Harmer Lab:Stacey Harmer

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I am a professor in the Department of Plant Biology at the University of California, Davis. My lab studies the molecular basis of circadian rhythms and how these daily rhythms affect plant physiology.

Contact Info

Mailing Address
Stacey L. Harmer
Department of Plant Biology, LS 1002
University of California, Davis
One Shields Ave
Davis, CA 95616
2129 Life Sciences
(530) 752-8101 (voice)
(530) 752-5410 (fax)
2123 Life Sciences
(530) 752-8102
slharmer at ucdavis.edu


  • 1998-2002 Postdoctoral Scholar, The Scripps Research Institute. Advisor: Steve Kay.
  • 1998 Ph.D. University of California, San Francisco. Advisor: Tony DeFranco.
  • 1991 B. A. University of California, Berkeley.


One of the great things about living in Davis is that we are close to the Sierra Nevada. For photos of some of my adventures there and elsewhere, please visit this simple photo gallery. Another wonderful thing about Davis is the twice-weekly Farmer's Market.


  • Atamian, H.S., Creux, N.M., Brown, E.A., Garner, A.G., Blackman, B.K., and Harmer, S.L. (2016) Circadian regulation of sunflower heliotropism, floral orientation, and pollinator visits. Science. 353(6299):587-90. doi: 10.1126/science.aaf9793 [1]
  • Atamian, H.S. and Harmer, S.L. (2016) Circadian regulation of hormone signaling and plant physiology. Plant Mol Biol. 91(6):691-702. doi: 10.1007/s11103-016-0477-4 [2]
  • Jones, M.A., Hu, W., Litthauer, S., Lagarias, J.C. and Harmer, S.L. (2015) A constitutively active allele of phytochrome B maintains circadian robustness in the absence of light. Plant Physiology. 169(1):814-25. doi: 10.1104/pp.15.00782 [3]
  • Brady, S.M., Burow, M., Busch, W., Carlborg, O., Denby, K.J., Glazebrook, J., Hamilton, E.S., Harmer, S.L., Haswell, E.S., Maloof, J.N., Springer, N.M., Kliebenstein, D.J. (2015) Reassess the t Test: Interact with All Your Data via ANOVA. Plant Cell. 2015 Jul 28. pii: tpc.15.00238 [4]
  • Vandenbrink, J.P., Brown, E.A., Harmer, S.L., Blackman, B.K. (2014) Turning heads: the biology of solar tracking in sunflower. Plant Sci. 224:20-6. doi: 10.1016/j.plantsci.2014.04.006 [5]
  • Anver, S., Roguev, A., Zofall, M., Krogan, N.J., Grewal, S.I.S., and Harmer, S.L. (2014) Yeast X-Chromosome Associated Protein 5 (Xap5) Functions with H2A.Z to Suppress Aberrant Transcripts. EMBO reports. pii: e201438902. [6]
  • Hsu, P.Y. and Harmer, S.L. (2014) Global profiling of the circadian transcriptome using microarrays. Methods in Molecular Biology. 1158:45-56 [7]
  • Hsu, P.Y. and Harmer, S.L. (2013) Wheels within wheels: the plant circadian system. Trends in Plant Science [8]
  • Hsu, P.Y., Devisetty, U.K., and Harmer, S.L. (2013) Cycling activators control the evening arm of the plant clock. eLife. 2:e00473. [9]
  • Zemach, A., Kim, M.Y., Hsieh, P.-H., Coleman-Derr,D., Eshed-Williams, L., Thao, K, Harmer, S.L., and Zilberman, D. (2013) The Arabidopsis nucleosome remodeler DDM1 allows DNA methyltransferases to access H1-containing heterochromatin. Cell 153(1):193-205 [10]
