The ethylene signal components identi?ed so far in rice include receptors, EIN2 and EIN3 homologues, and CTR1, RTE1, EBF1/2 and EIN5 homologues that are reversely conservative in dicotyledon Arabidopsis and monocotledon rice. But based on the observation of ein2 and eil1 mutants of rice that there is not strong phenotypic difference between the two mutants and wild type, we conclude that ethylene signal in rice might be more intricate than in Arabidopsis. Given that rice uses ethylene to control many processes that do not exist in Arabidopsis, it seems that new mechanisms may exist in rice ethylene signal pathway. This article reviewed ethylene signal pathways, regulation of ethylene signal, and ethylene responses in rice plants.
[1] Yau C P, et al. Differential expression of three genes encoding an ethylene receptor in rice during development, and in response to indole-3-acetic acid and silver ions[J]. J Exp Bot., 2004,55(397): 547-556.
[2] Kendrick M D, et al. Ethylene signaling: new levels of complexity and regulation[J]. Curr Opin Plant Biol., 2008,11(5): 479-485.
[3] Itoh J, et al. Rice plant development: from zygote to spikelet[J]. Plant Cell Physiol., 2005,46(1): 23-47.
[4] Jun S H, et al. OsEIN2 is a positive component in ethylene signaling in rice[J]. Plant Cell Physiol., 2004,45(3): 281-289.
[5] Mao C, et al. OsEIL1, a rice homolog of the Arabidopsis EIN3 regulates the ethylene response as a positive component[J]. Plant Mol Biol., 2006,61(1-2): 141-152.
[6] Rzewuski G, et al. Ethylene biosynthesis and signaling in rice[J]. Plant Science, 2008, 175(12):32-42.
[7] Watanabe H, et al. Cloning of a cDNA encoding an ETR2-like protein (Os-ERL1) from deep water rice (Oryza sativa L.) and increase in its mRNA level by submergence, ethylene, and gibberellin treatments[J]. J Exp Bot., 2004,55(399): 1 145-1 148.
[8] Stock J B, et al. Signal transduction in bacteria[J]. Nature, 1990,344(6 265): 395-400.
[9] Wuriyanghan H, et al. The ethylene receptor ETR2 delays floral transition and affects starch accumulation in rice[J]. Plant Cell, 2009,21(5): 1 473-1 494.
[10] Kanacher T, et al. A GAF-domain-regulated adenylyl cyclase from Anabaena is a self-activating cAMP switch[J]. EMBO J, 2002,21(14): 3 672-3 680.
[11] Alonso J M, et al. Genome-wide insertional mutagenesis of Arabidopsis thaliana[J]. Science, 2003,301(5633): 653-657.
[12] Resnick J S, et al. REVERSION-TO-ETHYLENE SENSITIVITY1, a conserved gene that regulates ethylene receptor function in Arabidopsis[J]. Proc Natl Acad Sci USA, 2006,103(20): 7 917-7 922.
[13] Adams-Phillips L, et al. Evidence that CTR1-mediated ethylene signal transduction in tomato is encoded by a multigene family whose members display distinct regulatory features[J]. Plant Mol Biol., 2004,54(3): 387-404.
[14] Olmedo G, et al. ETHYLENE-INSENSITIVE5 encodes a 5'-->3' exoribonuclease required for regulation of the EIN3-targeting F-box proteins EBF1/2[J]. Proc Natl Acad Sci USA, 2006,103(36): 13 286-13 293.
[15] Gagne J M, et al. Arabidopsis EIN3-binding F-box 1 and 2 form ubiquitin-protein ligases that repress ethylene action and promote growth by directing EIN3 degradation[J]. Proc Natl Acad Sci USA, 2004, 101(17): 6803-6808.
[16] Potuschak T, et al. EIN3-dependent regulation of plant ethylene hormone signaling by two arabidopsis F box proteins: EBF1 and EBF2[J]. Cell, 2003,115(6): 679-689.
