An integrated genomics approach to define niche establishment by rhodococcus fascians

Plant Physiology

Volumn 149, Issue 3, 2009, Pages 1366-1386

  Depuydt, Stephen ^a,b   Trenkamp, Sandra ^c   Fernie, Alisdair R ^c   Elftieh, Samira ^d   Renou, Jean Pierre ^d   Vuylsteke, Marnik ^a,b   Holsters, Marcelle ^a,b   Vereecke, Danny ^a,b  

^a DEPARTMENT OF PLANT SYSTEMS BIOLOGY   (Belgium)

^b GHENT UNIVERSITY   (Belgium)

^c UNIVERSITY OF POTSDAM   (Germany)

^d CEMAGREF   (France)

Author keywords

[No Author keywords available]

Indexed keywords

ACTINOBACTERIA; ARABIDOPSIS; ARABIDOPSIS THALIANA; BACTERIA (MICROORGANISMS); POSIBACTERIA; RHODOCOCCUS; RHODOCOCCUS FASCIANS;

EID: 63549142921     PISSN: 00320889     EISSN: 15322548     Source Type: Journal    
DOI: 10.1104/pp.108.131805     Document Type: Article

References (111)

1. Atkin OK, Millar AH, Gardeström P, Day DA (2000) Photosynthesis, carbohydrate metabolism and respiration in leaves of higher plants. In RC Leegood, TD Sharkey, S von Caemmerer, eds, Photosynthesis: Physiology and Metabolism, Advances in Phytosynthesis and Respiration, Vol 9. Kluwer Academic Publishers, Dordrecht, The Netherlands, pp 153-175
2. Berger S, Papadopoulos M, Schreiber U, Kaiser W, Roitsch T (2004) Complex regulation of gene expression, photosynthesis and sugar levels by pathogen infection in tomato. Physiol Plant 122: 419-428
3. Berger S, Sinha AK, Roitsch T (2007) Plant physiology meets phytopathology: plant primary metabolism and plant-pathogen interactions. J Exp Bot 58: 4019-4026
4. Biemelt S, Sonnewald U (2006) Plant-microbe interactions to probe regulation of plant carbon metabolism. J Plant Physiol 163: 307-318
5. Boboye B (2004) Degradation of trehalose by rhizobia and characteristics of a trehalose-degrading enzyme isolated from Rhizobium species NGR234. J Appl Microbiol 97: 256-261
6. Boyes DC, Zayed AM, Ascenzi R, McCaskill AJ, Hoffman NE, Davis KR, Görlach J (2001) Growth stage-based phenotypic analysis of Arabidopsis: a model for high throughput functional genomics in plants. Plant Cell 13: 1499-1510
7. Brazma A, Hingamp P, Quackenbusch J, Sherlock G, Spellman P, Stoeckert C, Aach J, Ansorge W, Ball CA, Causton HC, et al (2001) Minimum information about a microarray experiment (MIAME): toward standards for microarray data. Nat Genet 29: 365-371
8. Brenner WG, Romanov GA, Köllmer I, Bürkle L, Schmülling T (2005) Immediate-early and delayed cytokinin response genes of Arabidopsis thaliana identified by genome-wide expression profiling reveal novel cytokinin-sensitive processes and suggest cytokinin action through transcriptional cascades. Plant J 44: 314-333
9. Brodmann D, Schuller A, Ludwig-Müller J, Aeschbacher RA, Wiemken A, Boller T,Wingler A (2002) Induction of trehalase in Arabidopsis plants infected with the trehalose-producing pathogen Plasmodiophora brassicae. Mol Plant Microbe Interact 15: 693-700
10. Chou H-M, Bundock N, Rolfe SA, Scholes JD (2000) Infection of Arabidopsis thaliana leaves with Albugo candida (white blister rust) causes a reprogramming of host metabolism. Mol Plant Pathol 1: 99-113
11. Cona A, Rea G, Angelini R, Federico R, Tavladoraki P (2006) Functions of amine oxidases in plant development and defence. Trends Plant Sci 11: 80-88
12. Cornelis K, Maes T, Jaziri M, Holsters M, Goethals K (2002) Virulence genes of the phytopathogen Rhodococcus fascians show specific spatial and temporal expression patterns during plant infection. Mol Plant Microbe Interact 15: 398-403