  • Hu, W., Franklin, K.A., Sharrock, R.A., Jones, M.A., Harmer, S.L., and Lagarias, J.C. (2013) Unanticipated regulatory roles for Arabidopsis phytochromes revealed by null mutant analysis. Proceedings of the National Academy of the Sciences, 110(4) 1542-1547. [11]
  • Hsu, P.Y. and Harmer, S.L. (2012) Circadian phase has profound effects on differential expression analysis. PLoS One, 7(11):e49853 [12]
  • Jones, M.A., Williams, B.A., McNicol J., Simpson, C.G., Brown, J.W., and Harmer, S.L. (2012) Mutation of Arabidopsis SPLICEOSOMAL TIMEKEEPER LOCUS 1 causes circadian clock defects. Plant Cell, 24(10):4066-82 [13]
  • Nozue, K., Harmer. S.L., and Maloof, J.N. (2011) Genomic analysis of circadian clock-, light-, and growth-correlated genes reveals PIF5 as a modulator of auxin signaling in Arabidopsis. Plant Physiology, 156(1): 357-372 [14]
  • Rawat, R., Takahashi, N., Hsu, P.Y., Jones, M.A., Schwartz, J., Salemi, M.R., Phinney, B.S., and Harmer, S.L. (2011) REVEILLE8 and PSEUDO-REPONSE REGULATOR5 form a negative feedback loop within the Arabidopsis circadian clock. PLoS Genetics, 7(3): e1001350. [15]
  • Kerwin, R.E., Jimenez-Gomez, J.M., Fulop, D., Harmer, S.L., Maloof, J.N., Kliebenstein, D.J. (2011) Network QTL mapping of circadian clock outputs identifies metabolic pathway to clock linkages in Arabidopsis. Plant Cell, 23(2):471-85 [16]
  • Jones, M.A., and Harmer, S.L. (2011) JMJD5 functions in concert with TOC1 in the Arabidopsis circadian system. Plant Signaling & Behavior 6(3): 1-4. [17]
  • Jones, M.A., Covington, M.F., Ditacchio, L., Vollmers, C., Panda, S., Harmer, S.L. (2010) Jumonji domain protein JMJD5 functions in both the plant and human circadian systems. Proceedings of the National Academy of the Sciences 107(50): 21623-21628. [18]
  • Ellison ,C.T., Vandenbussche, F., Van Der Straeten, D., Harmer, S.L. (2011) XAP5 CIRCADIAN TIMEKEEPER regulates ethylene responses in aerial tissues of Arabidopsis. Plant Physiology, 155(2):988-999. [19]
  • Harmer, S.L. (2010). Plant biology in the fourth dimension. Plant Physiology, 154:467 - 470. [20]
  • Rawat, R., Schwartz, J., Jones, M.A., Sairanen, I., Cheng, Y., Andersson, C.R., Zhao, Y., Ljung, K., and Harmer, S.L. (2009). REVEILLE1, a Myb-like transcription factor, integrates the circadian clock and auxin pathways. Proceedings of the National Academy of the Sciences 106(39) 16883-16888. [21]
  • Harmer, S.L. (2009). The circadian system in higher plants. Annual Review of Plant Biology 60: 357 – 77. [22]
  • Covington, M.F., Maloof, J.N., Straume, M., Kay, S.A., and Harmer, S.L. (2008). Global transcriptome analysis reveals circadian regulation of key pathways in plant growth and development. Genome Biology, 9(8):R130. [23]
  • Martin-Tryon, E.L., and Harmer, S.L. (2008). XAP5 CIRCADIAN TIMEKEEPER coordinates light signals for proper timing of photomorphogenesis and the circadian clock in Arabidopsis. Plant Cell 20(5):1244-59. [24]
  • Gong, W., He, K., Covington, M.F., Dinesh-Kumar, S.P., Snyder, M., Harmer, S.L., Zhu, X., and Deng, X.P. (2008). The development of protein microarrays and their applications in DNA-protein and protein-protein interaction analyses of Arabidopsis transcription factors. Molecular Plant, 1(1): 27 – 41. [25]
