Jul 21, 2024  
2002-2003 Graduate Catalog 
2002-2003 Graduate Catalog [ARCHIVED CATALOG]


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The Department of Biological Sciences offers graduate study leading to the following M.S. and Ph.D. degrees. Independent research under the direction of a faculty advisor is a major component of all of these programs (excepting certain of the Masters degrees which have a non-thesis or literature-research option).

Doctor of Philosophy

  • Biological Sciences
  • Ecology and Environmental Sciences
  • Plant Science
  • Zoology

Master of Science

  • Botany and Plant Pathology
  • Ecology and Environmental Sciences
  • Entomology
  • Zoology

Research Specializations

Graduate-degree candidates conduct research under the guidance of the Biological Sciences faculty. The expertise of the faculty covers a broad spectrum, ranging from molecular and cell biology, through system- and organism-level biology, to ecology; and it applies to a diversity of organisms from protists and lower plants and invertebrate animals through vascular plants and vertebrates. By choosing a faculty advisor, graduate applicants can associate themselves with any of a number of research specializations:

Animal Behavior and Behavioral Ecology, including chronobiology, feeding behavior, foraging, host plant selection, reproductive behavior, behavior and endocrinology of birds, migration and predator-prey interactions.

Applied Biology, including biochemistry and biology of wood decay, biological control and insect pest management, fisheries, plant pathology, and toxicology.

Botany, Plant Biology, Mycology, including plant and fungal systematics, molecular and morphological phylogeny, reproductive biology, quantitative morphology, plant ecology, marine algal ecology, plant paleoecology, plant physiology, physiology and ecology of nitrogen fixation, microscopy of zoosporic fungi, mycology, and physiology and molecular biology of fungal pathogens.

Developmental and Cell Biology, including cell biology and physiology of bone cells, developmental genetics, embryology, and neurobiology.

Ecology and Environmental Biology, including aquatic, community, insect, plant, and wetland ecology; biogeochemistry; biodiversity; conservation biology; limnology; population dynamics; population modeling; and Quaternary paleoecology.

Entomology, including insect ecology and biodiversity, insect pathology, biological control and insect pest management, ecology of aquatic insects, forest entomology, and predator- prey interactions, pollination ecology, computer simulation of insect population dynamics.

Fisheries Biology, including ecology and behavior of fishes, toxicology, fish population ecology, ecology of intertidal fishes, salmonid biology, and aquaculture.

Freshwater Biology, including toxicology, ecology and behavior of fishes, stream and river ecology, paleolimnology, and wetland biology.

Genetics and Molecular Biology, including behavioral genetics, molecular systematics, pathogen-plant interactions, and plant virology.

Plant Pathology, including biochemistry and biology of wood decay, and pest management.

Physiology and Physiological Ecology, including environmental physiology of marine invertebrates, fungal physiology, insect-plant interactions, pathogen-plant interactions, nitrogen fixation, endocrine physiology and systemic physiology.

Systematics and Evolution, including microevolution, phylogenetics of plants, fungi, and invertebrates, comparative morphology, plant anatomy.

Special Options

The department is also associated with the Institute for Quaternary and Climate Studies with which students may arrange cooperative programs of study.

Students of genetics may choose, as an option, study in a Ph.D. program on mammalian genetics offered in cooperation with The Jackson Laboratory. Thesis work may be conducted at The Jackson Laboratory; the doctorate is awarded by the University.

Training in applied fishery science is provided through the Maine Cooperative Fish and Wildlife Research Unit, operated at the University under an agreement among the University, the Biological Resources Division of the U. S. Geological Survey, the Wildlife Management Institute, and the Maine Department of Inland Fisheries and Wildlife. Also, the Migratory Fish Research Institute supports basic research on fishes.


Key to the Department’s research efforts are several facilities providing equipment, space and professional personnel. Among equipment available for graduate-student use, for example, are automated DNA-sequencing equipment, laser confocal and electron microscopes, digital imaging equipment, gas liquid chromatographs, scintillation counters and controlled-environment chambers. Aquatic laboratories for raising fishes and invertebrates, greenhouses, The University of Maine herbarium, an on-campus arboretum, and numerous sites for field research on both managed and natural habitats in marine, freshwater, and terrestrial ecosystems are easily accessible. Sites managed by the Maine Agricultural and Forest Experiment Station include the Blueberry Hill Research Farm in Jonesboro, the Organic Blueberry Research Site in Whitneyville, the Aroostook Potato Research Farm in Presque Isle, The Rogers Sustainable Agriculture Research Farm in Stillwater, and the Demeritt and Penobscot Experimental Forests in Orono and Bradley. Marine research facilities are available through the University’s Ira C. Darling Center at Walpole, Maine; through the Huntsman Marine Science Center at St. Andrews, New Brunswick, Canada; and through the Mount Desert Island Biological Laboratory at Salsbury Cove, Maine.