13. Cornelis K, Ritsema T, Nijsse J, Holsters M, Goethals K, Jaziri M (2001) The plant pathogen Rhodococcus fascians colonizes the exterior and interior of the aerial parts of plants.Mol PlantMicrobe Interact 14: 599-608
14. Crespi M, Messens E, Caplan AB, Van Montagu M, Desomer J (1992) Fasciation induction by the phytopathogen Rhodococcus fascians depends upon a linear plasmid encoding a cytokinin synthase gene. EMBO J 11: 795-804
15. Crespi M, Vereecke D, TemmermanW, VanMontagu M, Desomer J (1994) The fas operon of Rhodococcus fascians encodes new genes required for efficient fasciation of host plants. J Bacteriol 176: 2492-2501
16. CroweML, Serizet C, Thareau V, Aubourg S, Rouze' P, Hilson P, Beynon J, Weisbeek P, Van Hummelen P, Reymond P, et al (2003) CATMA: a complete Arabidopsis GST database. Nucleic Acids Res 31: 156-158
17. D'Agostino IB, Deruére J, Kieber JJ (2000) Characterization of the response of the Arabidopsis response regulator gene family to cytokinin. Plant Physiol 124: 1706-1717
18. Delessert C, Wilson IW, Van Der Straeten D, Dennis ES, Dolferus R (2004) Spatial and temporal analysis of the local response to wounding in Arabidopsis leaves. Plant Mol Biol 55: 165-181
19. Dempsey DA, Shah J, Klessig DF (1999) Salicylic acid and disease resistance in plants. Crit Rev Plant Sci 18: 547-575
20. de O Manes C-L, Beeckman T, Ritsema T, Van Montagu M, Goethals K, Holsters M (2004) Phenotypic alterations in Arabidopsis thaliana plants caused by Rhodococcus fascians infection. J Plant Res 117: 139-145
21. de O Manes C-L, Van Montagu M, Prinsen E, Goethals K, Holsters M (2001) De novo cortical cell division triggered by the phytopathogen Rhodococcus fascians in tobacco. Mol Plant Microbe Interact 14: 189-195
22. Depuydt S, De Veylder L, Holsters M, Vereecke D (2009) Eternal youth, the fate of developing Arabidopsis leaves upon Rhodococcus fascians infection. Plant Physiol 149: 1387-1398
23. Depuydt S, Doležal K, Van Lijsebettens M, Moritz T, Holsters M, Vereecke D (2008) Modulation of the hormone setting by Rhodococcus fascians results in ectopic KNOX activation in Arabidopsis. Plant Physiol 146: 1267-1281
24. Desomer J, Dhaese P, Van Montagu M (1988) Conjugative transfer of cadmium resistance plasmids in Rhodococcus fascians strains. J Bacteriol 170: 2401-2405
25. Doehlemann G, Wahl R, Vranes M, de Vries RP, Kämper J, Kahmann R (2008) Establishment of compatibility in the Ustilago maydis/maize pathosystem. J Plant Physiol 165: 29-40
26. Downes BP, Crowell DN (1998) Cytokinin regulates the expression of a soybean b-expansin gene by a post-transcriptional mechanism. Plant Mol Biol 37: 437-444
27. Ehness R, Ecker M, Godt DE, Roitsch T (1997) Glucose and stress independently regulate source and sink metabolism and defense mechanisms via signal transduction pathways involving protein phosphorylation. Plant Cell 9: 1825-1841
28. Essmann J, Schmitz-Thom I, Schön H, Sonnewald S, Weis E, Scharte J (2008) RNA interference-mediated repression of cell wall invertase impairs defense in source leaves of tobacco. Plant Physiol 147: 1288-1299
29. Fernie AR, Carrari F, Sweetlove LJ (2004) Respiratory metabolism: glycolysis, the TCA cycle and mitochondrial electron transport. Curr Opin Plant Biol 7: 254-261
30. Gadjev I, Vanderauwera S, Gechev TS, Laloi C, Minkov IN, Shulaev V, Apel K, Inze' D, Mittler R, Van Breusegem F (2006) Transcriptomic footprints disclose specificity of reactive oxygen species signaling in Arabidopsis. Plant Physiol 141: 436-445
31. Galston AW, Sawhney RK (1990) Polyamines in plant physiology. Plant Physiol 94: 406-410
