Department of Biology

CLAY J. CARTER, Associate Professor


B.S., Biotechnology, 1997, Minnesota State University, Mankato
Ph.D. Biochemistry, 2002, Iowa State University

Office: 252C SSB
Phone: 218-726-7347



PLANT MOLECULAR AND CELLULAR BIOLOGY. I am primarily interested in the biochemistry of plant nectars and the molecular basis of nectar secretion. Other areas of interest include plant defense against pathogens and membrane trafficking in plants.

Selected Recent Publications

Lin I, Chen L-Q, Sosso D, Gase K, Kim S-G, Kessler D, Klinkenberg PM, Gorder MK, Qu X-Q, Hou B-H, Carter CJ, Baldwin IT, Frommer WB (in revision) Nectar secretion requires sucrose phosphate synthases and the sugar transporter SWEET9, Nature

Bender RL, Fekete ML, Klinkenberg PM, Hampton MW, Bauer B, Malecha M, Lindgren K, Maki J, Perera MADN, Nikolau BJ, Carter CJ (2013) PIN6 is required for nectary auxin response and short stamen development, Plant Journal, 74: 893-904

Bender R, Klinkenberg P, Jiang Z, Bauer B, Karypis G, Nguyen N, Perera MADN, Nikolau BJ, Carter CJ (2012) Functional genomics of nectar production in the Brassicaceae. FLORA, 7: 491-496

Xu, W.W., Carter, C. (2010) Parallel multiplicity and error discovery rate (EDR) in microarray experiments. BMC Bioinformatics, 11: 465

Hampton, M., Xu, W.W., Kram, B.W., Chambers, E., Ehrnriter, J., Gralewski, J.H., Joyal, T., Carter, C.J. (2010) Identification of differential gene expression in Brassica rapa nectaries through expressed sequence tag analysis. PLoS ONE, 5: e8782. (undergraduate researchers underlined)

Agee A.E., Surpin, M., Sohn, E.J., Girke, T., Rey, A.R., Kram, B.W., Carter, C., Wentzell, A.M., Kliebenstein, D.J., Jin, H.C., Park, O.K., Jin, H., Hicks, G.R., Raikhel, N. (2010) MODIFIED VACUOLE PHENOTYPE1 is an Arabidopsis myrosinase-associated protein involved in endomembrane protein trafficking. Plant Physiology, 152: 120-32

Ruhlmann, J.M., Kram, B.W., Carter, C.J. (2010) CELL WALL INVERTASE 4 is required for nectar production in Arabidopsis. Journal of Experimental Botany, 61: 395-404

Kram, B.W., Carter, C.J. (2009) Arabidopsis thaliana as a model for functional nectary analysis. Sexual Plant Reproduction, 22: 235-246

Kram, B.W.*, Xu, W.W.*, Carter, C.J. (2009) Uncovering the Arabidopsis thaliana nectary transcriptome: investigation of differential gene expression in floral nectariferous tissues. BMC Plant Biol 9: 92 (*equal contribution)

Kram, B.W., Bainbridge, E.A., Perera, M.A.D., Carter C. (2008) Identification, cloning and characterization of a GDSL lipase secreted into the nectar of Jacaranda mimosifolia. Plant Mol Biol 68: 173-183. (undergraduate researcher underlined)

Sohn, E.J., Rojas-Pierce, M., Pan, S., Carter, C., Serrano-Mislata, A., Madueño, F., Rojo, E., Surpin, M., Raikhel, N.V. (2007) The shoot meristem identity gene TFL1 is involved in flower development and trafficking to the protein storage vacuole. Proc Natl Acad Sci USA 104: 18801-18806.

Robert, S., Zouhar, J., Carter, C., Raikhel, N. (2007) Isolation of intact vacuoles from Arabidopsis rosette leaf-derived protoplasts. Nature Protoc 2: 259-262.

Carter, C., Healy, R., O’Tool, N.M., Naqvi, S.M., Ren, G., Park, S., Beattie, G.A., Horner, H.T., Thornburg, R.W. (2007) Tobacco nectaries express a novel NADPH oxidase that is implicated in the defense of floral reproductive tissues against microorganisms. Plant Physiol 143: 389-399.

Carter, C., Shafir, S., Vaknin, L., Palmer, R.G., Thornburg R. (2006) A novel role for proline in plant floral nectars. Naturwissenschaften93:72-79.

Surpin, M.*, Rojas-Pierce, M.*, Carter, C.*, Hicks, G.R.*,Vasquez, J., Raikhel, N.V. (2005) The power of chemical genomics to study the link between endomembrane system components and gravitropic response. Proc Natl Acad Sci USA 102: 4902-4907. (*equal contribution)

Carter, C.J.*, Pan, S.*, Zouhar, J.*, Avila, E.L., Girke, T., Raikhel, N.V. (2004) The vegetative vacuole proteome of Arabidopsis thaliana reveals predicted and unexpected proteins. Plant Cell16: 3285-3303. (*equal contribution)

* indicates equal contribution towards manuscript


  • BIOL 2101 Cell Biology
  • BIOL 3990 Plant Cell & Molecular Biology
  • BIOL 5232 Molecular Biology Lab
  • BIOL 4235 Biotechnology

Current Graduate Students

  • Mengyuan Jia
  • David Ongaro
  • Lisa Wiesen
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