木芙蓉花色形成的表型研究
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摘要
木芙蓉(Hibiscus mutabilis)又命芙蓉、拒霜花,为锦葵科、木槿属,落叶灌木或小乔木。本研究以不同花色木芙蓉为材料运用测色法,紫外分光光度法,化学显色法,植物成分分析等方法,从花色表型测定与分析,理化因子对类黄酮呈色的影响,木芙蓉花色素的化学分析等方面对木芙蓉花色的形成进行了研究。为木芙蓉品种分类以及新的花色品种的培育奠定基础,同时,为进化生物学研究提供参考依据。本试验主要结果如下:
     1.使用目视测色和色差仪测色两种方法对木芙蓉3个品种的花色进行评价,经综合分析将木芙蓉花色分成红色(red group)、白色(white group)、粉色(pink group)三个色系。运用色差仪对木芙蓉的花色进行了测量。结果发现,木芙蓉不同花色品种的亮度(L~*)和红度(a~*)表现出显著的负相关关系,即随着红度的降低,花色的亮度越来越高;L~*与黄度(b~*)表现出显著的正相关关系,这说明随着黄度的提高,花色的亮度也相应地增加。
     2.本试验研究了温度、光照、酸碱性和金属离子对木芙蓉花色素在离体条件下呈色的影响,结果发现:①55℃以下的温度对类黄酮呈色的影响不明显,只有红色、粉色品种花青素随温度升高略有降解;②光照可以明显影响红色品种花青素的呈色,花青素在光下容易降解,使颜色变浅,且光照越强降解幅度越大;白色系品种的黄酮类化合物对光照稳定;③pH值对红色系花青素的呈色影响较大。在pH值3.8缓冲体系中花青素具有最大的吸光度,之后随着溶液pH值的升高颜色变浅、λmax红移,部分品种花青素在pH>7.0时变绿,说明花青素的结构已经被破坏;pH值对白色系品种影响不大,仅pH>7.0时产生浑浊,同时白色系品种类黄酮变黄;④Fe~(3+)、Mg~(2+)、Al~(3+)、Ca~(2+)对红色系‘牡丹红’花青素有增色作用,金属离子对白色系‘醉芙蓉’白色花瓣黄酮类化合物无明显影响。
     3.在本研究中,使用紫外-可见光谱对木芙蓉花瓣中存在的类胡萝卜素、黄酮类化合物和花青素三类色素进行了分析,结果发现:①黄酮类化合物的特征吸收峰在331nm和271nm附近,存在于所有的木芙蓉品种中,由于它的光吸收特性,因此大部分化合物表现为无色或者很淡的黄色;②类胡萝卜素的特征吸收峰是442nm和471nm,属于可见光区,所以大多会表现出黄色到橙黄等花色。类胡萝卜素存在于红色和粉红木芙蓉花瓣中,表现出黄色或者与其他类型花色素共同形成花色;③花青素的特征吸收峰在525和252nm附近,在525nm处的光吸收,使花青素表现出非常鲜艳的花色,花青素存在于红色系和粉色系花瓣中。
     4.用紫外分光光度法分别测量了木芙蓉三个品种花瓣的总黄酮、花青素和类胡萝卜素的含量。结果发现:不同色素种类和含量的差异是形成花色色系的直接原因,总黄酮、总花青素和总类胡萝卜素的含量与亮度(L~*)呈负相关关系;红色和粉红系的主要色素花青素与红度(a~*)呈显著的正相关关系,而白色系的主要色素类胡萝卜素与黄度(b~*)同样也呈显著的正相关关系。总黄酮和总花青素的含量通常与总类胡萝卜素呈现负相关,而黄酮类化合物作为花青素的前体物质,与花青的含量呈现显著的正相关关系。
Hibiscus mutabilis belongs to Malvaceae,deciduous shrub or subtree.lt has a long history of cultivation in home and a broad. The research was based on the various flower color of Hibiscus mutabilis from Hu Nan. The methods of different color measurement (checked using normal eye and spectrophotometer),observation by visible spectra,specific color reactions and chromatography were used in this dissertation. The most studies focused on the three aspects:the first was the measurement and analysis of flower color; the second the effect of Physical and chemical factors on flower coloration:the third was the phytochemistry of flower pigments of Hibiscus mutilis petal. These data is useful for planting and breeding of the Hibiscus mutabilis.The results are indicated as follows:
     1.The flower color of 3 Hibiscus mutabilis cultivars was checked using normal eye and spectrophotometer.Lightness(L~*) and chromatic component (a~* and b~*)of these cultivars are obtained by spectrophotometer.According to the CIELAB color coordin-ate,all cultivars were divided into three groups:red, white,and pink based on the Q cluster analysis.In the same time,the system of flower color measurement of Hibiscus mutabilis was discussed.The digitization of flower color is very important for the studies of flower color of ornamental plants. Analysis of the CIELAB color coordinate of flower color,the relationship of significant negative correlations between a* and L~*was founded.That is,from green to red of flower color,the less lightness the flower has. But the significant positive correlation between b~* and L~* was showed in white, pink groups and red groups respectively.Those results show that a~* and b~* are the important factor for lightness and flower coloration.
