Paul G. Greenwood

Professor of Biology and Dr. Charles C. and Pamela W. Leighton Research Fellow; Senior Associate Provost and Dean of Faculty, 2011-2016

Box 4770

Phone: 207-859-5732
Fax: 207-859-5705
Mailing Address:
4770 Mayflower Hill
Waterville, Maine 04901-8847
Greenwood, Paul G.


B.A. Knox College 1980 M.S. Florida State University 1983 Ph.D. Florida State University 1987

Areas of Expertise

  • Cell biology
  • Invertebrate zoology
  • Cnidarians and nematocysts

Current Research

Our research centers around the development, discharge physiology, and biochemistry of nematocysts, the stinging structures common to jellyfish, sea anemones, corals, and related organisms. One project is the isolation and characterization of a unique calcium-binding protein that is a component of the nematocyst venom. Another project is an investigation of how predators of cnidarians protect themselves chemically and physically from nematocyst discharge. A third project uses molecular markers to study intraspecific and interspecific variation in several populations of the sea anemone Metridium.


  • Greenwood, P. G. 2009. Acquisition and use of nematocysts by cnidarian predators. Toxicon 54: 1065-1070.
  • LaRiviere, F. J., A. G. Newman,* M. L. Watts,* S. Q. Bradley,* J. E. Juskewitch,* P. G. Greenwood, J. T. Millard. 2009. Quantitative PCR analysis of diepoxybutane and epihalohydrin damage to nuclear versus mitochondrial DNA. Mutation Research 664: 48-54.
  • Greenwood, P. G., K. Garry,* A. Hunter* and M. Jennings*. 2004. Adaptable defense: a nudibranch mucus inhibits nematocyst discharge and changes with prey type. Biological Bulletin 206: 113-120.
  • Greenwood, P. G., I. M. Balboni* and C. Lohmann*. 2003. A sea anemone's environment affects discharge of its isolated nematocysts. Comparative Biochemistry and Physiology 134A: 275-281.
  • Greenwood, P. G. and J. Johnson* . 1999(abstract). The effects of neurotransmitters on nematocyst discharge in the sea anemone Metridium senile. Mol. Biol. Cell 10: 229a.
  • Greenwood, P. G. and M. Yunes* . 1997. How calcium is held in nematocysts: molecular modeling of poly g-glutamic acids and detection of a specific calcium binding protein in nematocysts of Metridium senile Linnaeus, 1761. In (den Hartog, J.P., ed) The Proceedings of the 6th International Conference on Coelenterate Biology, Museum of Natural History, Leiden, The Netherlands. pp. 209-214.
  • Greenwood, P. G. and L. K. Garrity* . 1991. Discharge of nematocysts isolated from aeolid nudibranchs. Hydrobiologia 216/217: 671-677.
  • Greenwood, P. G., L. A. Johnson* and R. N. Mariscal. 1989. Depletion of ATP in suspensions of isolated cnidae: A possible role of ATP in the maturation and maintenance of anthozoan cnidae. Comparative Biochemistry and Physiology 93A: 761-765.

    * denotes Colby student

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