title: Mechanisms of signal transduction by ethylene: overlapping and non-overlapping signalling roles in a receptor family.
abstract: The plant hormone ethylene regulates growth and development as well as responses to biotic and abiotic stresses. Over the last few decades, key elements involved in ethylene signal transduction have been identified through genetic approaches, these elements defining a pathway that extends from initial ethylene perception at the endoplasmic reticulum to changes in transcriptional regulation within the nucleus. Here, we present our current understanding of ethylene signal transduction, focusing on recent developments that support a model with overlapping and non-overlapping roles for members of the ethylene receptor family. We consider the evidence supporting this model for sub-functionalization within the receptor family, and then discuss mechanisms by which such a sub-functionalization may occur. To this end, we consider the importance of receptor interactions in modulating their signal output and how such interactions vary in the receptor family. In addition, we consider evidence indicating that ethylene signal output by the receptors involves both phosphorylation-dependent and phosphorylation-independent mechanisms. We conclude with a current model for signalling by the ethylene receptors placed within the overall context of ethylene signal transduction.
authors.affiliation: Department of Biological Sciences , Dartmouth College , Hanover, NH 03755 , USA ; Department of Biochemistry , Quaid-i-azam University , Islamabad 45320 , Pakistan.
authors.forename: Samina N
authors.initials: SN
authors.lastname: Shakeel
authors.forename: Xiaomin
authors.initials: X
authors.lastname: Wang
authors.forename: Brad M
authors.initials: BM
authors.lastname: Binder
authors.forename: G Eric
authors.initials: GE
authors.lastname: Schaller
keywords: Arabidopsis; ethylene; ethylene receptors; histidine kinase; hormone signalling; sub-functionalization; two-component system
references.citation: Plant Physiol. 2012 Dec;160(4):1968-84
references.pmid: 23043080
references.citation: BMC Biol. 2012 Feb 20;10:9
references.pmid: 22348804
references.citation: Ann Bot. 2008 Jan;101(2):229-48
references.pmid: 17956854
references.citation: PLoS One. 2010 Jan 08;5(1):e8640
references.pmid: 20062808
references.citation: Annu Rev Biochem. 2000;69:183-215
references.pmid: 10966457
references.citation: J Exp Bot. 2004 Mar;55(397):547-56
references.pmid: 14754915
references.citation: Mol Biosyst. 2011 Sep;7(9):2637-50
references.pmid: 21713283
references.citation: EMBO J. 2003 Mar 17;22(6):1282-8
references.pmid: 12628921
references.citation: Mol Plant. 2010 Sep;3(5):882-9
references.pmid: 20591837
references.citation: Plant Physiol. 1976 Sep;58(3):268-71
references.pmid: 16659660
references.citation: Cell. 1993 Feb 12;72(3):427-41
references.pmid: 8431946
references.citation: Plant Physiol. 2012 Jun;159(2):682-95
references.pmid: 22467798
references.citation: Nature. 2008 Feb 14;451(7180):789-95
references.pmid: 18273012
references.citation: J Exp Bot. 2008;59(4):965-72
references.pmid: 18349053
references.citation: Plant Cell. 1990 Jun;2(6):513-23
references.pmid: 2152173
references.citation: Proc Natl Acad Sci U S A. 2003 Jan 7;100(1):352-7
references.pmid: 12509505
references.citation: Mol Plant. 2008 Mar;1(2):380-7
references.pmid: 19825547
references.citation: Plant Physiol. 2004 Jun;135(2):660-7
references.pmid: 15208412
references.citation: Plant Physiol. 2002 Aug;129(4):1557-67
references.pmid: 12177468
