臺灣博碩士論文加值系統

中文摘要
在種子發育晚期，乾物質不再累積，細胞含水量開始降低，種子也逐漸進入休眠狀態。有許多新的蛋白質在這階段才開始合成與累積在細胞中，其功能可能與種子的發芽、休眠或耐旱等機制有關。這些新的蛋白質中，一種被稱為晚胚蛋白質（Late embryogenesis abundant proteins）的蛋白質已廣泛的在原核生物、真菌、植物、以及動物界等物種的細胞中發現。根據不同的分類標準，晚胚蛋白被分為六群（利用胺基酸組成的特性與保守性序列）或至少四群（利用保守性胺基酸殘基）。晚胚蛋白的功能至今仍不清楚，但從基因表現型式、定位分析、蛋白質化學特性、以及轉殖基因的研究，研究人員認為晚胚蛋白可能是一種乾旱保護劑（desiccation protectant），作為促進植物細胞抵抗缺水環境的工具之一。在正儲型種子獲得乾化耐受性（desiccation tolerance）的研究中，晚胚蛋白與非還原醣（non-reducing sugars）或許是最重要，也可能是研究最多的關鍵因子。晚胚蛋白與非還原醣的交互作用已被建議是增加種子乾化耐受性的方式，但詳細的作用機制仍有許多未知處有待研究。
在本論文中，將針對三種不同型態的第四群晚胚殖系，即GmPM1、GmPM16與GmPM28殖系，以分子生物學、細胞生物學、生物化學與生物物理學等方式，探討第四群晚胚基因的表現，以及其對應蛋白質的生理功能。本論文的第一部份為GmPM16蛋白質特性的研究。GmPM16蛋白質為一種具小分子量、高等電點，以及特殊的胺基酸殘基分佈型式的鹼性蛋白質。北方式墨點與數位北方式（digital Northern）分析法指出GmPM16轉錄物主要表現在種子。原位雜交（in situ hybridization）分析則發現GmPM16轉錄物主要累積在4 DPD種子的子葉葉肉細胞，但沒有在子葉維管束組織中發現。圓二色（circular dichroism，CD）及傅立葉紅外線轉換（Fourier transform infrared，FTIR）光譜分析指出GmPM16蛋白質在水溶液狀態下為不規則的鬆散結構，並存在有限的規則性二級結構。GmPM16蛋白質在水溶液狀態下無法與還原醣類產生交互作用，但可在SDS或TFE之類的-螺旋促進劑溶液中折疊出大量的-螺旋結構。另一方面，GmPM16蛋白質在完全乾燥的情況下也能誘導出大量的-螺旋結構。GmPM16蛋白質與非還原醣在脫水的情況下所形成的玻璃質態（glassy matrix），具有較高的璃質態轉換溫度（Tg）及較低的波溫係數（wavenumber–temperature coefficient，WTC）。這個結果顯示GmPM16蛋白質與非還原醣發生緊密的交互作用。因此，蛋白質/非還原醣玻璃質可能在脫水種子中扮演降低細胞損害的角色。
本論文的第二部份是針對另外兩個大豆第四群晚胚蛋白質，分別是鹼性的GmPM1與酸性的GmPM28蛋白質。數位北方式分析法顯示GmPM1轉錄物主要表現在種子及逆境處理的營養器官，而GmPM28則為具種子專一性的基因。流體動力及折疊狀態的計算結果顯示，高親水性晚胚蛋白質屬於自然非折疊蛋白質（natively unfolded proteins）的一種。CD與FTIR光譜分析也證明GmPM1與GmPM28蛋白質在水溶液中為不規則的鬆散結構，但提高溫度時光譜顯示蛋白質出現進一步的解鬆。CD差光譜（difference spectra）顯示GmPM1與GmPM28蛋白質在低溫下具有左旋性伸展螺旋（left-handed extended helical），或稱為poly (L-proline)-type II（PPII）結構。兩種蛋白在水溶液狀態下無法與還原醣類產生交互作用，但可以在SDS或TFE溶液中折疊出-螺旋結構。另一方面，GmPM1與GmPM28蛋白質在完全乾燥的情況下也能誘導出-螺旋結構。兩種蛋白質分別與非還原醣在脫水的情況下所形成的玻璃質態，與非還原醣玻璃質態相較，有較高的璃質態轉換溫度及較低的波溫係數；但與GmPM16蛋白質/非還原醣玻璃質態比較，其玻璃質態的緊密度較低。以上結果顯示這三種不同型態的第四群晚胚蛋白質均屬於自然非折疊蛋白質，在疏水或脫水環境下能形成規則性的二級結構。此外，第四群晚胚蛋白質都能與非還原醣所構成的緊密玻璃質態，這可能是第四群晚胚蛋白質的一般特性。
我們的結論認為大豆第四群晚胚基因具有複雜的調控機制，其對應的蛋白質則屬於自然非折疊蛋白質。第四群晚胚蛋白質在疏水或脫水環境下重新折疊出大量有規則性，如-螺旋結構的二級結構，此外，第四群晚胚蛋白質與非還原醣在脫水環境下所形成的玻璃質態顯示兩者有密切的交互作用。晚胚蛋白質在玻璃質態中可能扮演一種類似船錨的角色，並使玻璃質態更為緊密。鑑於以上結果，根據蛋白質的「結構－功能」範式，我們認為第四群晚胚蛋白質可能依據細胞的含水狀態而具有雙重的功能。

