Petunia pp 269-299 | Cite as

The Genetics of Flower Color

  • Giambattista Tornielli
  • Ronald Koes
  • Francesca Quattrocchio


With nearly a century of excellent research on the biochemistry and inheritance of color, and the corresponding development of incredible genetic resources, Petunia has offered perhaps the best genetic system for molecular analysis of flower color. The knowledge and materials available to the Petunia geneticist, together with the tools of genetic engineering, have allowed for the isolation and characterization of a large number of genes affecting flower color, including those encoding biosynthetic enzymes, regulators of their expression, and vacuolar function. Here we summarize current knowledge about the genes responsible for the amazing diversity of colors and color patterns observable in the genus Petunia and propose some evolutionary implications of these findings.


Flower Color Anthocyanin Biosynthesis Anthocyanin Pathway Vacuolar Acidification Vacuolar Lumen 
These keywords were added by machine and not by the authors. This process is experimental and the keywords may be updated as the learning algorithm improves.


  1. Alfenito, M.R., Souer, E., Goodman, C.D., Buell, R., Mol, J., Koes, R. and Walbot, V. (1998) Functional complementation of anthocyanin sequestration in the vacuole by widely divergent glutathione S-transferases. Plant Cell 10, 1135–1149.CrossRefPubMedGoogle Scholar
  2. Ando, T., Takahashi, M., Nakajima, T., Toya, Y., Watanabe, H., Kokubun, H. and Tatsuzawa, F. (2004) Delphinidin accumulation is associated with abnormal flower development in Petunias. Phytochem. 65, 2219–2227.CrossRefGoogle Scholar
  3. Baumann, K., Perez-Rodriguez, M., Bradley, D., Venail, J., Bailey, P., Hailing, J., Koes, R., Roberts, K. and Martin, C. (2007) Control of cell and petal morphogenesis by R2R3 MYB transcription factors. Development 134, 1691–1701.CrossRefPubMedGoogle Scholar
  4. Beld, M., Martin, C., Huits, H., Stuitje, A.R. and Gerats, A.G.M. (1989) Flavonoid synthesis in Petunia hybrida: Partial characterization of dihydroflavonol 4-reductase genes. Plant Mol. Biol. 13, 491–502.CrossRefPubMedGoogle Scholar
  5. Britsch, L. and Grisebach, H. (1986) Purification and characterization of (2S)-flavanone 3-hydroxylase from Petunia hybrida. Eur. J. Biochem. 156, 569–577.CrossRefPubMedGoogle Scholar
  6. Britsch, L., Ruhnau-Brich, B. and Forkmann, G. (1992). Molecular cloning, sequence analysis and in vitro expression of flavanone 3ß-hydroxylase from Petunia hybrida. J. Biol. Chem. 267, 5380–5387.PubMedGoogle Scholar
  7. Brugliera, F., Holton, T.A., Stevenson, T.W., Farcy, E., Lu, C.Y. and Cornish, E.C. (1994) Isolation and characterization of a cDNA clone corresponding to the Rt locus of Petunia hybrida. Plant J. 5, 81–92.CrossRefPubMedGoogle Scholar
  8. Brugliera, F., Barri-Rewell, G., Holton, T.A. and Mason, J.G. (1999) Isolation and characterization of a flavonoid 3'-hydroxylase cDNA clone corresponding to the Ht1 locus of Petunia hybrida. Plant J. 19, 441–451.Google Scholar
  9. Brugliera, F., Tull, D., Holton, T.A., Karan, M., Treloar, N., Simpson, K., Skurczynska, J. and Mason, J.G. (2000) Introduction of a cytochrome b5 enhances the activity of flavonoid 3',5'-hydroxylase (a cytochrome P450) in transgenic carnation. Intl. Plant Molec. Biol. Rep. 18 (Supp.).Google Scholar
  10. Davies, K.M., Bloor, S.J., Spiller, G.B. and Deroles, S.C. (1998) Production of yellow colour in flowers: Redirection of flavonoid biosynthesis in Petunia. Plant J. 13, 259–266.CrossRefGoogle Scholar
