Gene Symbol: ZTL
Description: Galactose oxidase/kelch repeat superfamily protein
Alias: ADAGIO 1, ADO1, FKF1-LIKE PROTEIN 2, FKL2, LKP1, LOV KELCH PROTEIN 1, MSF19.2, MSF19_2, ZEITLUPE, Galactose oxidase/kelch repeat superfamily protein
Species: thale cress
Products:     ZTL

Top Publications

  1. Kim W, Fujiwara S, Suh S, Kim J, Kim Y, Han L, et al. ZEITLUPE is a circadian photoreceptor stabilized by GIGANTEA in blue light. Nature. 2007;449:356-60 pubmed
    ..Rhythmic expression of the Arabidopsis thaliana F-box protein ZEITLUPE (ZTL) is necessary to sustain a normal circadian period by controlling the proteasome-dependent degradation of a ..
  2. Kim W, Hicks K, Somers D. Independent roles for EARLY FLOWERING 3 and ZEITLUPE in the control of circadian timing, hypocotyl length, and flowering time. Plant Physiol. 2005;139:1557-69 pubmed
    ..b>ZEITLUPE (ZTL) is a clock-related F-box protein, and altered ZTL expression causes fluence rate-dependent circadian period ..
  3. Fujiwara S, Wang L, Han L, Suh S, Salome P, McClung C, et al. Post-translational regulation of the Arabidopsis circadian clock through selective proteolysis and phosphorylation of pseudo-response regulator proteins. J Biol Chem. 2008;283:23073-83 pubmed publisher
    ..The more highly phosphorylated forms of PRR5 and TOC1 interact best with the F-box protein ZTL (ZEITLUPE), suggesting a mechanism to modulate their proteolysis...
  4. Martin Tryon E, Kreps J, Harmer S. GIGANTEA acts in blue light signaling and has biochemically separable roles in circadian clock and flowering time regulation. Plant Physiol. 2007;143:473-86 pubmed
    ..Using a clock-regulated luciferase reporter, we isolated new alleles of ZEITLUPE, LATE ELONGATED HYPOCOTYL, and GIGANTEA (GI)...
  5. Somers D, Schultz T, Milnamow M, Kay S. ZEITLUPE encodes a novel clock-associated PAS protein from Arabidopsis. Cell. 2000;101:319-29 pubmed
    ..This screen identified mutations at a locus, ZEITLUPE (ZTL), that lengthen the free-running period of clock-controlled gene transcription and cell expansion, and alter ..
  6. Baudry A, Ito S, Song Y, Strait A, Kiba T, Lu S, et al. F-box proteins FKF1 and LKP2 act in concert with ZEITLUPE to control Arabidopsis clock progression. Plant Cell. 2010;22:606-22 pubmed publisher
    Regulation of protein turnover mediated by ZEITLUPE (ZTL) constitutes an important mechanism of the circadian clock in Arabidopsis thaliana...
  7. Mas P, Kim W, Somers D, Kay S. Targeted degradation of TOC1 by ZTL modulates circadian function in Arabidopsis thaliana. Nature. 2003;426:567-70 pubmed
    ..Here we provide genetic and molecular evidence for a role of ZEITLUPE (ZTL) in the targeted degradation of TIMING OF CAB EXPRESSION 1 (TOC1) in Arabidopsis thaliana (thale cress)...
  8. Kevei E, Gyula P, Hall A, Kozma Bognar L, Kim W, Eriksson M, et al. Forward genetic analysis of the circadian clock separates the multiple functions of ZEITLUPE. Plant Physiol. 2006;140:933-45 pubmed
    ..of these loops remain to be identified; none of the known components is completely understood, including ZEITLUPE (ZTL), a gene implicated in regulated protein degradation...
  9. Kiba T, Henriques R, Sakakibara H, Chua N. Targeted degradation of PSEUDO-RESPONSE REGULATOR5 by an SCFZTL complex regulates clock function and photomorphogenesis in Arabidopsis thaliana. Plant Cell. 2007;19:2516-30 pubmed
    ..We demonstrated that the Pseudo-Receiver (PR) domain of PRR5 interacts directly with the F box protein ZEITLUPE (ZTL) in vitro and in vivo...

