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研究生：	梁哲語 Liang, Che-Yu
論文名稱：	評估紫藍色蝴蝶蘭顏色生成之原因 Assessment of violet-blue color formation in Phalaenopsis orchids
指導教授：	陳虹樺 Chen, Hong-Hwa
學位類別：	碩士 Master
系所名稱：	生物科學與科技學院 - 生命科學系 Department of Life Sciences
論文出版年：	2017
畢業學年度：	105
語文別：	英文
論文頁數：	57
中文關鍵詞：	酸鹼值、蝴蝶蘭、紫羅蘭-藍色、花青素、金屬離子、矢車菊素、飛燕草素、DpF3’5’H 、PeAHA 、PeF3’H 、PeMYB2 、PhF3’5’H
外文關鍵詞：	anthocyanin, cyanidin, delphinidin, DpF3’5’H, metal ions, PeAHA, PeF3’H, PeMYB2, pH, PhF3’5’H, Phalaenopsis, violet-blue
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蝴蝶蘭 (Phalaenopsis spp.) 為全世界重要的經濟花卉作物。蝴蝶蘭花卉中涵蓋許多的顏色，包括紅-紫色 (red-purple)、紫色 (purple)、紫-紫羅蘭色 (purple-violet)、紫羅蘭色 (violet)及紫羅蘭-藍色 (violet-blue)。然而紫羅蘭-藍色的蝴蝶蘭相較其他顏色的蝴蝶蘭是非常稀少，所以蘭花育種者及科學家都非常有興趣育種紫羅蘭-藍色的蝴蝶蘭，且期望育出真正藍色的蝴蝶蘭。花青素(anthocyanin)、液胞中酸化程度 (vacuolar acidification levels) 及金屬離子 (metal ions)為三個影響花色形成的主要因子。本篇論文擬分析紫羅蘭-藍色的蝴蝶蘭顏色生成原因，並將透過暫時性表現 (transient overexpression) 異質的 F3’5’H (flavonoid 3’,5’-hydroxylase) 以確認能否使飛燕草素 (delphinidin) 在蝴蝶蘭中累積並形成藍色。結果顯示從紫色到紫羅蘭-藍色的蝴蝶蘭中都呈現以矢車菊素為基底的花青素 (cyanidin-based anthocyanin)，然而飛燕草素在這些顏色的蝴蝶蘭中皆不存在。這與在不同顏色的蝴蝶蘭中的 PeF3’H (Phalaenopsis equestris flavonoid 3’-hydroxylase) 基因皆為高表現而 PhF3’5’H (Phalaenopsis hybrid flavonoid 3’,5’-hydroxylase) 基因幾乎不表現之明顯差異的基因表現結果一致。透過暫時性表現飛燕草的 DpF3’5’H (Delphinium hybrid flavonoid 3’,5’-hydroxylase ) 在紫色蝴蝶蘭栽培種 P. OX Honey ‘OX1372’ 無法明顯累積飛燕草素，然而增加暫時性表現PeMYB2 則可以有 8 % 飛燕草的累積，這表示單獨暫時性表現飛燕草的DpF3’5’H 並無法增加飛燕草素的累積。進一步測試飛燕草素的累積，暫時性表現 DpF3’5’H 及 PeMYB2 在白色蝴蝶蘭栽培種-P. Sogo Yukidian V3 中，呈現了高達 53.6% 以飛燕草素為基質的花青素(delphinidin-based anthocyanin) 累積在白花蝴蝶蘭中並呈現了新穎的藍色。然而在蝴蝶蘭中暫時性表現蝴蝶蘭的 PhF3’5’H 及 PeMYB2 顯示 PhF3’5’H 幾乎喪失了飛燕草素累積的能力。透過序列分析受質辨識位六號 (substrate recognition site 6, SRS6)，發現 PhF3'5'H 的 SRS6 序列中的第八號位胺基酸為纈安酸 (valine, V)，另外在第五及十號位胺基酸為脯胺酸 (proline, P)，而其他物種的 F3'5'H 的 SRS6 序列中第八號位則為丙胺酸 (alanine, A) 或絲氨酸 (serine,S)、第五號位及十號位胺基酸則分別為苯丙胺酸 (phenylalanine, F) 及榖氨醯胺 (Glutamine, Q)，顯示蝴蝶蘭的 PhF3'5'H 中 SRS6 序列與其他物種的 F3'5'H 有明顯差異。進一步以分子入坞預測 (molecular docking prediction)，發現以naringenin (N), dihydroquercetin (DHQ) 及 eriodictyol (E) 為受質，在DpF3'5'H中，受質的B-ring 芳香環 (aromatic ring)結合方向朝向SRS6，而PhF3'5'H呈現相反的受質結合方向，即B-ring 芳香環 (aromatic ring) 結合方向遠離 SRS6。此外，紫羅蘭-藍色的蝴蝶蘭的花瓣具有較高的酸鹼值 (pH5.33~5.54)，而紫色蝴蝶蘭花瓣則具有較低的酸鹼值 (pH4.77~5.04)。分離鑑定了五個氫離子傳輸蛋白的基因，分別命名為 PeAHA1~PeAHA5 ，其中 PeAHA1 及 PeAHA5 在不同顏色中具差異性的基因表現的結果，即在紫羅蘭-藍色的蝴蝶蘭有較低的 PeAHA5 基因表現，而在紫色蝴蝶蘭則有較高的 PeAHA5 基因表現。PeAHA1 則和 PeAHA5 相反的基因表現模式。推測這二個基因與紫羅蘭-藍色的蝴蝶蘭有較高的酸鹼值而紫色蝴蝶蘭則具有較低的酸鹼值呈現高度的相關性。進一步，在紫色及紫羅蘭-藍的蝴蝶蘭中，測量了五種金屬離子（鎂、鋁、鈣、鐵及鋅離子）的濃度及五種金屬離子與花青素的莫耳數比 (molar ratio)。在紫羅蘭-藍的蝴蝶蘭中，鋁離子、鈣離子及鐵離子與花青素的莫耳數比分別為紫色蝴蝶蘭的 3.6 倍、2.5 倍及3 倍，但鎂離子及鋅離子與花青素的莫耳數比則無明顯差異。綜合上述，液胞中酸化的不足及含較高的鋁離子、鈣離子及鐵離子與花青素的莫耳數比可能是造成紫羅蘭色-藍色的蝴蝶蘭形成的主因，但並非因為飛燕草的累積所致。

