中文摘要
糖類(carbohydrate)和氮素(nitrogen)的代謝相互關聯而共同地影響植物的各種生理功能，因此我們藉探討二者對細胞內基因表現的調控方式，以進一步瞭解植物的生長發育及對環境適應的機制。
首先探討糖調控基因表現的分子機制，以水稻懸浮培養細胞為材料，利用differential screening的方法從細胞缺糖4小時的cDNA library中獲得一群受糖抑制表現的基因(sugar down-regulated, SD genes)；而獲自日本水稻基因組分析實驗室的一群與細胞生長有關的基因，經北方墨點法實驗則證明它們都是受糖誘導表現的基因(sugar up-regulated, SU genes)。再分別以nuclear run-on assay的方法，或培養液中加入actinomycin D抑制RNA轉錄的方法，已經確定這些受糖誘導或抑制表現的基因，是受調控於轉錄或轉錄後的階層（transcriptional or posttranscriptional levels），而這種調控作用對同一群屬於SD或SU的基因也具有一致性。
接著我們探討糖的訊息傳遞途徑，以葡萄糖、蔗糖或葡萄糖的類似物3-OMG (3-O-methylglucose)及6-dG (6-deoxyglucose)處理水稻懸浮培養細胞，證明可以被磷酸化及代謝的糖，如葡萄糖及果糖(由蔗糖分解而來)均能誘導SU基因並抑制SD基因的表現；然而不能被磷酸化及代謝的糖，如3-OMG及6-dG，則無法誘導SU基因或抑制SD基因的表現。而糖代謝的中間產物pyruvate會抑制某些SD基因的表現，但SU基因的表現則不受影響，這些結果顯示糖訊息傳遞途徑與糖代謝途徑有重疊之處。利用蛋白質激(protein kinase)的抑制劑staurosporine及蛋白質去磷酸( protein phosphatases 1及2A )的抑制劑okadaic acid 處理水稻懸浮培養細胞，證明蛋白質的磷酸化作用及去磷酸化作用也參與糖調控SU及SD基因的表現。In-gel kinase活性分析顯示糖訊息可能經由一種38-KD的蛋白質激傳遞，最後抑制SD基因的表現。而缺糖訊息的傳遞則可能與55-、66-及68-KD的一組蛋白質激的活性有關，最後促進SD基因的表現。
穀類種子發芽時，儲存於胚乳中的養分會被分解為糖，氨基酸或較小的胜以供給幼苗的生長利用。已知cysteine proteinase (CysP)是分解胚乳中儲藏性蛋白質最主要的酵素。為了要進一步瞭解CysP在水稻生理上所扮演的角色及其基因表現的調控機制。我們已經從水稻的genomic library中篩選得到一個CysP基因OsEP3A (Oryza sativa endoproteinase 3A)。從DNA序列分析及RT-PCR的結果顯示OsEP3A不含intron。利用南方墨點法證明它在水稻基因組中為單一基因，且至少由5～7個類似的基因組成一個基因族。北方墨點法顯示OsEP3A基因主要表現於發芽的種子及老化葉片中；而缺氮的水稻懸浮培養細胞也會誘導OsEP3A mRNA的大量累積。利用農桿菌(Agrobacterium)的基因轉殖技術，將OsEP3A啟動子/gusA的嵌合基因轉殖到水稻細胞，證明啟動子的活性的確受到氮素所調控。以GUS染色法分析發芽中的轉殖種子，也顯示啟動子的活性受到植物發育時期，組織特異性及賀爾蒙所調控。
為了進一步探討雙子葉植物的種子於萌芽期間，α-amylase及CysP基因表現的差異，以瞭解兩者在雙子葉植物種子萌芽時的重要性。我們以空心菜為材料，發現在乾燥種子中便已累積大量的α-amylase及CysP mRNAs，當種子發芽初期兩種酵素的基因又會被誘導表現；其中胚乳的表現量在發芽第0.5天時達最高量，此後兩者mRNA的累積量均快速下降，到第3.5天時幾乎消失。然而在胚軸及子葉中則偵測不到α-amylase基因的表現，而CysP mRNA在萌芽胚軸中的累積情形與在胚乳中類似，在子葉中的表現則於萌芽第0.5天時逐漸被誘導表現到第2.5天時達最高量，之後其mRNA的累積量便快速的減少。由此證明CysP在雙子葉空心菜種子萌芽時似乎擔任較重要的角色。
碳素及氮素是構成細胞的主要成分，兩者的代謝緊密的相互關連並影響植物的各種生理功能。綜合上述的結果，證明糖及氮素調控著許多不同基因的表現，而其訊息首先可能經由kinase來傳遞，再藉著一連串的訊息傳遞階梯，最後以協同調控的方式導致目標基因的活化及抑制作用，以便達到碳/氮平衡來維持細胞最佳之代謝機轉。目前許多研究報告推測細胞是以全面性的調控(general control)方式來維持碳素及氮素代謝的平衡，這也是將來值得研究的方向之一。

英文摘要
In plants, carbohydrate and nitrogen metabolism are tightly-linked physiological functions and are essential for growth and development. The aim of the present study is to elucidate the mechanisms of sugar and nitrogen regulation of gene expression, which may help us better understand how plants respond to developmental and environmental cues during their life cycle.
