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Gene Review

ZTL  -  adagio protein 1

Arabidopsis thaliana

Synonyms: ADAGIO 1, ADO1, FKF1-LIKE PROTEIN 2, FKL2, LKP1, ...
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High impact information on ZTL

  • Map-based cloning of ZTL identified a novel 609 amino acid polypeptide consisting of an amino-terminal PAS domain, an F box and six carboxy-terminal kelch repeats [1].
  • Both yeast two-hybrid and in vitro binding studies show that there is a physical interaction between ADO1 and the photoreceptors CRY1 and phyB [2].
  • We found that a loss-of-function ado1 mutant is altered in both gene expression and cotyledon movement in circadian rhythmicity [2].
  • A third member of the ZTL gene family was identified in the Arabidopsis genome and was named LKP2 (for LOV kelch protein2) [3].
  • This proteolysis is proteasome dependent, implicating ZTL itself as substrate for ubiquitination [4].

Biological context of ZTL

  • This uncoupling of ztl phenotypes indicates that interactions of ZTL protein with multiple factors must be disrupted to generate the full ztl mutant phenotype [5].
  • Other mutants recovered include 11 new ztl alleles encompassing mutations in each of the ZTL protein domains [5].
  • Mutation of the kelch repeat domain affected protein binding at both the LOV/PAS and the F-box domains, indicating that interaction among ZTL domains leads to the strong phenotypes of kelch mutations [5].
  • The LKP1 LOV is very similar to the LOV domains in NPH1, a plasma membrane-associated blue light receptor kinase that regulates phototropism (Huala, E., Oeller, P.W., Liscum, E., Han, I-S., Larsen, E. & Briggs, W.R. (1997) Science, 278, 2120-2123) [6].
  • Transgenic plants with a beta-glucuronidase (GUS) reporter gene driven by a 1.5 kb LKP1 promoter display strong GUS activity in leaves [6].

Anatomical context of ZTL

  • The action of phytochromes and cryptochromes in photoperiodism is augmented by ZEITLUPE (ZTL) and FLAVIN-BINDING, KELCH REPEAT, F-BOX (FKF1) acting as accessory photoreceptors on entrainment and interaction, respectively [7].
  • The GFP-associated fluorescence in 35S:GFP-LKP1 plants is observed in nuclei and cytosol, indicating that LKP1 is a new nucleo-cytoplasmic factor that influences flowering time in the long day pathway of Arabidopsis [6].

Regulatory relationships of ZTL

  • Our results show that the TOC1-ZTL interaction is important in the control of TOC1 protein stability, and is probably responsible for the regulation of circadian period by the clock [8].

Other interactions of ZTL

  • LKP2 interacted not only with itself but also with other members of the family, LKP1 and FKF1 [9].
  • Here we show that the effects of ZTL and ELF3 on circadian clock function and early photomorphogenesis are additive [10].
  • ZTL overexpression does not delay flowering through changes in GIGANTEA or FLAVIN-BINDING, KELCH REPEAT, F-BOX levels, but through a ZTL-mediated reduction in CO expression [10].
  • Targeted degradation of TOC1 by ZTL modulates circadian function in Arabidopsis thaliana [8].
  • Interaction with PHYB was unaffected by mutation of any ZTL domain [5].

Analytical, diagnostic and therapeutic context of ZTL

  • Here we use a newly characterized Arabidopsis suspension cell culture to establish that the rhythmic changes in the levels of the clock-associated F-box protein, ZTL, are posttranscriptionally controlled through different circadian phase-specific degradation rates [4].


  1. ZEITLUPE encodes a novel clock-associated PAS protein from Arabidopsis. Somers, D.E., Schultz, T.F., Milnamow, M., Kay, S.A. Cell (2000) [Pubmed]
  2. An Arabidopsis circadian clock component interacts with both CRY1 and phyB. Jarillo, J.A., Capel, J., Tang, R.H., Yang, H.Q., Alonso, J.M., Ecker, J.R., Cashmore, A.R. Nature (2001) [Pubmed]
  3. A role for LKP2 in the circadian clock of Arabidopsis. Schultz, T.F., Kiyosue, T., Yanovsky, M., Wada, M., Kay, S.A. Plant Cell (2001) [Pubmed]
  4. Circadian phase-specific degradation of the F-box protein ZTL is mediated by the proteasome. Kim, W.Y., Geng, R., Somers, D.E. Proc. Natl. Acad. Sci. U.S.A. (2003) [Pubmed]
  5. Forward genetic analysis of the circadian clock separates the multiple functions of ZEITLUPE. Kevei, E., Gyula, P., Hall, A., Kozma-Bognár, L., Kim, W.Y., Eriksson, M.E., Tóth, R., Hanano, S., Fehér, B., Southern, M.M., Bastow, R.M., Viczián, A., Hibberd, V., Davis, S.J., Somers, D.E., Nagy, F., Millar, A.J. Plant Physiol. (2006) [Pubmed]
  6. LKP1 (LOV kelch protein 1): a factor involved in the regulation of flowering time in arabidopsis. Kiyosue, T., Wada, M. Plant J. (2000) [Pubmed]
  7. Light signals and flowering. Thomas, B. J. Exp. Bot. (2006) [Pubmed]
  8. Targeted degradation of TOC1 by ZTL modulates circadian function in Arabidopsis thaliana. Más, P., Kim, W.Y., Somers, D.E., Kay, S.A. Nature (2003) [Pubmed]
  9. Identification of ASK and clock-associated proteins as molecular partners of LKP2 (LOV kelch protein 2) in Arabidopsis. Yasuhara, M., Mitsui, S., Hirano, H., Takanabe, R., Tokioka, Y., Ihara, N., Komatsu, A., Seki, M., Shinozaki, K., Kiyosue, T. J. Exp. Bot. (2004) [Pubmed]
  10. Independent roles for EARLY FLOWERING 3 and ZEITLUPE in the control of circadian timing, hypocotyl length, and flowering time. Kim, W.Y., Hicks, K.A., Somers, D.E. Plant Physiol. (2005) [Pubmed]
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