光环境影响花青素合成途径中相关基因表达的机制
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摘要
本研究是以光敏感型津田芜菁和光不敏感型赤丸芜菁为试材，利用Northern杂交技术，对花青素生物合成途径中与花青素合成相关的主要结构基因PAL、CHS、CHI和AS及光信号传递相关的基因PHYA、COP1、CIP7和HY5的表达进行了检测。Northern分析采用的RNA样品，是从上述两品种的光处理和暗处理的肉质根部周皮层中提取的总RNA。采用的探针中，PHYA、COP1、CIP7、HY5及CHS基因的探针是利用芜菁的肉质根周皮层的总RNA合成，PAL和CHI基因的探针是利用草原龙胆花瓣的总RNA合成，AS基因的探针是利用草莓果实的总RNA合成。另外，利用HPLC液相分析系统，对两品种的光处理和暗处理样品中花青素和主要糖分(果糖、葡萄糖、蔗糖和麦芽糖)的含量进行了检测分析。对与花青素积累有明显相关的CHS基因进行了克隆。
通过实验得到如下结论：
1，光不敏感型的赤丸芜菁肉质根周皮中，光处理和暗处理都有大量花青素合成。而光敏感型的津田芜菁肉质根周皮中，只有光处理有大量花青素合成，暗处理几乎无花青素合成。
2，两品种肉质根及叶片等各组织的中葡萄糖的含量较果糖、蔗糖和麦芽糖的含量高，但葡萄糖的积累与花青素形成之间无明显相关性。两品种试材的肉质根肉质部分中四种糖的总含量高于其它部分，而叶片中含量极少。津田芜菁周皮中的果糖含量，光处理明显高于暗处理，且随光照处理时间的延长果糖含量逐渐提高，并以较高的含量稳定下来；其它三种糖含量光处理前后无明显变化。
3，CHS基因的表达，在光敏感型津田芜菁的光处理与暗处理之间有明显的差异；对津田芜菁地下根暗处理部分再进行光照处理，随光照处理的时间延长，CHS基因的表达量也逐渐提高，说明CHS基因的表达受光诱导，且表达量与花青素的积累呈正相关。在光不敏感型赤丸芜菁的不同光处理的试材中CHS基因的表达无明显的差异。因此，我们认为在光敏感型试材中CHS基因的表达对花青素的积累起关键的作用。AS基因的表达量与花青素的合成量相关，且在光敏感型津田芜菁中，AS基因的表达受光诱导。而光信号传递因子中PHYA的表达在两种品种试材中存在差异，只有光敏感型津田芜菁暗处理的试材中有表达。COP1和HY5基因在两种品种试材中，光处理与暗处理都表达，暗处理表达量比光处理少。CIP7的表达量则在两种品种试材间及光处理与暗处理中都无明显差异。
Two cultivars of Brassica campestris L. ssp. rapifera, light-independent 'Akamaru' turnip and light-dependent 'Tsuda' turnip, were used as test material in this experiment. The expression patterns of PAL, CHS, CHI, AS, PHYA, COP1, CIP7 and HY5 were analyzed by Northern Hybridization. PAL, CHS, CHI and AS are the principal structural genes correlated with anthocyanin biosynthesis in anthocyanin biosynthesis pathway; PHYA, COP1, CIP7 and HY5 are the genes correlated with light signal transmission in anthocyanin biosynthesis pathway. Total RNA samples used in Northern blotting were extracted from the light-exposed and light-interrupted cortex tissue of tuberous root of the two turnip cultivars. The probes of CHS, PHYA, COP1, CIP7 and HY5 genes were synthesized by total RNA extracted from the cortex tissue of tuberous root of turnip, the probes of PAL and CHI were synthesized by total RNA extracted from the petal tissue of Eustoma, and the probes of AS were synthesized by total RNA extracted from the fruit tissu
e of strawberry. The content of anthocyanin and main sugars (fructose, glucose, sucrose and maltose) in the light-exposed and light-interrupted samples of the two turnip cultivars were analyzed by HPLC. The CHS gene which is obvious correlated with anthocyanin accumulation was cloned as well.
The results are as follows:
1. A large quantity of anthocyanin was synthesized in both of the light-exposed and light-interrupted cortex tissue of tuberous root of light-independent 'Akamaru' turnip, but only in the light-exposed cortex tissue of tuberous root of light-dependent 'Tsuda' turnip. There was almost no anthocyanin synthesized in light-interrupted cortex tissue of tuberous root of light-independent 'Tsuda' turnip.