Phalaenopsis spp. is an important cash crop worldwide. There are abundant flower colors in Phalaenopsis ranging from red-purple, purple, purple-violet, violet, and violet-blue hues. However, violet-blue color is less bred than the other colors, so breeders and geneticists are very interested in breeding violet-blue orchids, and expect to produce ‘true blue’ orchids. Anthocyanin, vacuolar pH and metal ions are three major factors influencing flower color. This study aims to identify the factors causing violet-blue color in Phalaenopsis flowers as well as analyze whether delphinidin accumulation and blue pigmentation formation can be achieved by transient overexpression of heterologous F3’5’H (flavonoid 3’,5’-hydroxylase) in Phalaenopsis. Results showed that cyanidin-based anthocyanin was majorly accumulated in various colors of Phalaenopsis flowers ranged from purple to violet-blue colors, while delphinidin was not detected. This was concomitant with the fact that higher expression of PeF3’H (Phalaenopsis equestris flavonoid 3’-hydroxylase) and little to no expressions of PhF3’5’H (Phalaenopsis hybrid flavonoid 3’,5’-hydroxylase) in various colors of Phalaenopsis. Transient overexpression of DpF3’5’H from Delphinium hybrid in purple flower P. OX Honey ‘OX1372’ resulted limited delphinidin accumulation. However, transient overexpression of both DpF3’5’H and PeMYB2 resulted 8% delphinidin accumulation. It suggests that only overexpression of DpF3’5’H was not sufficient for delphinidin accumulation in transient assay. Furthermore, upto 53.6% delphinidin accumulation and novel blue-hue were resulted in white flower P. Sogo Yukidian V3 cultivar by transient overexpression of both DpF3’5’H and PeMYB2. In contrast, transient overexpression of both PhF3’5’H and PeMYB2 did not result delphinidin accumulation, indicating that PhF3’5’H was lack of the ability of delphinidin accumulation. Sequence analysis showed that the substrate recognition site 6 (SRS6) of PhF3’5’H has valine (V) at position 8, and with proline (P) at both positions 5 and 10, while the F3’5’Hs from other plants have alanine (A) or serine (S) at position 8, and phenylalanine (F) at position 5 and glutamine (Q) at position 10. Prediction of molecular docking of the substrates including naringenin, dihydroquercetin and eriodictyol showed their binding direction of aromatic rings (B-ring) were oriented toward SRS6 of DpF3'5'H, while the substrate binding direction is away from the SRS6 of PhF3'5'H. In addition, the pH values of violet-blue and purple Phalaenopsis flowers were ranged from 5.33-5.54 and 4.77-5.04, respectively. To understand the proton transport, five PeAHAs were identified from OrchidBase, namely PeAHA1~PeAHA5. Among them, PeAHA5 showed lower expression in the violet-blue cultivars than in the purple cultivars, while PeAHA1 showed a converse expression profiles. This may be correlated to the higher vacuolar pH in violet-blue color of Phalaenopsis than that in both purple and white color ones. Furthermore, concentration of five metal ions including Mg2+, Al3+, Ca2+, Fe3+ and Zn2+ were measured in white, purple, extreme deep-purple and violet-blue Phalaenopsis flowers. The molar ratio of Al3+, Ca2+ and Fe3+ metal elements to anthocyanin were 3.6 X, 2.5 X, and 3.0 X higher in violet-blue color than those in the purple color Phalaenopsis, respectively. In contrast, there were no significant differences in the molar ratio of Mg2+ and Zn2+ metal elements to anthocyanin among various flower colors. Conclusively, violet-blue color formation in Phalaenopsis was caused by the absence of vacuolar acidification as well as the higher molar ratio of Al3+, Ca2+ and Fe3+ metal elements to cyanidin-based anthocyanin than that in purple color Phalaenopsis, but not due to the formation of delphinidin.

