Volume 181 THE Number 1 BIOLOGICAL BULLETIN ne Biological LIBRA AUG22 1991 Woods Hole, Mass. AUGUST, 1991 Published by the Marine Biological Laboratory THE Marine Biological Laboratory ' LIBRARY AUG 2 2 1991 Woods Hole, Mass. BIOLOGICAL BULLETIN PUBLISHED BY THE MARINE BIOLOGICAL LABORATORY Associate Editors PETER A. V. ANDERSON, The Whitney Laboratory, LIniversity of Florida DAVID EPEL, Hopkins Marine Station, Stanford University J MALCOLM SHICK, University of Maine, Orono Editorial Board GEORGE J. AUGUSTINE, University of Southern RUDOLF A. RAFF, Indiana University California KENSAL VAN HOLDE, Oregon State University Louis LEIBOVITZ, Marine Biological Laboratory STEVEN VOGEL, Duke University Editor: MICHAEL J. GREENBERG, The Whitney Laboratory, University of Florida Managing Editor: PAMELA L. CLAPP, Marine Biological Laboratory AUGUST, 1991 Printed and Issued by LANCASTER PRESS, Inc. PRINCE & LEMON STS. LANCASTER, PA THE BIOLOGICAL BULLETIN THE BIOLOGICAL BULLETIN is published six times a year by the Marine Biological Laboratory, MBL Street, Woods Hole, Massachusetts 02543. Subscriptions and similar matter should be addressed to Subscription Manager, THE BIOLOGICAL BUL- LETIN, Marine Biological Laboratory, Woods Hole, Massachusetts 02543. Single numbers, $25.00. Sub- scription per volume (three issues), $72.50 ($145.00 per year for six issues). Communications relative to manuscripts should be sent to Michael J. Greenberg, Editor-in-Chief, or Pamela L. Clapp, Managing Editor, at the Marine Biological Laboratory, Woods Hole, Massachusetts 02543. Telephone: (508) 548-3705, ext. 428. FAX: 508-540-6902. POSTMASTER: Send address changes to THE BIOLOGICAL BULLETIN, Marine Biological Laboratory, Woods Hole, MA 02543. Copyright (E) 1991, by the Marine Biological Laboratory Second-class postage paid at Woods Hole, MA, and additional mailing offices. ISSN 0006-3 185 INSTRUCTIONS TO AUTHORS The Biological Bulk-tin accepts outstanding original research reports of general interest to biologists throughout the world. Papers are usually of intermediate length (10-40 manuscript pages). A limited number of solicited review papers may be ac- cepted after formal review. A paper will usually appear within four months after its acceptance. Very short, especially topical papers (less than 9 manuscript pages including tables, figures, and bibliography) will be pub- lished in a separate section entitled "Research Notes." A Re- search Note in The Biological Bulletin follows the format of similar notes in Nature It should open with a summary para- graph of 150 to 200 words comprising the introduction and the conclusions. The rest of the text should continue on without subheadings, and there should be no more than 30 references. 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Comp Physiol., not J. Cell. Comp. Physiol '.) E. Unusual words in journal titles should be spelled out in full, rather than employing new abbreviations invented by the author. For example, use Ril I 'isindafjelags Islendinga with- out abbreviation. F. All single word journal titles in full (e.g. I'eliger, Ecology, Brain). G. The order of abbreviated components should be the same as the word order of the complete title (i.e. Proc. and Trans. placed where they appear, not transposed as in some BIOLOGICAL ABSTRACTS listings). H. A few well-known international journals in their pre- ferred forms rather than WORLD LIST or USASI usage (e.g. Na- ture, Science, Evolution NOT Nature, Loud., Science, N.Y.; Evolution. Lancaster, Pa.) 6. Reprints, page proofs, and charges. Authors receive their first 100 reprints (without covers) free of charge. Additional re- prints may be ordered at time of publication and normally will be delivered about two to three months after the issue date. Authors (or delegates for foreign authors) will receive page proofs of articles shortly before publication. They will be charged the current cost of printers' time for corrections to these (other than corrections of printers' or editors' errors). Other than these charges for authors' alterations. The Biological Bulletin does not have page charges. The Marine Biological Laboratory Ninety-Third Report for the Year 1990 One-Hundred and Third Year Officers of the Corporation Denis M. Robinson, Honorary Chairman of the Board of Trustees Prosser Gifford, Chairman of the Board of Trustees Harlyn O. Halvorson, President of the Corporation and Director Robert D. Manz, Treasurer Kathleen Dunlap, Clerk of the Corporation Contents . Report of the President and Director Report of the Treasurer . Financial Statements Report of the Librarian 17 Educational Programs Summer Courses . . 