2. Glucose' s content was more abundant than that of fructose, sucrose and maltose in tuberous root and leaf tissues of the two cultivars, but there were no obvious correlations between glucose accumulation and anthocyanin biosynthesis. The content of the four sugars in tuberous tissue was higher than that in other tissues of tuberous root of the two cultivars. There were little sugars in leaf tissues. The content of fructose in the light-exposed cortex tissue of tuberous root of 'Tsuda' turnip was obviously higher than that in light-interrupted cortex tissue. The content of fructose was
increased gradually along with the prolongation of light-exposing time and stabled at high content level. There were no evident changes for the content of other three sugars after light-exposing.
3. There was distinct difference for the expression patterns of CHS gene in light-exposed and light-interrupted tissues of light-dependent 'Tsuda' turnip. Light-exposing the tuberous root of 'Tsuda' turnip grown under dark condition, the expression quantity of CHS gene was gradually increased along with the prolongation of light-exposing time. This indicated that the expression of CHS gene was induced by light, and the expression quantity was positively correlated with accumulation of anthocyanin. There was no distinct difference for the expression patterns of CHS gene in light-exposed and light-interrupted tissues of light-independent 'Akamaru' turnip. Therefore, we think that the expression of CHS gene played a key role in accumulation of anthocyanin in light-dependent materials. The expression quantity of AS gene was correlated with synthesis quantity of anthocyanin. The expression of AS gene was induced by light in light-dependent 'Tsuda' turnip. The expression of light signal transmission factor PHYA i
s different in the two cultivars. PHYA gene expressed only in light-interrupted part of light-dependent 'Tsuda' turnip. In the two cultivars, COP1 and HY5 expressed both in the light-exposed and light-interrupted parts of the two cultivars, but the expression quantity in light-exposed parts was higher than that in light-interrupted parts. The expression of CIP7 had no obvious difference in light-exposed and light-interrupted parts of two cultivars.
通过实验得到如下结论：
1，光不敏感型的赤丸芜菁肉质根周皮中，光处理和暗处理都有大量花青素合成。而光敏感型的津田芜菁肉质根周皮中，只有光处理有大量花青素合成，暗处理几乎无花青素合成。
2，两品种肉质根及叶片等各组织的中葡萄糖的含量较果糖、蔗糖和麦芽糖的含量高，但葡萄糖的积累与花青素形成之间无明显相关性。两品种试材的肉质根肉质部分中四种糖的总含量高于其它部分，而叶片中含量极少。津田芜菁周皮中的果糖含量，光处理明显高于暗处理，且随光照处理时间的延长果糖含量逐渐提高，并以较高的含量稳定下来；其它三种糖含量光处理前后无明显变化。
3，CHS基因的表达，在光敏感型津田芜菁的光处理与暗处理之间有明显的差异；对津田芜菁地下根暗处理部分再进行光照处理，随光照处理的时间延长，CHS基因的表达量也逐渐提高，说明CHS基因的表达受光诱导，且表达量与花青素的积累呈正相关。在光不敏感型赤丸芜菁的不同光处理的试材中CHS基因的表达无明显的差异。因此，我们认为在光敏感型试材中CHS基因的表达对花青素的积累起关键的作用。AS基因的表达量与花青素的合成量相关，且在光敏感型津田芜菁中，AS基因的表达受光诱导。而光信号传递因子中PHYA的表达在两种品种试材中存在差异，只有光敏感型津田芜菁暗处理的试材中有表达。COP1和HY5基因在两种品种试材中，光处理与暗处理都表达，暗处理表达量比光处理少。CIP7的表达量则在两种品种试材间及光处理与暗处理中都无明显差异。
Two cultivars of Brassica campestris L. ssp. rapifera, light-independent 'Akamaru' turnip and light-dependent 'Tsuda' turnip, were used as test material in this experiment. The expression patterns of PAL, CHS, CHI, AS, PHYA, COP1, CIP7 and HY5 were analyzed by Northern Hybridization. PAL, CHS, CHI and AS are the principal structural genes correlated with anthocyanin biosynthesis in anthocyanin biosynthesis pathway; PHYA, COP1, CIP7 and HY5 are the genes correlated with light signal transmission in anthocyanin biosynthesis pathway. Total RNA samples used in Northern blotting were extracted from the light-exposed and light-interrupted cortex tissue of tuberous root of the two turnip cultivars. The probes of CHS, PHYA, COP1, CIP7 and HY5 genes were synthesized by total RNA extracted from the cortex tissue of tuberous root of turnip, the probes of PAL and CHI were synthesized by total RNA extracted from the petal tissue of Eustoma, and the probes of AS were synthesized by total RNA extracted from the fruit tissu
e of strawberry. The content of anthocyanin and main sugars (fructose, glucose, sucrose and maltose) in the light-exposed and light-interrupted samples of the two turnip cultivars were analyzed by HPLC. The CHS gene which is obvious correlated with anthocyanin accumulation was cloned as well.
