THE 2011 BULLETIN EDITORIAL COMMITTEE Editor Dr. J.B. Claiborne Managing Editor Michael P. McKernan Dr. J.B. Claiborne, Chair Dr. Elizabeth Crockett Dr. David H. Evans Dr. Raymond Henry Dr. Karl Karnaky Dr. David Miller Dr. Antonio Planchart Dr. Robert L. Preston Dr. Alice Villalobos Published by the Mount Desert Island Biological Laboratory June 2011 $10.00 THE BULLETIN VOLUME 50, 2011 Mount Desert Island Biological Laboratory Salisbury Cove, Maine 04672 TABLE OF CONTENTS Introduction ii Memorial - Raymond Rappaport, Ph.D. iv Memorial - David W. Towle, Ph.D. vii Report Titles x Reports 1-90 Officers and Trustees 92 Scientific Personnel 95 Summer Fellowship Recipients 102 Seminars, Workshops, Conferences, Courses 105 Publications 120 Author Index 122 Species Index 124 Keyword Index 125 Funding Index 126 THE MOUNT DESERT ISLAND BIOLOGICAL LABORATORY CONNECTING SCIENCE, ENVIRONMENT, AND HEALTH INTRODUCTION The Mount Desert Island Biological Laboratory (MDIBL) is an independent, non-profit biological and biomedical research facility and international center for comparative physiology, environmental health sciences, marine functional genomics, and studies of regeneration. The Laboratory is located on the north shore of Mount Desert Island, overlooking the gulf of Maine about 120 miles northeast of the Portland near the mouth of the Bay of Fundy. The island, well known for Acadia National Park, provides a variety of habitats including shallow and deep saltwater, a broad intertidal zone, saltwater and freshwater marshes, freshwater lakes and streams, forests and meadows. The Laboratory is among the oldest cold-water research facilities in the Eastern United States, and its unique site provides an outstanding environment for studying the physiology of marine and freshwater flora and fauna. During 2010, the scientific personnel included 74 doctoral level scientists (including 60 Investigators), plus 111 students, fellows, and technical staff, representing 79 institutions in 23 US states, Canada, Czech Republic, Germany, Portugal, the Netherlands, and the United Kingdom. HISTORY AND ORGANIZATION MDIBL was founded in 1898 at South Harpswell, Maine by J.S. Kingsley of Tufts University. The Wild Gardens of Acadia donated its present site at Salisbury Cove, and relocation was completed in 1921. The Wild Gardens of Acadia, a land-holding group headed by George B. Dorr and John D. Rockefeller, Jr., who was instrumental in the founding of Acadia National Park. In 1914, the Laboratory was incorporated under the laws of the State of Maine as a non-profit scientific and educational institution. Founded as a teaching laboratory, MDIBL is now a center for marine research and education that attracts investigators and students from across the U.S. and around the world. Since the pioneering work of H.W. Smith, E.K. Marshall and Roy P. Forster on various aspects of renal and osmoregulatory physiology of local fauna, the Laboratory has become known worldwide as a center for investigations in electrolyte and transport physiology, developmental biology, electrophysiology and marine molecular biology. The Mount Desert Island Biological Laboratory is owned and operated by the Board of Trustees and Members of the Corporation; at present, there are 265 members. Officers of the Corporation - Chair, Vice-Chair, Director, Secretary, Treasurer, Clerk - and an Executive Committee are elected from among the Trustees. The Chair and Executive Committee oversee and promote long-range goals of the Laboratory. The Director, with the aid of a full-time Administrative Director, staff and an external Board of Scientific Advisors is responsible for implementing the scientific, educational and public service activities of the Laboratory. APPLICATIONS AND FELLOWSHIPS Research space is available for the entire summer season (June 1 - September 30) or for shorter research visits during this time frame. Applications for the coming summer must be submitted by early January each year. Investigators are invited to use the year-round facilities at other times of the year, but such plans should include prior consultation with the MDIBL office concerning available facilities and specimen supply. A number of fellowships and scholarships are available to research scientists, undergraduate faculty and students, and high school students. These funds may be used to cover the cost of laboratory rent, housing and supplies. Stipends are granted with many of the student awards. Applicants for fellowships for the coming summer research period are generally due in early January. For further information on research fellowships, please contact: Dr. Patricia H. Hand Administrative Director Mount Desert Island Biological Laboratory P.O. Box 35 Salisbury Cove, Maine 04672 Tel. (207) 