More Information


  1. Para A, Farré E, Imaizumi T, Pruneda Paz J, Harmon F, Kay S. PRR3 Is a vascular regulator of TOC1 stability in the Arabidopsis circadian clock. Plant Cell. 2007;19:3462-73 pubmed
    ..PRR3 was able to bind to TOC1 in yeast and in plants and to perturb TOC1 interaction with ZEITLUPE (ZTL), which targets TOC1 for proteasome-dependent degradation...
  2. Díaz Camino C, Risseeuw E, Liu E, Crosby W. A high-throughput system for two-hybrid screening based on growth curve analysis in microtiter plates. Anal Biochem. 2003;316:171-4 pubmed
  3. Wang L, Fujiwara S, Somers D. PRR5 regulates phosphorylation, nuclear import and subnuclear localization of TOC1 in the Arabidopsis circadian clock. EMBO J. 2010;29:1903-15 pubmed publisher
    ..g. timeless/period and clock/cycle), suggesting evolutionary convergence of a conserved mechanism across kingdoms. ..
  4. Han L, Mason M, Risseeuw E, Crosby W, Somers D. Formation of an SCF(ZTL) complex is required for proper regulation of circadian timing. Plant J. 2004;40:291-301 pubmed
    ..In Arabidopsis a novel F-box protein, ZEITLUPE (ZTL), plays an important role in the control of the free-running period of the circadian clock...
  5. Fukamatsu Y, Mitsui S, Yasuhara M, Tokioka Y, Ihara N, Fujita S, et al. Identification of LOV KELCH PROTEIN2 (LKP2)-interacting factors that can recruit LKP2 to nuclear bodies. Plant Cell Physiol. 2005;46:1340-9 pubmed
    ..COL1 and other CO/COL family proteins could also interact with LKP1/ZTL, LKP2 or FKF1. The LKP2-binding site in CO or COL1 was near the center of each protein...
  6. Miyazaki Y, Takase T, Kiyosue T. ZEITLUPE positively regulates hypocotyl elongation at warm temperature under light in Arabidopsis thaliana. Plant Signal Behav. 2015;10:e998540 pubmed publisher
    ..Proteins of the flavin-binding KELCH repeat F-box 1 / LOV KELCH protein 2 / ZEITLUPE family are positive regulators of hypocotyl elongation under red light in Arabidopsis...
  7. Yasuhara M, Mitsui S, Hirano H, Takanabe R, Tokioka Y, Ihara N, et al. Identification of ASK and clock-associated proteins as molecular partners of LKP2 (LOV kelch protein 2) in Arabidopsis. J Exp Bot. 2004;55:2015-27 pubmed
    ..LKP2 interacted not only with itself but also with other members of the family, LKP1 and FKF1...
  8. Cha J, Kim J, Kim T, Zeng Q, Wang L, Lee S, et al. GIGANTEA is a co-chaperone which facilitates maturation of ZEITLUPE in the Arabidopsis circadian clock. Nat Commun. 2017;8:3 pubmed publisher
    ..Through an unknown mechanism GIGANTEA stabilizes the F-box protein ZEITLUPE, a key regulator of the circadian clock...
  9. Takase T, Nishiyama Y, Tanihigashi H, Ogura Y, Miyazaki Y, Yamada Y, et al. LOV KELCH PROTEIN2 and ZEITLUPE repress Arabidopsis photoperiodic flowering under non-inductive conditions, dependent on FLAVIN-BINDING KELCH REPEAT F-BOX1. Plant J. 2011;67:608-21 pubmed publisher
    LOV KELCH PROTEIN2 (LKP2), ZEITLUPE (ZTL)/LOV KELCH PROTEIN1 (LKP1) and FLAVIN-BINDING KELCH REPEAT F-BOX1 (FKF1) constitute a family of Arabidopsis F-box proteins that regulate the circadian clock...
  10. Song Y, Estrada D, Johnson R, Kim S, Lee S, MacCoss M, et al. Distinct roles of FKF1, Gigantea, and Zeitlupe proteins in the regulation of Constans stability in Arabidopsis photoperiodic flowering. Proc Natl Acad Sci U S A. 2014;111:17672-7 pubmed publisher
    ..Here we demonstrated that the FKF1-interacting proteins Gigantea (GI) and Zeitlupe (ZTL) are involved in CO stability regulation...