First, to study the mechanisms that control sucrose-dependent gene expression, we performed differential screening of a cDNA library constructed from poly(A)+ mRNA prepared from 4 h sucrose-starved rice cells, and obtained six genes whose expression was sugar down-regulated (SD genes). Meanwhile, Northern blot analysis demonstrated that expression of several cDNAs, which encode proteins known to be required for cell growth, were sugar up-regulated (SU genes). Nuclear run-on transcription analysis and an assay with inhibition of mRNA transcription with actinomycin D demonstrated that expression of SU and SD genes in cultured rice suspension cells were differentially and coordinatelly regulated by sucrose at both transcriptional and posttranscriptional levels.
Next, to study the sugar signal transduction pathway, rice suspension cells were treated with sucrose, glucose, or glucose analogs. The results showed that glucose by itself or glucose and fructose hydrolyzed from sucrose all activated the expression of SU genes and suppressed the expression of SD genes. In constrast, the glucose analogs, 3-OMG (3-O-methylglucose) and 6-dG (6-deoxyglucose), which cannot be phosphorylated and metabolized by cells, did not activate the expression of all the SU genes but suppressed the expression of certain SD genes in sucrose-starved sells. Pyruvate suppressed the expression of certain SD genes, which suggests that the sugar signal transduction pathway overlap the metabolic pathway for gene repression. Study with protein kinase and protein phosphatase inhibitors, staurosporine and okadaic acid, respectively, demonstrated that protein phosphorylation and dephosphorylation were involved in sugar-regulated expression of SU and SD genes. In-gel kinase activity assays revealed that a 38-KD protein kinase activity might be involved in the suppression of the SD genes by sugars, while a group of 55-, 66- and 68-KD protein kinase activity might be involved in the activation of the SD genes under sugar starvation.
During the germination of cereal grains, sugar, amino acids, and small peptides derived from hydrolysis of endosperm nutrients are taken up by the embryos to support the growth of seedling. Cysteine proteinases (CysP) play a major role in the degradation of endosperm storage proteins. To investigate the significance of CysP expression in rice physiology and the regulatory mechanism of CysP expression, we isolated the genomic clones OsEP3A (Oryza sativa endoproteinase 3A). Genomic Southern blot analysis revealed that OsEP3A was a single copy gene and there were at least 5 to 7 genes encoding OsEP3A homologs in the rice genome. Northern blot anaylsis revealed that OsEP3A gene expression was preferentially expressed in germinating seeds and senescing leaves of rice. In cultured rice suspension cells, the expression of OsEP3A was up-regulated by nitrogen starvation. The OsEP3A promoter was linked to the coding sequence of a reporter gene, gusA, encoding b-glucuronidase (GUS), and stably introduced into the rice genome through an Agrobacterium transformation-mediated transformation system. The OsEP3A promoter was sufficient to confer the nitrogen metabolic regulation of GUS expression in cultured suspension cells. Histochemical study also indicated that the OsEP3A promoter was sufficient to confer the temporal, spatial and hormonal regulation of GUS expression in germinating seeds.
We also studied the expression pattern of a-amylase and CysP in the germinating seeds of a dicot plants, water spinach (Ipomoea auuatica Forsk). Accumulation of a-amylase and CysP mRNAs was aboundant in mature dry seeds. After the onset of germination, levels of both mRNAs in endosperm were enhanced and reached at peaks at day 0.5, then decreased rapidly to a very low level at day 3.5. However, no a-amylase mRNA was detected in axises and cotyledons of germinating seeds. The expression patterns of CysP mRNA were similar in both axises and endosperms during seeds germination. The accumulation of CysP mRNA in cotyledons increased gradually 0.5 day after imbibition, reached a peak at day 2.5, and then decreased sharply.
In plants, carbon and nitrogen are the two most important elements in maintaining cell structure and life. These two elements control expression of a variety of genes related to growth and metabolism. As cells are capable of maintaining a balance in ratio of carbon to nitrogen, suggesting that the synthesis and utilization of these two elements are tightly controled. It would be interesting to study in future whether there exist general mechanisms for sugar and nitrogen regulation of gene expression, and whether the sugar and nitrogen signal transduction pathways cross-talk in respond to changes in physiological and environmental conditions.

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