌ๊ฑด 221 5.3.3.2. -(์–ด)์„œ ๊ตฌ๋ฌธ 226 5.3.3.3. ์˜๋ฏธํ•ญ์˜ ์†์„ฑ ์ „ํ™˜ ๋˜๋Š” ์ผ์น˜ ์—ฌ๋ถ€ 228 5.3.4. ์ •๋ฆฌ: [V1+์–ด+V2{์ฃผ๋‹ค}]์˜ ์‚ฌ๊ฑด ๊ตฌ์กฐ ํ•ฉ์„ฑ ์œ ํ˜• 230 6. ํ•œ๊ตญ์–ด ์—ฐ์‡„๋™์‚ฌ ๊ตฌ๋ฌธ์˜ ๋…ผํ•ญ ๊ตฌ์กฐ์™€ ์‚ฌ๊ฑด ๊ตฌ์กฐ์˜ ์ƒ๊ด€์„ฑ 234 6.1. ๋…ผํ•ญ ๊ตฌ์กฐ์™€ ์‚ฌ๊ฑด ๊ตฌ์กฐ์˜ ๋Œ€์‘ ๊ด€๊ณ„ 234 6.2. [V1+์–ด+V2{๊ฐ€๋‹ค/์˜ค๋‹ค}] ์—ฐ์‡„๋™์‚ฌ ๊ตฌ๋ฌธ์˜ ๋…ผํ•ญ ๊ตฌ์กฐ์™€ ์‚ฌ๊ฑด ๊ตฌ์กฐ์˜ ์ƒ๊ด€์„ฑ 238 6.2.1. ํ•ฉ์„ฑ ๋ฐฉ์‹์ด ์ผ์น˜ํ•˜๋Š” ๋ถ€๋ฅ˜ 240 6.2.2. ํ•ฉ์„ฑ ๋ฐฉ์‹์ด ์ผ์น˜ํ•˜์ง€ ์•Š๋Š” ๋ถ€๋ฅ˜ 241 6.3. [V1+์–ด+V2{์ฃผ๋‹ค}] ์—ฐ์‡„๋™์‚ฌ ๊ตฌ๋ฌธ์˜ ๋…ผํ•ญ ๊ตฌ์กฐ์™€ ์‚ฌ๊ฑด ๊ตฌ์กฐ์˜ ์ƒ๊ด€์„ฑ 243 6.3.1. ํ•ฉ์„ฑ ๋ฐฉ์‹์ด ์ผ์น˜ํ•˜๋Š” ๋ถ€๋ฅ˜ 245 6.3.2. ํ•ฉ์„ฑ ๋ฐฉ์‹์ด ์ผ์น˜ํ•˜์ง€ ์•Š๋Š” ๋ถ€๋ฅ˜ 247 6.4. ์ •๋ฆฌ 248 7. ๊ฒฐ๋ก  251 ์ฐธ ๊ณ  ๋ฌธ ํ—Œ 255 Appendix 267 Abstract 274Docto

    ๊ตญํšŒ์˜ ์˜ˆ์‚ฐ์•ˆ ์‹ฌ์‚ฌ๊ณผ์ •์— ์žˆ์–ด์„œ ์˜์‚ฌ๊ฒฐ์ • ์š”์ธ ์—ฐ๊ตฌ

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    ํ•™์œ„๋…ผ๋ฌธ (์„์‚ฌ)-- ์„œ์šธ๋Œ€ํ•™๊ต ๋Œ€ํ•™์› : ํ–‰์ •ํ•™๊ณผ(์ •์ฑ…ํ•™์ „๊ณต), 2017. 2. ๊ธˆํ˜„์„ญ.๊ตญํšŒ์˜ ์˜ˆ์‚ฐ์‹ฌ์˜๋Š” ๊ตญ๋ฏผ์˜ ๋Œ€ํ‘œ๊ธฐ๊ด€์ธ ์ž…๋ฒ•๋ถ€์— ์˜ํ•ด ํ–‰์ •๋ถ€๊ฐ€ ์ œ์ถœํ•œ ์˜ˆ์‚ฐ์•ˆ์„ ๊ตญ๊ฐ€์  ์ฐจ์›์—์„œ ์‹ฌ์˜โ€ค์˜๊ฒฐํ•˜๋Š” ๊ฒƒ์œผ๋กœ ๊ตญํšŒ์˜์›, ์˜ˆ์‚ฐ๋‹น๊ตญ, ์ •๋ถ€๋ถ€์ฒ˜ ๋ฐ ํ•ด๋‹น ์˜ˆ์‚ฐ์•ˆ์„ ๋‘˜๋Ÿฌ์‹ผ ์ดํ•ด๊ด€๊ณ„์ธ๋“ค์ด ์ฐธ์—ฌํ•˜๊ณ  ์žˆ๋‹ค. ์ด ๊ณผ์ •์—์„œ ๊ฐ€์žฅ ํฐ ์˜ํ–ฅ๋ ฅ์„ ๋ฐœํœ˜ํ•˜๋Š” ์ฃผ์ฒด๋Š” ๊ตญ๋ฏผ์œผ๋กœ๋ถ€ํ„ฐ ๋Œ€ํ‘œ์„ฑ์„ ๋ถ€์—ฌ๋ฐ›๊ณ  ์˜ˆ์‚ฐ์‹ฌ์˜์— ์ฐธ์—ฌํ•˜๋Š” ๊ตญํšŒ์˜์›์ด๋‹ค. ๊ทธ๋Ÿฌ๋‚˜ ๊ตญํšŒ์˜์›์€ ์ „๋ฌธ์„ฑ๋ณด๋‹ค๋Š” ๊ฐ๊ณ„๊ฐ์ธต์˜ ๋Œ€ํ‘œ์„ฑ์ด ์šฐ์„ ์‹œ๋˜๊ณ , ์„ ๊ฑฐ๋ฅผ ํ†ตํ•ด 4๋…„๋งˆ๋‹ค ์ฃผ๊ธฐ์ ์œผ๋กœ ๊ต์ฒด๋˜๊ธฐ ๋•Œ๋ฌธ์— ํŠน์ • ์ •์ฑ…์˜์—ญ์— ๋Œ€ํ•œ ์ „๋ฌธ์ ์ธ ์ง€์‹๊ณผ ๊ฒฝํ—˜์„ ์ถ•์ ํ•จ์— ์žˆ์–ด ๊ทผ๋ณธ์ ์ธ ํ•œ๊ณ„๊ฐ€ ์žˆ๋‹ค. ์ด๋Ÿฌํ•œ ์˜ˆ์‚ฐ๊ณผ์ •์—์„œ์˜ ๋ณธ์งˆ์ ์ธ ์ •์น˜์„ฑ์„ ๊ทน๋ณตํ•˜๊ณ  ์ „๋ฌธ์ ์ด๊ณ  ๊ฐ๊ด€์ ์ธ ์˜ˆ์‚ฐ์•ˆ ์‹ฌ์‚ฌ๋ฅผ ์ง€์›ํ•˜๊ธฐ ์œ„ํ•œ ์ œ๋„์  ์žฅ์น˜๋กœ์„œ ๊ตญํšŒ๋Š” ๊ตญํšŒ ์˜ˆ์‚ฐ๊ฒฐ์‚ฐํŠน๋ณ„์œ„์›ํšŒ ์ˆ˜์„์ „๋ฌธ์œ„์›๊ณผ ๊ตญํšŒ์˜ˆ์‚ฐ์ •์ฑ…์ฒ˜์™€ ๊ฐ™์€ ๊ตญํšŒ ์ง€์› ์กฐ์ง์„ ๋‘๊ณ  ์žˆ๋‹ค. ์ด๋“ค ์กฐ์ง์€ ๊ตญํšŒ์˜์›์—๊ฒŒ ์˜ˆ์‚ฐ์•ˆ ์‹ฌ์‚ฌ์— ํ•„์š”ํ•œ ์žฌ์ •์ •๋ณด๋ฅผ ์ œ๊ณตํ•˜๊ณ , ํšŒ์˜ ๋ณด์ขŒ ๋ฐ ๋ณด๊ณ ์„œ ์ž‘์„ฑโ€ค๋ฐœ๊ฐ„ ๋“ฑ์„ ํ†ตํ•ด ์งโ€ค๊ฐ„์ ‘์ ์œผ๋กœ ์˜ˆ์‚ฐ๊ณผ์ •์— ์ฐธ์—ฌํ•˜๊ณ  ์žˆ๋‹ค. ์ด๋“ค ์ง€์›์กฐ์ง์€ ๊ตญํšŒ์˜์›์ด๋‚˜ ํ–‰์ •๋ถ€์ฒ˜์™€ ๊ฐ™์ด ์ง์ ‘์ ์œผ๋กœ ์˜ˆ์‚ฐ์˜ ๊ณผ์ • ์ „๋ฉด์— ๋‚˜์„œ ์˜ˆ์‚ฐ์•ˆ์„ ํŽธ์„ฑํ•˜๊ฑฐ๋‚˜ ์‹ฌ์‚ฌํ•  ์ˆ˜ ์žˆ๋Š” ๊ถŒํ•œ์€ ์—†์ง€๋งŒ, ๊ตญํšŒ์˜์›์˜ ์ง€์›์กฐ์ง ํ™œ์šฉ์—ฌ๋ถ€์™€ ์ •๋„์— ๋”ฐ๋ผ ์˜ˆ์‚ฐ๊ณผ์ •์—์„œ ์œ ์˜๋ฏธํ•œ ์—ญํ• ์„ ํ•  ์ˆ˜ ์žˆ๋‹ค. ๊ทธ๋ ‡์ง€๋งŒ ๊ตญํšŒ์˜ ํ•ต์‹ฌ์ ์ธ ์˜ˆ์‚ฐ์•ˆ ์‹ฌ์˜๊ณผ์ •์€ ๋น„๊ณต๊ฐœ๋กœ ์ด๋ฃจ์–ด์ ธ ๊ตฌ์ฒด์ ์ธ ์ž๋ฃŒ์— ๋Œ€ํ•œ ์ ‘๊ทผ์ด ์šฉ์ดํ•˜์ง€ ์•Š๊ณ , ์ด๋“ค ์ง€์›์กฐ์ง์˜ ์˜ํ–ฅ๋ ฅ์€ ๊ณต์‹์ ์œผ๋กœ ํŒŒ์•…ํ•˜๊ธฐ ์–ด๋ ต๊ธฐ ๋•Œ๋ฌธ์— ์˜ˆ์‚ฐ์•ˆ ์‹ฌ์‚ฌ๊ณผ์ •์—์„œ ์ง€์›์กฐ์ง์ด ๋ณธ๋ž˜์˜ ์ทจ์ง€์— ๋”ฐ๋ผ ๊ธฐ๋Šฅ์„ ๋‹คํ•˜๊ณ  ์žˆ๋Š”์ง€๋ฅผ ํ™•์ธํ•˜๊ธฐ๊ฐ€ ์‰ฝ์ง€ ์•Š๋‹ค. ๋ณธ ๋…ผ๋ฌธ์€ ์‹ค์งˆ์ ์œผ๋กœ ์˜ˆ์‚ฐ์•ˆ์˜ ์‹ฌ์˜๊ฐ€ ์ด๋ฃจ์–ด์ง€๋Š” ๊ตญํšŒ ์˜ˆ์‚ฐ๊ฒฐ์‚ฐํŠน๋ณ„์œ„์›ํšŒ ์˜ˆ์‚ฐ์•ˆ์กฐ์ •์†Œ์œ„์›ํšŒ์˜ ์‹ฌ์‚ฌ ๊ธฐ์ดˆ์ธ ์ด๋ฅธ ๋ฐ”, ์†Œ์œ„์ž๋ฃŒ์™€ ๊ตญํšŒ์ง€์›์กฐ์ง์˜ ๋ฐœ๊ฐ„๋ณด๊ณ ์„œ ๋“ฑ์„ ๋ฐ”ํƒ•์œผ๋กœ ํ•œ ์„ธ๋ถ€์‚ฌ์—… ๋‹จ์œ„์˜ ์ฆ๊ฐ ์˜์‚ฌ๊ฒฐ์ • ๋ถ„์„์„ ํ†ตํ•ด ๊ตญํšŒ์˜์›์˜ ์˜ˆ์‚ฐ์•ˆ ์ฆ๊ฐ๊ฒฐ์ •์— ์˜ํ–ฅ์„ ๋ฏธ์น˜๋Š” ์š”์ธ๊ณผ ๊ตญํšŒ์ง€์›์กฐ์ง์ด ์˜ˆ์‚ฐ์•ˆ ์‹ฌ์‚ฌ๊ณผ์ •์— ์žˆ์–ด ์ œ ๊ธฐ๋Šฅ์„ ์ˆ˜ํ–‰ํ•˜๊ณ  ์žˆ๋Š”์ง€ ์—ฌ๋ถ€๋ฅผ ํ™•์ธํ•ด ๋ณด๊ณ ์ž ํ•˜์˜€๋‹ค. ์ด๋ฅผ ์œ„ํ•ด ๋‹คํ•ญ ๋กœ์ง€์Šคํ‹ฑ ํšŒ๊ท€๋ถ„์„์„ ํ†ตํ•ด์„œ ๊ตญํšŒ์˜์›์˜ ๋‹นํŒŒ์„ฑ, ์ง€์—ญ์„ฑ, ๋Œ€ํ‘œ์„ฑ๊ณผ ๊ตญํšŒ์ง€์›์กฐ์ง์˜ ์˜๊ฒฌ ์œ ๋ฌด์— ๋”ฐ๋ฅธ ๊ตญํšŒ์˜ ์˜ˆ์‚ฐ์•ˆ ์‹ฌ์‚ฌ ์ดํ›„ ์˜ˆ์‚ฐ์•ˆ ์ฆ๊ฐ ์ˆ˜์ •์—ฌ๋ถ€๋ฅผ ์‚ดํŽด๋ณด์•˜๋‹ค. ๋…๋ฆฝ๋ณ€์ˆ˜๋กœ์„œ ๊ตญํšŒ์˜์›์˜ ๋‹นํŒŒ์„ฑ, ์ง€์—ญ์„ฑ ๋ฐ ๋Œ€ํ‘œ์„ฑ์€ ์˜ˆ์‚ฐ์•ˆ์กฐ์ •์†Œ์œ„์›ํšŒ์—์„œ ์„ธ๋ถ€์‚ฌ์—…์— ๋Œ€ํ•ด ์ฆ๊ฐ์˜๊ฒฌ์„ ์ œ์‹œํ•œ ์˜์›์„ ์—ฌ๋‹น ๋ฐ ์•ผ๋‹น, ์ง€์—ญ๊ตฌ ๋ฐ ๋น„๋ก€๋Œ€ํ‘œ, ๋™์ผ ์ƒ์ž„์œ„ ๋ฐ ๊ทธ ์™ธ ์ƒ์ž„์œ„ ์†Œ์†์˜์›์œผ๋กœ ๊ตฌ๋ถ„ํ•˜์—ฌ ๊ฐ๊ฐ์˜ ์ฐจ์ด๋กœ ์ธก์ •ํ•˜์˜€๊ณ , ๊ตญํšŒ์ง€์›์กฐ์ง์˜ ์˜ํ–ฅ๋ ฅ์€ ์„ธ๋ถ€์‚ฌ์—…์— ๋Œ€ํ•œ ์ง€์›์กฐ์ง์˜ ์ฆ์•ก๋ฐ˜๋Œ€ ์˜๊ฒฌ ์ œ์‹œ์—ฌ๋ถ€๋กœ ์ธก์ •ํ•˜์˜€๋‹ค. ๋ถ„์„ ๊ฒฐ๊ณผ, ํ•ด๋‹น ์‚ฌ์—…์— ๋Œ€ํ•˜์—ฌ ์ฆ์•ก์˜๊ฒฌ์„ ์ œ์‹œํ•œ ์—ฌ๋‹น์˜์›์˜ ์ˆ˜๊ฐ€ ์•ผ๋‹น์˜์›์˜ ์ˆ˜๋ณด๋‹ค ๋งŽ์„์ˆ˜๋ก ์˜ˆ์‚ฐ์•ˆ ์œ ์ง€์— ๋น„ํ•ด ์ฆ์•ก ์ˆ˜์ •๋  ๊ฐ€๋Šฅ์„ฑ์ด ์ •์ ์ธ ๊ฒƒ์œผ๋กœ ๋‚˜ํƒ€๋‚ฌ์œผ๋ฉฐ, ๊ตญํšŒ์ง€์›์กฐ์ง์˜ ์ฆ์•ก๋ฐ˜๋Œ€์˜๊ฒฌ์ด ์—†๋Š” ๊ฒฝ์šฐ๋ณด๋‹ค ์˜๊ฒฌ์„ ์ œ์‹œํ•œ ๊ฒฝ์šฐ ์˜ˆ์‚ฐ์•ˆ ์œ ์ง€๋ณด๋‹ค๋Š” ๊ฐ์•ก ์ˆ˜์ •๋  ๊ฐ€๋Šฅ์„ฑ์ด ๋†’์€ ๊ฒƒ์œผ๋กœ ๋‚˜ํƒ€๋‚ฌ๋‹ค. ์ฆ๋ช…๋œ ๊ฐ€์„ค์˜ ๊ฒฐ๊ณผ๋กœ ์•Œ ์ˆ˜ ์žˆ๋Š” ๊ฒƒ์€ ์˜ˆ์‚ฐ์•ˆ์กฐ์ •์†Œ์œ„์›ํšŒ์˜ ์„ธ๋ถ€์‚ฌ์—… ๋‹จ์œ„ ์˜ˆ์‚ฐ์•ˆ ์‹ฌ์‚ฌ์˜ ์ฆ๊ฐ ๊ฒฐ์ •์— ์žˆ์–ด ์˜์›๋“ค์€ ์†Œ์† ์ •๋‹น์— ์˜ํ–ฅ์„ ๋ฐ›๊ณ  ์žˆ๊ณ , ๊ฐ์•ก์‚ฌ์—…์— ์žˆ์–ด์„œ ๊ตญํšŒ์˜์›์˜ ์ฆ๊ฐ ๊ฒฐ์ •์— ๋Œ€ํ•œ ํ•ฉ๋ฆฌ์„ฑ์˜ ๊ทผ๊ฑฐ๋กœ ์ง€์›์กฐ์ง์˜ ์˜๊ฒฌ์ด ํ™œ์šฉ๋˜๊ณ  ์žˆ๋‹ค. ์ด๋Š” ๊ตญํšŒ์ง€์›์กฐ์ง์ด ์ง์ ‘์ ์ธ ์˜ˆ์‚ฐ์•ˆ ์‹ฌ์‚ฌ์ฃผ์ฒด๋กœ์„œ ์ฆ๊ฐ ๊ฒฐ์ •์„ ํ•  ์ˆ˜๋Š” ์—†์ง€๋งŒ ๊ฐ„์ ‘์ ์œผ๋กœ๋Š” ๊ตญํšŒ์˜์›์˜ ์˜์‚ฌ๊ฒฐ์ •์— ์˜ํ–ฅ์„ ์ฃผ๋Š” ์š”์ธ์œผ๋กœ ์ž‘์šฉํ•˜์—ฌ ์˜ˆ์‚ฐ์•ˆ ์‹ฌ์‚ฌ๊ณผ์ •์—์„œ ์œ ์˜๋ฏธํ•œ ์—ญํ• ์„ ํ•˜๊ณ  ์žˆ์Œ์„ ๋‚˜ํƒ€๋ƒ„์„ ๋ณด์—ฌ์ค€๋‹ค๊ณ  ํ•  ์ˆ˜ ์žˆ๋‹ค.์ œ1์žฅ. ์„œ๋ก  1 ์ œ1์ ˆ. ์—ฐ๊ตฌ๋ชฉ์  ๋ฐ ํ•„์š” 1 ์ œ2์ ˆ. ์—ฐ๊ตฌ์˜ ๋Œ€์ƒ๊ณผ ๋ฒ”์œ„ 4 ์ œ3์ ˆ. ์—ฐ๊ตฌ์˜ ๋ฐฉ๋ฒ• 5 ์ œ2์žฅ. ์ด๋ก ์  ๋…ผ์˜์™€ ์„ ํ–‰์—ฐ๊ตฌ ๊ฒ€ํ†  7 ์ œ1์ ˆ. ์ด๋ก ์  ๋…ผ์˜ 7 1. ๊ตญํšŒ์˜์›์˜ ์˜์‚ฌ๊ฒฐ์ •์— ๊ด€ํ•œ ์ด๋ก  7 2. ๊ตญํšŒ์ง€์›์กฐ์ง์— ๊ด€ํ•œ ์ด๋ก  11 ์ œ2์ ˆ. ์„ ํ–‰์—ฐ๊ตฌ์˜ ๊ฒ€ํ†  16 1. ๊ตญํšŒ์˜ ์˜ˆ์‚ฐ์‹ฌ์˜์— ๊ด€ํ•œ ์š”์ธ ์—ฐ๊ตฌ 16 2. ๊ตญํšŒ์ง€์›์กฐ์ง์˜ ์˜ํ–ฅ๋ ฅ์— ๊ด€ํ•œ ์—ฐ๊ตฌ 19 ์ œ3์žฅ. ๊ตญํšŒ์˜ ์˜ˆ์‚ฐ์•ˆ ์‹ฌ์‚ฌ 21 ์ œ1์ ˆ. ๊ตญํšŒ์˜ ์˜ˆ์‚ฐ์•ˆ ์‹ฌ์‚ฌ ์ ˆ์ฐจ 21 ์ œ2์ ˆ. ๊ตญํšŒ์˜ ์˜ˆ์‚ฐ์•ˆ ์‹ฌ์‚ฌ ๊ด€๋ จ ์ง€์›์กฐ์ง 29 1. ์œ„์›ํšŒ ์ˆ˜์„์ „๋ฌธ์œ„์› 29 2. ๊ตญํšŒ์˜ˆ์‚ฐ์ •์ฑ…์ฒ˜ 31 ์ œ4์žฅ. ์—ฐ๊ตฌ์„ค๊ณ„ ๋ฐ ๋ถ„์„๋ฐฉ๋ฒ• 34 ์ œ1์ ˆ. ์—ฐ๊ตฌ์˜ ๋ถ„์„ํ‹€๊ณผ ๊ฐ€์„ค 34 ์ œ2์ ˆ. ์—ฐ๊ตฌ๋ณ€์ˆ˜ 38 1. ์ข…์†๋ณ€์ˆ˜ 38 2. ๋…๋ฆฝ๋ณ€์ˆ˜ 39 3. ํ†ต์ œ๋ณ€์ˆ˜ 43 ์ œ3์ ˆ. ์ž๋ฃŒ์˜ ์ˆ˜์ง‘ ๋ฐ ๋ถ„์„๋ฐฉ๋ฒ• 45 ์ œ5์žฅ. ์—ฐ๊ตฌ๊ฒฐ๊ณผ 49 ์ œ1์ ˆ. ๊ธฐ์ˆ ํ†ต๊ณ„ ๋ถ„์„ 49 ์ œ2์ ˆ. ๋‹คํ•ญ๋กœ์ง€์Šคํ‹ฑ ํšŒ๊ท€๋ถ„์„ 51 ์ œ3์ ˆ. ๋ถ„์„๊ฒฐ๊ณผ 53 1. ์˜์›์˜ ์˜ํ–ฅ์š”์ธ์— ๋Œ€ํ•œ ๋ถ„์„๊ฒฐ๊ณผ 53 2. ์ง€์›์กฐ์ง์˜ ์˜ํ–ฅ์š”์ธ์— ๋Œ€ํ•œ ๋ถ„์„๊ฒฐ๊ณผ 54 ์ œ6์žฅ. ๊ฒฐ๋ก  56 ์ œ1์ ˆ. ์—ฐ๊ตฌ๊ฒฐ๊ณผ์˜ ์š”์•ฝ 56 ์ œ2์ ˆ. ์˜์˜์™€ ํ•œ๊ณ„ 57 1. ์˜์˜ 57 2. ํ•œ๊ณ„ 60 ์ฐธ๊ณ ๋ฌธํ—Œ 62 Abstract 69Maste