288-3605 Fax. (207) 288-2130 [email protected] Students should contact: Michael McKernan Director of Education and Conferences [email protected] ACKNOWLEDGEMENTS The Mount Desert Island Biological Laboratory is indebted to the National Institutes of Health and National Science Foundation and for substantial support. Funds for building renovations and new construction continue to permit the Laboratory to expand and upgrade its research and teaching facilities. Individual research projects served by the Laboratory are funded by private and government agencies, and all of these projects have benefited from the NSF and NIH grants to the Laboratory. For supporting our educational initiative, MDIBL acknowledges the National Science Foundation Research Experience for Undergraduates, National Institute of Environmental Health Sciences Short Term Educational Experience in Research program, Maine IDeA Network for Biomedical Research Excellence (NCRR/NIH), Cserr/Grass Foundation, Milbury Fellowship Fund, Northeast Affiliate of the American Heart Association, Cystic Fibrosis Foundation, Blum/Halsey Fellowship, Stanley Bradley Fund, Stan and Judy Fund, Adrian Hogben Fund, Bodil Schmidt-Nielsen Fellowship Fund, Maine Community Foundation, the Hearst Foundation, the Betterment Fund and many local businesses and individuals. Ray Rappaport with his sand dollar collecting scoop at MDIBL in 2000 (left) and a first division sea urchin embryo undergoing cytokinesis (right) labeled for the microtubules of the mitotic apparatus (top) and the activated myosin (bottom) of the furrow-generating contractile ring. Raymond Rappaport, Ph.D. 1922 – 2010 “That form ever follows function.”- Louis Sullivan. One of the founders of modern American architecture, Louis Sullivan, was famous for emphasizing the crucial nature of the connection between form and function. Raymond Rappaport, an accomplished amateur architect in his own right, built a distinguished career employing this foundational principle in the study of cytokinesis, the final stage of cell division where the two daughter cells physically separate. Ray performed elegant experiments employing hand-fashioned tools to alter the geometry of dividing sand dollar embryos in order to gain fundamental insights into the basic relationship between form and function underpinning cytokinesis. Ray grew up in North Bergen, New Jersey and enrolled in Bethany College in West Virginia. His education there was interrupted by service during World War II in both the Army Anti-Aircraft and Medical Corps. He finished his undergraduate studies at Columbia University and then went on to complete a masters degree at the University of Michigan in 1948. While at Michigan he met and married his life partner Barbara Nolan who was also enrolled in a masters degree program. From Michigan he went on to attend Yale University where he earned a Ph.D. in 1952. That same year he was hired as an assistant professor at Union College, a small liberal arts institution in Schenectady, New York where he spent 35 years teaching and doing research. Most summers since 1948 Ray and Barbara spent from May to September at the Mount Desert Island Biological Laboratory. In his over 60 years of affiliation with MDIBL Ray served in a number of important capacities including Director, President of the Corporation, member of the Board of Trustees, and as a full-time Senior Scientist following his retirement from Union in 1987. He also contributed to the facilities of the MDIBL through the architectural design of several MDIBL cottages and laboratory buildings as well as the cooperative dining hall. In the field of cell biology his work came to be identified with the MDIBL in the same way that in the field of nephrology the MDIBL is identified with the work of Homer Smith and E.K. Marshall. Over a career spanning five decades Ray studied cytokinesis mainly in the large and clear embryos of the sand dollar Echinarachnius parma. Each summer season Ray and Barbara would scoop up animals from a rowboat in Emory Cove and place them in a sea water table. Ray would then sex these animals by injecting just a segment with the spawning inducer potassium chloride and in this way separate males and females into different portions of the sea water table. These animals would then be partially shed throughout the season and ultimately returned to Emory Cove in the early autumn. His operations in the lab reflected the deep sense of sustainability evident in his use of animals. It was difficult to find any plastic item in Ray’s lab as everything, from watch glasses to petri dishes to test tube to flasks, was made of glass and therefore reusable. It was also difficult to find paper towels as Ray instead had dozens of cloth towels – as he often would say it