蝴蝶蘭的香味性狀一直是育種者的目標之一。大葉蝴蝶蘭為培育香味蝴蝶蘭的重要親本，其花部香味的主要成分為單萜類。過去的研究成果已經選殖及分析大葉蝴蝶蘭單萜類生合成之關鍵酵素PbGDPS。本研究為了探討影響蝴蝶蘭中單萜類合成的調控機制，首先建構具香味的大葉蝴蝶蘭花部轉錄體，並與不具香味的台灣阿嬤蝴蝶蘭花部轉錄分析比較後，確認參與單萜類生合成之酵素基因表現量差異乃是影響蝴蝶蘭中單萜類合成的關鍵因子。在影響PbGDPS基因表現量的因子中，先行分析反式元件。利用轉錄體的表現量聚合分析，在約2,300個轉錄因子中，選出了八個在大葉蝴蝶蘭中具有高表現量的轉錄因子，其中五個能夠反式激活參與大葉蝴蝶蘭單萜類合成的酵素基因，包含PbbHLH4、PbbHLH6、PbbZIP4、PbERF1與PbNAC1。進一步在台灣阿嬤蝴蝶蘭中的異位暫時表現結果中，PbbHLH4能最有效提升GDPS與單萜類合成酵素MTPS7的基因表現，並提升近千倍的單萜類合成量，而其他四個的效果較為微弱。顯示PbbHLH4是影響蝴蝶蘭單萜類合成的主要的反式因子。在探討影響GDPS表現的順式因子方面，藉由分析十二個具香味與不具香味蝴蝶蘭的GDPS啟動子序列，發現一個雙重覆片段僅存於在具香味蝴蝶蘭的GDPS啟動子。利用系列刪除實驗確認此雙重覆片段乃為決定GDPS啟動子活性的關鍵順式因子。進一步，利用阿拉伯芥的順式元件篩選酵母單雜交文庫實驗，分離出會結合上此雙重覆片段的轉錄因子，AtbZIP18。並在大葉蝴蝶蘭的花部轉錄體中，篩選出在DNA結合區域與AtbZIP18具有83%相似度的PbbZIP4。PbbZIP4能夠反式激活具雙重覆片段的GDPS啟動子，卻無法激活不具有雙重覆片段的GDPS啟動子序列。PbbZIP4在台灣阿嬤蝴蝶蘭中的異位暫時表現能有效促進單萜類物質的產生。顯示此雙重覆片段是一個影響GDPS表現與單萜類合成的重要順式元件。最後，在探討影響蝴蝶蘭單萜類香味合成的外在與內在因子實驗，發現外在的光因子與內在的生物週期韻律因子皆會影響蝴蝶蘭的單萜類花香釋放。本研究結果不僅能使我們對單萜類花香調控有更進一步的了解，亦能應用在蝴蝶蘭產業，創造出更具吸引力的香味蘭花。

Floral scent trait is one of the important novel targets for orchid breeders. Phalaenopsis bellina has a charming floral scent and is widely used as a breeding parent for the scent phenotype. The main floral volatiles in P. bellina are monoterpenes. Previous studies have cloned and analyzed PbGDPS, the key enzyme involved in the monoterpenes biosynthesis in P. bellina. To study the regulation of floral scent biosynthesis in Phalaenopsis orchids, floral transcriptome libraries of scented P. bellina were constructed and compared to that of the scentless P. aphrodite. Significant differential expression of GDPS between P. bellina and P. aphrodite, indicating that the elevated GDPS expression in the scented P. bellina is critical for its monoterpene accumulation. To identify the upstream factors regulating GDPS in Phalaenopsis orchids, I focused on the trans-factor first. By comparative transcriptome analysis, eight candidate transcription factors (TFs) out of 2360 TFs were showed to enhance differential expression between the two transcriptomes. Among them, five TFs, including PbbHLH4, PbbHLH6, PbbZIP4, PbERF1, and PbNCA1, showed transactivation for monoterpene biosynthesis genes. Ectopic transient expression of these TFs in the scentless P. aphrodite revealed that PbbHLH4 most profoundly induces the monoterpene phenotype with a ~1000-fold increase of monoterpenoid production, while the other 4 showed minor effects. These results indicate its major role for monoterpene biosynthesis in Phalaenopsis orchids. Next, I examined the cis-factor affecting the expression of GDPS. By analyzing the GDPS promoter fragments isolated from 12 scented and scentless orchids, a dual repeat cis-element is present only in the Phalaenopsis orchids emitting monoterpenes. Serial deletions of the GDPS promoter fragment demonstrates that this dual repeat is crucial for its promoter activities. Screening the Arabidopsis TFs cDNA library by using yeast one-hybrid assay reveals that AtbZIP18 is able to bind the dual repeat. PbbZIP4 is isolated in the transcriptome of P. bellina with an 83% identity in the DNA binding region. PbbZIP4 can only transactivate the GDPS promoter fragment containing the complete or near-complete dual repeat. In addition, ectopic transient expression of PbbZIP4 induces production of monoterpenes in the scentless orchid. These results indicate that the dual repeat is significant for GDPS gene expression, and thus subsequent monoterpene biosynthesis in Phalaenopsis orchids. Furthermore, by exploring the external and internal signals regulating the floral monoterpene emission, it is found to be under the control of light and circadian rhythm in Phalaenopsis orchids. Taken together, this study not only leads to a better understanding of the floral scents regulation in Phalaenopsis orchids, but also assist the marker-based selection as well as the molecular breeding of scented orchids.

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