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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). ]]
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'''''FLORICAULA/LEAFY'' (''FLO/LFY'')''' genes encode a plant specific transcription factor family that controlling floral fate of reproductive phase. <ref name="Weigel1992"> Weigel, D., Alvarez, J., David R., Yanofsky, M.F. & Meyerowitz, E.M. 1992. ''LEAFY'' controls floral meristem identity in Arabidopsis. Cell 69 :843-859, http://dx.doi.org/10.1016/0092-8674(92)90295-N.</ref><ref name="Coen1990"> Coen, E.S., Romero, J.M., Doyle, S., Elliot, R., Murphy, G. & Carpenter, R. 1990. ''Floricaula'': a homeotic gene required for flower development in Antirrhinum majus. Cell 63: 1311–1322 http://dx.doi.org/10.1016/0092-8674(90)90426-F</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. The 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 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>.
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'''''FLORICAULA/LEAFY'' (''FLO/LFY'')''' genes encode a plant specific transcription factor family that controlling floral fate of reproductive phase. <ref name="Weigel1992"> Weigel, D., Alvarez, J., David R., Yanofsky, M.F. & Meyerowitz, E.M. 1992. ''LEAFY'' controls floral meristem identity in Arabidopsis. Cell 69 :843-859, http://dx.doi.org/10.1016/0092-8674(92)90295-N.</ref><ref name="Coen1990"> Coen, E.S., Romero, J.M., Doyle, S., Elliot, R., Murphy, G. & Carpenter, R. 1990. ''Floricaula'': a homeotic gene required for flower development in Antirrhinum majus. Cell 63: 1311–1322 http://dx.doi.org/10.1016/0092-8674(90)90426-F</ref>. In the plant model system ''Arabidopsis thaliana '', ‘’LFY’’ also acts upstream of floral homeotic genes to modulate floral 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 downstream genes by binding to promoter regions. 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. The 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 recognizing and binding to these promoters has yet to be elucidated until the first structure report about two DNA-protein complex： <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 structure about LFY homolog in mosses provided new insights of structural basis of how LEAFY evolved 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>.

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This Sandbox is Reserved from 01/04/2014, through 30/06/2014 for use in the course "510042. Protein structure, function and folding" taught by Prof Adrian Goldman, Tommi Kajander, Taru Meri, Konstantin Kogan and Juho Kellosalo at the University of Helsinki. This reservation includes Sandbox Reserved 923 through Sandbox Reserved 947.
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Contents

Evolution of DNA binding domain of LEAFY: from angiosperms to mosses

Introduction

Figure 1. Schematic presentation LEAFY regulatory roles in controlling floral organ identity (A) and the ABC model in Arabidopsis thaliana (B).
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. [1][2]. In the plant model system Arabidopsis thaliana , ‘’LFY’’ also acts upstream of floral homeotic genes to modulate floral organ identity. [3] LFY activates the downstream genes by binding to promoter regions. LFY can directly bind to the promoter to APELATA1 (AP1), while co-regulators UNUSUAL FLORAL ORGANS (UFO) [4] and WUSCHEL (WUS)[5] are required for increment of binding affinity to promoter regions of APELATA3 (AP3) and AGAMOUS (AG), respectively. The exact mechanism how LFY recognizing and binding to these promoters has yet to be elucidated until the first structure report about two DNA-protein complex： and [6] . 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[7] . Recently, a new structure about LFY homolog in mosses provided new insights of structural basis of how LEAFY evolved by changing DNA binding activity[8].


Structure of LFY binding with AP1 and AG promoter region (PDB entry: 2VY1/2VY2)

Drag the structure with the mouse to rotate

LEAFY Evolution

Image:Structure course.007.jpg
Figure 4. A summary of LEAFY evolution by substation in three residues of DNA binding domain. Superposition of

Reference

  1. ↑ 1.0 1.1 1.2 Weigel, D., Alvarez, J., David R., Yanofsky, M.F. & Meyerowitz, E.M. 1992. LEAFY controls floral meristem identity in Arabidopsis. Cell 69 :843-859, http://dx.doi.org/10.1016/0092-8674(92)90295-N.
  2. ↑ 2.0 2.1 Coen, E.S., Romero, J.M., Doyle, S., Elliot, R., Murphy, G. & Carpenter, R. 1990. Floricaula: a homeotic gene required for flower development in Antirrhinum majus. Cell 63: 1311–1322 http://dx.doi.org/10.1016/0092-8674(90)90426-F
  3. ↑ 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
  4. ↑ 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
  5. ↑ 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. The Plant Cell 15: 1296-1309. http://dx.doi.org/10.1105/tpc.009548
  6. ↑ 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
  7. ↑ 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
  8. ↑ 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
  9. ↑ Siriwardana, N. S. & Lamb, R. S. 2012. A conserved domain in the N-terminus is important for LEAFY dimerization and function in Arabidopsis thaliana. The Plant Journal 71: 736–749. http://dx.doi.org/10.1111/j.1365-313X.2012.05026.x8
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