  11. de Vetten, N., Quattrocchio, F., Mol, J. and Koes, R. (1997) The an11 locus controlling flower pigmentation in Petunia encodes a novel WD-repeat protein conserved in yeast, plants and animals. Genes Dev. 11, 1422–1434.CrossRefPubMedGoogle Scholar
  12. de Vetten, N., ter Horst, J., van Schaik, H.-P., den Boer, B., Mol, J. and Koes, R. (1999) A cytochome b5 is required for full activity of flavonoid 3'5'-hydroxylase, a cytochrome P450 involved in the formation of blue flower colors. Proc. Natl. Acad. Sci., USA 96, 778–783.Google Scholar
  13. de Vlaming, P., Cornu, A., Farcy, E., Gerats, A.G.M., Maizonnier, D., Wiering, H. and Wijsman, H.J.W. (1984) Petunia hybrida: A short description of the action of 91 genes, their origin and their map location. Plant Mol. Biol. Rep. 2, 21–42.CrossRefGoogle Scholar
  14. Dixon, R.A., Blyden, E.R., Robbins, M.P., van Tunen, A.J. and Mol, J.N.M. (1988) Comparative biochemistry of chalcone isomerases. Phytochem. 27, 2801–2808.CrossRefGoogle Scholar
  15. Dooner, H.K., Robbins, T. and Jorgensen, R. (1991) Genetic and developmental control of anthocyanin biosynthesis. Annu. Rev. Genet. 25, 173–199.CrossRefPubMedGoogle Scholar
  16. Forkmann, G., de Vlaming, P., Spribille, R., Wiering, H. and Schram, A.W. (1986) Genetic and biochemical studies on the conversion of dihydroflavonols to flavonols in flowers of Petunia hybrida. Z. Naturforsch. 41c, 179–186.Google Scholar
  17. Forkmann, G. and Martens, S. (2001) Metabolic engeneering and applications of flavonoids. Curr. Opin. Biotech. 12, 155–160.CrossRefPubMedGoogle Scholar
  18. Froemel, S., de Vlaming, P., Stotz, G., Wiering, H., Forkmann, G. and Schram, A.W. (1985) Genetic and biochemical studies on the conversion of flavanones to dihydroflavonols in flowers of Petunia hybrida Theor. Appl. Genet. 70, 561–568.CrossRefGoogle Scholar
  19. Gerats, A.G.M., De Vlaming, P., Doodeman, M. and Schram, A.W. (1982) Genetic control of the conversion of dihydroflavonols into flavonols and anthocyanins in flowers of Petunia hybrida. Planta 155, 364–368.CrossRefGoogle Scholar
  20. Gerats, A.G.M., Wallroth, M., Donker-Koopman, W., Groot, S.P.C. and Schram, A.W. (1983) The genetic control of the enzyme UDP glucose flavonoid 3-O-glucosyltransferase in flowers of Petunia hybrida. Theor. Appl. Genet. 65, 349–352.CrossRefGoogle Scholar
  21. Gerats, A.G.M., Farcy, E., Wallroth, M., Groot, S.P.C. and Schram, A. (1984) Control of anthocyanin biosynthesis in Petunia hybrida by multiple allelic series of the genes An1 and An2. Genetics 106, 501–508.PubMedGoogle Scholar
  22. Heller, W. and Forkmann, G. (1988) Biosynthesis. In: J.B. Harborne (Ed.), The Flavonoids. Chapman and Hall, London, pp. 399–425.Google Scholar
  23. Holton, T.A., Brugliera, F. and Tanaka, Y. (1993a) Cloning and expression of flavonol synthase from Petunia hybrida. Plant J. 4, 1003–1010.Google Scholar
  24. Holton, T.A., Brugliera, F., Lester, D.R., Tanaka, Y., Hyland, G.D., Menting, J.G.T., Lu, C.-Y., Farcy, E., Stevenson, T.W. and Cornish, E.C. (1993b) Cloning and expression of cytochrome P450 genes controlling flower colour. Nature 336, 276–279.Google Scholar
  25. Holton, T.A. and Cornish, E.C. (1995) Genetics and biochemistry of anthocyanin biosynthesis. Plant Cell 7, 1071–1083.CrossRefPubMedGoogle Scholar
  26. Hondo, T., Yoshida, K., Nakagawa, A., Kawai, T., Tamura, H. and Goto, T. (1992) Structural basis of blue-color development in flower petals from Commelina communis, Nature 358, 515–518.CrossRefGoogle Scholar