19 Short Courses . Summer Research Programs Principal Investigators .... Other Research Personnel ... 30 Library Readers Institutions Represented 34 Year-Round Research Programs Honors Board of Trustees and Committees Laboratory Support Staff Members of the Corporation Life Members Regular Members 54 Associate Members 64 Certificate of Organization 68 Articles of Amendment 68 Bylaws 68 Report of the President and Director I began last year's report by discussing the Laboratory's needs and reminding the Corporation of the pressing need for a new Marine Resources Center capable of providing a reliable, healthy, and genetically denned supply of marine organisms. I closed that discussion of the long-awaited, many-times-planned MRC with the hope that this important facility would become a reality "before too many more director's reports are filed." I begin this director's report, my fourth, the Laboratory's ninety-third (covering its one-hundred third year) with the very good news that the new MRC will indeed be a reality very soon. As I write this report (in the spring of 1991), the building is going up outside my window. Where we used to have a carpenters' shop and a parking lot, we now have pilings on top of which the walls of a state-of-the-art facility for holding marine animals are rising. The key step toward the MRC in 1990 was the passage, in October, of a Federal appropriation bill that included $4.75 million toward the MRC. Coupled with earlier Federal support, this brought funding for the MRC and related projects to a total of $8. 95 million. Part of the planning for the new MRC involved making alternate plans for parking and a re-location of the carpenters' shop. The plan at the close of 1990 is to build a new carpenters' shop on the MBL campus next to the Broderick House and to put a new, off-site parking lot in the MBL Woods off Oyster Pond Road near Memorial Circle. In the next director's report, I fully expect to be able to report that these auxiliary projects are completed and that the new MRC is up and very close to going on-line. While preparations for construction of the MRC occupied much of our attention in 1990, we simultaneously continued to plan for an Advanced Studies Laboratory (ASL), which together with the MRC will constitute the Marine Biomedical Institute for Advanced Studies. At the close of 1990, the new ASL was in the second of three design phases. The construction of the new Marine Resources Center and the progress toward a new Advanced Studies Laboratory are important steps toward ensuring a bright future for the Laboratory, but other long- standing and well-documented needs remain. To address these additional needs, the Trustees at their June 22 meeting initiated a long-range development program. The Executive Committee approved the use of a development consultant to help us review our readiness for mounting a major fund raising campaign. The consultant. Browning Associates, presented a cogent analysis of the Laboratory, which will help us greatly expand our development operation in 1991 and beyond. Research At the urging of the External Scientific Advisory Committee, we established in 1990 a Scientific Council to function in an advisory capacity to the Executive Committee. The Council is charged with guiding the Laboratory in: the development of scientific and educational programs the use of scientific resources the evaluation and promotion of scientific staff the recruitment of new scientific positions the initiation of institutional grant proposals. The council is to work in conjunction with existing committees, such as the Research Space Committee and the Instruction Committee. Council members are appointed by the director, with the approval of the Executive Committee. The council is to include up to three members of the year-round 2 Annual Report Aerial view of the MBL's Marine Resources Center. Photo by Robert Colder. scientific community, up to three members of the summer community, two non-MBL scientists, the MBL director who serves e.\ officio. and an c.