The results are as follows:
1. A large quantity of anthocyanin was synthesized in both of the light-exposed and light-interrupted cortex tissue of tuberous root of light-independent 'Akamaru' turnip, but only in the light-exposed cortex tissue of tuberous root of light-dependent 'Tsuda' turnip. There was almost no anthocyanin synthesized in light-interrupted cortex tissue of tuberous root of light-independent 'Tsuda' turnip.
2. Glucose' s content was more abundant than that of fructose, sucrose and maltose in tuberous root and leaf tissues of the two cultivars, but there were no obvious correlations between glucose accumulation and anthocyanin biosynthesis. The content of the four sugars in tuberous tissue was higher than that in other tissues of tuberous root of the two cultivars. There were little sugars in leaf tissues. The content of fructose in the light-exposed cortex tissue of tuberous root of 'Tsuda' turnip was obviously higher than that in light-interrupted cortex tissue. The content of fructose was
increased gradually along with the prolongation of light-exposing time and stabled at high content level. There were no evident changes for the content of other three sugars after light-exposing.
3. There was distinct difference for the expression patterns of CHS gene in light-exposed and light-interrupted tissues of light-dependent 'Tsuda' turnip. Light-exposing the tuberous root of 'Tsuda' turnip grown under dark condition, the expression quantity of CHS gene was gradually increased along with the prolongation of light-exposing time. This indicated that the expression of CHS gene was induced by light, and the expression quantity was positively correlated with accumulation of anthocyanin. There was no distinct difference for the expression patterns of CHS gene in light-exposed and light-interrupted tissues of light-independent 'Akamaru' turnip. Therefore, we think that the expression of CHS gene played a key role in accumulation of anthocyanin in light-dependent materials. The expression quantity of AS gene was correlated with synthesis quantity of anthocyanin. The expression of AS gene was induced by light in light-dependent 'Tsuda' turnip. The expression of light signal transmission factor PHYA i
s different in the two cultivars. PHYA gene expressed only in light-interrupted part of light-dependent 'Tsuda' turnip. In the two cultivars, COP1 and HY5 expressed both in the light-exposed and light-interrupted parts of the two cultivars, but the expression quantity in light-exposed parts was higher than that in light-interrupted parts. The expression of CIP7 had no obvious difference in light-exposed and light-interrupted parts of two cultivars.
引文
1．王曼，王小菁．监光、紫外光的受体及其对CHS表达诱导的研究．植物学通报，2000，19(3)：265-271
2．历采虹，邵宏波．高等植物花器官特异性基因研究．北方园艺．1999，126：7-8
3．包满珠．植物花青素基因的克隆及应用——文献综述．园艺学报．1997，24(3)：219-284
4．历彩虹，邵宏波．高等植物花器官特异性基因研究．北方园艺．1999，126：7-8
5．刘虎俊．富士果皮中色素的测定．林业科技通讯．1998，21-23
6．刘青林，陈俊愉．观赏植物花器官主要观赏性状的遗传与改良——文献综述．园艺学报．1998，25(1)：81-86
7．刘青林，陈俊愉．观赏植物花器官主要观赏性状的遗传与改良．园艺学报．1998，25(1)：81-86
8．孙大业，郭艳林，马力耕．细胞信号转导(第二版)．北京，北京科学出版社，1998
9．吕英民，张大鹏．果实发育过程中糖的积累．植物生理学通讯．2000，36(3)：258-265
10．池方，李树人，田红星．光照对套袋苹果花青素含姑的影响．河北农业大学学报．1997，31(2)：174-177
11．许智宏，刘春明．植物发育的分子机理．北京，北京科学出版社。1998
12．何奕昆，代庆阳，苏学辉．雁来红叶色转变与超微结构及色素含量的关系．四川师范学院学报．1995，16(3)：195-198
13．余叔文，汤章城．植物生理与分子生物学(第二版)．北京，北京科学出版社．1998
14．吴少华，张大生．花青素生成相关基因dfr研究进展．福建林学院学报．2002，22(2)：189-19
15．宋光泉，古练权．荔枝果皮花色素苷的光分解研究进展．中山大学学报．1998，38(5)：50-55
16．宋光泉，古练权．荔枝果皮花色素苷的光分解研究．中山大学学报．1999，38(5)：58-60
17．束怀瑞．果树栽培生理学．北京，中国农业出版社．1993
18．汪政科，彭镇华．观赏植物基因工程研究进展．林业科学研究．2000，13(1)：97-102
19．沈同，王镜岩主编．生物化学(第二版)．北京，高等教育出版社．1990
20．苏焕然．花卉基因工程研究进展．北方园艺．1996，09：26-27