Applicants are encouraged to identify an area of research interest and a potential advisor at the time of application; they should feel free to contact members of the faculty to discuss possible research projects even before submission of the application. A research project is a central part of both the M.S. and Ph.D. degrees.

All applicants will be automatically considered for teaching or research assistantships. Many students are supported by research grants to individual faculty members; interested students should contact faculty members directly for further information on grant-supported assistantships.
Additional information is available from the Graduate Coordinator, Department of Biological Sciences, 5751 Murray Hall, Orono, ME 04469-5751, (207) 581-2540, E-mail: umbiosci@maine.edu, www.umesci.maine.edu/biology/.

Graduate Faculty

Christopher S. Campbell, Ph.D. (Harvard, 1980), Professor and Chair. Plant systematics, molecular phylogeny, reproductive biology, and quantitative morphology.

Susan J. Hunter, Ph.D. (Pennsylvania State University, 1980), Associate Professor and Associate Chair. Cell biology and physiology of bone cells and developing extracellular matrix; role of growth factors in skeletal physiology.

A. Randall Alford, Ph.D. (Louisiana State, 1980), Professor. Insect physiology, behavior, and chemical ecology.

Andrei Alyokhin, Ph.D. (University of Massachusetts, Amherst, 1999), Assistant Professor. Insect behavior and ecology, integrated pest management, biological control.

Seanna L. Annis, Ph.D. (University of Guelph, 1995), Assistant Professor. Physiological, molecular, and field studies of fungal pathogens of plants and animals.

Christopher S. Cronan, Ph.D. (Dartmouth College, 1978), Professor. Biogeochemistry; plant ecology; ecosystem ecology.

Ronald B. Davis, Ph.D. (Cornell, 1961), Professor. Limnology and wetland/peatland ecology; late Quaternary paleoecology, including paleolimnology, pollen, and diatom analysis.

Harold B. Dowse, Ph.D. (New York University, 1971), Professor. Genetic and molecular analysis of cardiac pacemakers, biology of oscillating systems, mathematical analysis and modeling of biological systems.

Francis A. Drummond, Ph.D. (University of Rhode Island, 1986), Professor. Insect quantitative ecology, pest management, population dynamics, simulation modeling, biostatistics, and pollination ecology.

William E. Glanz, Ph.D. (University of California, Berkeley, 1977), Associate Professor. Community and foraging ecology and social behavior of mammals and birds; evolution and biogeography of North and South American rodents.

Eleanor Groden, Ph.D. (Michigan State University, 1989), Associate Professor. Insect ecology, insect pathology, biological control.

Terry A. Haines, Ph.D. (Michigan State University, 1971), Professor. Aquatic toxicology and environmental effects of pollutants.

Rebecca Holberton, Ph.D. (State University of New York, Albany, 1991), Associate Professor. The endocrine basis of bird ecology and behavior; reproductive biology, bird migration and conservation.

Alexander D. Huryn, Ph.D. (University of Georgia, 1986), Associate Professor. Aquatic insects; stream and river ecology.

George L. Jacobson, Jr., Ph.D. (University of Minnesota, 1975), Professor. Plant ecology and paleoecology; long-term history and dynamics of vegetation and climate in the western hemisphere.

Jody J. Jellison, Ph.D. (Oregon State University, 1983), Professor. Immunochemical characterization of wood decay; metal metabolism in fungi.

Michael T. Kinnison, Ph.D. (University of Washington, 1999), Assistant Professor. Microevolution, aquatic ecology, population and conservation genetics, fish ecology (including salmonids).

John R. Moring, Ph.D. (University of Washington, 1973), Professor. Fish population ecology; salmonid biology and aquaculture; intertidal ecology; logging problems.

John M. Ringo, Ph.D. (University of California, Davis, 1973), Professor. Genetic analysis of behavior; cardiac rhythms in Drosophila; animal signaling and reproductive isolation; evolution of behavior.

Christa R. Schwintzer, Ph.D. (University of Michigan, 1969), Professor. Physiological ecology with emphasis on nitrogen fixation and wetland plants.

Stylianos M. Tavantzis, Ph.D. (Pennsylvania State University, 1980), Professor. Molecular genetics of virulence in plant pathogenic fungi; biological control of fungal plant pathogens.

John D. Tjepkema, Ph.D. (University of Michigan, 1971), Professor. Whole-plant physiology; physiology and ecology of nitrogen fixation.

Mary S. Tyler, Ph.D. (University of North Carolina, 1975), Professor. Developmental biology; organogenesis in vertebrates; morphogenesis in Drosophila; educational multimedia materials.

Seth Tyler, Ph.D. (University of North Carolina, 1975), Professor. Invertebrate biology; electron and fluorescence microscopy; phylogeny of lower invertebrates, especially meiofauna.

Robert L. Vadas, Ph.D. (University of Washington, 1968), Professor. Marine and algal ecology; recruitment; plant-herbivore and predator-prey interactions.