\ officio executive secretary. The first council is composed of: year-round MBL scientists Drs. John Hobbie (vice- chairman), Mitchell Sogin, and Felix Strumwasser; summer MBL scientists Drs. Barbara Ehrlich, Gerald Fischbach, and Joseph Sanger, Dr. Holger Jannasch. Woods Hole Oceanographic Institution, and Dr. Howard Hiatt, Brigham and Women's Hospital. Dr. Leslie Garrick serves as ex officio executive secretary, and, as director of the MBL, I serve as ex officio chairman of the council. The council met throughout 1990 to evaluate scientists for appointment and promotion and to set directions for future scientific development and expansion. They completed a draft of one position paper on Laboratory directions in cellular and developmental biology. The council plans to prepare additional position papers on environmental sciences/ ecology, microbiology, molecular evolution, neurobiology, and plant sciences. These documents are targeted for completion in 1991 and 1992. Instruction The instruction program continued in 1990 with its unanimously acclaimed courses. One new short course was added: Pathogenesis of Neuroimmunologic Diseases, co-directed b\ J. Murdoch Ritchie, Yale LJniversity, and Byron H. Waksman, Harvard University and New York University. At the urging of the External Scientific Advisory Committee, we have been taking a careful look at the cost of the courses. The Instruction Committee looked at cost containment in 1990, and, while there is as yet no consensus on how to proceed, it is clear that we will have to do something about spiraling cost increases in this age of decreasing federal support for advanced training. New fellowships in 1990 included the Nikon Fellowship: a Bernard Davis Fellowship for studies in microbiology or molecular evolution; the Daniel S. Grosch Scholarship Fund for studies in environmental toxicology; and the Porter Fellowships for Minority Students/Investigators for work in physiology. A list of fellowship recipients appears later in this issue. Library The Library made continued progress toward applying computing technology to library services. As described in the Report of the Librarian, Jane Fessenden, the MBL Librarian of 29 years, retired in 1990. In December, Dr. David Stonehill accepted the position of Director of the MBL/WHOI Library and Scientific Information Research Center. Dr. Stonehill has been a national leader in the development of modern information services, having worked for Report of the President and Director 3 Jelle Atema, the new director of the Boston University Marine Program. Photo by Judith Anderson. NASA, a number of leading universities, and most recently the President of the United States. Governance In August, Dr. Prosser GifTord announced his intention to step down from the chairmanship of the MBL Board of Trustees. Dr. Gifford led the MBL for 14 years, through a period of unprecedented change and growth. In a little short of a decade and a half, he worked with 4 of the 1 1 directors the Laboratory has had in its 103 year existence. Under his leadership, the Laboratory's facilities were significantly upgraded, the year-round science program grew substantially, the educational program maintained its character, the library prospered and began an exciting modernization program, and the administrative and development staffs were strengthened. Neurobiologist Dr. Jelle Atema assumed the directorship of the Boston University Marine Program in August. He replaces Dr. Rudi Strickler. who had directed BUMP since 1987. Dr. Atema has energetic plans for BUMP's graduate and undergraduate programs, and his long acquaintance with the MBL bodes well for our partnership with Boston University. In their February meeting, the Trustees approved a new Long Range Financial Planning Committee charged with reporting to the Trustees and/or the Executive Committee on the existence and appropriateness of long-range financial planning mechanisms of the laboratory. Responsibility for planning remains with the administration of the laboratory, while the new committee is to assure that the financial risks of growth have been properly anticipated and planned for. The committee is to include two at-large trustees, two corporate trustees, and the treasurer, e.