    ๋ถ„์—ด์„ฑ ํšจ๋ชจ์—์„œ ์ฒ  ํก์ˆ˜ ์–ต์ œ ์ „์‚ฌ์ธ์ž Fep1์˜ ํŠน์„ฑ ๋ถ„์„

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    ํ•™์œ„๋…ผ๋ฌธ (๋ฐ•์‚ฌ)-- ์„œ์šธ๋Œ€ํ•™๊ต ๋Œ€ํ•™์› : ์ƒ๋ช…๊ณผํ•™๋ถ€ ๋ฏธ์ƒ๋ฌผํ•™๊ณผ, 2017. 2. ๋…ธ์ •ํ˜œ.Iron is an important cofactor for a wide variety of proteins involved in the major life processes, such as respiration and tricarboxylic acid cycle, DNA replication and repair, and nitrogen fixation. The essential redox-active property of iron that enables facile switch between ferric (Fe3+) and ferrous iron (Fe2+) also renders toxicity via generating reactive oxygen species. Therefore, most organisms are equipped with regulated mechanisms to maintain optimal intracellular levels of iron. In the fission yeast Schizosaccharomyces pombe, two repressors Php4 and Fep1 regulate iron-dependent expression of genes for iron usage/storage and acquisition, respectively. The iron-sensing depends on the CGFS-type monothiol glutaredoxin Grx4 that binds Fe-S cluster in a homodimer or in a heterodimer with a BolA-type protein Fra2. Under iron-rich condition, Grx4 binding to Php4 causes its cytosolic sequestration, resulting in the induction of iron usage/storage genes. Grx4 also binds Fep1 regardless of iron availability. Under iron-starved condition, Grx4 and Fra2 inhibit Fep1 repressor activity, resulting in derepression of iron acquisition genes. Fep1, a GATA-family transcription factor, binds to the promoter regions of genes for iron acquisition, such as reductive iron import (fio1+, frp1+), siderophore transport (str1+, str2+, str3+), and vacuolar transport (abc3+), to avoid iron overload under iron-rich conditions. The N-terminal DNA-binding domain of Fep1 contains four conserved cysteines located between the two zinc finger motifs. It has been demonstrated that the N-terminal 241 aa residues of Fep1 can bind to the target promoters in vivo under iron-replete condition. Through this N-terminal domain, Fep1 is known to interact with the monothiol glutaredoxin domain of Grx4 under iron-starved condition. Under iron-starved condition, Fep1 is released from binding to its target sites, inducing iron uptake genes. Fep1 orthologs are well conserved, especially in the DNA binding domain, across filamentous fungi, such as Ustilago maydis (Urbs1), Neurospora crassa (SRE), Histoplasma capsulatum (Sre1), Aspergillus spp. (SREA), Candida albicans (Sfu1), and Cryptococcus neoformans (Cir1). Even though the function and interaction partners of Fep1 have been elucidated extensively in S. pombe, the molecular basis by which Fep1 is inactivated under iron starvation remains unknown. In this study, to elucidate the mechanism behind iron-sensing by Fep1, I pursued biochemical and spectroscopic analyses of Fep1, in full length and truncated forms, as isolated or reconstituted proteins, with the wild type or substituted cysteine mutations. Evidences are presented