21．陈俊伟，张良诚，张上隆．果实中的糖分积累机理．植物生理学通讯．2000，36(6)：497-503
22．周音，张智奇，殷丽青等．观赏植物基因工程研究进展及存在的问题．上海农业学报．2000，16(1)：90-96
23．周爱琴，祝军，生吉萍等．苹果花青素形成与PAL活性及蛋白质含量的关系．中国农业大学学报．1997，2(3)：97-99
24．孟繁静．植物花发育的分子生物学．北京，中国农业出版社．2000
25．洪宁，童哲．光敏色素在植物个体发育中的作用．植物生理学通讯．1998，34(6)：417
-422
26．郧忠亮．花卉作物的花色基因工程．北方园艺．1994，96：37-38
27．金波．一品红花色的探讨．园艺学报．1994，21(1)：87-90
28．胡桂兵，高飞飞，陈大成等．套袋对荔枝果皮中的色素、酚类物质和内源激素的影响．华南农业大学学报．2001，22(1)：31-34
29．赵宗方，宋亭华，高红胜．巨峰葡萄色素发育若干规律．江苏农学院学报．1989，10(4)：17-21
30．徐绍颖．植物生长调节剂与果树生产．上海，上海科技出版社．1987
31．阎隆飞，张玉麟．分子生物学(第二版)．北京，中国农业大学出版社．1997
32．黄卫东，原永兵，彭宜本．温带果树结实生理．北京，中国农业大学出版社．1994
33．曾骧．果树生理学．北京，中国农业大学出版社．1992
34．程林梅，张一萍，孙毅。草莓生物技术研究进展．植物学通报．2002，19(1)：39-43
35．潘增光，范晖，束怀瑞．苹果果实花青素形成与乙烯释放的关系．植物生理学通讯．1995，31(5)：338-340
36．潘增光，王国宾，李奎明等．新红星苹果果实着色期几种色素含量变化及其相关性．植物生理学通讯．1996，32(5)：347-349
37. Anju Singh, M. Tamil Selvi and Rameshwar shrma. Sunlight-induced anthocyanin. Journal of Experimental Botany, 1999, (150) 339: 1619-1625
38. Asako lida, Toru K azuoka et al. A zinc finger protein RHL41 mediates the light acclimatization response in Arabidopsis. The Plant Joumal, 2000, 24(2): 191-203s
39. Axel alesander hirner, Hanns Uirich seitz. Isoforms of chalcone synthase in Daucus carota L. and their differential expression in organs from the european wild carrot and in ultraviolet-A-irradiated cell cultures. Planta. 2000 (2) 10: 933-998
40. Chory J, Chatterjee-M, Cook-R-K, Elich-T. From Seed germination to flowering, light controls plant development via the pigment. Biological Abstracts. 1997/01-1997/06
41. Christoph Poppe, Uta sweere, Helge Drumm-herrel and Eberhard schafer. The blue light receptor cryptchrome 1 can act independently of phytochrome A and B in Arabidopsis thaliana. The Plant Journal. 1998, 16(4): 465-471
42. David Jackson, Keith Roberts and Cathie Martin. Temporal and spatial control of expression of anthocyanib biosynthetic genes in developing flowers of antirrhinum majus. The Plant Journal 1992, 2(4): 425-434
43. Dongsun Cao, Yun Lin, and Chi-Lien Cheng. Genetic interactions between the Chlorate-resistanrt mutant cr88 and the photomorphogentic mutants copl and hy5. The Plant Cell. 2000, 12: 199-210
44. Enriqueta Moyano, Ignacioportero-robles et al. A fruit-specific putative
dihydroflavonol 4-Reductase Gene is differentially expressed in strawbrrry during the ripening process. Plant Physiol. 1998, 117: 711-716
45. Hou Yongmin, Von Albrecht G et al. A new class of Arabidopsis constomorphogenic genes involved in regulating cotyledon development. Biological Abstracts. 1993/01-1993/06
46. Jennie A. Jackson, Gareth I. Jenkins. Extension-growth responses and expression of flavonoid biosynthesis genes in the Arabidopsis hy4 mutant. Planta. 1995, 197: 233-239
47. Keiko U. Torii, Timony W. McNellis and Xing-Xing Deng. Functional dissection of its three structural modules in light control of Seeding development. The EMBO Journal. 1998, (17): 19 5577-5587
48. Kwok-Shing-F, Piekos-Barry et al. A complement of ten essential and pleiotropic Arabidopsis COP/DET/FUS genes is necessary for repression of photomorphagenesis in darkness. Biological Abstracts. 1996/01-1996/06
49. Kwok-Shing-F, Solano-Roberto, Tsuge-Tomohiko, Chamovitz-Daniel-A et al. Arabidopsis homologs of a c-Jun coactivator are present both in monomeric from znd in the COP9 complex, and their abundance is differentially affected by the pleiotropic cop/det/fus mutations. Biological Abstracts. 1999/01-1999/06
50. Lay-Hong Ang, sudip Chattopadhyay et al. Molecular interaction between COP1and HY5 Defines a Regulatory switch for light control of Arabidopsis development. Molecular Cell. 1998,1,213-222