     2.The color expression of flavonoids in white cultivars is not sensitive to temperature, light,and pH value.The color expression of anthoeyanins of red and pink cultivars is stable under 55℃in methanol and sensitive to light and pH value. Anthoeyanins degradate significantly in light.The red color of anthoeyanins in some cultivars fades gradually and has a bathoehromiec shift ofλ_(vis-max):as The increase of pH value in solution.some metal ions, such as Fe~(3+),Mg~(2+),Al~(3+),Ca~(2+)are positive to red cultivar'mu dan hong' .Metal ions do not affect flavonoids in cultivar'zui fu rong',obviously. These results emphasize that anthocyanins play significant roles in red and pink cultivars of the Hibiscus mutabilis.
     3. The flower pigment sarevery important for flower coloration.The study by UV-visible spectra indicate that flower pigments of Hibiscus mutabilis are consist of carotenoids,anthocyanins and other flavon oids.The characteristic absorption peak of carotenoid was 440nm and 470nm,the anthoeyanin was 531nm,and the other Flavonoid was 331nm and 271nm. There is only flavone in white color flower;the red and pink flower contains anthoe-yanlns and other flavonoids.
     4.To understand the quantitative effect of major pigment s on flower color variation in Hibiscus mutabilis cultivars,flavornols (FL),carotenoids (CA), chlorophylls(CH) and anthoeyanins(AN) were extracted from the flowers and analyzed both qualitative-ly and quantitatively with UV-visible spectral.The results indicated that all of the cultivars contain flavonol,which are colorless and act as the copigment for some color groups.The anthoeyanins were responsible for flower colors of the red,and pink, its content is important for the flower coloration.The relationship of significant negative correlations between total anthoeyanins(TA),total carotenoid (TC), total flavonoids (TF) and L~* was founded.The significant positive correlations between TA and a~*,TC and b~*were founded respectively.The content of TC is negatively related to TF and TA,but the content of TF is positively related to TA.At last,according to the content of flower pigments,all cultivars were classified into five groups:FLgroup,containing flavonols;FL,AN group,anthocyanins and flavonols;FL,CA group flavonols and carotenoids;FL,AN,CA group, anthocyanins, flavonols and carotenoids; FL, CH, CA, group,flavonols,chlorophylls and carotenoids.The classification according to flower color and pigments is correlative and this will be benefited for cultivar classification of Hibiscus mutabilis.
引文
[1]中国植物志编辑委员会.湖南植物志[M].湖南科技出版社.
    [2]邾枝花,夏伦祝.木芙蓉研究进展[J].安徽医药,2005,9(5):324~325.
    [3]符诗聪,张凤华,史炜镔,等.木芙蓉有效组分的抗炎实验研究初步报道[J].上海第二医科大学学报,2001,21(1):14~16.
