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== Introduction ==
== Introduction ==
[[Image:Organ_identity.jpg|600px|left|thumb| Figure 1. Schematic presentation LEAFY regulatory roles in controlling floral organ identity (A) and the ABC model in ''Arabidopsis thaliana'' (B). ]]
[[Image:Organ_identity.jpg|600px|left|thumb| Figure 1. Schematic presentation LEAFY regulatory roles in controlling floral organ identity (A) and the ABC model in ''Arabidopsis thaliana'' (B). ]]
'''''FLORICAULA/LEAFY'' (''FLO/LFY'')''' genes encode a plant specific transcription factor family that controlling floral fate of reproductive phase. <ref name="Weigel1992"> Detlef Weigel, John Alvarez, David R. Smyth, Martin F. Yanofsky, Elliot M. Meyerowitz, LEAFY controls floral meristem identity in Arabidopsis. Cell 69 :843-859, http://dx.doi.org/10.1016/0092-8674(92)90295-N.</ref> In the plant model system ''Arabidopsis thaliana '', ‘’LFY’’ also acts upstream of floral homeotic genes to modulate organ identity. <ref name= ''Irish2010'' > Irish, V. F. (2010), The flowering of Arabidopsis flower development. The Plant Journal, 61: 1014–1028.  http://dx.doi.org/10.1111/j.1365-313X.2009.04065.x </ref>  LFY activates the organ identity genes by binding to promoter regions of floral organ identity genes. LFY can directly bind to the promoter to '''''APELATA1'' (''AP1'')''', while co-regulators '''''UNUSUAL FLORAL ORGANS'' (''UFO'')''' <ref name= ''Chae2008'' >Chae, E., Tan, Q.K., Hill, T.A. & Irish, V.F. 2008. An Arabidopsis F-box protein acts as a transcriptional co-factor to regulate floral development. Development 135:1235-45  http://dx.doi.org/10.1242/dev.015842</ref> and '''''WUSCHEL'' (''WUS'')'''<ref name= ''Hong2003'' >HONG, R.L., HAMAGUCHI, L., BUSCH, M.A. and WEIGEL, D. (2003). Regulatory elements of the floral homeotic gene AGAMOUS identified by phylogenetic footprinting and shadowing. Plant Cell 15: 1296-1309. http:/ / dx. doi. org/ 10. 1105/ tpc. 009548</ref> are required for increment of binding affinity to promoter regions of '''''APELATA3'' (''AP3'')''' and '''''AGAMOUS'' (''AG'')''', respectively. The exact mechanism how LFY binds to these promoters has yet to be well elucidated until the first structure report about <scene name='57/579703/Ap1-dimer/1'>LFY-pAP1</scene> and <scene name='57/579703/2vy2_assembly/1'>LFY-pAG</scene><ref name= ''Hames2008'' >Hames, C., Ptchelkine, D., Grimm, C., Thevenon, E., Moyroud, E., Gérard, F. Martiel, J.L., Benlloch, R., Parcy, F. & Müller, C.W. 2008. Structural basis for LEAFY floral switch function and similarity with helix-turn-helix proteins. EMBO Journal 27:2628-2637. http://dx.doi.org/10.1038/emboj.2008.184
'''''FLORICAULA/LEAFY'' (''FLO/LFY'')''' genes encode a plant specific transcription factor family that controlling floral fate of reproductive phase. <ref name="Weigel1992"> Detlef Weigel, John Alvarez, David R. Smyth, Martin F. Yanofsky, Elliot M. Meyerowitz, LEAFY controls floral meristem identity in Arabidopsis. Cell 69 :843-859, http://dx.doi.org/10.1016/0092-8674(92)90295-N.</ref> In the plant model system ''Arabidopsis thaliana '', ‘’LFY’’ also acts upstream of floral homeotic genes to modulate organ identity. <ref name= ''Irish2010'' > Irish, V. F. (2010), The flowering of Arabidopsis flower development. The Plant Journal, 61: 1014–1028.  