51. L.H.J. Kerckhoffs, P.M. Kelmenson, M.E.L. schreuder, et al. Characterization of the gene encoding the apoprotein of phytochrome B2 in tomato, and identification of molecular lesions in two mutant alleles. Mol Gen Genet. 1999, 261: 901-907
52. Margaret Ahmad and Anthony R. Cashmore. The blue-light receptor cryptochrome 1 shows functional dependence on phytochrome A or phytochrome B in Arabidopsis thaliana. The Plant Journal. 1997, 11(3): 421-427
53. Margaret Ahmad, Jose A. Jarillok, and Anthony R, Cashmore. Chimerc proteins between cry1 and cry2 Arabidopsis blue light photoreceptors indicate overlapping functions and varying protein stability. The Plant Cell. 1998, 10: 197-207
54. Mary L..Durbin, Bonnie McCaig and Michael T. Clegg. Molecular evolution of the chalcone synthase multigene family in the morning glory genome. Plant Molecular Biology. 2000, 42: 79-92
55. Mayer-Raphael, Raventos_Dora, Chua-Nam-Hai. Det1, cop1, and cop9 mutations. Biological Abstracts. 1997/01-1997/06
56. Melanie Kaltenbach, Gudrun schroder, Elmon schmelzer, Verena Lutz and Joachim schroder. Flavonoid hydroxylase from Catharanthus roseus: cDNA, heterologous expression, enzyme properties and cell-type specific expression in plants. The Plant Journal. 1999, 19(2),: 183-193
57. Michal Oren-shamir, Ada Levi-Nissim. UV-light effect on the leaf pigmentation of cotinus coggygria 'Royal Purple' Scientia Horticulturae 1997, 71: 59-66
58. Mundt-Kirsten-E, porte-Joanne, Murray-Johanne-M, et al. The COP9/signalosome complex is conserved in fission yeast and has a role in s Phsase. Biological Abstracts. 2000/01-2000/06
59. Naoki Yamamoto, Xing-Wang Deng. Protein nucleocytoplasmic transport and its light regulation in plants. Genes to Cells. 1999, 4, 489-500
60. Partt LH. Cordonnier MM. Phytochrome and Photoregulation Plants. Academic Press. 1987, 83.
61. R. Chaturvedi, R. shyam and P.V. sane. steady state levels of D1protein and psbA thanscript during UV-B inactivation of photosystem Ⅱ in wheat. Biochemistry and Molecular Biology Internation. 1998, (44)5: 925-932
62. Rhonda L. Feinbaum, Gisela storz, and Frederick Mausubel. High intensity and blue light regulated expression of chimeric chalcone synthase genes in transgenic Arabidopsis thaliana plants. Mol Gen genet. 1991, 226: 449-456
63. Roldan-Marta, Gomes-mena-Concepcion, Ruiz-Garcia-Leonor, salinas-Julio, Martinez-zapater-Jose-M sucrose availability on the aerial part of the plant promotes morphogenesis and flowering of Arabidopsis in the dark. Biological Abstracts. 2000/01-2000/06
64. Ronald E. Koes, Francesca Quattrocchio and Joseph N.M. Mol. The flavonoid biosynthetic pathway in plants: function and evolution. Bioessays. 1994, 16. 2
65. Saneyuki Kawabata, Yuhua Li, et al. Role of Photoreceptor - and sugar-mediated Reactions in Light Dependent Anthocyanin Production in Lily and stock Flowers. J. Japan. Soc. Hort. Sci. 2002
66. Serino-giovanna, Tsuge-Tomohiko, Kwok-shing, Matsui-minami, Wei-Ning, Deng-Xing, Wang. Arabidopsis COP8 and fus4 mutations define the same gene that encode subunit 4 of the COP9 signalosome. Biological Abstracts. 2000/01-2000/06
67. Staub-jeffrey-M, Wei-Ning; Deng-Xing-Wang. Evidence for FUS6 as a component of the nuclear-localized COP9 complex in Arabidopsis. Biological Abstracts. 1997/01-1997/01
68. Szekeres-Miklos, Nemeth-Kinga et al. Brassinasteroids rescue the deficiency of CYP90 a cytochrome p450 controlling cell elongation and de-dtiolation in
Arabidopsis. Biological Abstracts. 1996/01-1996/06
69. Tim Kunkel, Gunther Neuhaus, Alfred Batschauer, Nam-Hai Chua and Eberhard schafer. Functional analysis of yeast-derived phytochrome A and B phycocyanobilin adducts. The Plant Journal. 1996, 10(4): 625-636