    [4]张慧娟,马 景,姚莉韵等.木芙蓉叶有效组分致突变与急性毒性实验研究[J].广西科学,2002,9(1):53~56.
    [5]沈钦海,陈国民等.木芙蓉对四氯化碳大鼠急性肝损伤的保护作用[J].湖北中医杂志,2004,21(10):3~4.
    [6]姚莉韵,陆阳,陈泽乃.木芙蓉叶化学成分研究[J].中草药,2003,34(3):201~203
    [7]唐玉贵,陈尔.10个木芙蓉品种在南宁的引种试验[J].西部林业科学,2008,37(4):80~82.
    [8]宋娟娟,庄东红.木槿属几种植物的染色体数目及其倍性的研究[J].热带亚热带植物学报,2001,9(3):213~216.
    [9]陈有民.园林树木学[M].北京:中国林业出版社,1990.
    [10]林浩然,郑幼兰,陈仁通等.木芙蓉治疗滴虫性阴道炎及霉菌性阴道炎的实验和临床研究[J].医学研究通讯,1990,19(10):22~5.
    [11]程金水编著.园林植物遗传育种学[M].北京:中国林业出版社,2000:23~32.
    [12]安田齐著.花色的生理生化[M].北京:中国林业出版社,1989.
    [13]梁祖霞(1998)花的色彩之谜.科技潮,12(3):58~59.
    [14]余树勋,吴应祥(1993)花卉词典.农业出版社,北京,336.
    [15]孟繁静(2000)植物花发育的分子生物学.中国农业出版社,北京,225~269.
    [16]苏焕然(1996)花色与色素.生物学杂志,13(5):46~47.
    [17]徐文辉,高海卿,陈华进(2000)菊花某些性状遗传规律的初步探讨.浙江林学院学报,17(1):37~41.
    [18]包满珠,1997.植物花青素基因的克隆进展及其应用-文献综述[J].园艺学报,24(3):279~284.
    [19]汪政科,彭镇华,2000.观赏植物基因工程研究进展[J].林业科学研究,13(1):97~102
    [20]林泉,色素基因的表达和调控.见:许智宏,刘春明主编.植物发育的分子机理[M],北京:科学出版社.1998.107~119.
    [21]徐昌杰,张上隆,2000.植物类胡萝卜素的生物合成及其调控[J].植物生理学通讯,36(1):64~70.
    [22]傅荣昭,马江生,曹光诚等,1995.观赏植物色香形基因工程研究进展-文献综述[J].园艺学报,22(4):381~387.
    [23]Mol J,Holton T A, Koes R E,1995. Genetic engineering of commercial traits of floral crops [J].Trend Biotechnol,13:350~355.
    [24]Meyer P,Heidmann I,Forkmann G,et al,1987. Anewpetunia flower color generated by transformation of a mutantwith a maize gene[J].Nature,330:677~687.
    [25]Van der krol A R, Lenting R J,Veenstra J G,et al,1988. An antisense chalone synthase gene in transgenetic plants inhibits flowerpigmentation [J].Nature,333:860~869.
    [26]LuC Y, Nugent G, Terse W R,et al,1991. Agrobacterium-mdicatd tannsformation of carnation (Dianthus carryophyllus L.) [J].Bio/Technology,9:864~868.
    [27]Aanhane T, Affrs D R, Lorigene B VF,et al,1995. First rDNA flower to debut in Australia this summer.Biotechnology News[J],15(14):3-7.
    [28]Robinson KEP, Firoozabady E,1993. Transformation of floriculture crops [J].SciHorticul,55: 83~99.
    [29]颜华,宋云,李羽云等,1997.查尔酮异构酶基因的克隆序列分析及在大肠杆菌中的表达[J].植物学报,39(1030~1034).
    [30]傅荣昭,马江生,曹光诚等,1995.观赏植物色香形基因工程研究进展-文献综述[J].园艺学报,22(4):381~387.
    [31]储文华,1996.用根癌农杆菌在丝石竹进行基因转化[J].上海师范大学学报(自然科学版),25(4):90~91.