\ officio. Treasurer Robert Manz chaired the new committee, which began by making a comprehensive review of the new MRC. In August, the Long Range Financial Planning Committee gave the Trustees a favorable report on the financial planning for the MRC. That favorable report was an important part of the briefing that led to the Trustees' decision to proceed toward construction of the new MRC. Personnel A group of MBL employees, including service, maintenance, and clerical workers, voted in February to affiliate with the Hospital Workers Union, Local 767. Collective bargaining negotiations began almost immediately after the election, and with the union and management both negotiating in a spirit of cooperation and mutual respect, we were able to negotiate our first contract in an impressively short time. A fair, reasonable, and workable contract was signed in July, and the work of the Laboratory proceeded without disruption. Long-time manager of marine resources John Valois retired in November after 40 years at the MBL. In addition to running the collecting operation, Mr. Valois has served for years as a spokesman for marine biology and has appeared regularly in newspapers and on television extolling the virtues of marine animals as models for research. On his departure, Edward Enos was promoted to Superintendent of Marine Resources. Mr. Enos has been a collector for many years and was eminently well-prepared to take the reins of the department. MBI, Science Writing Fellows in the Ilands-On Laboratory Course. Photo by James Hrynyshyn. 4 Annual Report The Biological Bulletin Under the editorship of Michael J. Greenberg, The Biological Bulletin continued to publish well-presented reports of outstanding research that is of general interest to biologists throughout the world. In 1990, Dr. Greenberg announced that all page charges would be dropped, and that authors would be offered 100 free reprints for publishing in the journal. Most importantly, he noted in his 1990 report to the corporation, a carefully prepared manuscript can now appear in print as soon as 3.6 months after its submission. In fact, some manuscripts meeting the criteria of the new Research Notes section which features brief communications of high quality and currency may appear in print even sooner. All of these initiatives have been highlighted in the journal's newsletter. The Biological Board. The newsletter, which is published "occasionally" by the editorial staff, was created to highlight and promote the articles appearing in. and the policies of, the journal. Dr. Greenberg has also been attempting to adjust the mix of papers appearing in the Bulletin so that the journal more closely reflects research here at the MBL. To assist in this effort. Dr. Greenberg has recruited three Associate Editors to aid in the review and solicitation of manuscripts. Drs. J. Malcolm Shick (University of Maine, Orono), Peter A. V. Anderson (The Whitney Laboratory), and David Epel (Hopkins Marine Station) will serve four-year terms as Associate Editors in their respective fields of physiology and metabolism, neurobiology, and developmental biology. Science Writing Fellowships In its fifth year, the MBL Science Writing Fellowships Program evolved a one-week, hands-on course in cellular and molecular biology for science writers. Co-directed by Dr. Robert Goldman, Northwestern University Medical School, and Boyce Rensberger, The Washington Post, the course began with an introduction to cells and a microscopy demonstration, and ended five days later with the writers cloning and sequencing DNA. The course, which will be expanded to include a neurobiology component in 1991, is open to all science writers and serves as an orientation for the MBL Science Writing Fellows. Directorship of the Science Writing Program has passed from founding director James Shreeve to Dr. Byron Waksman. Mr. Shreeve will continue to serve on the program's advisory board. The Tokyo String Quartet. Pholo by Christian Steiner. Public programs We continued to offer a few modest programs for our non-scientist neighbors on the Upper Cape. In July we held a public symposium on Science and Public Policy. The keynote address was delivered by Massachusetts Senator John Kerry, who urged the science community to become more involved in the very political process of forging a coherent national science policy. The symposium was followed by the second annual MBL Chamber Music Concert, featuring the Tokyo String Quartet. The first annual Falmouth Forum concluded in early 1990 with presentations by Dr. Allen Counter on Black Arctic explorer Matthew Henson, Anne Hawley on the government as patron of the arts, a musical portrait of Harry Truman by David McCullough, and a panel discussion on energy and the environment. The