70. Timothy A. Holton, Edwina C. Cornish. Genetics and biochemistry of anthocyanin biosynthesis. The Plant Cell. 1995,7: 1071-1083
71. Toshihiro Toguri, Naoyuki Umemoto, Osamu Kobayashi and Takeshi Ohtani. Activation of anthocyanin synthesis genes by white light in eggplant hypocotyls tissues, and identification of an inducible P-450 cDNA. Plant Molecular Biology. 1993, 23: 933-946
72. Waiters-Robin-6, Gogers-jennifer-J-M, Shephard-Freya, Horton-Peter. Acclimation of Arabidopsis thaliana to the light environment: the role of photoreceptors. Biological Abstracts. 2000/01-2000/06
73. Yoshiharu Y. Yamamoto, minami Matsui, Lay-Hong Ang, and Xing-Wang. Role of a COP1 interactive protein in mediating light-regulated gene expression in Arabidopsis. The Plant Cell. 1998,10: 1083-1094
74. Zhou-D-X., Kim-Y-J, Li-Y-F, Carol-P, Mache_R. COP1b, an isoform of COP1 generated by alternative splicing, has a negative effect on COP1 function in regulating light-dependent seeding development in Arabidopsis. Biological Abstracts. 1998/01-1998/06
2．历采虹，邵宏波．高等植物花器官特异性基因研究．北方园艺．1999，126：7-8
3．包满珠．植物花青素基因的克隆及应用——文献综述．园艺学报．1997，24(3)：219-284
4．历彩虹，邵宏波．高等植物花器官特异性基因研究．北方园艺．1999，126：7-8
5．刘虎俊．富士果皮中色素的测定．林业科技通讯．1998，21-23
6．刘青林，陈俊愉．观赏植物花器官主要观赏性状的遗传与改良——文献综述．园艺学报．1998，25(1)：81-86
7．刘青林，陈俊愉．观赏植物花器官主要观赏性状的遗传与改良．园艺学报．1998，25(1)：81-86
8．孙大业，郭艳林，马力耕．细胞信号转导(第二版)．北京，北京科学出版社，1998
9．吕英民，张大鹏．果实发育过程中糖的积累．植物生理学通讯．2000，36(3)：258-265
10．池方，李树人，田红星．光照对套袋苹果花青素含姑的影响．河北农业大学学报．1997，31(2)：174-177
11．许智宏，刘春明．植物发育的分子机理．北京，北京科学出版社。1998
12．何奕昆，代庆阳，苏学辉．雁来红叶色转变与超微结构及色素含量的关系．四川师范学院学报．1995，16(3)：195-198
13．余叔文，汤章城．植物生理与分子生物学(第二版)．北京，北京科学出版社．1998
14．吴少华，张大生．花青素生成相关基因dfr研究进展．福建林学院学报．2002，22(2)：189-19
15．宋光泉，古练权．荔枝果皮花色素苷的光分解研究进展．中山大学学报．1998，38(5)：50-55
16．宋光泉，古练权．荔枝果皮花色素苷的光分解研究．中山大学学报．1999，38(5)：58-60
17．束怀瑞．果树栽培生理学．北京，中国农业出版社．1993
18．汪政科，彭镇华．观赏植物基因工程研究进展．林业科学研究．2000，13(1)：97-102
19．沈同，王镜岩主编．生物化学(第二版)．北京，高等教育出版社．1990
20．苏焕然．花卉基因工程研究进展．北方园艺．1996，09：26-27
21．陈俊伟，张良诚，张上隆．果实中的糖分积累机理．植物生理学通讯．2000，36(6)：497-503
22．周音，张智奇，殷丽青等．观赏植物基因工程研究进展及存在的问题．上海农业学报．2000，16(1)：90-96
23．周爱琴，祝军，生吉萍等．苹果花青素形成与PAL活性及蛋白质含量的关系．中国农业大学学报．1997，2(3)：97-99
24．孟繁静．植物花发育的分子生物学．北京，中国农业出版社．2000
25．洪宁，童哲．光敏色素在植物个体发育中的作用．植物生理学通讯．1998，34(6)：417
-422
26．郧忠亮．花卉作物的花色基因工程．北方园艺．1994，96：37-38
27．金波．一品红花色的探讨．园艺学报．1994，21(1)：87-90
28．胡桂兵，高飞飞，陈大成等．套袋对荔枝果皮中的色素、酚类物质和内源激素的影响．华南农业大学学报．2001，22(1)：31-34
29．赵宗方，宋亭华，高红胜．巨峰葡萄色素发育若干规律．江苏农学院学报．1989，10(4)：17-21
30．徐绍颖．植物生长调节剂与果树生产．上海，上海科技出版社．1987
31．阎隆飞，张玉麟．分子生物学(第二版)．北京，中国农业大学出版社．1997
32．黄卫东，原永兵，彭宜本．温带果树结实生理．北京，中国农业大学出版社．1994
33．曾骧．果树生理学．北京，中国农业大学出版社．1992
34．程林梅，张一萍，孙毅。草莓生物技术研究进展．植物学通报．2002，19(1)：39-43
35．潘增光，范晖，束怀瑞．苹果果实花青素形成与乙烯释放的关系．植物生理学通讯．1995，31(5)：338-340
36．潘增光，王国宾，李奎明等．新红星苹果果实着色期几种色素含量变化及其相关性．植物生理学通讯．1996，32(5)：347-349
37. Anju Singh, M. Tamil Selvi and Rameshwar shrma. Sunlight-induced anthocyanin. Journal of Experimental Botany, 1999, (150) 339: 1619-1625
38. Asako lida, Toru K azuoka et al. A zinc finger protein RHL41 mediates the light acclimatization response in Arabidopsis. The Plant Joumal, 2000, 24(2): 191-203s
39. Axel alesander hirner, Hanns Uirich seitz. Isoforms of chalcone synthase in Daucus carota L. and their differential expression in organs from the european wild carrot and in ultraviolet-A-irradiated cell cultures. Planta. 2000 (2) 10: 933-998
40. Chory J, Chatterjee-M, Cook-R-K, Elich-T. From Seed germination to flowering, light controls plant development via the pigment. Biological Abstracts. 1997/01-1997/06
41. Christoph Poppe, Uta sweere, Helge Drumm-herrel and Eberhard schafer. The blue light receptor cryptchrome 1 can act independently of phytochrome A and B in Arabidopsis thaliana. The Plant Journal. 1998, 16(4): 465-471