    [32]Paeeh K.Colour development in flowers [J].Annual Review of Plant Physiology,1955,6: 273~298.
    [33]白新祥,戴思兰,菊花花色研究进展[C]中国园艺学会观赏园艺专业委员会.中国观赏园艺研究进展2004.北京:中国林业出版社,2004a:131~137.
    [34]Tanaka Y,TsudaS,Kusumi T.MetaLbolic engineering to modify flower color [J].Plant Cell physiology,1998,11:1119~1126.
    [35]何小玲,王金发.1998.观赏花卉的品质基因及其基因工程问题[J].植物生理学通讯,34(6):462~466.
    [36]郑志亮,1994.花卉作物的花色基因工程[J].北方园艺,21(3):37~38.
    [37]哈本著.黄酮类化合物[M].北京:科学出版社,1983.
    [38]Kobayashi N,Schmidt J,Nimtz M,Wray V,Schliemann W.Betalains from Christmas cactus [J] Phytoehemistry,2000,54:419-26.
    [39]Yoshida K,Yki T K,Kameda K,Kondo T.Sepal color variation of Hydrangea macrophylla and vacuolar pH measured with a protonselective microelectrode [J].plant Cell physioog-y,2003(a),44(3):262~268.
    [40]Markhama K R,Ryana K G,Gouldb K S,Rickardsc G K.Cell wall sited flavonoids in lisianthus flower petals [J].Phytochemistry,2000(a),54(7):681-687.
    [41]Yasuda H.Studies on "bluing effecf"in the petals of red rose II.observation on the development of the tannin body in the upper epidermal cells of bluing petals [J]. Cytologia, 1974,39:107~112.
    [42]Hemleben V,Anthoeyanin carrying structures in specific genotypes of Matthiola incana R.Br.Z [J].Naturibrsch,1981,36c:925-927.
    [43]Asen S,Stewart R N,Norris K H.Anthoeyanin,flavonol copiglments,and pH responsible for larkspur flower colour [J].Phytochemistry,1975a,14:2677-2682.
    [44]Markam K R,Gould K S,Winefield C S,Mitchell K A,Bloor S J,Boase M R.Anthoeyanic vacuolar inclusions-their nature and significance in flower colouration [J]. Phytoehemistry, 2000(b)55:327~336.
    [45]Gonnet J E.origin of the color of Cv.Rhapsody in blue rose and some other so-called"blue"roses[J].J.Agrie.Food Chem,2003,51:4990-4994.
    [46]Iwashina T.The Structure and Distribution of the Flavonoids in Plants[J] Journal of Plant Researeh,2000,113:287~299.
    [47]谭仁祥著.植物成分分析[M].北京:科学出版社,2002,486~495.
    [48]Anderseno M.Markam K R.Flavonoids:chemistry,biochemistry,and applications [M]. FL: CRC,Taylor&Franeis,2006.
    [49]Grotewold E.The Geneties and Biochemistry of Floral Pigments[J].Annual Review of Plant Biology,2006,57:761~780.
    [50]Cook N Cand Samman S.Flavonoids-chemistry, metabolism, cardioprotective effects,and dietary sources.Nutritional Bioehemistry,1996,7:66~76.
    [51]Williams C A,Grayer R J.Anthoeyanlns and other flavonoids[J].Natural Product report,2004, 21:539~573.
    [52]高锦明著.植物化学[M].北京:科学出版社,2003:166~169.
    [53]Forkmann G.Flavonoids as flower pigments the formation of the natural spectrum and its extension by genetic engineering [J].Plant Breeding,1991,106:1~26.
    [54]Freitas V,Mateus N.Chemical transformations of anthoeyanins yielding a variety of colours (Review) [J].Environ Chem Lett,2006,4:175~183.
    [55]Holton T A,Cornish E C.Geneties and Bioehemistry of Anthoeyanin Biosyn thesis[J].The Plant Cell,1995,7:1071~1083.
    [56]惠柏棣编著.类胡萝卜素化学和生物化学[M].北京:中国轻工业出版社,2005.