  • Covington, M.F. and Harmer, S.L (2007) The circadian clock regulates auxin signaling and responses in Arabidopsis. PLoS Biology, 5(8): e227 [26]
  • Nozue, K., Covington, M.F., Duek, P.D., Lorrain, S., Fankhauser, C., Harmer, S.L., and Maloof, J.N. (2007) Rhythmic growth explained by coincidence between internal and external cues. Nature, 448:358-61. [27]
  • Walley, J.W., Coughlan, S., Hudson, M.E., Covington, M.F., Kaspi, R., Banu, G., Harmer, S.L., and Dehesh, K. (2007) Mechanical stress induces biotic and abiotic stress responses via a novel cis-element. PLoS Genetics, 3(10): e172. [28]
  • Oliverio, K.A., Crepy, M., Martin-Tryon, E.L., Milich, R., Harmer, S.L., Putterill, J., Yanovsky, M., Casal, J.J. (2007) GIGANTEA regulates phytochrome A-mediated photomorphogenesis independently of its role in the circadian clock. Plant Physiology 144:495-502 [29]
  • Martin-Tryon, E.L., Kreps, J.A., Harmer, S.L. (2007). GIGANTEA acts in blue light signaling and has biochemically separable roles in circadian clock and flowering time regulation. Plant Physiology 143:473-486. [30]
  • Harmer, S. L. and Kay, S. A. (2005). Positive and negative factors confer phase-specific circadian regulation of transcription in Arabidopsis. Plant Cell 17:1926-1940. [31]
  • Farre, E. M., Harmer, S. L., Harmon, F. G., Yanovsky, M. J., and Kay, S. A. (2005). Overlapping and distinct roles of PRR7 and PRR9 in the Arabidopsis circadian clock. Current Biology 15:47-54. [32]
  • Alabadi, D., Mas, P., Yanovsky, M., Harmer, S. L., and Kay, S. A. (2002). Critical role for CCA1 and LHY in maintaining circadian rhythmicity in Arabidopsis. Current Biology 12:757-761. [33]
  • Harmer, S. L., Panda, S., and Kay, S. A. (2001). Molecular bases of circadian rhythms. Annual Review of Cell and Developmental Biology 17:215-253. [34]
  • Harmer, S. L., Hogenesch, J. B., Straume, M., Chang, H. S., Han, B., Zhu, T., Wang, X., Kreps, J. A., and Kay, S. A. (2000). Orchestrated transcription of key pathways in Arabidopsis by the circadian clock. Science 290:2110-2113. [35]
  • Harmer, S.L., and DeFranco, A.L. (1999). The src homology domain 2-containing inositol phosphatase SHIP forms a ternary complex with Shc and Grb2 in antigen receptor-stimulated B lymphocytes. The Journal of Biological Chemistry 274:12183-12191. [36]
  • Harmer, S.L., and DeFranco, A.L. (1997). Shc contains two Grb2 binding sites needed for efficient formation of complexes with SOS in B lymphocytes. Molecular and Cellular Biology 17:4087-4095. [37]
  • Biemann, H. P., Harmer, S.L., and Koshland, D. E. Jr. (1996). An aspartate/insulin receptor chimera mitogenically activates fibroblasts. The Journal of Biological Chemistry 271: 27927-27930. [38]
  • Crowley, M. T., Harmer, S. L., and DeFranco, A.L. (1996). Activation-induced association of a 145-kDa tyrosine-phosphorylated protein with Shc and Syk in B lymphocytes and macrophages. The Journal of Biological Chemistry 271: 1145-1152. [39]
  • Bradfisch, G. A., and Harmer, S. L. (1990). Ω-conotoxin GVIA and nifedipine inhibit the depolarizing action of the fungal metabolite, destruxin B on muscle from the tobacco budworm (Heliothis virescens). Toxicon 28:1249-1254 [40]