42. David Jackson, Keith Roberts and Cathie Martin. Temporal and spatial control of expression of anthocyanib biosynthetic genes in developing flowers of antirrhinum majus. The Plant Journal 1992, 2(4): 425-434
43. Dongsun Cao, Yun Lin, and Chi-Lien Cheng. Genetic interactions between the Chlorate-resistanrt mutant cr88 and the photomorphogentic mutants copl and hy5. The Plant Cell. 2000, 12: 199-210
44. Enriqueta Moyano, Ignacioportero-robles et al. A fruit-specific putative
dihydroflavonol 4-Reductase Gene is differentially expressed in strawbrrry during the ripening process. Plant Physiol. 1998, 117: 711-716
45. Hou Yongmin, Von Albrecht G et al. A new class of Arabidopsis constomorphogenic genes involved in regulating cotyledon development. Biological Abstracts. 1993/01-1993/06
46. Jennie A. Jackson, Gareth I. Jenkins. Extension-growth responses and expression of flavonoid biosynthesis genes in the Arabidopsis hy4 mutant. Planta. 1995, 197: 233-239
47. Keiko U. Torii, Timony W. McNellis and Xing-Xing Deng. Functional dissection of its three structural modules in light control of Seeding development. The EMBO Journal. 1998, (17): 19 5577-5587
48. Kwok-Shing-F, Piekos-Barry et al. A complement of ten essential and pleiotropic Arabidopsis COP/DET/FUS genes is necessary for repression of photomorphagenesis in darkness. Biological Abstracts. 1996/01-1996/06
49. Kwok-Shing-F, Solano-Roberto, Tsuge-Tomohiko, Chamovitz-Daniel-A et al. Arabidopsis homologs of a c-Jun coactivator are present both in monomeric from znd in the COP9 complex, and their abundance is differentially affected by the pleiotropic cop/det/fus mutations. Biological Abstracts. 1999/01-1999/06
50. Lay-Hong Ang, sudip Chattopadhyay et al. Molecular interaction between COP1and HY5 Defines a Regulatory switch for light control of Arabidopsis development. Molecular Cell. 1998,1,213-222
51. L.H.J. Kerckhoffs, P.M. Kelmenson, M.E.L. schreuder, et al. Characterization of the gene encoding the apoprotein of phytochrome B2 in tomato, and identification of molecular lesions in two mutant alleles. Mol Gen Genet. 1999, 261: 901-907
52. Margaret Ahmad and Anthony R. Cashmore. The blue-light receptor cryptochrome 1 shows functional dependence on phytochrome A or phytochrome B in Arabidopsis thaliana. The Plant Journal. 1997, 11(3): 421-427
53. Margaret Ahmad, Jose A. Jarillok, and Anthony R, Cashmore. Chimerc proteins between cry1 and cry2 Arabidopsis blue light photoreceptors indicate overlapping functions and varying protein stability. The Plant Cell. 1998, 10: 197-207
54. Mary L..Durbin, Bonnie McCaig and Michael T. Clegg. Molecular evolution of the chalcone synthase multigene family in the morning glory genome. Plant Molecular Biology. 2000, 42: 79-92
55. Mayer-Raphael, Raventos_Dora, Chua-Nam-Hai. Det1, cop1, and cop9 mutations. Biological Abstracts. 1997/01-1997/06
56. Melanie Kaltenbach, Gudrun schroder, Elmon schmelzer, Verena Lutz and Joachim schroder. Flavonoid hydroxylase from Catharanthus roseus: cDNA, heterologous expression, enzyme properties and cell-type specific expression in plants. The Plant Journal. 1999, 19(2),: 183-193
57. Michal Oren-shamir, Ada Levi-Nissim. UV-light effect on the leaf pigmentation of cotinus coggygria 'Royal Purple' Scientia Horticulturae 1997, 71: 59-66
58. Mundt-Kirsten-E, porte-Joanne, Murray-Johanne-M, et al. The COP9/signalosome complex is conserved in fission yeast and has a role in s Phsase. Biological Abstracts. 2000/01-2000/06
59. Naoki Yamamoto, Xing-Wang Deng. Protein nucleocytoplasmic transport and its light regulation in plants. Genes to Cells. 1999, 4, 489-500