series was well enough received by local audiences that the MBL Associates brought it back for the winter of 1990- 9 1 . Marshall Goldman began the second Falmouth Forum series in November with a presentation on Perestroika, and in December actress Julie Harris read from Lucifer's Child, her new one-woman play. Harlyn O. Halvorson Report of the Treasurer The year 1990 was one of building for the Laboratory. Total support and revenues increased from $16.2 to $17.1 million due primarily to the initiation of construction of the Marine Resources Center, supported by a grant from the federal government; $1.4 million was received and expended on this project in 1990. Gifts received decreased approximately $1.6 million; 1989 was the last year of the receipt of significant funds from the Howard Hughes Foundation multi-year grant for the Library and the Education programs. Dining hall revenues rose $170,000 due to increases in food service functions, attendees, and meal card prices. Investment income grew $200,000 from 1989 to 1990 due to increases of long- and short-term investments and the length of time that they were available to the Laboratory during the respective years. Recovery of indirect costs related to research and instruction grew by approximately $150,000, all of which was attributable to the growth of MBL sponsored research. Other operating revenues were essentially unchanged from 1989. Total expenses grew approximately $700,000, from $15 million in 1989 to $15.7 million in 1990. The most significant increases were in Research Services and Plant Operations; the former is due primarily to the expansion of the operations of the Mass Spectrometry Laboratory, the Instrument Development Laboratory, and the Protein and Nucleic Acid Chemistry Center operated jointly with the Woods Hole Oceanographic Institution; the latter is due primarily to increased utility, insurance, and labor costs, as well as significant expenses involving the removal of asbestos and other repairs and renovations of our housing. It is also worth noting that the Laboratory was able to offer approximately $100,000 more scholarships, fellowships, and stipends in 1990 than in 1989, a 35% increase. While our aim is to provide much more, the trend is very heartening, as we have almost doubled the amount of scholarships, fellowships, and stipends from 1988. The Current Unrestricted Fund ended the year with an excess of support and revenues over expenses of $93,407. This combines the results of the Housing and Dining Auxiliary with all other Operations. The Housing and Dining Auxiliary had an excess of support and revenues over expenses of $267,761, of which $65,000 was used for scheduled repayment of debt owed on the Memorial Circle cottages and $199,863 was transferred to the Repairs and Replacements Reserve for housing. More sobering and troublesome, however, is that the operations of the Laboratory without the Housing and Dining operation experienced a deficiency of revenues to expenses of $174,354. The single greatest reason for this was our inability to meet budgeted goals for unrestricted current gifts. The balance sheet shows that we ended 1990 with a significant degree of liquidity as cash and short-term investments accounted for approximately two-thirds of current assets. Investments at market grew by approximately $400,000. Essentially all of that was attributable to gifts for endowment, as the net impact of the market on the valuation of our investments was a decline of less than $100,000. Land, buildings and equipment, net of accumulated depreciation, grew by almost $1 million due primarily to the beginning of construction of the Marine Resources Center. When completed, the cost of this project is expected to total $9 million. The operating budget for 1991 reflects a determination to maintain the current level of services at the Laboratory in the most cost-effective manner possible, while we redouble our efforts to generate more funds to support operations. Our greatest single challenge is still the need to find a cost-effective way to provide services that are primarily 6 Annual Report used during the summer months. The strategy of the Laboratory for the last 20 years has been to increase the size of the year-round program to provide a more stable financial base for the Laboratory. The Ecosystems Center has been the most notable success to date, and we trust that this will be a model for other