  27. Huits, H. (1993) Mutable Genes and Control of Flower Pigmentation in Petunia hybrida. Vrije Universiteit, Amsterdam, The Netherlands.Google Scholar
  28. Huits, H.S.M., Gerats, A.G.M., Kreike, M.M., Mol, J.N.M. and Koes, R.E. (1994) Genetic control of dihydroflavonol 4-reductase gene expression in Petunia hybrida. Plant J. 6, 295–310.CrossRefPubMedGoogle Scholar
  29. Jonsson, L.M.V., de Vlaming, P., Wiering, H., Aarsman, M.E.G. and Schram, A.W. (1983) Genetic control of anthocyanin-O-methyltransferase in flowers of Petunia hybrida, Theor. Appl. Genet. 66, 349.CrossRefGoogle Scholar
  30. Jonsson, L.M.V., Aarsman, M.E.G., Poulton, J.E. and Schram, A.W. (1984a) Properties and genetic control of four methyltransferases involved in methylation of anthocyanins in flowers of Petunia hybrida. Planta 160, 174.Google Scholar
  31. Jonsson, L.M.V., Aarsman, M.E.G., Van Diepen, J., Smit, N. and Schram, A.W. (1984b) Properties and genetic control of anthocyanidin 5-0-glucosyltransferase in flowers of Petunia hybrida. Planta 160, 341–347.Google Scholar
  32. Kitamura, S., Shikazono, N. and Tanaka, A. (2004) TRANSPARENT TESTA 19 is involved in the accumulation of both anthocyanins and proanthocyanidins in Arabidopsis. Plant J. 37, 104–114.CrossRefPubMedGoogle Scholar
  33. Kitamura, S. (2006) Transport of Flavonoids. In: E. Grotewold (Ed.), The Science of Flavonoids. Springer, NY, pp. 123–146.CrossRefGoogle Scholar
  34. Koes, R.E., Spelt, C.E., Mol, J.N.M. and Gerats, A.G.M. (1987) The chalcone synthase multigene family of Petunia hybrida (V30): Sequence homology, chromosomal localization and evolutionary aspects. Plant Mol. Biol. 10, 375–385.CrossRefGoogle Scholar
  35. Koes, R.E. (1988) Genes involved in flavonoid biosynthesis in Petunia hybrida: The chalcone synthase multigene family. Vrije Universiteit, Amsterdam, The Netherlands.Google Scholar
  36. Koes, R.E., Spelt, C.E. and Mol, J.N.M. (1989a) The chalcone synthase multigene family of Petunia hybrida (V30): Differential, light-regulated expression during flower development and UV light induction. Plant Mol. Biol. 12, 213–225.Google Scholar
  37. Koes, R.E., Spelt, C.E., Van den Elzen, P.J.M. and Mol, J.N.M. (1989b) Cloning and molecular characterization of the chalcone synthase multigene family of Petunia hybrida. Gene 81, 245–257.Google Scholar
  38. Koes, R.E., Van Blokland, R., Quattrocchio, F., Van Tunen, A.J. and Mol, J.N.M. (1990) Chalcone synthase promoters in Petunia are active in pigmented and unpigmented cell types. Plant Cell 2, 379–392.CrossRefPubMedGoogle Scholar
  39. Koes, R.E., Quattrocchio, F. and Mol, J.N.M. (1994) The flavonoid biosynthetic pathway in plants: Function and evolution. BioEssays 16, 123–132.CrossRefGoogle Scholar
  40. Koes, R., Verweij, C.W. and Quattrocchio, F. (2005) Flavonoids: A colorful model for the regulation and evolution of biochemical pathways. Trends Plant Sci. 5, 236–242.CrossRefGoogle Scholar
  41. Koseki, M., Goto, K., Masuta, C. and Kanazawa, A. (2005) The star-type color pattern in Petunia hybrida "Red Star” flowers is induced by sequence-specific degradation of chalcone synthase RNA. Plant Cell Physiol. 6, 1879–1883.CrossRefGoogle Scholar
  42. Kroon, J., Souer, E., de Graaff, A., Xue, Y., Mol, J. and Koes, R. (1994) Cloning and structural analysis of the anthocyanin pigmentation locus Rt of Petunia hybrida: Characterization of insertion sequences in two mutant alleles. Plant J. 5, 69–80.CrossRefPubMedGoogle Scholar
  43. Kroon, A.R. (2004) Transcription Regulation of the Anthocyanin Pathway in Petunia hybrida. Vrije Universiteit, Amsterdam, The Netherlands.Google Scholar