32. Garcia-Brugger A, Lamotte O, Vandelle E, Bourque S, Lecourieux D, Poinssot B,Wendehenne D, PuginA (2006) Early signaling events induced by elicitors of plant defenses. Mol Plant Microbe Interact 19: 711-724
33. Gartemann K-H, Abt B, Bekel T, Burger A, Engemann J, Flügel M, Gaigalat L, Goesmann A, Gräfen I, Kalinowski J, et al (2008) The genome sequence of the tomato-pathogenic actinomycete Clavibacter michiganensis subsp. michiganensis NCPPB382 reveals a large island involved in pathogenicity. J Bacteriol 190: 2138-2149
34. Gechev TS, Van Breusegem F, Stone JM, Denev I, Laloi C (2006) Reactive oxygen species as signals that modulate plant stress responses and programmed cell death. Bioessays 28: 1091-1101
35. Goethals K, Vereecke D, Jaziri M, Van Montagu M, Holsters M (2001) Leafy gall formation by Rhodococcus fascians. Annu Rev Phytopathol 39: 27-52
36. Hammond-Kosack KE, Jones JDG (1996) Resistance gene-dependent plant defense responses. Plant Cell 8: 1773-1791
37. Herbers K, Meuwly P, Frommer WB, Me'traux J-P, Sonnewald U (1996) Systemic acquired resistance mediated by the ectopic expression of invertase: possible hexose sensing in the secretory pathway. Plant Cell 8: 793-803
38. Herbers K, Sonnewald U (1998) Molecular determinants of sink strength. Curr Opin Plant Biol 1: 207-216
39. Herbers K, Takahata Y,Melzer M, Mock H-P, Hajirezaei M, Sonnewald U (2000) Regulation of carbohydrate partitioning during the interaction of potato virus Y with tobacco. Mol Plant Pathol 1: 51-59
40. Hilson P, Allemeersch J, Altmann T, Aubourg S, Avon A, Beynon J, Bhalerao RP, Bitton F, Caboche M, Cannoot B, et al (2004) Versatile gene-specific sequence tags for Arabidopsis functional genomics: transcript profiling and reverse genetics applications. Genome Res 14: 2176-2189
41. Jameson P (2000) Cytokinins and auxins in plant-pathogen interactions- an overview. Plant Growth Regul 32: 369-380
42. Johnson R, Guderian RH, Eden F, Chilton M-D, Gordon MP, Nester EW (1974) Detection and quantitation of octopine in normal plant tissue and in crown gall tumors. Proc Natl Acad Sci USA 71: 536-539
43. Jones JDG, Dangl JL (2006) The plant immune system. Nature 444: 323-329
44. Jubault M, Hamon C, Gravot A, Lariagon C, Delourme R, Bouchereau A, Manzanares-Dauleux MJ (2008) Differential regulation of root arginine catabolism and polyamine metabolism in clubroot-susceptible and partially resistant Arabidopsis genotypes. Plant Physiol 146: 2008-2019
45. Jwa N-S, Agrawal GK, Tamogami S, Yonekura M, Han O, Iwahashi H, Rakwal R (2006) Role of defense/stress-related marker genes, proteins and secondary metabolites in defining rice self-defense mechanisms. Plant Physiol Biochem 44: 261-273
46. Kocal N, Sonnewald U, Sonnewald S (2008) Cell wall-bound invertase limits sucrose export and is involved in symptom development and inhibition of photosynthesis during compatible interaction between tomato and Xanthomonas campestris pv vesicatoria. Plant Physiol 148: 1523-1536
47. Korves TM, Bergelson J (2003) A developmental response to pathogen infection in Arabidopsis. Plant Physiol 133: 339-347
48. Kusano T, Berberich T, Tateda C, Takahashi Y (2008) Polyamines: essential factors for growth and survival. Planta 228: 367-381
49. Laloi C, Mestres-Ortega D, Marco Y, Meyer Y, Reichheld J-P (2004) The Arabidopsis cytosolic thioredoxin h5 gene induction by oxidative stress and itsW-box-mediated response to pathogen elicitor. Plant Physiol 134: 1006-1016
50. Lamb C, Dixon RA (1997) The oxidative burst in plant disease resistance. Annu Rev Plant Physiol Plant Mol Biol 48: 251-275
51. Laval V, Koroleva OA, Murphy E, Lu C, Milner JJ, Hooks MA, Tomos AD (2002) Distribution of actin gene isoforms in the Arabidopsis leaf measured in microsamples from intact individual cells. Planta 215: 287-292