that Fep1 binds iron, in the form of Fe-S cluster. Spectroscopic and biochemical analyses of as isolated and reconstituted Fep1 suggest that the dimeric Fep1 binds Fe-S clusters. Iron to acid-labile sulfide stoichiometry of purified Fep1-N238 was roughly 1 : 1 in both before and after chemical reconstitution. Furthermore, the reconstitution of the His-tagged Fep1-N238 requires not only iron donor but also sulfide donor, indicating that Fep1 binds [Fe-S] clusters. The mutation study revealed that the cluster-binding depended on the conserved cysteines located between the two zinc fingers in the DNA binding domain. EPR analyses revealed [Fe-S]-specific peaks indicative of mixed presence of [2Fe-2S], [3Fe-4S], or [4Fe-4S]. Overall, the finding that Fep1 is an Fe-S protein fits nicely with the model that the Fe-S-trafficking Grx4 senses intracellular iron environment and modulates the activity of Fep1.CHAPTER I. INTRODUCTION 1 I.1. Iron homeostasis and disease 2 I.2. Regulation of iron homeostasis in Schizosaccaromyces pombe 3 I.2.1. Fep1, the iron-regulatory GATA-type repressor 3 I.2.2. Php4, a key regulator for iron economy 6 I.2.3. Roles for Grx4 in iron homeostasis in S. pombe 8 I.3. CGFS monothiol glutaredoxins 10 I.3.1. Mitochondrial monodomain Grxs in the maturation of Fe-S protein 13 I.3.2. Nucleo-cytosolic multidomain Grxs in iron metabolism 14 I.4. CGFS monothiol glutaredoxins and BolA proteins 18 I.4.1. Evolutive conservation of the Grx-BolA interaction 18 I.4.2. Roles of Grx3/4 and Fra2 in iron homeostasis in S. cerevisiae 20 I.4.3. Role of Grx4 and Fra2 in iron homeostasis in S. pombe 23 I.5. Biology of Schizosaccharomyces pombe 24 I.5.1. The early research and phylogeny of S. pombe 24 I.5.2. Life cycle of S. pombe 25 I.5.3. Genomic information of S. pombe 27 I.6. Aims of this study 30 CHAPTER II. MATERIALS AND METHODS 32 II.1. Strains and plasmids construction 33 II.2. Protein expression and purification 33 II.3. Transformation of Escherichia coli and Yeast 33 II.4. Western blot analysis 34 II.5. Preparation of apoprotein 34 II.6. Fe-S cluster reconstitution in vitro 35 II.7. Assays for quantification of iron, sulfide and protein 35 II.7.1. Determination of iron concentration 35 II.7.2. Determination of sulfide concentration 36 II.7.3. Determination of protein concentration 36 II.8. EPR spectroscopy 36 CHAPTER III. RESULTS & DISCUSSION 38 III. 1. Purification and UV-visible absorption spectroscopy of full-length Fep1 39 III. 1.1. Purification of full-length Fep1 39 III. 1.2. UV-visible absorption spectroscopy of full-length Fep1 43 III.2. Fep1-N238 binds an Fe-S cluster via four conserved cysteine residues 44 III.2.1. Purification and UV-visible absorption spectroscopy of Fep1-N238 44 III. 2.2. Purification and UV-visible absorption spectroscopy of Fep1 cysteine mutants 53 III. 2.3. Both iron and sulfide are required to reconstitute iron-bound Fep1 56 III. 2.4. Iron and acid labile sulfide content of purified Fep1-N238 56 III. 3. EPR analyses of Fep1-N238 62 CHAPTER IV. CONCLUSION 67 REFERENCES 72 ๊ตญ๋ฌธ์ดˆ๋ก 80Docto