    [57]Kost H P.Handbook of Chromatography Plant pigments volumel[M]:Fat-Soluble Pigments. New York:CRC Press,Inc.1991.
    [58]Paeker L.Methods in Enzymology, Volume 213, Part A,chemistry, separation, quantitation and antioxidation [M].Academic Press,INC.1992.
    [59]朱长甫,陈星,王英典.植物类胡萝卜素生物合成及其相关基因在基因工程中的应用[J].植物生理与分子生物学学报,2004,30(6):609~618.
    [60]Nielsen K,Deroles S C,Markham K R,Bradley M J,Podivinsky E,Mansonl D.Antisense flavono. thase alters copigmentation and flower color in lisianthus[J].Molecular Breeding, 2002,9;217~229.
    [61]陶俊,张上隆,徐昌杰,安新民,张良诚.类胡萝卜素素的相关基因及其基因工程[J].生物工程学报,2002,18(3):276~281.
    [62]Strack D,Wray V(1989) Anthocyanins.In JB Harbornee,ed,Method in Plant Biochemistry voll. AcademicPress,London,PP.2000,26:325~356.
    [63]王长泉,刘涛,王宝山.植物甜菜素研究进展[J].植物学通报,2006,23(3):302~311.
    [64]Kobayashi N,Schmidt J,Nimtz M,Wray V,Schliemann W.Betalains from Christmas cactus[J] Phytoehemistry,2000,54:419~26.
    [65]Sehliemann W,Cai Y,Degenkolb T,Sehlnidt J,Corke H.Betalains of Celosia argentea [J]. Phytoehemistry,2001,58:159~165.
    [66]Kugler F,StintZing F C,Carle R.Charaeterisation of betalain patterns of differently coloured inflorescenees from Gomphrena globosa Land Bougainvillea sp.by HPLC-DAD-ESI-MS[J] Anal Bioanal Chem,2007,387:637-648.
    [67]Jurd L,Asen S.The formation of metal and coopiglment complexes of cyaniding 3-glucoside [J] Phytoehemsitry,1966,5:1273~1281.
    [68]Stewart R N,Noris K H,Asen S.Microspectrophotometric measurement of pH and pH effect on color of petal epidermal cells[J].Phytoehemistry,1975,14:837~942.
    [69]Mazza G,Brouillard R.The mechhasm of copigmentation of anthoeyanins in aqueous soluti-ons [J].Phytoehemistry,1990,29:1097~1102.
    [70]Mol J,Comish E,Mason J.Novel colored flowers[J].Current Opininon in Bioteeh,1999,10:198-201.
    [71]Tanaka S F,Inagald Y,Yamaguchi T,Saito N,Iida S. Color-enchaneing protein in blue petals [J]. Nature,2000,407:581.
    [72]Brouillard R Origin of the exceptional colour stability of the Zebrina anthocyanin Phytochem-istry,1981,20:143~145.
    [73]Stewart R N,Noris K H,Asen S.Microspectrophotometric measurement of pH and pH effect on color of petale epidemal cells[J].Phytoehemistry,1975,14:837~942.
    [74]Broulllard R,The in vivo expression of anthoeyanin colour in p lants [J]. Phytoehemistry,1983, 22:1311~1323.
    [75]Mol J,Grotewold E,Koes R E.How genes Paint flowers and seeds [J].Trends in Plant science, 1998,3:212~217.
    [76]安田齐著.花色之谜[M].北京:中国林业出版社,1989.
    [77]Yamaguchi T,Tanaka F S,Inagaki Y,Saito N,Sakakibara K Y,Tanaka YKusumi T,Iida S. Genes eneoding the vacular Na~+/H~+exchanger and flower coloration[J].Plant and cell Physiology, 2001,42:451~61.
    [78]Weiss D,Halevy AH(1989) Stamens and gibberellin in the regulation of corolla pigmentation and growth in Petunia hybrida Planta 179;89~96.
    [79]Giuliano G,Bartley GE,Scolnik PA(1993)Regulation of carotenoid biosynthesis during tomato development. The Plant Cell,5379~5387
    [80]Hayashi T,Ueda M,Isobe M,Goto T.A new molecular mechanism of blue color development with protocyanin,a supramolecular pigment from conflower cyanus [J].Tetralledron Letters, 1998,39(45):8307~8310.
    [81]Kondo T K,Yoshida A,Nakagawa T. Struetural basis of blue-color development in fiower petals from Commelina communis[J].Nature,1992,358:515~518.
    [82]Kondo T,oyama K,Yoshida K.Chiral Molecular Recognition on Formation of a Metal oan tho-eyanin:A Supramolecular Metal Complex Pigment from Blue Flowers of salvia Patens[J]. Angewandte Chemie International Edition,2001,40(5):894~897.
    [83]Takeda K,Tominago S.The anthocyanin in blue flowers of centaurea cyanus [J].Bot May Tokyo,1983,96:359.
    [84]Takeda M,Itoi H.Bluieing of sepal color of Hydrangea macrophylla [J]. Phytochemistry,1989, 24:2251-2254.
    [85]Yasuda H.Studies on" bluing effect" in the Petals of red rose.observation on the develoPment Of the tannin body in the upper epidermal cells of bluing Petals [J]. Cytologia,1974,39:107-112.
    [86]Tanaka Y,Katsumoto Y,Brugliera F,Mason J.Genetic engineering in floriculture [J].Plant cell, tissue and organ culture,2005,80:1~24.
    [87]Tucher A O,Maciarello M J,Tueker S S.A suivey of color charts for biologieal description [J]. Taxon,1991,40:201~214.
    [88]Voss D H.Relating colorimeter measurement of Plant color to the Ro yal Horticultural Society ColourChart[J].Hortseienee,1992,27:1256~1260.
    [89]R.H.S.The Royal Horticultural Soeiety Colour Chart.2001,London.
    [90]本恩.颜色技术原理[M].李小梅,马如,陈立荣等译.北京:化学工业出版社,2002,50~145.
    [91]苏焕然(1996)花色与色素.生物学杂志,13(5):46~47.
    [92]胡晓丹,谢笔钧,王建中.金盏菊花黄色素的性质研究[J].北京林业大学学报,2001,23(5):35~38.
    [93]赵昶灵,郭维明,陈俊愉.梅花花色色素种类和含量的初步研究[J].北京林业大学学报,2004,26(2):68~73.
    [94]王重庆编著.高级生物化学实验教程M.北京:北京大学出版社1994:167~168.
    [95]白新祥,戴思兰.反义RNA技术在花色育种中的应用[J],植物学通报,2005,22(3):284~291.
    [96]安田齐著.花色之谜[M].北京:中国林业出版社,1989.
    [97]Dyrby M,Westergard N,StaPelfeldt H.Light and heat sensitivity of red cabbage extract in soft drink model systems[J].Food chemistry,2001,72:431-437.
    [98]Clement J S,Mabry T J.Pigment evolution in the Caryophyllales:A systematic overview [J]. Bot.Aeta,1996,109:360~367.
    [99]Preece T F,Wileox H J.Stage of growth of the chrysanthemum flower jnt Pathology,1966, 71~72.
    [100]宋光泉,古练权.荔枝果皮花色素营的光分解研究.中山大学学报,1999,38:58~60.
    [101]刘虎俊.富士果皮中色素的测定.林业科技通讯,1998,21~23.
    [102]ShvartsM,BorochorA,WeissD.Low temperature enhances petunia flower pigmentation and iduces chalcone synthase gene expression [J].Physiologia Plantarum,1977,99(1):67~72.
    [103]Mazza G, Brouillard R-The mechemism of copigmentation of anthocyanins in aqueous solutions[J].Phytochem,1990,29:1097~1102.
    [104]Baranac T M,Petranovic N A,Dimitric-Markovic J M,Spectrophotometric study of anthocy-anin copigmentation reactions [J].Agr and Food Chem,1996,44:1333~1336.