52. Lisec J, Schauer N, Kopka J, Willmitzer L, Fernie AR (2006) Gas chromatography mass spectrometry-based metabolite profiling in plants. Nat Protoc 1: 387-396
53. Loria R, Kers J, Joshi M (2006) Evolution of plant pathogenicity in Streptomyces. Annu Rev Phytopathol 44: 467-487
54. Luo Y, LiW-M, Wang W (2008) Trehalose: protector of antioxidant enzymes or reactive oxygen species scavenger under heat stress? Environ Exp Bot 63: 378-384
55. Lurin C, Andre's C, Aubourg S, Bellaoui M, Bitton F, Bruyére C, Caboche M, Debast C, Gualberto J, Hoffmann B, et al (2004) Genome-wide analysis of Arabidopsis pentatricopeptide repeat proteins reveals their essential role in organelle biogenesis. Plant Cell 16: 2089-2103
56. Maere S, Heymans K, Kuiper M (2005) BiNGO: a Cytoscape plugin to assess enrichment of Gene Ontology categories in Biological Networks. Bioinformatics 21: 3448-3449
57. Maes T, Vereecke D, Ritsema T, Cornelis K, Ngo Thi Thu H, Van Montagu M, Holsters M, Goethals K (2001) The att locus of Rhodococcus fascians strain D188 is essential for full virulence on tobacco through the production of an autoregulatory compound. Mol Microbiol 42: 13-28
58. Maxwell K, Johnson GN (2000) Chlorophyll fluorescence-a practical guide. J Exp Bot 51: 659-668
59. Mittler R, Vanderauwera S, Gollery M, Van Breusegem F (2004) The reactive oxygen gene network in plants. Trends Plant Sci 9: 490-498
60. Miyawaki K, Matsumoto-Kitano M, Kakimoto T (2004) Expression of cytokinin biosynthetic isopentenyltransferase genes in Arabidopsis: tissue specificity and regulation by auxin, cytokinin, and nitrate. Plant J 37: 128-138
61. Mogensen JE,Wimmer R, Larsen JN, Spangfort MD, Otzen DE (2002) The major birch allergen, Bet v 1, shows affinity for a broad spectrum of physiological ligands. J Biol Chem 277: 23684-23692
62. Müller J, Aeschbacher RA, Wingler A, Boller T, Wiemken A (2001) Trehalose and trehalase in Arabidopsis. Plant Physiol 125: 1086-1093
63. Müller J, Boller T, Wiemken A (1995) Trehalose and trehalase in plants: recent developments. Plant Sci 112: 1-9
64. Murgia I, Briat JF, Tarantino D, Soave C (2001) Plant ferritin accumulates in response to photoinhibition but its ectopic overexpression does not protect against photoinhibition. Plant Physiol Biochem 39: 797-805
65. Navarro L, Dunoyer P, Jay F, Arnold B, Dharmasiri N, Estelle M, Voinnet O, Jones JDG (2006) A plant miRNA contributes to antibacterial resistance by repressing auxin signaling. Science 312: 436-439
66. Navarro L, Zipfel C, Rowland O, Keller I, Robatzek S, Boller T, Jones JDG (2004) The transcriptional innate immune response to flg22: interplay and overlap with Avr gene-dependent defense responses and bacterial pathogenesis. Plant Physiol 135: 1113-1128
67. Nielsen TH, Krapp A, Röper-Schwarz U, Stitt M (1998) The sugarmediated regulation of genes encoding the small subunit of Rubisco and the regulatory subunit of ADP glucose pyrophosphorylase is modified by phosphate and nitrogen. Plant Cell Environ 21: 443-454
68. Noguchi K, Yoshida K (2008) Interaction between photosynthesis and respiration in illuminated leaves. Mitochondrion 8: 87-99
69. Nunes-Nesi A, Sweetlove LJ, Fernie AR (2007) Operation and function of the tricarboxylic acid cycle in the illuminated leaf. Physiol Plant 129: 45-56
70. Oxborough K, Baker NR (1997) An instrument capable of imaging chlorophyll a fluorescence from intact leaves at very low irradiance and at cellular and subcellular levels of organization. Plant Cell Environ 20: 1473-1483