    ๊ฑด๋ฌผ ๊ณต๊ธฐ์œ ๋™ํŠน์„ฑ์„ ๊ณ ๋ คํ•œ ์นจ๊ธฐ ์ €๊ฐ๋ฐฉ์•ˆ ํ‰๊ฐ€์— ๊ด€ํ•œ ์—ฐ๊ตฌ

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    ํ•™์œ„๋…ผ๋ฌธ (์„์‚ฌ)-- ์„œ์šธ๋Œ€ํ•™๊ต ๋Œ€ํ•™์› : ๊ฑด์ถ•ํ•™๊ณผ, 2015. 2. ์—ฌ๋ช…์„.๋ณธ ์—ฐ๊ตฌ์—์„œ๋Š” ์ค‘๊ทœ๋ชจ ์‚ฌ๋ฌด์†Œ ๊ฑด๋ฌผ์˜ ์นจ๊ธฐ์˜ ์˜ํ–ฅ์š”์ธ์„ ๊ณ ๋ คํ•œ ๊ณ„์ ˆ๋ณ„ ๊ฑด๋ฌผ ๋‚ด ๊ณต๊ธฐ์œ ๋™ ๋ถ„์„์„ ํ†ตํ•ด ๊ทธ ํŠน์„ฑ์„ ๊ทœ๋ช…ํ•˜๊ณ  ์นจ๊ธฐ๋กœ ์ธํ•œ ๊ฑด๋ฌผ์˜ ๊ณต๊ธฐ์œ ๋™์œผ๋กœ ์ธํ•œ ๋ฌธ์ œ์  ๊ด€๋ฆฌ ๋ฐ ๊ฐœ์„ ์„ ์œ„ํ•ด ์ฒด๊ณ„์ ์ด๊ณ  ํšจ์œจ์ ์ธ ์ €๊ฐ๋ฐฉ์•ˆ ํ‰๊ฐ€๋ฐฉ๋ฒ•์„ ์ˆ˜๋ฆฝํ•˜์˜€๋‹ค. ์ด๋ฅผ ์œ„ํ•ด ๊ธฐ์กด๋ฌธํ—Œ ๊ณ ์ฐฐ์„ ํ†ตํ•ด ๊ธฐ์กด ์นจ๊ธฐ ์ €๊ฐ๋ฐฉ์•ˆ์˜ ํŠน์ง• ๋ฐ ์ ์šฉ๋ฐฉ๋ฒ•์˜ ํ•œ๊ณ„์ ์„ ๋ถ„์„ํ•˜๊ณ  ๋Œ€ํ‘œ์ ์ธ ๊ณต๊ธฐ์œ ๋™ ์˜ํ–ฅ์ธ์ž๋ฅผ ๋„์ถœํ•˜์˜€๋‹ค. ๋„์ถœ๋œ ๊ธฐ์กด ์—ฐ๊ตฌ์˜ ํ•œ๊ณ„์ ์„ ๋ณด์™„ ๋ฐ ์˜ํ–ฅ์ธ์ž๋ฅผ ๊ณ ๋ คํ•˜์—ฌ ์ค‘๊ทœ๋ชจ ์‚ฌ๋ฌด์†Œ ๊ฑด๋ฌผ์˜ ๊ฑด๋ฌผ ์™ธํ”ผ๋ฅผ ํ†ตํ•œ ๊ฑด๋ฌผ ๋‚ด ๊ณต๊ธฐ์œ ๋™, ํ™•์‚ฐ ํŠน์„ฑ์— ๋ฏธ์น˜๋Š” ์˜ํ–ฅ ๋ถ„์„์„ ์ˆ˜ํ–‰ํ•˜์—ฌ ์—ฐ๊ตฌ๋ฅผ ์ง„ํ–‰ํ•˜์˜€๋‹ค. ๋ณธ ์—ฐ๊ตฌ์˜ ๊ฒฐ๊ณผ๋ฅผ ์š”์•ฝํ•˜๋ฉด ๋‹ค์Œ๊ณผ ๊ฐ™๋‹ค.์ œ 1 ์žฅ ์„œ ๋ก  1.1 ์—ฐ๊ตฌ์˜ ๋ฐฐ๊ฒฝ ๋ฐ ๋ชฉ์  1 1.2 ์—ฐ๊ตฌ์˜ ๋ฒ”์œ„ ๋ฐ ๋ฐฉ๋ฒ• 3 ์ œ 2 ์žฅ ๊ฑด๋ฌผ์˜ ๊ณต๊ธฐ์œ ๋™์— ๊ด€ํ•œ ์˜ˆ๋น„์  ๊ณ ์ฐฐ 7 2.1 ๊ฑด๋ฌผ์˜ ๊ณต๊ธฐ์œ ๋™์— ๊ด€ํ•œ ์ด๋ก ์  ๊ณ ์ฐฐ 7 2.1.1 ๊ณต๊ธฐ์œ ๋™ ๊ธฐ๋ณธ์ด๋ก  7 2.1.2 ๊ณต๊ธฐ์œ ๋™ ์š”์†Œ 10 2.2 ๊ธฐ์กด ์—ฐ๊ตฌ๋ฌธํ—Œ ๋ถ„์„์„ ํ†ตํ•œ ์นจ๊ธฐ๊ฐ€ ๊ฑด๋ฌผ์— ๋ฏธ์น˜๋Š” ์˜ํ–ฅ ๋ฐ ๋ฌธ์ œ์  ๋„์ถœ 13 2.3 ์†Œ๊ฒฐ 15 ์ œ 3 ์žฅ ๊ฑด๋ฌผ์˜ ๊ณต๊ธฐ์œ ๋™ ๋ชจ๋ธ ํ•ด์„ ๋ฐฉ๋ฒ• ๊ณ ์ฐฐ 17 3.1 ๊ฑด๋ฌผ ๊ณต๊ธฐ์œ ๋™ ํ•ด์„๋ชจ๋ธ์˜ ๊ฒ€ํ†  17 3.1.1 ๊ณต๊ธฐ์œ ๋™ ํ•ด์„ ๋ชจ๋ธ 18 3.1.2 ๋„คํŠธ์›Œํฌ ๋ชจ๋ธ 21 3.1.3 ๊ณต๊ธฐ์œ ๋™ ๊ฒฝ๋กœ(Airflow Path)์˜ ์ •์˜ 25 3.2 ๊ฑด๋ฌผ ๊ณต๊ธฐ์œ ๋™ ๋ชจ๋ธ 29 3.2.1 ๋Œ€์ƒ๊ฑด๋ฌผ ๊ฐœ์š” ๋ฐ ์ธก์ • 29 3.2.2 ๋ชจ๋ธ๋ง ๋ฐ ๊ฒ€์ฆ 35 3.3 ์†Œ ๊ฒฐ 37 ์ œ 4 ์žฅ ๊ฑด๋ฌผ์˜ ์นจ๊ธฐ์™€ ์˜ํ–ฅ์ธ์ž ๋ถ„์„ 39 4.1 ๊ฐœ์š” 39 4.2 ๋Œ€์ƒ๊ฑด๋ฌผ์˜ ๊ณต๊ธฐ์œ ๋™ ํŠน์„ฑ 40 4.3 ๊ฑด๋ฌผ์—์„œ์˜ ๊ณต๊ธฐ์œ ๋™ ์˜ํ–ฅ์ธ์ž ๋ถ„์„ 41 4.3.1 ์˜ํ–ฅ์ธ์ž ์ •์˜ 41 4.3.2 ํ™˜๊ฒฝ๊ด€๋ จ ์ธ์ž 42 4.3.3 ๊ฑด์ถ•๊ด€๋ จ ์ธ์ž 44 4.4 ์†Œ๊ฒฐ 46 ์ œ 5 ์žฅ ๋Œ€์ƒ๊ฑด๋ฌผ ๊ตฌ์„ฑ์š”์†Œ์˜ ์นจ๊ธฐ๋Ÿ‰๊ณผ ์˜ํ–ฅ์ธ์ž ๋ถ„์„ 47 5.1 ๊ฐœ์š” 47 5.2 ๋ถ„์„์ง€์  ์„ ์ • 48 5.3 ์นจ๊ธฐ๋Ÿ‰๊ณผ ์˜ํ–ฅ์ธ์ž์˜ ์ƒ๊ด€๊ด€๊ณ„ ๋ถ„์„ 51 5.3.1 ์™ธ๊ธฐ์˜จ 55 5.3.2 ํ’ํ–ฅ 56 5.3.3 ํ’์† 57 5.4 ๊ฑด๋ฌผ ๋‚ด ํ™•์‚ฐ ๋ถ„์„ 58 5.5 ๊ฑด๋ฌผ์˜ ์นจ๊ธฐ ์ €๊ฐ๋ฐฉ์•ˆ ํ‰๊ฐ€ 62 5.6 ์†Œ๊ฒฐ 65 ์ œ 6 ์žฅ ๊ฒฐ ๋ก  67 ์ฐธ๊ณ  ๋ฌธํ—Œ 69 ABSTRACT 73Maste

    ๊ธˆ์œต์ž์œ ํ™”์˜ ์ •์น˜๊ฒฝ์ œ-2000๋…„๋Œ€ ๋ง๋ ˆ์ด์‹œ์•„์˜ ์„ ๋ณ„์  ๊ธˆ์œต์ž์œ ํ™”์ •์ฑ… ์‚ฌ๋ก€์—ฐ๊ตฌ-

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    ํ•™์œ„๋…ผ๋ฌธ (์„์‚ฌ)-- ์„œ์šธ๋Œ€ํ•™๊ต ๋Œ€ํ•™์› : ์ •์น˜ํ•™๊ณผ, 2012. 8. ์ž„ํ˜œ๋ž€.