中文摘要	I
Abstract	III
致謝	V
Contents	VI
List of Tables	IX
List of Figures	X
List of Appendix Figures	XII
Abbreviations	XIII
1. Introduction	1
1.1. Phalaenopsis orchids	1
1.1.1	Economic importance of Phalaenopsis	1
1.1.2 Breeding of blue color flower in Phalaenopsis	1
1.2 Importance of flower color in higher plants	2
1.2.1 Pigment components of flower color	2
1.2.2 Anthocyanin	2
1.2.3 Flower of blue color	2
1.3 Biological roles of anthocyanin	2
1.3.1 The biosynthesis pathway of anthocyanin	2
1.3.2 F3’H and F3’5’H in the anthocyanin biosynthesis pathway	3
1.3.3 Genetic engineering for producing blue color flowers	3
1.3.4 PeMYB2 transcription factor in regulation of floral pigmentation in Phalaenopsis spp.	4
1.4 Biological roles of H+ concentration in vacuole	4
1.4.1 pH value and flower color-shift	4
1.4.2 Transporter on tonoplast for the regulation of vacuolar acidification	5
1.5 Biological roles of vacuolar metal ions	5
1.5.1 Structure of metalloanthocyanin	5
1.5.2 Metal ions and blue color formation	6
2. Purpose	7
3. Materials and methods	8
3.1 Plant materials	8
3.2 Definition of flower colors	8
3.3 Extraction of anthocyanin	9
3.4 High-performance liquid chromatography (HPLC)	9
3.5 RNA extraction and reverse transcription to cDNA	10
3.6 5’-RACE and 3’-RACE	10
3.7 Quantitative real-time PCR	10
3.8 Sequence alignment and phylogenetic analysis	11
3.9 Molecular modeling and docking	11
3.10 Transient overexpression of heterologous F3’5’Hs and PeMYB2 in Phalaenopsis flowers	12
3.11 pH value measurement and visible absorption spectra of corolla homogenates	12
3.12 Metal ion measurement of the colorful petals	13
4. Results	14
4.1 Anthocyanin	14
4.1.1 Cyanidin-based anthocyanin is majorly accumulated in various colors of Phalaenopsis spp.	14
4.1.2 PeF3’H and PhF3’5’H belong to cytochrome 75B and 75A family, respectively	14
4.1.3 Strong expression of PeF3’H was concomitant to cyanidin-based anthocyanin in various Phalaenopsis spp.	15
4.1.4 Accumulation of delphinidin-based anthocyanin and blue pigmentation formation was rescued from transient overexpression both of DpF3’5’H and PeMYB2	15
4.1.5 Substrate recognition site 6 (SRS6) of PeF3’H and PhF3’5’H and prediction of substrate docking	16
4.1.6 Expression of F3’H and F3’5’H by transcriptomic analysis	17
4.2 Vacuolar pH value	17
4.2.1 Violet-blue color cultivars have higher vacuolar pH than purple and white cultivars	17
4.2.2 Absorption spectra of anthocyanin extract from Phalaenopsis	18
4.2.3 Five PeAHAs belong to H+-ATPase 3A family	18
4.2.4 Expression profiles of PeAHA1~PeAHA5 genes	19
4.3 Metal ions	19
4.3.1 Positive correlation between concentration of Mg2+ increased and cyanidin-based anthocyanin accumulation	19
5. Discussion	21