71. Payne RW, Arnold GM (2002) GenStat Release 6.1 ReferenceManual Part 3: Procedure Library PL14. VSN International, Hemel Hempstead, UK
72. Penna S (2003) Building stress tolerance through over-producing trehalose in transgenic plants. Trends Plant Sci 8: 355-357
73. Pertry I, Va' clavI'kova' K, Depuydt S, Galuszka P, SpI'chal L, Temmerman W, Stes E, Schmülling T, Kakimoto T, Van Montagu M, et al (2009) Identification of Rhodococcus fascians cytokinins and their modus operandi to reshape the plant. Proc Natl Acad Sci USA 106: 929-934
74. Putnam ML, Miller ML (2007) Rhodococcus fascians in herbaceous perennials. Plant Dis 91: 1064-1076
75. Quilliam RS, Swarbrick PJ, Scholes JD, Rolfe SA (2006) Imaging photosynthesis in wounded leaves of Arabidopsis thaliana. J Exp Bot 57: 55-69
76. Raghavendra AS, Padmasree K (2003) Beneficial interactions of mitochondrial metabolism with photosynthetic carbon assimilation. Trends Plant Sci 8: 546-553
77. Rashotte AM, Carson SDB, To JPC, Kieber JJ (2003) Expression profiling of cytokinin action in Arabidopsis. Plant Physiol 132: 1998-2011
78. Riou-Khamlichi C, Menges M, Healy JMS, Murray JAH (2000) Sugar control of the plant cell cycle: differential regulation of Arabidopsis D-type cyclin gene expression. Mol Cell Biol 20: 4513-4521
79. Robert-Seilaniantz A, Navarro L, Bari R, Jones JDG (2007) Pathological hormone imbalances. Curr Opin Plant Biol 10: 372-379
80. Robinette D, Matthysse AG (1990) Inhibition by Agrobacterium tumefaciens and Pseudomonas savastanoi of development of the hypersensitive response elicited by Pseudomonas syringae pv. phaseolicola. J Bacteriol 172: 5742-5749
81. Roitsch T (1999) Source-sink regulation by sugar and stress. Curr Opin Plant Biol 2: 198-206
82. Roitsch T, Balibrea ME, Hofmann M, Proels R, Sinha AK (2003) Extracellular invertase: key metabolic enzyme and PR protein. J Exp Bot 54: 513-524
83. Roitsch T, Ehneß R (2000) Regulation of source/sink relations by cytokinins. Plant Growth Regul 32: 359-367
84. Roitsch T, Gonza'lez M-C (2004) Function and regulation of plant invertases: sweet sensations. Trends Plant Sci 9: 606-613
85. Rolland F, Baena-Gonzalez E, Sheen J (2006) Sugar sensing and signaling in plants: conserved and novel mechanisms. Annu Rev Plant Biol 57: 675-709
86. Salvucci ME (2008) Association of Rubisco activase with chaperonin-60b: a possible mechanism for protecting photosynthesis during heat stress. J Exp Bot 59: 1923-1933
87. Scharte J, Schön H, Weis E (2005) Photosynthesis and carbohydrate metabolism in tobacco leaves during an incompatible interaction with Phytophthora nicotianae. Plant Cell Environ 28: 1421-1435
88. Schauer N, Semel Y, Roessner U, Gur A, Balbo I, Carrari F, Pleban T, Perez-Melis A, Bruedigam C, Kopka J, et al (2006) Comprehensive metabolic profiling and phenotyping of interspecific introgression lines for tomato improvement. Nat Biotechnol 24: 447-454
89. Scholes JD, Lee PJ, Horton P, Lewis DH (1994) Invertase: understanding changes in the photosynthetic and carbohydrate metabolism of barley leaves infected with powdery mildew. New Phytol 126: 213-222
90. Simo'n-Mateo C, Depuydt S, de Oliveira Manes CL, Cnudde F, Holsters M, Goethals K, Vereecke D (2006) The phytopathogen Rhodococcus fascians breaks apical dominance and activates axillary meristems by inducing plant genes involved in hormone metabolism. Mol Plant Pathol 7: 103-112
91. Smeekens S (2000) Sugar-induced signal transduction in plants. Annu Rev Plant Physiol Plant Mol Biol 51: 49-81