MBL sponsored programs. The principal financial benefit of this strategy is the potential to achieve efficiencies in the cost of administration and services by spreading them across a broader base. We recognize that these will only be achieved by focusing on ways to be as efficient as possible. The principal financial cost of this strategy is the capital cost of the laboratory building and equipment to house the scientists and their science, as well as the reserve or endowment funds necessary to attract scientists. Through the federal funding to date of the MBIAS project, we are well on our way towards funding the necessary buildings. The next significant challenge will be generating the necessary program support funds. This strategy presumes the continued excellence of the summer research and education programs. Our education program shows many signs of health and vitality. Courses are oversubscribed; we have had renewed success in obtaining grants, and funding remains in place to support the overhead costs of the program for the next few years. Our single greatest challenge at the moment is the renewal of this support either through endowment or renewed medium-term funding. Our summer research program shows signs of financial stress. Applications, occupancy, and revenues have been flat to marginally declining for the last couple of years. It is clear that reductions in federal funding have placed a greater budgetary burden on scientists seeking to come to the MBL in the summer, and some have had to withdraw. The Laboratory has long subsidized the cost of a summer stay for all scientists from its discretionary resources the summer laboratory fee falls significantly short of the fully allocated cost of the space but our ability to do so is limited. We can selectively support a few scientists, principally young investigators, and we have had some success recently with the establishment of two Nikon fellowships for young summer scientists, but we need to do much more. It is clear that increased attention needs to be paid to the summer research program to continue its financial vitality. We need to attract more applicants; we need to find more funds to allow young scientists to become familiar with the MBL; and we need to be as efficient as possible in offering and delivering the services that make a stay at the MBL the most productive scientific experience possible. Financially, your Laboratory lives on the edge. On the positive side, we have continued to demonstrate our ability to attract funds from the federal government, from foundations, and from individuals. We are in the process of significantly upgrading our physical plant, to make the Laboratory an even more attractive place to do science. Our housing budget is currently generating the funds necessary to assure that we will be able to continue to maintain and upgrade those facilities. Our endowment has continued to grow. On the negative side, our endowment is not large enough to provide a financial cushion for any of our major programs with the significant exception of the Ecosystems Center. We have limited funds to recruit scientists whether for summer or year-round programs. Our support services do not cover their costs through charges for their services, and operations aside from Housing and Dining are in deficit. To move off the edge we will need to do three things: More clearly define, articulate, and communicate the MBL's special contribution to science and society Significantly improve our generation of capital and operating support through fund raising Redefine for ourselves what constitutes the most effective delivery of services in the most efficient manner. Robert D. Manz Financial Statements Coopers &Lybrand certified public accountants REPORT OF INDEPENDENT ACCOUNTANTS To the Trustees of Marine Biological Laboratory Woods Hole. Massachusetts We have audited the accompanying balance sheet of Marine Biological Laboratory as of December 3 1 , 1990 and the related statement of support, revenues, expenses and changes in fund balances for the year then ended. We previously examined and reported upon the financial statements of the Laboratory for the year ended December 31, 1989, which condensed statements are presented for comparative purposes only. These financial statements are the responsibility of the Laboratory's management. Our responsibility is to express an opinion on these financial statements based