5.1 “True blue” color Phalaenopsis flower could be created by genetic engineering of anthocyanin biosynthesis pathway	21
5.2 F3’5’H was expressed in roots in addition to in floral organs	22
5.3 The same anthocyanin compound but with different vacuolar pH values between blue and red hue flowers	22
5.4 Vacuolar acidification is regulated by proton pump	22
5.5 Color-shift from violet-blue to purple color might be regulated from the divergence of molar ratio of Fe3+ to cyanidin-based anthocyanin in Phalaenopsis	23
6. Conclusion and perspectives	24
7. References	25
Table 1. Primers used in this study.	30
Table 2. List of concentration of metal elements and anthocyanin and their molar ratio.	32

Figure 1. Color definition according to Royal Horticultural Society Color Chart (RHSCC).	33
Figure 2. Flowers used in this study, and their definition of color code.	34
Figure 3. HPLC analysis of anthocyanin compound.	35
Figure 4. Phylogenetic tree of F3’5’H and F3’H.	36
Figure 5. Quantitative RT-PCR analysis of F3’H and F3’5’H.	37
Figure 6. HPLC of anthocyanin compound in P. OX Honey ‘OX1372’ with transient overexpression of PhF3’5’H or DpF3’5’H.	38
Figure 7. HPLC of anthocyanin compound in P. OX Honey ‘OX1372’ with transient overexpression PhF3’5’H or DpF3’5’H with the presence of PeMYB2.	39
Figure 8. HPLC of anthocyanin compound in P. Sogo Yukidian ‘V3’ with transient overexpression both PhF3’5’H or DpF3’5’H with the presence of PeMYB2.	40
Figure 9. Multiple alignment of deduced amino acid sequences of F3’5’H and F3’H from P. Purple Martin.	41
Figure 10. Multiple alignment of substrate recognition site 6 (SRS6) with F3’Hs and F3’5’Hs.	42
Figure 11. Multiple alignment of SRS1, SRS2, SRS4, SRS5 and SRS6 between DpF3’5’H and PhF3’5’H.	43
Figure 12. Substrate docking of F3’5’H and its substrates.	44
Figure 13. Expression of both of F3’5’H and F3’H in various tissues of Phalaenopsis with FPKM values in transcriptomics analysis.	45
Figure 14 . Expression of both of F3’5’H and F3’H in various organs of Phalaenopsis and Vanilloideae by detection of FPKM and RPKM values in transcriptomics analysis.	46
Figure 15. Assessment of pH value in petal crude extracts of various colors cultivars of Phalaenopsis.	47
Figure 16. Absorption spectra of anthocyanin extract from P. Big Chili.	48
Figure 17. Absorption spectra of anthocyanin extract from P. OX Honey ‘OX1372’.	49
Figure 18. Absorption spectra of anthocyanin extract at different time points.	50
Figure 19. Phylogenetic tree of PeAHA1-PeAHA5.	51
Figure 20. Quantitative RT-PCR analysis of five P-type H+ ATPase.	52
Appendix 1. Flower color of various flowering development stages of violet-blue P. Kenneth Schubert and P. Purple Martin	53
Appendix 2. Six native species used as parents for breeding blue-hue Phalaenopsis	54
Appendix 3. The three major classes of plant pigments	55
Appendix 4 . The anthocyanin biosynthesis pathway	56
Appendix 5. Anthocyanin production and qRT-PCR analysis of expression profile analysis in the flowers in transient assay of overexpression PeMYB2	57

                                    

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