  44. Maizonnier, D. and Moessner, A. (1979) Localization of the linkage groups on the seven chromosomes of the Petunia hybridagenome. Genetica 51(2), 143–148.CrossRefGoogle Scholar
  45. Matsubara, K., Kodama, H., Kokubun, H., Watanabe, H. and Ando, T. (2005) Two novel transposable elements in a cytochrome P450 gene govern anthocyanin biosynthesis of commercial petunias. Gene 358, 121–126.CrossRefPubMedGoogle Scholar
  46. Meyer, P., Heidmann, I., Forkmann, G. and Saedler, H. (1987) A new petunia flower colour generated by transformation of a mutant with a maize gene. Nature 330, 677–678.CrossRefPubMedGoogle Scholar
  47. Mo, Y., Nagel, C. and Taylor, L.P. (1992) Biochemical complementation of chalcone synthase mutants defines a role for flavonols in functional pollen. Proc. Natl. Acad. Sci., USA 89, 7213–7217.Google Scholar
  48. Mol, J.N.M., Schram, A.W., de Vlaming, P., Gerats, A.G.M., Kreuzaler, F., Hahlbrock, K., Reif, H.J. and Veltkamp, E. (1983) Regulation of flavonoid gene expression in Petunia hybrida: Description and partial characterization of a conditional mutant in chalcone synthase gene expression. Molec. Gen. Genet.192, 424–429.CrossRefGoogle Scholar
  49. Mol, J., Cornish, E., Mason, J. and Koes, R. (1999) Novel coloured flowers. Curr. Opin. Biotech. 10, 198–201.CrossRefPubMedGoogle Scholar
  50. Mueller, L.A., Goodman, C.D., Silady, R.A. and Walbot, V. (2000) AN9, a Petunia glutathione S-transferase required for anthocyanin sequestration, is a flavonoid-binding protein. Plant Physiol. 123, 1561–1570.CrossRefPubMedGoogle Scholar
  51. Mur, L. (1995) Characterization of Members of the myb Gene Family of Transcription Factors from Petunia hybrida. Vrije Universiteit, Amsterdam, The Netherlands.Google Scholar
  52. Nakajima, J., Tanaka, Y., Yamazaki, M. and Saito, K. (2001) Reaction mechanism from leucoanthocyanidin to anthocyanidin 3-glucoside, a key reaction for coloring in anthocyanin biosynthesis. J. Biol. Chem. 276, 25797–25803.CrossRefPubMedGoogle Scholar
  53. Nakajima, T., Matsubara, K., Kodama, H., Kokubun, H., Watanabe, H. and Ando, T. (2005) Insertion and excision of a transposable element governs the red floral phenotype in commercial Petunia. Theor. Appl. Genet. 110, 1038–1043.CrossRefPubMedGoogle Scholar
  54. Napoli, C., Lemieux, C. and Jorgensen, R. (1990) Introduction of a chimeric chalcone synthase gene into petunia results in reversible co-suppression of homologous genes in trans. Plant Cell 2, 279–289.CrossRefPubMedGoogle Scholar
  55. Napoli, C.A., Fahy, D., Wang, H.Y. and Taylor, L.P. (1999) white anther: A petunia mutant that abolishes pollen flavonol accumulation, induces male sterility and is complemented by a chalcone synthase transgene. Plant Physiol. 120, 615–622.CrossRefPubMedGoogle Scholar
  56. Noda, K., Glover, B.J., Linstead, P. and Martin, C. (1994) Flower colour intensity depends on specialized cell shape controlled by a Myb-related transcription factor. Nature 369, 661–664.CrossRefPubMedGoogle Scholar
  57. Oud, J.S.N., Schneiders, H., Kool, A.J. and van Grinsven, M.Q.J.M. (1995) Breeding of transgenic orange Petunia hybrida varieties. Euphyt. 85, 403–409.CrossRefGoogle Scholar
  58. Pollak, P.E., Vogt, T., Mo, Y. and Taylor, L.P. (1993) Chalcone synthase and flavonol accumulation in stigmas and anthers of Petunia hybrida. Plant Physiol. 102, 925–932.PubMedGoogle Scholar
  59. Quattrocchio, F., Wing, J.F., Leppen, H.T.C., Mol, J.N.M. and Koes, R.E. (1993) Regulatory genes controlling anthocyanin pigmentation are functionally conserved among plant species and have distinct sets of target genes. Plant Cell 5, 1497–1512.CrossRefPubMedGoogle Scholar