references.citation: FEBS Lett. 2000 May 26;474(1):29-32
references.pmid: 10828445
references.citation: Cell. 1999 Apr 30;97(3):383-93
references.pmid: 10319818
references.citation: Comput Appl Biosci. 1994 Dec;10(6):685-6
references.pmid: 7704669
references.citation: J Biol Chem. 2012 Jul 27;287(31):26094-103
references.pmid: 22692214
references.citation: J Biol Chem. 2007 Aug 24;282(34):24752-8
references.pmid: 17595158
references.citation: Proc Natl Acad Sci U S A. 1998 May 12;95(10):5812-7
references.pmid: 9576967
references.citation: J Biol Chem. 2002 May 31;277(22):19861-6
references.pmid: 11916973
references.citation: Plant J. 2008 Jan;53(2):275-86
references.pmid: 17999643
references.citation: Arabidopsis Book. 2002;1:e0071
references.pmid: 22303216
references.citation: Mol Plant. 2008 Mar;1(2):308-20
references.pmid: 19825542
references.citation: Plant Physiol. 2006 Dec;142(4):1690-700
references.pmid: 17071649
references.citation: Plant J. 2008 Apr;54(1):129-40
references.pmid: 18182027
references.citation: Plant Physiol. 1995 Jun;108(2):597-607
references.pmid: 7610160
references.citation: Annu Rev Plant Biol. 2006;57:837-58
references.pmid: 16669784
references.citation: J Biol Chem. 2004 Nov 19;279(47):48734-41
references.pmid: 15358768
references.citation: Proc Natl Acad Sci U S A. 2003 Mar 4;100(5):2992-7
references.pmid: 12606727
references.citation: Science. 2012 Oct 19;338(6105):390-3
references.pmid: 22936567
references.citation: BMC Plant Biol. 2007 Jan 15;7:3
references.pmid: 17224067
references.citation: Biochem J. 2009 Oct 23;424(1):1-6
references.pmid: 19769567
references.citation: J Exp Bot. 2009;60(13):3923-33
references.pmid: 19648171
references.citation: Science. 1988 Aug 26;241(4869):1086-9
references.pmid: 17747490
references.citation: Plant J. 2007 Aug;51(3):458-67
references.pmid: 17655616
references.citation: Cell Res. 2012 Nov;22(11):1613-6
references.pmid: 23070300
references.citation: Mol Cell Proteomics. 2006 Apr;5(4):749-57
references.pmid: 16340016
references.citation: Plant Cell. 2004 Dec;16(12):3386-99
references.pmid: 15539472
references.citation: AoB Plants. 2012;2012:pls031
references.pmid: 23119138
references.citation: Plant Physiol. 2002 Dec;130(4):1983-91
references.pmid: 12481081
references.citation: Proc Natl Acad Sci U S A. 1995 May 9;92(10):4129-33
references.pmid: 11607538
references.citation: Plant Cell Physiol. 2009 Sep;50(9):1636-50
references.pmid: 19608714
references.citation: J Biol Chem. 2008 Aug 29;283(35):23801-10
references.pmid: 18577522
references.citation: Proc Natl Acad Sci U S A. 2006 May 16;103(20):7917-22
references.pmid: 16682642
references.citation: Plant Physiol. 2012 Sep;160(1):488-97
references.pmid: 22797658
references.citation: J Biol Chem. 2010 Dec 24;285(52):40706-13
references.pmid: 20952388
references.citation: Curr Opin Plant Biol. 2009 Oct;12(5):548-55
references.pmid: 19709924
references.citation: Proc Natl Acad Sci U S A. 1998 Jun 23;95(13):7825-9
references.pmid: 9636235
references.citation: Plant J. 2005 Mar;41(5):651-9
references.pmid: 15703053
references.citation: Genes Dev. 1998 Dec 1;12(23):3703-14
references.pmid: 9851977
references.citation: Plant Physiol. 2012 Jul;159(3):1263-76
references.pmid: 22566492
references.citation: Plant Cell. 1998 Aug;10(8):1321-32
references.pmid: 9707532
references.citation: Plant J. 2003 Jan;33(2):221-33
references.pmid: 12535337
references.citation: Proc Natl Acad Sci U S A. 2012 Nov 20;109(47):19486-91
references.pmid: 23132950
references.citation: Science. 1999 Feb 12;283(5404):996-8
references.pmid: 9974395