http://dx.doi.org/10.1111/j.1365-313X.2009.04065.x </ref>  LFY activates the organ identity genes by binding to promoter regions of floral organ identity genes. LFY can directly bind to the promoter to '''''APELATA1'' (''AP1'')''', while co-regulators '''''UNUSUAL FLORAL ORGANS'' (''UFO'')''' <ref name= ''Chae2008'' >Chae, E., Tan, Q.K., Hill, T.A. & Irish, V.F. 2008. An Arabidopsis F-box protein acts as a transcriptional co-factor to regulate floral development. Development 135:1235-45  http://dx.doi.org/10.1242/dev.015842</ref> and '''''WUSCHEL'' (''WUS'')'''<ref name= ''Hong2003'' >HONG, R.L., HAMAGUCHI, L., BUSCH, M.A. and WEIGEL, D. (2003). Regulatory elements of the floral homeotic gene AGAMOUS identified by phylogenetic footprinting and shadowing. Plant Cell 15: 1296-1309. http://dx.doi.org/10.1105/tpc.009548</ref> are required for increment of binding affinity to promoter regions of '''''APELATA3'' (''AP3'')''' and '''''AGAMOUS'' (''AG'')''', respectively. The exact mechanism how LFY binds to these promoters has yet to be well elucidated until the first structure report about <scene name='57/579703/Ap1-dimer/1'>LFY-pAP1</scene> and <scene name='57/579703/2vy2_assembly/1'>LFY-pAG</scene><ref name= ''Hames2008'' >Hames, C., Ptchelkine, D., Grimm, C., Thevenon, E., Moyroud, E., Gérard, F. Martiel, J.L., Benlloch, R., Parcy, F. & Müller, C.W. 2008. Structural basis for LEAFY floral switch function and similarity with helix-turn-helix proteins. EMBO Journal 27:2628-2637. http://dx.doi.org/10.1038/emboj.2008.184
</ref> . Among land plants, FLO/LFY homologs share a highly conserved DNA binding region that a hypothesis claimed substitution in this domain might result in the functional divergence<ref name= ''Maize2005'' >MAIZEL, A., BUSCH, M.A., TANAHASHI, T., PERKOVIC, J., KATAO, M., HASEBE, M. and WEIGEL, D. (2005). The floral regulator LEAFY evolves by substitutions in the DNA binding domain. Science 308: 260-263. http://dx.doi.org/10.1126/science.1108229</ref> . Recently, a new study (ref 7) provided new insights of structural basis of LEAFY evolution by changing DNA binding activity<ref name= ''Sayou2014'' >Sayou, C., Monniaux, M., Nanao, M.H., Moyroud, E., Brockington, S.F., Thévenon, E., Chahtane, H., Warthmann, N., Melkonian, M., Zhang, Y., Wong, G., Weigel, D., Parcy, F. and Dumas, R. 2014. A Promiscuous Intermediate Underlies the Evolution of LEAFY DNA Binding Specificity Science  343: 645-648 http://dx.doi.org/10.1126/science.1248229</ref>.  
</ref> . Among land plants, FLO/LFY homologs share a highly conserved DNA binding region that a hypothesis claimed substitution in this domain might result in the functional divergence<ref name= ''Maize2005'' >MAIZEL, A., BUSCH, M.A., TANAHASHI, T., PERKOVIC, J., KATAO, M., HASEBE, M. and WEIGEL, D. (2005). The floral regulator LEAFY evolves by substitutions in the DNA binding domain. Science 308: 260-263. http://dx.doi.org/10.1126/science.1108229</ref> . Recently, a new study (ref 7) provided new insights of structural basis of LEAFY evolution by changing DNA binding activity<ref name= ''Sayou2014'' >Sayou, C., Monniaux, M., Nanao, M.H., Moyroud, E., Brockington, S.F., Thévenon, E., Chahtane, H., Warthmann, N., Melkonian, M., Zhang, Y., Wong, G., Weigel, D., Parcy, F. and Dumas, R. 2014. A Promiscuous Intermediate Underlies the Evolution of LEAFY DNA Binding Specificity Science  343: 645-648 http://dx.doi.org/10.1126/science.1248229</ref>.