    [105]Anna B, Alicjazk Jan O.The effects of heating, UV-irradiation and storage on stability of the anthocyanin-polyphenol copigment complex[J].Food Chem,2003,18:349~355.
    [106]Dong A,Cevalloscasals,Luis C Z.Stability of anthocyanin-based aqueous extracts of Andean purple corn and red-fleshed sweet potato compared to synthetic and natural colorants [J].Food Chem,2004,86:69~77.
    [107]Donald K,Dougal,l David C.Anthocyanins from wild carrot suspension cultures acylated with supplied carboxylic·acid[J]-Carbohydate Research,1998,310:77~189.
    [108]Mazza G,Brouillard R.The mechemism of copigmentation of anthocyanins in aqueous solutions [J].Phytochem,1990,29:1097~1102.
    [109]马自超,吴伟志.有关花青素色素发色机理研究介绍[J].中国野生植物,1989(3):5~7.
    [110]柯雅星,陈慧君.木芙蓉花治疗痈的疗效观察[J].中国社区医师,2002,24(3):99~102.
    [111]keda K.Mettakko-anthocyanins Ⅱ Further experiments of synthesizing crystalline blue metalloanthocyanins using various kinds of bivalentmetals[J].Proceedings of the Japan Academy,1997.
    [112]杨少勇,安银岭,樊国盛.蓝色花植物花色素的着色机理[J].北京林业大学学报,2003(5):53:257~261.
    [113]LipphardtS,BrettcshjoneiderR,Kreuzaler F.UV-Inducide transient expression in parsley protoplasts identifies regulatory ciselements of chimeric Antirrhinum majus chalconesy-nthase gene [J].EMBO J,1997 (7):4027~4033.
    [114]朱新贵,邓捷,郭勇.玫瑰茄悬浮细胞合成花青素的光效应研究[J].工业微生物,1999(1):25~27.
    [115]陈孝泉著.植物化学分类学[M].北京:高等教育出版社,1990:55~63.
    [116]马卡姆.黄酮类化合物结构鉴定技术[M].北京:科学出版社,1990:42~58.
    [117]Nair,M.,Asweeter pill to swallow,Chem.Brit.1999,35,16.
    [118]Chen ZW,Hu YZ,WuHH, JiangHD. Synthesis and biological evaluation of flavonoids as vasorelaxant agents [J].Bioor-ganic& Medicinal Chemistry Letters,2004 (14):3949~3952.
    [119]张华峰,王瑛,黄宏文.类黄酮生物合成途径的进化及其在淫羊藿中的研究展望[J].中草药.
    [120]白新祥.菊花花色形成的表型分析[D].北京林业大学学士论文,2007.
    [121]白新祥,胡可,戴思兰,王亮生.不同花色菊花品种花色素成分的初步分析[J].北京林业大学学报,2006,28(5):84~89.
    [122]Karl H D.A rapid method for the extraction and quantitative of total anthocyanin of cranber-ry fruit [J].Agriculture food ehemistry,1978,26(6):1452-1453
    [123]Sakata Y,Aoki N,Tsunematsu S.Petal colouration and pigmentation of tree peony bred and selected in Daion Island(Shimane prefecture)[J].Journal Japanese Society Horticulturae Science,1995,64(2):351~357.
    [124]何小颜著.花与中国文化[M].北京:人民出版社,1999:219~226.
    [125]黄济明著.花卉育种知识[M].北京:中国林业出版社,1987:65~71.
    [126]李燕梅,易莲爱,温平,赵资智,朱永亮,张竞.菊花电子束辐射诱变育种的研究[J].核农学通报,1992,13(3):113~115.
    [127]蒋甲福,陈发棣,郭维明.小菊杂种一代部分性状的遗传与变异[J].南京农业大学学报,2003,26(2):11~15.
    [128]栗茂腾,余龙江,王丽梅,刘建民,雷呈.菊花花色遗传及花色嵌合体发现[J].遗传,2005,27(6):948~952.

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