  60. Quattrocchio, F., Wing, J.F., van der Woude, K., Mol, J.N.M. and Koes, R. (1998) Analysis of bHLH and MYB-domain proteins: Species-specific regulatory differences are caused by divergent evolution of target anthocyanin genes. Plant J. 13, 475–488.CrossRefPubMedGoogle Scholar
  61. Quattrocchio, F., Wing, J., van der Woude, K., Souer, E., de Vetten, N., Mol, J. and Koes, R. (1999) Molecular analysis of the anthocyanin2 gene of Petunia and its role in the evolution of flower color. Plant Cell 11, 1433–1444.CrossRefPubMedGoogle Scholar
  62. Quattrocchio, F., Verweij, W., Kroon, A., Spelt, C., Mol, J. and Koes, R. (2006) PH4 of Petunia is an R2R3-MYB protein that activates vacuolar acidification through interactions with Basic-Helix-Loop-Helix transcription factors of the anthocyanin pathway. Plant Cell 18, 1274–1291.CrossRefPubMedGoogle Scholar
  63. Que, Q., Wang, H. and Jorgensen, R.A. (1998) Distinct patterns of pigment suppression are produced by allelic sense and antisense chalcone synthase transgenes in Petunia flowers. Plant J. 13, 401–409.CrossRefGoogle Scholar
  64. Reif, J.H., Wiesbach, U., Deumling, B. and Saedler, H. (1985) Cloning and analysis of two genes for chalcone synthase from Petunia hybrida. Mol Gen. Genet. 199, 208–215.Google Scholar
  65. Shimada, Y., M., O., Nakano-Shimada, R., Okinaka, Y., Kiyokawa, S. and Kikuchi, Y. (2001) Genetic engineering of the anthocyanin biosynthetic pathway with flavonoid-3',5'-hydroxylase: specific switching of the pathway in Petunia. Plant Cell Rep. 20, 456–462.CrossRefGoogle Scholar
  66. Snowden, K.C. and Napoli, C.A. (1998) PsI: a novel Spm-like transposable element from Petunia hybrida. Plant J. 14, 43–54.CrossRefPubMedGoogle Scholar
  67. Sompornpailin, K., Makita, Y., Yamazaki, M. and Saito, K. (2002) A WD-repeat-containing putative regulatory protein in anthocyanin biosynthesis in Perilla frutescens. Plant Mol. Biol. 50, 485–495.CrossRefPubMedGoogle Scholar
  68. Spelt, C., Quattrocchio, F., Mol, J. and Koes, R. (2000) anthocyanin1 of Petunia encodes a basic-Helix Loop Helix protein that directly activates structural anthocyanin genes. Plant Cell 12, 1619–1631.CrossRefPubMedGoogle Scholar
  69. Spelt, C., Quattrocchio, F., Mol, J. and Koes, R. (2002) ANTHOCYANIN1 of Petunia controls pigment synthesis, vacuolar pH, and seed coat development by genetically distinct mechanisms. Plant Cell 14, 2121–2135.CrossRefPubMedGoogle Scholar
  70. Stafford, A.H. (1974) Possible multiple enzyme complexes regulating the formation of C6–C3 phenolic compounds and lignuins in higher plants. Rec.Adv.Phytochem. 8, 53–79.Google Scholar
  71. Stafford, H. (1990) Flavonoid Metabolism. CRC Press, Boca Raton, Florida.Google Scholar
  72. Stam, M.E. (1997). Post-transcriptional Silencing of Flower Pigmentation Genes in Petunia hybrida by (Trans)gene Repeats. Vrije Universiteit, Amsterdam.Google Scholar
  73. Stam, M., Mol, J.N.M. and Kooter, J.M. (1997) The silence of genes in transgenic plants. Ann. Bot. 79, 3–12.CrossRefGoogle Scholar
  74. Tanaka, Y., Katsumoto, Y., Brugliera, F. and Mason, J. (2005) Genetic engineering in floriculture. Plant Cell, Tiss. Organ. Cult. 80, 1–24.CrossRefGoogle Scholar
  75. Taylor, L.P. and Jorgensen, R. (1992) Conditional male fertility in chalcone synthase-deficient Petunia. J. Hered. 83, 11–17.Google Scholar