Abstract
Late embryogenesis abundant (LEA) proteins are synthesized in seeds at a late stage of development. During this stage, the newly synthesized seed protein profile is very different from that of the mid-development stage and is associated with desiccation tolerance. LEA proteins have been widely reported from organisms including prokaryotes and fungus, and in plant and animal kingdoms and classified into 6 groups, on the basis of amino acid sequence and conserved motifs, or 4 groups, by computer analysis with the POPP package. Gene expression pattern, localization analysis, protein chemical properties, and transgenic plant analysis has led to the proposal that LEA proteins may play a protective role in cell survival under desiccation. In higher plants, anhydrobiosis (orthodox) seeds may maintain germination ability for decades or even centuries under favorable conditions. Desiccation tolerance usually refers to tolerance to anhydrobiosis. The presence of LEA proteins and non-reducing sugars may be the most important and probably best-known key factors in acquisition of desiccation tolerance in seeds. The interaction between LEA proteins and sugars is suggested to be involved in enhanced desiccation tolerance. However, further investigations are required. For this objective, the soybean LEA 4 proteins were used as model system to investigate protein characteristics and the relation between LEA proteins and sugars.

The first part of this thesis is the characterization of GmPM16 protein, a soybean LEA 4 protein. The protein has a low molecular weight, high pI value, and an unusual amino acid residue distribution along the protein. Northern blot and digital Northern analysis indicate that GmPM16 transcripts are expressed mainly in seeds. In situ hybridization study reveals that the transcripts are detected in cotyledon mesophyll cells but not in the vascular system of mature or pod-dried soybean seeds. Circular dichroism (CD) analysis and Fourier transform infrared (FTIR) spectroscopy revealed that GmPM16 proteins in solution are highly disordered, possessing only a small portion of -helical structures, and the proteins in sodium dodecyl sulfate (SDS) or trifluoroethanol (TFE) solutions exhibit abundant -helical structures. Proteins and non-reducing sugars do not interact in the aqueous state. However, GmPM16 proteins undergo dehydration-induced conformational changes and adopt high amount -helical structures. Drying the mixture of proteins and non-reducing sugars results in the formation of a glassy state with relatively high glass transition temperature (Tg) and low wavenumber-temperature coefficient (WTC) values. It is thus suggested that GmPM16 proteins interact with sugar and form tightly glassy matrixes in the dry state. The protein-sugar glasses may play a role in reducing cellular damage in drying seeds by changing protein conformation as well as forming tight cellular glassy matrices.
The second part in this thesis is the characterization of 2 soybean LEA 4 proteins, including basic GmPM1 protein and acidic GmPM28 protein. Digital Northern analysis indicates that GmPM1 transcripts are expressed mainly in seeds and abiotic-stressed vegetative tissues, while GmPM28 transcripts are expressed specifically in seeds. The hydrodynamic data and calculation of folding status indicate that the hydrophilic LEA proteins belong to natively unfolded proteins. CD and FTIR spectra indicate that both proteins in solution are highly unfolded. However, CD spectra collected at different temperatures illustrate that the protein exists in equilibrium between 2 extended conformational states, such as unordered and left-handed extended helical or poly (L-proline)-type II structures. Both proteins adopt a largely ordered conformation in the presence of -helical promoting co-solvents or in the dry state but proteins and nonreducing sugars do not interact in the aqueous state. Drying a mixture of GmPM1 or GmPM28 proteins and nonreducing sugars results in the formation of a glassy matrix with higher Tg and lower WTC values. All 3 different types of soybean LEA 4 proteins studied adopt dehydration-induced -helical conformations and interact with sugars in the dry state, which indicates that the changes may be common characteristics of LEA 4 proteins.
We conclude that soybean Lea 4 genes contain complicated expression patterns and their proteins are members of natively unfolded proteins. LEA 4 proteins give hydrophobic- and dehydration-induced conformational changes, from highly disordered to highly -helical structures, and interact with sugars to form a protein-sugar glassy matrix in the dry state. High Tg and low WTC values suggest that these proteins may be anchored to compact the cellular bioglasses. According to the protein structure-function paradigm, it is suggested that LEA 4 proteins may play bipartite roles depending on the aqueous status of cells.

List of abbreviations---------------------------------------------------------------------- 1
Chinese abstract--------------------------------------------------------------------------- 2
Abstract------------------------------------------------------------------------------------ 5
Chapter 1 Introduction------------------------------------------------------------------- 8
Anhydrobiosis---------------------------------------------------------------- 8
Acquisition of desiccation tolerance in seeds---------------------------- 9
Aims of this thesis----------------------------------------------------------- 13
References-------------------------------------------------------------------- 14
Chapter 2 Literature Review------------------------------------------------------------ 18
Introduction------------------------------------------------------------------- 18
The classification of LEA proteins---------------------------------------- 19
The regulation of Lea genes------------------------------------------------ 28
Anhydrobiosis and LEA proteins------------------------------------------ 36
The structure of LEA proteins--------------------------------------------- 44
Conclusion-------------------------------------------------------------------- 51
Tables-------------------------------------------------------------------------- 52
References-------------------------------------------------------------------- 57
Chapter 3 Gene cloning and characterization of a soybean (Glycine max L.) LEA protein, GmPM16-----------------------------------------------------
78
Introduction------------------------------------------------------------------- 78
Materials and Methods------------------------------------------------------ 81
Results------------------------------------------------------------------------- 85
Discussion-------------------------------------------------------------------- 92
Tables-------------------------------------------------------------------------- 96
References-------------------------------------------------------------------- 98
Figure legends---------------------------------------------------------------- 103
Figures------------------------------------------------------------------------- 106
Chapter 4 Characterization of two soybean (Glycine max L.) LEA 4 proteins - CD and FTIR studies--------------------------------------------------------
114
Introduction------------------------------------------------------------------- 114
Materials and Methods------------------------------------------------------ 116
Results------------------------------------------------------------------------- 120
Discussion-------------------------------------------------------------------- 126
Tables-------------------------------------------------------------------------- 133
References-------------------------------------------------------------------- 135
Figure legends---------------------------------------------------------------- 141
Figures------------------------------------------------------------------------ 143
Chapter 5 General Discussion---------------------------------------------------------- 150
The effects of hexa His tag in recombinant group 4 LEA proteins--- 150
Desiccation tolerance and LEA proteins---------------------------------- 151
The expression of Lea 4 genes--------------------------------------------- 151
Role of LEA 4 proteins in dehydrating tissues--------------------------- 153
Role of LEA 4 proteins in anhydrobiotic tissues------------------------ 155
Conclusion and future work------------------------------------------------ 158
References-------------------------------------------------------------------- 160
Tables-------------------------------------------------------------------------- 164
Chapter 6 Appendix---------------------------------------------------------------------- 165
FTIR techniques------------------------------------------------------------- 165
Crystallization studies------------------------------------------------------- 166
NMR analysis---------------------------------------------------------------- 166
Transgenic plants------------------------------------------------------------ 167
Heterogenous expression--------------------------------------------------- 168
In vitro translation assay---------------------------------------------------- 168
The expression patterns of Arabidopsis Lea genes---------------------- 168
Tables-------------------------------------------------------------------------- 170
Figure legends---------------------------------------------------------------- 171
Figures------------------------------------------------------------------------ 172

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