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).
- Biological Sciences
- Ecology and Environmental Sciences
- Plant Science
- Botany and Plant Pathology
- Ecology and Environmental Sciences
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
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
Physiology and Physiological Ecology, including environmental physiology of
marine invertebrates, fungal physiology, insect-plant interactions,
pathogen-plant interactions, nitrogen fixation, endocrine physiology and
Systematics and Evolution, including microevolution, phylogenetics of plants,
fungi, and invertebrates, comparative morphology, plant anatomy.
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
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
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,
Christopher S. Campbell, Ph.D. (Harvard, 1980), Professor and Chair.
Plant systematics, molecular phylogeny, reproductive biology, and quantitative
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
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,
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
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;
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
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
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.
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.
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
William O. Bray, Ph.D. (Missouri, 1981), Professor of Mathematics and
Cooperating Professor of Biological Sciences. Classical analysis, harmonic
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,
Michael S. Greenwood, Ph.D. (Yale, 1969), Professor of Forest
Resources and Cooperating Professor of Biological Sciences. Tree physiology and
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
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
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