  76. Tsuda, S., Fukui, Y., Nakamura, N., Katsumoto, Y., Yonekura-Sakakibara, K., Fukuchi-Mizutani, M., Ohira, K., Ueyama, Y., Ohkawa, H., Holton, T.H., Kusumi, T. and Tanaka, Y. (2004) Flower color modification of Petunia hybrida commercial varieties by metabolic engineering. Plant Biotech. 21, 377–386.Google Scholar
  77. van der Krol, A.R., Lenting, P.E., Veenstra, J., Van der Meer, I.M., Koes, R.E., Gerats, A.G.M., Mol, J.N.M. and Stuitje, A.R. (1988) An anti-sense chalcone synthase gene in transgenic plants inhibits flower pigmentation. Nature 333, 866–869.CrossRefGoogle Scholar
  78. van der Krol, A.R., Mur, L.A., Beld, M., Mol, J.N.M. and Stuitje, A.R. (1990) Flavonoid genes in Petunia: Addition of a limited number of gene copies may lead to a suppression of gene expression. Plant Cell 2, 291–299.CrossRefPubMedGoogle Scholar
  79. van Houwelingen, A., Souer, E., Spelt, C., Kloos, D., Mol, J. and Koes, R. (1998) Analysis of flower pigmentation mutants generated by random transposon mutagenesis in Petunia hybrida. Plant J. 13, 39–50.PubMedGoogle Scholar
  80. van Tunen, A.J., Koes, R.E., Spelt, C.E., van der Krol, A.R., Stuitje, A.R. and Mol, J.N.M. (1988) Cloning of the two chalcone flavanone isomerase genes from Petunia hybrida: Coordinate, light-regulated and differential expression of flavonoid genes. EMBO J. 7, 1257–1263.PubMedGoogle Scholar
  81. van Tunen, A.J., Hartman, S.A., Mur, L.A. and Mol, J.N.M. (1989) Regulation of chalcone flavanone isomerase (CHI) gene expression in Petunia hybrida: The use of alternative promoters in corolla, anthers and pollen. Plant Mol. Biol. 12, 539–551.CrossRefGoogle Scholar
  82. van Tunen, A.J., Mur, L., Brouns, G.S., Rienstra, J.-D., Koes, R.E. and Mol, J.N.M. (1990) Pollen- and anther-specific chi promoters from Petunia: Tandem promoter regulation of the chiA gene. Plant Cell 2, 393–401.CrossRefPubMedGoogle Scholar
  83. Verweij, C.W. (2007) Vacuolar Acidification: Mechanism, Regulation and Function in Petunia Flowers. Vrije Universiteit, Amsterdam, The Netherlands.Google Scholar
  84. Verweij, W., Spelt, C., Di Sansebastiano, G.P., Vermeer, J., Reale, L., Ferranti, F., Koes, R.E. and Quattrocchio, F. (In press) A novel type of tonoplast localized H+-ATPase is required for vacuolar acidification and coloration of flowers and seeds. Nature Cell Biology.Google Scholar
  85. Weiss, D., Van der Luit, A.H., Kroon, J.T.M., Mol, J.N.M. and Kooter, J.M. (1993) The Petunia homologue of the Antirhinum majus candi gene and Zea mays A2 flavonoid genes: Homology to flavanone 3- hydroxylase and ethylene forming enzyme. Plant Mol. Biol. 22, 893–897.CrossRefPubMedGoogle Scholar
  86. Winkel, B.S.J. (2004) Metabolic channeling in plants. Annu. Rev. Plant Biol. 55, 85–107.CrossRefPubMedGoogle Scholar
  87. Yamazaky, M., Yamagishi, E., Gong, Z., Fukuchi-Mizutani, M., Fukui, Y., Tanaka, Y., Kusumi, T., Yamaguchi, M. and Saito, K. (2002) Two flavonoid glucosyltransferases from Petunia hybrida: Molecular cloning, biochemical properties and developmentally regulated expression. Plant Molec. Biol. 48, 401–411.CrossRefGoogle Scholar
  88. Ylstra, B., Busscher, J., Franken, J., Hollman, P.C.H., Mol, J.N.M. and van Tunen, A.J.J. (1994) Flavonols and fertilization in Petunia hybrida: Localization and mode of action during pollen tube growth. Plant J. 6, 201–212.CrossRefGoogle Scholar

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© Springer Science+Business Media, LLC 2009

Authors and Affiliations

  • Giambattista Tornielli
    • 1
  • Ronald Koes
  • Francesca Quattrocchio
  1. 1.Department of Developmental BiologyVrije Universiteit AmsterdamNederland

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