92. Swarbrick PJ, Schulze-Lefert P, Scholes J (2006) Metabolic consequences of susceptibility and resistance (race-specific and broad-spectrum) in barley leaves challenged with powdery mildew. Plant Cell Environ 29: 1061-1076
93. Temmerman W, Vereecke D, Dreesen R, Van Montagu M, Holsters M, Goethals K (2000) Leafy gall formation is controlled by fasR, an AraCtype regulatory gene, in Rhodococcus fascians. J Bacteriol 182: 5832-5840
94. Torres MA, Jones JDG, Dangl JL (2006) Reactive oxygen species signaling in response to pathogens. Plant Physiol 141: 373-378
95. Truman W, Torres de Zabala M, Grant M (2006) Type III effectors orchestrate a complex interplay between transcriptional networks to modify basal defence responses during pathogenesis and resistance. Plant J 46: 14-33
96. Tymowska-Lalanne Z, Kreis M (1998) Expression of the Arabidopsis thaliana invertase gene family. Planta 207: 259-265
97. Vandeputte O, Öden S, Mol A, Vereecke D, Goethals K, El Jaziri M, Prinsen E (2005) Biosynthesis of auxin by the Gram-positive phytopathogen Rhodococcus fascians is controlled by compounds specific to infected plant tissues. Appl Environ Microbiol 71: 1169-1177
98. Vandeputte O, Vereecke D, Mol A, Lenjou M, Van Bockstaele D, El Jaziri M, Baucher M (2007) Rhodococcus fascians infection accelerates progression of tobacco BY-2 cells into mitosis through rapid changes in plant gene expression. New Phytol 175: 140-154
99. Vargas WA, Pontis HG, Salerno GL (2008) New insights on sucrose metabolism: evidence for an active A/N-Inv in chloroplasts uncovers a novel component of the intracellular carbon trafficking. Planta 227: 795-807
100. Vereecke D, Burssens S, Simo'n-Mateo C, Inze' D, Van Montagu M, Goethals K, Jaziri M (2000) The Rhodococcus fascians-plant interaction: morphological traits and biotechnological applications. Planta 210: 241-251
101. Vereecke D, Cornelis K, Temmerman W, Holsters M, Goethals K (2002a) Versatile persistence pathways for pathogens of animals and plants. Trends Microbiol 10: 485-488
102. Vereecke D, Cornelis K, Temmerman W, Jaziri M, Van Montagu M, Holsters M, Goethals K (2002b) Chromosomal locus that affects the pathogenicity of Rhodococcus fascians. J Bacteriol 184: 1112-1120
103. Vogel G, Fiehn O, Jean-Richard-dit-Bressel L, Boller T, Wiemken A, Aeschbacher RA, Wingler A (2001) Trehalose metabolism in Arabidopsis: occurrence of trehalose and molecular cloning and characterization of trehalose-6-phosphate synthase homologues. J Exp Bot 52: 1817-1826
104. Walters DR (2003) Polyamines and plant disease. Phytochemistry 64: 97-107
105. Walters DR, McRoberts N (2006) Plants and biotrophs: a pivotal role for cytokinins? Trends Plant Sci 11: 581-586
106. Walters DR, Shuttleton MA (1985) Polyamines in the roots of turnip infected with Plasmodiophora brassicae Wor. New Phytol 100: 209-214
107. Wingler A (2002) The function of trehalose biosynthesis in plants. Phytochemistry 60: 437-440
108. Wolfinger RD, Gibson G,Wolfinger ED, Bennett L, Hamadeh H, Bushel P, Afshari C, Paules RS (2001) Assessing gene significance from cDNA microarray expression data via mixed models. J Comput Biol 8: 625-637
109. Wright DP, Baldwin BC, Shephard MC, Scholes JD (1995) Source-sink relationships in wheat leaves infected with powdery mildew. I. Alterations in carbohydrate metabolism. Physiol Mol Plant Pathol 47: 237-253
110. Wright DP, Read DJ, Scholes JD (1998) Mycorrhizal sink strength influences whole plant carbon balance of Trifolium repens L. Plant Cell Environ 21: 881-891
111. Zimmermann P, Hirsch-Hoffmann M, Hennig L, Gruissem W (2004) GENEVESTIGATOR: Arabidopsis microarray database and analysis toolbox. Plant Physiol 136: 2621-2632