[17] Chao Q, et al. Activation of the ethylene gas response pathway in Arabidopsis by the nuclear protein ETHYLENE- INSENSITIVE3 and related proteins[J]. Cell, 1997,89(7): 1 133-1 144.
[18] Hiraga S, et al. Involvement of two rice ETHYLENE INSENSITIVE3-LIKE genes in wound signaling[J]. Mol Genet Genomics, 2009,282(5): 517-529.
[19] Riechmann J L, et al. The AP2/EREBP family of plant transcription factors[J]. Biol Chem., 1998,379(6): 633-646.
[20] Nakano T, et al. Genome-wide analysis of the ERF gene family in Arabidopsis and rice[J]. Plant Physiol., 2006,140(2): 411-432.
[21] Ohme-Takagi M, et al. Ethylene-inducible DNA binding proteins that interact with an ethylene-responsive element[J]. Plant Cell, 1995,7(2): 173-182.
[22] Solano R, et al. Nuclear events in ethylene signaling: a transcriptional cascade mediated by ETHYLENE-INSENSITIVE3 and ETHYLENE-RESPONSE-FACTOR1[J]. Genes Dev., 1998,12(23): 3 703-3 714.
[23] Cheong Y H, et al. BWMK1, a rice mitogen-activated protein kinase, locates in the nucleus and mediates pathogenesis- related gene expression by activation of a transcription factor[J]. Plant Physiol., 2003,132(4): 1961-1972.
[24] Hu Y, et al. Overexpression of OsERF1, a novel rice ERF gene, up-regulates ethylene-responsive genes expression besides affects growth and development in Arabidopsis[J]. J Plant Physiol., 2008,165(16):1 717-1 725.
[25] Hattori Y, et al. The ethylene response factors SNORKEL1 and SNORKEL2 allow rice to adapt to deep water[J]. Nature, 2009,460(7258): 1 026-1 030.
[26] Perata P, et al. Submergence tolerance in rice requires Sub1A, an ethylene-response-factor-like gene[J]. Trends Plant Sci., 2007,12(2): 43-46.
[27] Kende H, et al. Deepwater rice: A model plant to study stem elongation[J]. Plant Physiol., 1998,118(4): 1 105-1 110.
[28] Jackson M B, et al. Physiological and molecular basis of susceptibility and tolerance of rice plants to complete submergence[J]. Ann Bot., 2003,91 Spec No.: 227-241.
[29] Alonso J M, et al. EIN2, a bifunctional transducer of ethylene and stress responses in Arabidopsis[J]. Science, 1999,284(5 423): 2 148-2 152.
[30] Pareek A, et al. Whole-genome analysis of Oryza sativa reveals similar architecture of two-component signaling machinery with Arabidopsis[J]. Plant Physiol., 2006,142(2): 380-397.
[31] Hoffmann-Benning S, et al. On the role of abscisic Acid and gibberellin in the regulation of growth in rice[J]. Plant Physiol., 1992,99(3): 1 156-1 161.
[32] Yang S H, et al. Characterization of genes encoding ABA 8'-hydroxylase in ethylene-induced stem growth of deepwater rice (Oryza sativa L.)[J]. Biochem Biophys Res Commun., 2006,350(3): 685-690.
[33] Steffens B, et al. Epidermal cell death in rice is regulated by ethylene, gibberellin, and abscisic acid[J]. Plant Physiol., 2005,139(2): 713-721.
[34] Steffens B, et al. Interactions between ethylene, gibberellin and abscisic acid regulate emergence and growth rate of adventitious roots in deepwater rice[J]. Planta, 2006,223(3): 604-612.
[35] Ikeda A, et al. The slender rice mutant, with constitutively activated gibberellin signal transduction, has enhanced capacity for abscisic acid level[J]. Plant Cell Physiol., 2002,43(9): 974-979.
[36] Mergemann H, et al. Ethylene induces epidermal cell death at the site of adventitious root emergence in rice[J]. Plant Physiol., 2000,124(2): 609-614.
[37] Lorbiecke R, et al. Adventitious root growth and cell-cycle induction in deepwater rice[J]. Plant Physiol., 1999,119(1): 21-30.