๋ณธ ๋…ผ๋ฌธ์˜ ๋ชฉ์ ์€ 2000๋…„๋Œ€ ๋ง๋ ˆ์ด์‹œ์•„์˜ ์„ ๋ณ„์  ๊ธˆ์œต์ž์œ ํ™”์ •์ฑ…์˜ ๋ฐฐ๊ฒฝ์„ ๋ง๋ ˆ์ด์‹œ์•„ ์ •์น˜๊ฒฝ์ œ์˜ ์ œ๋„์  ์œ ์‚ฐ๊ณผ ๊ตญ๋‚ด์ •์น˜์  ๋™ํ•™์„ ์ค‘์‹ฌ์œผ๋กœ ๊ณ ์ฐฐํ•˜๋Š” ๊ฒƒ์ด๋‹ค. ๋ง๋ ˆ์ด์‹œ์•„๋Š” ๋™์•„์‹œ์•„ ์œ„๊ธฐ ๋‹น์‹œ ์œ„๊ธฐ๋ฅผ ๊ฒฝํ—˜ํ•œ ๋‹ค๋ฅธ ๊ตญ๊ฐ€๋“ค๊ณผ๋Š” ๋‹ฌ๋ฆฌ IMF์˜ ์‹ ์ž์œ ์ฃผ์˜์  ์ •์ฑ…์„ ๊ฑฐ๋ถ€ํ•˜๊ณ  ์ž๋ณธํ†ต์ œ์ •์ฑ…๊ณผ ๊ณ ์ •ํ™˜์œจ์ œ๋กœ ํšŒ๊ท€ํ•˜์—ฌ ์ „ ์„ธ๊ณ„ ํˆฌ์ž์ž๋“ค๊ณผ ์ „๋ฌธ๊ฐ€๋“ค์˜ ๋น„๋‚œ์„ ๋ฐ›์•˜๋‹ค. ๊ทธ๋Ÿฐ๋ฐ ๋ง๋ ˆ์ด์‹œ์•„๋Š” ์˜ค๋Š˜๋‚  ์ „ ์„ธ๊ณ„์—์„œ ๊ฐ€์žฅ ๋ฐœ๋‹ฌ๋œ ์ด์Šฌ๋žŒ๊ธˆ์œต์‚ฐ์—…์˜ ์ค‘์‹ฌ์ง€๋กœ์„œ ์ฃผ๋ชฉ์„ ๋ฐ›๊ณ  ์žˆ๋‹ค. ๋ณธ ์—ฐ๊ตฌ๋Š” ๊ตญ๊ฐ€๋“ค์ด ๋™์ผํ•œ ์œ„๊ธฐ๋ฅผ ๊ฒฝํ—˜ํ•˜์˜€์Œ์—๋„ ๋ถˆ๊ตฌํ•˜๊ณ  ์ •์ฑ…๋Œ€์‘์—์„œ๋Š” ์™œ ์ฐจ์ด๋ฅผ ๋ณด์ด๋Š”๊ฐ€, ์œ„๊ธฐ ๋Œ€์‘์ •์ฑ…๊ณผ ๊ด€๋ จํ•˜์—ฌ ๋งŽ์€ ๋น„ํŒ์„ ๋ฐ›์•˜๋˜ ๋ง๋ ˆ์ด์‹œ์•„๊ฐ€ ์–ด๋–ป๊ฒŒ ์„ธ๊ณ„ ์ตœ๊ณ ์˜ ์ด์Šฌ๋žŒ๊ธˆ์œต์‹œ์žฅ์„ ์šด์˜ํ•˜๊ณ  ์žˆ๋Š”๊ฐ€์— ๋Œ€ํ•œ ์˜๋ฌธ์—์„œ๋ถ€ํ„ฐ ์ถœ๋ฐœํ•˜์˜€๋‹ค. ๋™์•„์‹œ์•„ ์œ„๊ธฐ ์ดํ›„ 2000๋…„๋Œ€ ๋ง๋ ˆ์ด์‹œ์•„ ์ •๋ถ€๋Š” ์„ ๋ณ„์  ๊ธˆ์œต์ž์œ ํ™”์ •์ฑ…์„ ์ถ”์ง„ํ•˜์˜€๋‹ค. ๋ง๋ ˆ์ด์‹œ์•„ ์ •๋ถ€๋Š” ๋‹ค๋ฅธ ๊ตญ๊ฐ€๋“ค๊ณผ๋Š” ๋‹ฌ๋ฆฌ ์ „๋ฉด์  ๊ธˆ์œต์ž์œ ํ™”๋ฅผ ์‹ค์‹œํ•˜๋Š” ๊ฒƒ์ด ์•„๋‹ˆ๋ผ ๊ธˆ์œต์ž์œ ํ™”์˜ ์†๋„์™€ ๋ฐฉํ–ฅ์„ ์กฐ์ ˆํ•ด ์„ ๋ณ„์ ์ด๊ณ  ์ ์ง„์ ์ธ ๊ธˆ์œต์ •์ฑ…์„ ์ถ”์ง„ํ•ด์•ผ ํ–ˆ์œผ๋ฉฐ, ์ด ๊ณผ์ •์—์„œ ์ •๋ถ€๊ฐ€ ์ด๋‹ˆ์…”ํ‹ฐ๋ธŒ๋ฅผ ๊ฐ–๊ณ  ๊ธˆ์œต์ž์œ ํ™”์˜ ์†๋„์™€ ๋ฐฉํ–ฅ์„ ์กฐ์ ˆ ํ•  ์ˆ˜ ์žˆ์—ˆ๋‹ค. ์ด๋Š” ์œ„๊ธฐ ๋‹น์‹œ ๋ฐœ์ƒํ–ˆ๋˜ ๊ตญ๋‚ด์ •์น˜์  ๋™ํ•™๊ณผ ๋ง๋ ˆ์ด์‹œ์•„ ์ •์น˜๊ฒฝ์ œ์˜ ์„ธ๊ฐ€์ง€ ์ œ๋„์  ์œ ์‚ฐ์ด ์ž‘๋™ํ•œ ๊ฒฐ๊ณผ๋‹ค. ๋ง๋ ˆ์ด์‹œ์•„์—์„œ๋Š” ์ผ๋ฐ˜๊ธˆ์œต๊ณผ ์ด์Šฌ๋žŒ๊ธˆ์œต์ด ๋™์‹œ์— ์šด์˜๋˜๊ณ  ์žˆ๋‹ค. ์ •๋ถ€๋Š” ๊ฐ ๋ถ€๋ฌธ์˜ ์ž์œ ํ™”์— ๊ฐ๊ธฐ ๋‹ค๋ฅธ ํƒœ๋„๋ฅผ ๋ณด์ด๋Š” ์ „๋žต์„ ์ทจํ•จ์œผ๋กœ์จ ์ •๋ถ€์˜ ๊ณ ์œ ํ•œ ์ •์น˜์  ์ดํ•ด๋ฅผ ์ˆ˜ํ˜ธํ•˜๊ณ  ์ง‘๊ถŒ์œ ์ง€์™€ ์ •๊ถŒ๊ณต๊ณ ํ™”๋ผ๋Š” ํ•ต์‹ฌ์  ๋ชฉํ‘œ๋ฅผ ๋‹ฌ์„ฑํ•ด์™”๋‹ค. 1969๋…„ ์ข…์กฑํญ๋™ ์ดํ›„ ์‚ฌํšŒ์•ˆ์ •๊ณผ ๊ฒฝ์ œ์„ฑ์žฅ์„ ์œ„ํ•ด ์ •์น˜์ง€๋„์ž์™€ ์ •๋ถ€์— ๊ฐ•๋ ฅํ•œ ๊ถŒํ•œ์ด ์ฃผ์–ด์ง€๋ฉด์„œ ์ง€์†๋˜์–ด ์™”๋˜ ๋ง๋ ˆ์ด ํŠนํ˜œ๋ถ„๋ฐฐ์ •์ฑ…, ๊ตญ๊ฐ€์ฃผ๋„์˜ ๊ฒฝ์ œ์„ฑ์žฅ์ „๋žต, ๊ตญ๊ฐ€์šฐ์œ„์˜ ์ •์น˜๊ตฌ๋„๋ผ๋Š” ์„ธ ๊ฐ€์ง€ ์ œ๋„์  ์œ ์‚ฐ์ด ๋ง๋ ˆ์ด์‹œ์•„์˜ ์ •์น˜๊ฒฝ์ œ์˜ ๊ทผ๊ฐ„์ด ๋˜์–ด์™”๋‹ค. ์ด๋Ÿฌํ•œ ์ œ๋„์  ์œ ์‚ฐ์ด ๋ง๋ ˆ์ด์‹œ์•„ ์ •๋ถ€๊ฐ€ ๊ณ ์œ ์˜ ์ •์ฑ…์  ์„ ํ˜ธ๋ฅผ ๊ฐ–๊ณ  ์ •์ฑ…๊ฒฐ์ •๊ณผ์ •์—์„œ ์ž์œจ์„ฑ์„ ๋ณด์žฅ๋ฐ›์œผ๋ฉฐ ์ฃผ์š”ํ•œ ํ–‰์œ„์ž๋กœ์„œ ์—ญํ• ์„ ์ˆ˜ํ–‰ํ•  ์ˆ˜ ์žˆ๋„๋ก ํ•˜์˜€์œผ๋ฉฐ, ์‚ฌํšŒ๋ถ€๋ฌธ์˜ ์ดํ•ด๋ฅผ ์ •๋ถ€์˜ ์ „๋žต์ ์ธ ์ดํ•ด์— ๋”ฐ๋ผ ์„ ๋ณ„ํ•˜์—ฌ ์ •์ฑ…์ˆ˜ํ–‰๊ณผ์ •์— ๋ฐ˜์˜ํ•  ์ˆ˜ ์žˆ๊ฒŒ ๋งŒ๋“ค์—ˆ๋˜ ๊ฒƒ์ด๋‹ค. ๋…๋ฆฝ ์ดํ›„ ํ•˜๋‚˜์˜ ์ •๋‹น์ด ๊ตญ๋ฏผ๋“ค์—๊ฒŒ ์••๋„์ ์ธ ์ง€์ง€๋ฅผ ๋ฐ›์œผ๋ฉฐ ํ†ต์น˜ํ•ด ์˜จ ๋ง๋ ˆ์ด์‹œ์•„ ์ •์น˜์ฒด์ œ์˜ ํŠน์„ฑ์ƒ ๋™์•„์‹œ์•„ ์œ„๊ธฐ ์ดํ›„ 1998๋…„๊ณผ 1999๋…„์— ๋ฐœ์ƒํ•˜์˜€๋˜ ๋ง๋ ˆ์ด์‹œ์•„ ๊ตญ๋‚ด์ •์น˜์  ๋ณ€๋™์€ ์ •๋ถ€์™€ ์ง‘๊ถŒ์—ฌ๋‹น์—๊ฒŒ ์ƒ๋‹นํ•œ ํƒ€๊ฒฉ์„ ์ž…ํ˜”๋‹ค. ์‹œ๋ฏผ๊ณผ ์•ผ๋‹น๋“ค์€ ๊ฒฝ์ œ์œ„๊ธฐ์™€ ์œ„๊ธฐ ๋Œ€์‘ ๋‹น์‹œ ์ ๋‚˜๋ผํ•˜๊ฒŒ ๋“œ๋Ÿฌ๋‚œ ์ •์‹ค์ฃผ์˜์™€ ๋ถ€์ •๋ถ€ํŒจ, ์•ˆ์™€๋ฅด ์ด๋ธŒ๋ผํž˜ ๋‹น์‹œ ๋ถ€์ด๋ฆฌ์ด์ž ์žฌ๋ฌด๋ถ€ ์žฅ๊ด€์— ๋Œ€ํ•œ ๊ฐ€ํ˜นํ•œ ์ฒ˜๋ถ„์— ๋ถˆ๋งŒ์„ ๊ฐ–๊ณ  ๊ฐœํ˜์šด๋™์„ ์ „๊ฐœํ•˜์˜€์œผ๋ฉฐ 1999๋…„ ์ด์„ ์—์„œ ์ด์Šฌ๋žŒ๊ธ‰์ง„์ฃผ์˜ ์•ผ๋‹น์ธ PAS์— ์ „๋ก€ ์—†๋Š” ์ง€์ง€๋ฅผ ๋ณด์—ฌ์คŒ์œผ๋กœ์จ ๋งˆํ•˜ํ‹ฐ๋ฅด ์ •๊ถŒ์— ๋Œ€ํ•œ ๋ถˆ๋งŒ๊ณผ ๊ฐœํ˜์— ๋Œ€ํ•œ ์š”๊ตฌ๋ฅผ ํ‘œ์ถœํ•˜์˜€๋‹ค. ํŠนํžˆ ์ด์Šฌ๋žŒ๊ธ‰์ง„์ฃผ์˜ ์•ผ๋‹น PAS์˜ ์•ฝ์ง„์€ ์ •๋ถ€์™€, ๊ณผ๊ฑฐ๋ถ€ํ„ฐ ์ด์Šฌ๋žŒ์  ๊ฐ€์น˜๋ฅผ ๋‘๊ณ  ๊ฒฝ์Ÿํ•ด์˜จ UMNO์˜ ์ง€์ง€๊ธฐ๋ฐ˜์— ์ถฉ๊ฒฉ์„ ๊ฐ€ํ–ˆ๋‹ค. PAS๋Š” ์ •๋ถ€์™€ ์ง‘๊ถŒ์—ฌ๋‹น์ธ UMNO์˜ ์„ธ์†์  ์ด์Šฌ๋žŒ์ฃผ์˜๋ฅผ ๊ฒฝ์ œ์œ„๊ธฐ์˜ ์ฃผ์š”ํ•œ ์›์ธ์œผ๋กœ ์ง€๋ชฉํ•˜๊ณ  ๊ฒฝ์ œ๋ฅผ ์ด์Šฌ๋žŒ์  ๋ฐฉ์‹์œผ๋กœ ์šด์šฉํ•ด์•ผ ํ•  ํ•„์š”์„ฑ์„ ์—ญ์„คํ•˜์˜€๋‹ค. ๋˜ํ•œ ๊ฐœํ˜์šด๋™์„ธ๋ ฅ์€ ์ •๋ถ€์˜ ๋ณดํ˜ธ์ฃผ์˜์ ์ด๊ณ  ํ์‡„์ ์ธ ์ •์ฑ… ํ•˜์— ํŠน๊ถŒ์  ์ง€์œ„๋ฅผ ๋ˆ„๋ ค์˜ค๋˜ ์†Œ์ˆ˜์˜ ๋ง๋ ˆ์ด ๋Œ€๊ธฐ์—…๊ฐ€๋“ค์˜ ๋ฐฉํƒ•ํ•œ ๊ธฐ์—…์šด์˜ ๋ฐ ๋ถˆํˆฌ๋ช…ํ•œ ๊ธฐ์—…์ง€๋ฐฐ๊ตฌ์กฐ์™€ ์ด๋“ค์„ ๋ณดํ˜ธํ•˜๊ธฐ ์œ„ํ•ด ์ •๋ถ€๊ฐ€ ์ทจํ–ˆ๋˜ ์ž๋ณธํ†ต์ œ์ •์ฑ…๊ณผ ๊ณ ์ •ํ™˜์œจ์ œ๋ฅผ ๋น„๋‚œํ•˜์˜€๋‹ค. ๊ทธ ์ค‘์—์„œ๋„ ์ฃผ์‹์„ ์†Œ์œ ํ•˜๊ณ  ์žˆ๋˜ ๋ง๋ ˆ์ด ์ค‘์‚ฐ์ธต๋“ค๊ณผ, ๋ง๋ ˆ์ด์‹œ์•„ ๊ตญ๋‚ด๊ธฐ์—…๊ณผ ํ•ฉ์ž‘ํˆฌ์ž๋ฅผ ํ•˜๊ฑฐ๋‚˜ ์ง์ ‘ํˆฌ์žํ˜•ํƒœ๋กœ ๊ฒฝ์ œํ™œ๋™์„ ํ•˜๊ณ  ์žˆ์—ˆ๋˜ ์™ธ๊ตญ์ธ ํˆฌ์ž์ž๋“ค์€ ์ •๋ถ€์˜ ์ž๋ณธํ†ต์ œ์ •์ฑ…์œผ๋กœ ์ธํ•œ ๋Œ€์™ธ์‹ ์ธ๋„ ํ•˜๋ฝ๊ณผ ์™ธ๊ตญํˆฌ์ž์ž๋ณธ ์œ ์ถœ์ด ์•ผ๊ธฐํ•œ ์ฃผ์‹์‹œ์žฅ์˜ ํ•˜๋ฝ์— ํฐ ์ดํ•ด๋ฅผ ๊ฐ–๊ณ  ์žˆ์—ˆ๋‹ค. ์ •๋ถ€๋Š” ๋ฐ˜๋Œ€์„ธ๋ ฅ์˜ ์ดํ•ด๋ฅผ ๊ณ ๋ คํ•˜์—ฌ ๊ฐœํ˜๊ณผ ๋ณ€ํ™”๋ฅผ ์ถ”์ง„ํ•˜์ง€ ์•Š์„ ์ˆ˜ ์—†์—ˆ๊ณ , ๋”ฐ๋ผ์„œ ์œ„๊ธฐ์˜ ํ•ต์‹ฌ์  ๋ฌธ์ œ์˜€๋˜ ๊ธˆ์œต๋ถ€๋ฌธ์— ๋Œ€ํ•œ ์žฌ์ •๋น„์™€ ๊ฒฝ์ œ๋ฅผ ํšŒ๋ณตํ•˜๊ธฐ ์œ„ํ•œ ์ƒˆ๋กœ์šด ์„ฑ์žฅ๋™๋ ฅ์œผ๋กœ์จ ๊ธˆ์œต๋ถ€๋ฌธ์„ ๋ฐœ์ „์‹œํ‚ค๋Š” ์ „๋žต์„ ์ทจํ•˜์˜€๋‹ค. ํŠนํžˆ ์ •๋ถ€๋Š” ์ด์Šฌ๋žŒ๊ธˆ์œต์‚ฐ์—… ๋ฐœ์ „์„ ์ฃผ์š”ํ•œ ์ •์ฑ…๋ชฉํ‘œ๋กœ ์„ค์ •ํ•˜๊ณ  2003๋…„ ํ›„๋ฐ˜๋ถ€ํ„ฐ ์ด์Šฌ๋žŒ๊ธˆ์œต๋ถ€๋ฌธ์— ๊ธ‰์ง„์ ์ด๊ณ  ๋น ๋ฅธ ์†๋„์˜ ์ž์œ ํ™”์ •์ฑ…์„ ์‹ค์‹œํ•˜์—ฌ ์ด์Šฌ๋žŒ๊ธˆ์œต์‚ฐ์—…์„ ์„ฑ์žฅ์‹œ์ผฐ๋‹ค. ์ด์Šฌ๋žŒ๊ธˆ์œต์‚ฐ์—…์˜ ์„ฑ์žฅ๊ณผ ์ž์œ ํ™”๋Š” ๋ง๋ ˆ์ด์‹œ์•„์˜ ๋‹ค์–‘ํ•œ ๊ตญ๋‚ด์ •์น˜ํ–‰์œ„์ž๋“ค์˜ ์„ ํ˜ธ๋ฅผ ๋งŒ์กฑ์‹œํ‚ฌ ์ˆ˜ ์žˆ๋‹ค๋Š” ์ธก๋ฉด์—์„œ ์ •๋ถ€๊ฐ€ ์ „๋žต์ ์œผ๋กœ ๊ณ ๋ คํ•œ ์‚ฌ์•ˆ์ด๋ผ๊ณ  ๋ถ„์„ํ•  ์ˆ˜ ์žˆ๋‹ค. ์ผ๋ฐ˜๊ธˆ์œต๋ถ€๋ฌธ์ด ์•„๋‹Œ ์ด์Šฌ๋žŒ๊ธˆ์œต๋ถ€๋ฌธ์„ ์ž์œ ํ™”์‹œ์ผœ ์„ฑ์žฅ์‹œํ‚ด์œผ๋กœ์จ ์ •๋ถ€๊ฐ€ ์ „๋žต์ ์œผ๋กœ ์–ป์„ ์ˆ˜ ์žˆ๋Š” ์ด์ต์€ ๋‹ค์–‘ํ•˜๋‹ค. ์ •๋ถ€๋Š” ์ ๊ทน์ ์ธ ์ž์œ ํ™” ์ •์ฑ…์„ ํ†ตํ•ด ์ด์Šฌ๋žŒ๊ธˆ์œต์‚ฐ์—…์„ ์„ฑ์žฅ์‹œํ‚ด์œผ๋กœ์จ ์ด์Šฌ๋žŒ์  ํ—Œ์‹ ์„ ์˜์‹ฌํ•˜๋Š” ์ด์Šฌ๋žŒ๊ธ‰์ง„์•ผ๋‹น๋“ค์˜ ๋น„๋‚œ์— ๋Œ€์ฒ˜ํ•  ์ˆ˜ ์žˆ์œผ๋ฉฐ, ์ด์Šฌ๋žŒ๊ธˆ์œต๊ธฐ๊ด€์— ๋Œ€ํ•œ ์‹ ๊ทœ์˜์—…ํ—ˆ๊ฐ€๊ถŒ์„ ๋ฐœ๋ถ€ํ•˜๊ณ  ์™ธ๊ตญ์ธ์ง€๋ถ„์†Œ์œ ํ•œ๋„ ๋น„์œจ์„ ์ฆ๋Œ€์‹œ์ผœ ์™ธ๊ตญํˆฌ์ž์ž๋ณธ์— ๋Œ€ํ•œ ๊ทœ์ œ๋ฅผ ์™„ํ™”ํ•จ์œผ๋กœ์จ ์™ธ๊ตญ์ธ ํˆฌ์ž์ž๋“ค์„ ์œ ์น˜ํ•˜๊ณ , ์ด์Šฌ๋žŒ๊ธˆ์œต์‚ฐ์—…์— ๋Œ€ํ•œ ํˆฌ์žํ™•๋Œ€๋ฅผ ํ†ตํ•ด ๊ณ ์šฉ์ฐฝ์ถœ๊ณผ ๊ฒฝ์ œ์„ฑ์žฅ์— ๊ธฐ์—ฌํ•  ์ˆ˜ ์žˆ์œผ๋ฉฐ, ๋ง๋ ˆ์ด ๊ธฐ์—…๋“ค๋ฟ๋งŒ ์•„๋‹ˆ๋ผ ํ™”์ธ ๊ธฐ์—…๋“ค์—๊ฒŒ ๋‹ค์–‘ํ•œ ์ž๊ธˆ์กฐ๋‹ฌ๊ตฌ์กฐ๋ฅผ ์ œ๊ณตํ•˜์—ฌ ์œ„ํ—˜๋ถ„์‚ฐ๊ณผ ์ž๊ธˆ์กฐ๋‹ฌ์„ ์šฉ์ดํ•˜๊ฒŒ ํ•  ์ˆ˜ ์žˆ๋‹ค. ๊ทธ๋ฆฌ๊ณ  ์ •๋ถ€๋Š” ์ผ๋ฐ˜๊ธˆ์œต๋ถ€๋ฌธ์˜ ๊ธˆ์œต์ž์œ ํ™”์—๋Š” ๊ธฐ์กด์˜ ๋ณดํ˜ธ์ฃผ์˜์ ์ธ ์ •์ฑ…์„ ์ง€์†์ ์œผ๋กœ ์œ ์ง€ํ•จ์œผ๋กœ์จ ๋ง๋ ˆ์ด ์ข…์กฑ์˜ ๋ถ€์™€ ์ด์ต์„ ๋ณดํ˜ธํ•˜๊ณ  ๋ง๋ ˆ์ด์˜ ์ˆ˜ํ˜ธ์ž๋กœ์„œ ์ •๋ถ€์˜ ์—ญํ• ์„ ๊ณต๊ณ ํžˆ ํ•˜๋ฉฐ ํ•ต์‹ฌ์ง€์ง€๊ธฐ๋ฐ˜์ธ ๋ง๋ ˆ์ด๋กœ๋ถ€ํ„ฐ์˜ ์ง€์ง€๋ฅผ ์ง€์†์‹œํ‚ฌ ์ˆ˜ ์žˆ๋‹ค. ๋™์•„์‹œ์•„ ์œ„๊ธฐ ์ดํ›„ 2000๋…„๋Œ€ ๋ง๋ ˆ์ด์‹œ์•„ ์ •๋ถ€๊ฐ€ ์ทจํ•œ ์„ ๋ณ„์  ๊ธˆ์œต์ž์œ ํ™”์ •์ฑ…์€ ๋ง๋ ˆ์ด์‹œ์•„์˜ ๋‹ค์–‘ํ•œ ๊ตญ๋‚ด์ •์น˜ํ–‰์œ„์ž๋“ค์˜ ์ดํ•ด์™€ ์ˆ˜์‹ญ ๋…„ ๊ฐ„ ๋ง๋ ˆ์ด์‹œ์•„ ์ •์น˜์˜ ํ•ต์‹ฌ ์ถ•์ธ ์ •๋ถ€์™€ UMNO์˜ ์••๋„์ ์ธ ์ •์น˜์  ์šฐ์œ„๋ฅผ ์ง€์†์‹œํ‚ฌ ์ˆ˜ ์žˆ๋Š” ์ •๋ถ€์˜ ์ „๋žต์ ์ธ ์„ ํƒ์ด์—ˆ๋‹ค๊ณ  ํ‰๊ฐ€ํ•  ์ˆ˜ ์žˆ์„ ๊ฒƒ์ด๋‹ค. ๋ณธ ์—ฐ๊ตฌ๋Š” ๋ง๋ ˆ์ด์‹œ์•„์˜ ์„ ๋ณ„์  ๊ธˆ์œต์ž์œ ํ™”์ •์ฑ…์˜ ๋ฐฐ๊ฒฝ์„ ๋™์•„์‹œ์•„ ์œ„๊ธฐ ์ดํ›„ ๋ฐœ์ƒํ•œ ๋ง๋ ˆ์ด์‹œ์•„์˜ ๊ตญ๋‚ด์ •์น˜์  ๋™ํ•™๊ณผ ๋™์‹œ์— ๋ง๋ ˆ์ด์‹œ์•„ ์ •์น˜๊ฒฝ์ œ์˜ ์ œ๋„์  ์œ ์‚ฐ์„ ํ•จ๊ป˜ ๊ณ ์ฐฐํ•จ์œผ๋กœ์จ ์„ธ๊ฐ€์ง€ ํ•จ์˜๋ฅผ ์ œ๊ณตํ•  ๊ฒƒ์œผ๋กœ ๊ธฐ๋Œ€ํ•œ๋‹ค. ์ฒซ์งธ, ๋น„๊ต์ •์น˜ํ•™์  ํ•จ์˜๋‹ค. ๋ง๋ ˆ์ด์‹œ์•„์˜ ๊ธˆ์œต์ž์œ ํ™”์ •์ฑ…์€ ๊ฐ™์€ ๋™์•„์‹œ์•„ ์œ„๊ธฐ๋ฅผ ๊ฒฝํ—˜ํ•œ ๊ตญ๊ฐ€๋“ค๊ณผ๋Š” ๊ตฌ๋ณ„๋˜๋Š” ์ ์ง„์ ์ด๊ณ  ์„ ๋ณ„์ ์ธ ํŠน์ง•์„ ๋ณด์ธ๋‹ค. ๊ทธ๋Ÿผ์—๋„ ๋ถˆ๊ตฌํ•˜๊ณ  ๋ง๋ ˆ์ด์‹œ์•„์˜ ๊ฒฝ์ œ์—์„œ ๊ธˆ์œต์‚ฐ์—…์ด ์ฐจ์ง€ํ•˜๋Š” ๋น„์ค‘์€ ํ•œ๊ตญ์ด๋‚˜ ์ธ๋„๋„ค์‹œ์•„๋ณด๋‹ค ๋†’๋‹ค. ๋˜ํ•œ ๋ง๋ ˆ์ด์‹œ์•„๋Š” ์˜ค๋Š˜๋‚  ๊ธ€๋กœ๋ฒŒ ์ด์Šฌ๋žŒ๊ธˆ์œต์‚ฐ์—…์„ ์„ ๋„ํ•˜๊ณ  ์žˆ๋‹ค. ๋ณธ ์—ฐ๊ตฌ๋Š” ๊ตญ๊ฐ€ ๊ฐ„ ์ •์ฑ…์  ์ฐจ์ด์˜ ๋™์ธ์„ ๋ฐํ˜€๋‚ด์–ด ๋น„๊ต์ •์น˜ํ•™์  ํ•จ์˜๋ฅผ ์ œ๊ณตํ•  ๊ฒƒ์ด๋‹ค. ๋‘˜์งธ, ์ด๋ก ์  ํ•จ์˜๋‹ค. ๋ณธ ์—ฐ๊ตฌ๋Š” ๋ง๋ ˆ์ด์‹œ์•„ ๊ธˆ์œต์ž์œ ํ™”์— ๋Œ€ํ•œ ๊ธฐ์กด ์—ฐ๊ตฌ๋“ค์ด ๊ฐ„๊ณผํ–ˆ๋˜ ์ œ๋„์  ์š”์†Œ๋ฅผ ๋ถ„์„์˜ ์ฃผ์š”ํ•œ ๋ณ€์ˆ˜๋กœ ๊ณ ๋ คํ•˜์˜€๋‹ค. ๋˜ํ•œ ์ œ๋„์  ์š”์†Œ๋ฅผ ๊ณ ๋ คํ•จ์œผ๋กœ์จ ์—ญ์‹œ ๊ธฐ์กด ์—ฐ๊ตฌ๋“ค์ด ๊ฐ„๊ณผํ–ˆ๋˜ ์ •๋ถ€๋ผ๋Š” ํ–‰์œ„์ž์˜ ์—ญํ• ๊ณผ ์ค‘์š”์„ฑ์„ ๋ฐœ๊ฒฌํ•˜์—ฌ ๋ง๋ ˆ์ด์‹œ์•„ ์ •์น˜๊ฒฝ์ œ์—์„œ ์ œ๋„์™€ ํ–‰์œ„์ž ๊ฐ„์˜ ์ƒํ˜ธ์ž‘์šฉ์„ ๋“œ๋Ÿฌ๋‚ด์—ˆ๋‹ค. ๋”ฐ๋ผ์„œ ๋ณธ ์—ฐ๊ตฌ๋Š” ์ œ๋„์  ์ ‘๊ทผ์˜ ์œ ์šฉ์„ฑ์„ ๊ฐ•์กฐํ•˜๊ณ  ๊ธฐ์กด ๋ง๋ ˆ์ด์‹œ์•„ ์ •์น˜๊ฒฝ์ œ์—์„œ ์ œ๋Œ€๋กœ ์ฃผ๋ชฉ ๋ฐ›์ง€ ๋ชปํ–ˆ๋˜ ์ •๋ถ€๋ผ๋Š” ํ–‰์œ„์ž๋ฅผ ์กฐ๋ช…ํ•˜๋Š” ์ด๋ก ์  ํ•จ์˜๋ฅผ ์ง€๋‹Œ๋‹ค. ๋งˆ์ง€๋ง‰์œผ๋กœ ๋ณธ ์—ฐ๊ตฌ๋Š” ์ •์ฑ…์  ํ•จ์˜๋ฅผ ์ œ๊ณตํ•œ๋‹ค. 1997๋…„ ๋™์•„์‹œ์•„ ์œ„๊ธฐ๋ฅผ ๊ฒฝํ—˜ํ•œ ๊ตญ๊ฐ€๋“ค์€ ์˜คํžˆ๋ ค ๋”์šฑ ๋น ๋ฅธ ์†๋„์™€ ๋„“์€ ๋ฒ”์œ„์—์„œ ๊ธˆ์œต์‹œ์žฅ๊ฐœ๋ฐฉ์„ ์ถ”์ง„ํ•ด์™”๋‹ค. ๋ฐ˜๋ฉด ๋ง๋ ˆ์ด์‹œ์•„ ์ •๋ถ€๋Š” ์„ ๋ณ„์ ์ด๊ณ  ์ ์ง„์ ์ธ ๊ธˆ์œต์ž์œ ํ™”์ •์ฑ…์„ ์ถ”์ง„ํ•˜์—ฌ ๊ธˆ์œต์ž์œ ํ™”์˜ ์†๋„์™€ ๋ฐฉํ–ฅ์„ ์กฐ์ ˆํ•ด์™”๋‹ค. ๊ทธ๋Ÿฐ๋ฐ ๋ง๋ ˆ์ด์‹œ์•„์˜ ๊ธˆ์œต๋ถ€๋ฌธ์€ ์ธ๋„๋„ค์‹œ์•„, ํ•œ๊ตญ, ํƒœ๊ตญ ๋“ฑ ๋ณด๋‹ค ๊ธˆ์œต์‹œ์žฅ์˜ ๋ฐœ๋‹ฌ, ์•ˆ์ •์„ฑ ์ธก๋ฉด์—์„œ ์ „ ์„ธ๊ณ„์ ์œผ๋กœ ๋†’์€ ์ˆ˜์ค€์˜ ํ‰๊ฐ€๋ฅผ ๋ฐ›๊ณ  ์žˆ๋‹ค. ๋ง๋ ˆ์ด์‹œ์•„ ์ •๋ถ€์˜ ๊ธˆ์œต์ž์œ ํ™”์ •์ฑ…์€ ๋‹ค๋ฅธ ๊ตญ๊ฐ€๋“ค์—๊ฒŒ ์ •์ฑ…์  ์‹œ์‚ฌ์ ์„ ์ œ๊ณตํ•  ๊ฒƒ์ด๋‹ค.์ œ 1 ์žฅ ์„œ ๋ก  1 ์ œ 1 ์ ˆ ๋ฌธ์ œ์ œ๊ธฐ 1 ์ œ 2 ์ ˆ ์—ฐ๊ตฌ๋Œ€์ƒ ๋ฐ ๋ฐฉ๋ฒ• 6 ์ œ 3 ์ ˆ ๋…ผ๋ฌธ์˜ ๊ตฌ์„ฑ 7 ์ œ 2 ์žฅ ๊ธฐ์กด์—ฐ๊ตฌ ๊ฒ€ํ†  ๋ฐ ์ด๋ก ์  ๋ถ„์„ํ‹€ 9 ์ œ 1 ์ ˆ ๊ธฐ์กด์—ฐ๊ตฌ ๊ฒ€ํ†  9 ์ œ 2 ์ ˆ ์—ฐ๊ตฌ๊ฐ€์„ค ๋ฐ ์ด๋ก ์  ๋ถ„์„ํ‹€ 15 ์ œ 3 ์žฅ ๋ง๋ ˆ์ด์‹œ์•„ ์ •์น˜๊ฒฝ์ œ์˜ ์ œ๋„์  ์œ ์‚ฐ 28 ์ œ 1 ์ ˆ ๋ง๋ ˆ์ด ํŠนํ˜œ๋ถ„๋ฐฐ์ •์ฑ… 28 ์ œ 2 ์ ˆ ๊ตญ๊ฐ€์ฃผ๋„์˜ ๊ฒฝ์ œ์„ฑ์žฅ์ „๋žต 33 ์ œ 3 ์ ˆ ๊ตญ๊ฐ€์šฐ์œ„์˜ ์ •์น˜๊ตฌ๋„ 36 ์ œ 4 ์žฅ ๋ง๋ ˆ์ด์‹œ์•„ ๊ธˆ์œต์ž์œ ํ™”์˜ ์ •์น˜์  ๋™ํ•™ 46 ์ œ 1 ์ ˆ ๋™์•„์‹œ์•„ ์œ„๊ธฐ์™€ ์ •์น˜๋ณ€๋™ 46 1. ์œ„๊ธฐ์˜ ์›์ธ๊ณผ ์ •์ฑ…๋ณ€๋™ 46 2. ๊ฐœํ˜์šด๋™๊ณผ ์ด์„  54 3. ์ •์น˜๋ณ€๋™๊ณผ ์ •๋ถ€์˜ ๋Œ€์‘ 62 ์ œ 2 ์ ˆ ๋ง๋ ˆ์ด์‹œ์•„ ๊ธˆ์œต์ž์œ ํ™”์˜ ์œ ํ˜• 71 1. ๊ธˆ์œต๋ถ€๋ฌธ๋งˆ์Šคํ„ฐํ”Œ๋žœ๊ณผ ์ž๋ณธ์‹œ์žฅ๋งˆ์Šคํ„ฐํ”Œ๋žœ 72 2. ์„ ๋ณ„์  ๊ธˆ์œต์ž์œ ํ™”: ๋ณดํ˜ธ์ฃผ์˜์—์„œ ์ ์ง„์  ์ž์œ ํ™”๋กœ 77 3. ๊ฒฝ์ œํšŒ๋ณต๊ณผ 2004๋…„ ์ด์„ ๊ฒฐ๊ณผ 90 ์ œ 5 ์žฅ ๊ฒฐ๋ก  94 ์ฐธ๊ณ ๋ฌธํ—Œ 98 Abstract 109Maste
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