Katherine E. Webster, Ph.D. (University of Wisconsin-Madison, 1998), Assistant Professor. Ecology of lakes and management of aquatic ecosystems.

Stephen A. Woods, Ph.D. (University of Massachusetts, 1989), Associate Professor. Forest entomology; insect ecology, and biodiversity.


Research and Associate Faculty

Young-mee Ahn, Ph.D. (Illinois, Urbana, 1996), Associate Research Assistant Professor. Mycology.

Melvin S. Fuller, Ph.D. (California, Berkeley, 1959), Associate Professor. Biology of freshwater and marine fungi.

Joyce E. Longcore, Ph.D. (Univeristy of Maine, 1991), Research Assistant Professor. Chytridio-mycete systematics and phylogeny; chytrid pathogen of amphibians.

Constance Stubbs, Ph.D. (University of Maine, 1992), Research Assistant Scientist. Pollination ecology, lichenology, mycology.


Cooperating Faculty

Susan H. Brawley, Ph.D. (California, Berkeley, 1978), Professor of Marine Sciences and Cooperating Professor of Biological Sciences. Developmental biology and physiology of fertilization; marine ecology; environmental control of reproduction.

William O. Bray, Ph.D. (Missouri, 1981), Professor of Mathematics and Cooperating Professor of Biological Sciences. Classical analysis, harmonic analysis.

Alfred Bushway, Ph.D. (Purdue , 1978), Professor of Food Science and Cooperating Professor of Biological Sciences. Natural products metabolism.

Rodney J. Bushway, Ph.D. (Texas A & M , 1977), Professor of Food Science and Cooperating Professor of Biological Sciences. Pesticide detection, natural toxicants.

Michael S. Greenwood, Ph.D. (Yale, 1969), Professor of Forest Resources and Cooperating Professor of Biological Sciences. Tree physiology and improvement.

Keith W. Hutchison, Ph.D. (Wisconsin-Madison, 1974), Professor of Biochemistry and Cooperating Professor of Biological Sciences. Molecular genetics of growth and development, genome flux.

Richard Jagels, Ph.D. (Illinois 1968), Professor of Forest Ecosystem Science and Cooperating Professor of Biological Sciences. Autecological effects on structure of woody plants.

Irving Kornfield, Ph.D. (SUNY, Stony Brook 1974), Professor of Marine Sciences and Cooperating Professor of Biological Sciences. Population biology, speciation theory, molecular systematics.

David H. Lambert, Ph.D. (Pennsylvania State , 1979), Associate Professor of Plant Pathology and Cooperating Associate Professor of Biological Sciences. The molecular biology of host-pathogen interactions, diseases of horticultural crops.

Alan R. Langille, Ph.D. (Pennsylvania State , 1967), Professor of Agronomy and Cooperating Professor of Biological Sciences. Hormonal regulation of potato tuberization, protoplast regeneration.

William H. Livingston, Ph.D. (Minnesota, 1985), Associate Professor of Forest Pathology and Cooperating Associate Professor of Biological Sciences. Disease, ectomycorrhizal, and ethylene effects on growth of conifers.

James D. McCleave, Ph.D. (Montana State, 1967), Professor of Marine Sciences and Cooperating Professor of Biological Sciences. Migratory and transport mechanisms of fishes; fisheries oceanography.

Kathleen Murray, Ph.D. (Massachusetts, 1990), State Entomologist and Cooperating Assistant Professor of Biological Sciences. Agricultural pest ecology, plant-insect interactions, use of natural products in pest management.

Paul Rawson, Ph.D. (South Carolina, 1996), Assistant Professor of Marine Sciences and Cooperating Assistant Professor of Biological Sciences. Quantitative and population genetics of marine invertebrates, molecular evolution.

J. Malcolm Shick, Ph.D. (Texas A&M University, 1974), Professor. Ecological physiology of marine invertebrates; biology of symbiotic cnidarians; UV photobiology and oxidative stress.

Bruce D. Sidell, Ph.D. (Illinois, 1975), Professor of Marine Sciences and Cooperating Professor of Biological Sciences. Physiology and biochemistry of fishes, mechanisms of metabolic compensation to environmental variables in fishes, physiology of Antarctic fish species.

Robert S. Steneck, Ph.D. (Johns Hopkins, 1983), Professor of Marine Sciences and Cooperating Professor of Biological Sciences. Ecology and evolutionary biology of benthic marine algae, invertebrates and communities. An emphasis on crustose coralline algae, herbivores and lobsters.

Rebecca J. Van Beneden, Ph.D. (The Johns Hopkins, 1983), Professor, Biochemistry and Marine Sciences, Cooperating Professor of Biological Sciences. Environmental toxicology: molecular mechanisms of carcinogenesis, comparative carcinogenesis, aquatic toxicology.

Michael E. Vayda, Ph.D. (Princeton, 1983), Professor of Biochemistry and Cooperating Professor of Biological Sciences. Plant stress responses, translational control, gene regulatory mechanisms, low-oxygen stress, cold adaptation.

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