  11. Jarillo J, Capel J, Tang R, Yang H, Alonso J, Ecker J, et al. An Arabidopsis circadian clock component interacts with both CRY1 and phyB. Nature. 2001;410:487-90 pubmed
    ..Here we describe ADAGIO1 (ADO1), a gene of Arabidopsis thaliana that encodes a protein containing a PAS domain...
  12. Kiyosue T, Wada M. LKP1 (LOV kelch protein 1): a factor involved in the regulation of flowering time in arabidopsis. Plant J. 2000;23:807-15 pubmed
    ..Here, we report a putative photo-regulatory factor sequence in LKP1 (LOV kelch protein 1)...
  13. Zhang B, Wang L, Zeng L, Zhang C, Ma H. Arabidopsis TOE proteins convey a photoperiodic signal to antagonize CONSTANS and regulate flowering time. Genes Dev. 2015;29:975-87 pubmed publisher
    ..In addition, TOE1 interacts with the LOV domain of FKF1 and likely interferes with the FKF1-CO interaction, resulting in partial degradation of the CO protein in the afternoon to prevent premature flowering. ..
  14. Johansson M, McWatters H, Bako L, Takata N, Gyula P, Hall A, et al. Partners in time: EARLY BIRD associates with ZEITLUPE and regulates the speed of the Arabidopsis clock. Plant Physiol. 2011;155:2108-22 pubmed publisher
    ..We show that EBI associates with ZEITLUPE (ZTL), known to act in the plant clock as a posttranslational mediator of protein degradation...
  15. Kim J, Geng R, Gallenstein R, Somers D. The F-box protein ZEITLUPE controls stability and nucleocytoplasmic partitioning of GIGANTEA. Development. 2013;140:4060-9 pubmed publisher
    ..Previous work has shown that the F-box protein ZEITLUPE (ZTL) and clock element GIGANTEA (GI) heterodimerize in the cytosol, thereby stabilizing ZTL...
  16. Kim T, Kim W, Fujiwara S, Kim J, Cha J, Park J, et al. HSP90 functions in the circadian clock through stabilization of the client F-box protein ZEITLUPE. Proc Natl Acad Sci U S A. 2011;108:16843-8 pubmed publisher
    ..Here we show that the Arabidopsis circadian clock-associated F-box protein ZEITLUPE (ZTL) is a unique client for cytoplasmic HSP90...
  17. Pudasaini A, Zoltowski B. Zeitlupe senses blue-light fluence to mediate circadian timing in Arabidopsis thaliana. Biochemistry. 2013;52:7150-8 pubmed publisher
    ..The Zeitlupe (ZTL), Flavin-Kelch-Fbox-1 (FKF1), and LOV-Kelch-Protein-2 (LKP2) proteins dictate measurement of the day length, ..
  18. Liu H, Wang Q, Liu Y, Zhao X, Imaizumi T, Somers D, et al. Arabidopsis CRY2 and ZTL mediate blue-light regulation of the transcription factor CIB1 by distinct mechanisms. Proc Natl Acad Sci U S A. 2013;110:17582-7 pubmed publisher
    ..Instead, two of the three light-oxygen-voltage (LOV)-domain photoreceptors, ZEITLUPE and LOV KELCH PROTEIN 2, but not FLAVIN-BINDING KELCH REPEAT 1, are required for the function and blue-light ..
  19. Pudasaini A, Shim J, Song Y, Shi H, Kiba T, Somers D, et al. Kinetics of the LOV domain of ZEITLUPE determine its circadian function in Arabidopsis. elife. 2017;6: pubmed publisher
    A LOV (Light, Oxygen, or Voltage) domain containing blue-light photoreceptor ZEITLUPE (ZTL) directs circadian timing by degrading clock proteins in plants...
  20. Saitoh A, Takase T, Kitaki H, Miyazaki Y, Kiyosue T. Gene expression profile of zeitlupe/lov kelch protein1 T-DNA insertion mutants in Arabidopsis thaliana: Downregulation of auxin-inducible genes in hypocotyls. Plant Signal Behav. 2015;10:e1071752 pubmed publisher
    ..b>ZEITLUPE (ZTL) or LOV KELCH PROTEIN1 (LKP1) is a positive regulator of warmth-induced hypocotyl elongation under white ..