60. Partt LH. Cordonnier MM. Phytochrome and Photoregulation Plants. Academic Press. 1987, 83.
61. R. Chaturvedi, R. shyam and P.V. sane. steady state levels of D1protein and psbA thanscript during UV-B inactivation of photosystem Ⅱ in wheat. Biochemistry and Molecular Biology Internation. 1998, (44)5: 925-932
62. Rhonda L. Feinbaum, Gisela storz, and Frederick Mausubel. High intensity and blue light regulated expression of chimeric chalcone synthase genes in transgenic Arabidopsis thaliana plants. Mol Gen genet. 1991, 226: 449-456
63. Roldan-Marta, Gomes-mena-Concepcion, Ruiz-Garcia-Leonor, salinas-Julio, Martinez-zapater-Jose-M sucrose availability on the aerial part of the plant promotes morphogenesis and flowering of Arabidopsis in the dark. Biological Abstracts. 2000/01-2000/06
64. Ronald E. Koes, Francesca Quattrocchio and Joseph N.M. Mol. The flavonoid biosynthetic pathway in plants: function and evolution. Bioessays. 1994, 16. 2
65. Saneyuki Kawabata, Yuhua Li, et al. Role of Photoreceptor - and sugar-mediated Reactions in Light Dependent Anthocyanin Production in Lily and stock Flowers. J. Japan. Soc. Hort. Sci. 2002
66. Serino-giovanna, Tsuge-Tomohiko, Kwok-shing, Matsui-minami, Wei-Ning, Deng-Xing, Wang. Arabidopsis COP8 and fus4 mutations define the same gene that encode subunit 4 of the COP9 signalosome. Biological Abstracts. 2000/01-2000/06
67. Staub-jeffrey-M, Wei-Ning; Deng-Xing-Wang. Evidence for FUS6 as a component of the nuclear-localized COP9 complex in Arabidopsis. Biological Abstracts. 1997/01-1997/01
68. Szekeres-Miklos, Nemeth-Kinga et al. Brassinasteroids rescue the deficiency of CYP90 a cytochrome p450 controlling cell elongation and de-dtiolation in
Arabidopsis. Biological Abstracts. 1996/01-1996/06
69. Tim Kunkel, Gunther Neuhaus, Alfred Batschauer, Nam-Hai Chua and Eberhard schafer. Functional analysis of yeast-derived phytochrome A and B phycocyanobilin adducts. The Plant Journal. 1996, 10(4): 625-636
70. Timothy A. Holton, Edwina C. Cornish. Genetics and biochemistry of anthocyanin biosynthesis. The Plant Cell. 1995,7: 1071-1083
71. Toshihiro Toguri, Naoyuki Umemoto, Osamu Kobayashi and Takeshi Ohtani. Activation of anthocyanin synthesis genes by white light in eggplant hypocotyls tissues, and identification of an inducible P-450 cDNA. Plant Molecular Biology. 1993, 23: 933-946
72. Waiters-Robin-6, Gogers-jennifer-J-M, Shephard-Freya, Horton-Peter. Acclimation of Arabidopsis thaliana to the light environment: the role of photoreceptors. Biological Abstracts. 2000/01-2000/06
73. Yoshiharu Y. Yamamoto, minami Matsui, Lay-Hong Ang, and Xing-Wang. Role of a COP1 interactive protein in mediating light-regulated gene expression in Arabidopsis. The Plant Cell. 1998,10: 1083-1094
74. Zhou-D-X., Kim-Y-J, Li-Y-F, Carol-P, Mache_R. COP1b, an isoform of COP1 generated by alternative splicing, has a negative effect on COP1 function in regulating light-dependent seeding development in Arabidopsis. Biological Abstracts. 1998/01-1998/06