references.citation: Science. 1995 Dec 15;270(5243):1809-11
references.pmid: 8525372
references.citation: DNA Res. 1997 Apr 28;4(2):161-8
references.pmid: 9205844
references.citation: Trends Plant Sci. 1999 Jul;4(7):269-274
references.pmid: 10407443
references.citation: Cell. 1998 Jul 24;94(2):261-71
references.pmid: 9695954
references.citation: Proc Natl Acad Sci U S A. 1998 Apr 28;95(9):5401-6
references.pmid: 9560288
references.citation: Plant Physiol. 2000 Aug;123(4):1449-58
references.pmid: 10938361
references.citation: Plant Cell. 2005 Nov;17(11):3007-18
references.pmid: 16227453
references.citation: Science. 1993 Oct 22;262(5133):539-44
references.pmid: 8211181
references.citation: Mol Plant. 2011 Jul;4(4):626-34
references.pmid: 21690206
references.citation: J Biol Chem. 2010 Nov 26;285(48):37263-70
references.pmid: 20876528
references.citation: Curr Biol. 2011 May 10;21(9):R320-30
references.pmid: 21549954
references.citation: Plant J. 2003 Jun;34(5):709-18
references.pmid: 12787251
references.citation: Plant Physiol. 2011 May;156(1):417-29
references.pmid: 21386032
references.citation: J Biol Chem. 2003 Sep 5;278(36):34725-32
references.pmid: 12821658
references.citation: PLoS One. 2011;6(9):e24173
references.pmid: 21912672
references.citation: Plant Physiol. 2002 Sep;130(1):10-4
references.pmid: 12226482
references.citation: Plant Cell. 2003 Sep;15(9):2032-41
references.pmid: 12953109
references.citation: Arabidopsis Book. 2008;6:e0112
references.pmid: 22303237
references.citation: EMBO J. 2004 Aug 18;23(16):3290-302
references.pmid: 15282545
references.citation: J Biol Chem. 1995 May 26;270(21):12526-30
references.pmid: 7759498
references.citation: Genetics. 1995 Mar;139(3):1393-409
references.pmid: 7768447
references.citation: Plant Physiol. 2009 Jun;150(2):547-51
references.pmid: 19369589
references.citation: Proc Natl Acad Sci U S A. 2006 May 16;103(20):7923-8
references.pmid: 16682641
references.citation: Proc Natl Acad Sci U S A. 2000 May 9;97(10):5663-8
references.pmid: 10792050
references.citation: Science. 1995 Sep 22;269(5231):1712-4
references.pmid: 7569898
references.citation: Annu Rev Microbiol. 2009;63:133-54
references.pmid: 19575571
references.citation: Ann Bot. 2005 May;95(6):901-15
references.pmid: 15753119
references.citation: Plant Physiol. 2004 Oct;136(2):2913-20
references.pmid: 15466220
references.citation: Bioessays. 2001 Jul;23(7):619-27
references.pmid: 11462215
references.citation: Anal Biochem. 2008 Jun 1;377(1):72-6
references.pmid: 18384742
references.citation: J Exp Bot. 2009;60(12):3311-36
references.pmid: 19567479
references.citation: Plant Physiol. 2004 Oct;136(2):2961-70
references.pmid: 15466228
references.citation: J Biol Chem. 2008 Oct 3;283(40):26996-7006
references.pmid: 18693252
references.citation: Science. 1999 Jun 25;284(5423):2148-52
references.pmid: 10381874
references.citation: Plant Physiol. 2007 Sep;145(1):75-86
references.pmid: 17644624
references.citation: FEBS Lett. 2000 Aug 4;478(3):227-32
references.pmid: 10930573
references.citation: Plant Physiol. 2006 Jun;141(2):587-97
references.pmid: 16617090
references.citation: FEBS Lett. 2007 Oct 30;581(26):5105-9
references.pmid: 17931631
references.citation: Plant Cell. 2000 Mar;12(3):443-55
references.pmid: 10715329
references.citation: Plant Physiol. 2006 Oct;142(2):492-508
references.pmid: 16891553
references.citation: Cell. 1997 Jun 27;89(7):1133-44
references.pmid: 9215635
references.citation: BMC Plant Biol. 2010 Apr 08;10:60
references.pmid: 20374664
references.citation: Plant Cell. 2009 May;21(5):1473-94
references.pmid: 19417056