on our audit. We conducted our audit in accordance with generally accepted auditing standards. Those standards require that we plan and perform the audit to obtain reasonable assurance about whether the financial statements are free of material misstatement. An audit includes examining, on a test basis, evidence supporting the amounts and disclosures in the financial statements. An audit also includes assessing the accounting principles used and significant estimates made by management, as well as evaluating the overall financial statement presentation. We believe that our audit provides a reasonable basis for our opinion. In our opinion, the financial statements referred to above present fairly, in all material respects, the financial position of Marine Biological Laboratory at December 31, 1990, and its support, revenues, expenses and changes in fund balances for the year then ended in conformity with generally accepted accounting principles. Our audit was conducted for the purpose of forming an opinion on the basic financial statements taken as a whole. The supplemental schedules of support, revenues, expense and changes in fund balances for current funds (Schedule I), endowment funds (Schedule II) and plant funds (Schedule III) as of December 31, 1990 are presented for purposes of additional analysis and are not a required part of the basic financial statements. Such information has been subjected to the auditing procedures applied in the audit of the basic financial statements and, in our opinion, is fairly stated, in all material respects, in relation to the basic financial statements taken as a whole. Boston, Massachusetts April 19. 1991 LjQOOtno t oCu O ^~ r~~- _^ fN iO r^ OO O r^j oo >o r~- __ rO t O n VO r-~. o <^ oo q P! S r-- S 3 oo r- o oo ON OO O OO r- p- P^ ON OO rn S rn >/-> r-- o -o C-N] ri u-, O ^ ^O iy~i O <^i NO ^r TT o r\ m O oo 2 VO ri oo O I ^T O^ (N d f*"i <^| Tj- ^- oo ^^ TJ- r^i oo ^ *o ON ri r- (N r*-i " ^" -"V u-i r^ mrr -* r-i (N O "* rn O O ON OO O oo ON ^ 00 ON m oo TJ- ^ u-, ui m m fN O <*"> ^^f OO . oo ON 00 ^H r-- o ^t P-- /% VD O oo o^ SO rn ON ON iO f^J */^ Tf ^O VO ON so /"> 1 oo r-l I/-, ^J- QV DC O 'T r~- _J ON P- Cl Vi ON VO 00 O ON 00 sO \o r- P- w~ O r~ ON __< r*^ ON O) ^O p- -< O "0 SO (N r~^ I 'O >o O ^O n r-i vi ON fN ON ^ ON cn ' ^^ ' 1-' vO p^T ro U-, TT <^l 2 ON ' - t a & Z. g 8 -ic 5 " "S < r^ W5 ABORATORY ;TS 990 Is for 1989) LIABILITIES AND FUND B Current portion of long-term debt Accounts payable and accrued ex] Deferred income Total current liabilities Mortgage and notes payable (Not( Deferred support (Note K) Q h I? i >, U ^ ' G 2 - - - = ( Total liabilities Current unrestricted fund balance Endowment funds: Quasi-endowment unrestricted Endowment, income for unrest Endowment, income for restric Quasi-endowment restricted - 1 UJ " $ X ' O -t o ^T oo oo * ON O l~~ -r ~ -. ^^ ^D so "/" tO rn it O OO O r- -5 U 1J . 0^7 ,C ^^ *! c 03 O < c ta i -J ^ S -J <- O CL io NO "/"i O O "^ ;> ON VO O O rn rn rn Os OO ON 00_ OO "X O Oo' ri ^-~ 1 Oa. < oa /< c > i U o &0 CQ rj O VO Tf- rn O ON fNJ n so O oo rN UJ -C Z~ oo ON Tf oo oo r- _ ON r~ O ON OO so vo OO O /"! ""' O g "^ Os ^f "/"i "^ O */"> r> O ^ /"> Tf OO rn NO S ? S ?, ^' CM ^^ -~, rn i/"t so o> 3 c3 "O C J^ > 3 > 8 = b 2-0 A U U C 09 O urren I |H a "S E ^ g 3 23 o t> '5 3 c -* o v,j ^ H t o 'C a 5 1 "| *- a V) t/5 O ^ ' ' -C IsJJ 8 cement res 5 o 1 o i/i 73 C/3 c U ^ C C. E C t/i r3 "^ JZ "3 J U C T3 C a C/5 T3 U c _ 2 C '- 5 V) OJ 5 c. c S 2 "n C 75 c D u Oi U a: c c o C ca a c c o 00 c c' a * I vi O % b 73 ? ^ ci o -c h- t- r oo rn oo oo rn x o __ ^ sO c/l _ TD - S ^ ^^dso^-S i C SO Os O O"> oo 2 U. O^f ooo ^ r~i C?N oo oo cj oo [^- OO Os s a r I 2 ^ iX r I - "* C so' in SO so 3 r-ici T T'^-f O^r^ooOu/^Vi VI C Tt Os v! so oo * Tf 9 a- 2 o u- r-ri r*-ioc oor-r-n^^Tj- O" O CJ ol O so rn p < ^ r i ri' r-' o ri r i fN r 1 ri rl rn o oC o\ Os' U i^ CN! rsi r4 (A (U Tf ^ ^ < $ _i c DC S ^^ ~ "^ z 2 P < UJ Q ^ | o w fe > o u IT* "S u 'C 2 I w fc-J flj O- 4J O 3 u S g S s C D LU > ' J^ 7- w r: 5 a: > 0:1 H o o ^ ri -^t so ri rj ^-' so' Tt r-- p Tf Os Os' T' i' O V) Os_ r- oo r*^ r- r^ r 4 s i i OJ BC oo ^- ri so S OS rn so Cs| oo' on c | 1 o O C"1I W~i/"1d | Y~i o-i r~ j Tf ^-t 1 C 1 ^O ""> o^ so OO ri OO r-4 SO sO_ Z ^ u 1 2 lipiE!ill DC o- r*-, SO sf SO -f T S c ^ sC' r-' LU D W o "O " ^ C o O Si "5 ~ o-2 o g c '-5 i! ' ~ "S '> 3 C | a tU OJ 3 3 ^ C/l C T3 \ C t ~ a C __ QJ 4J t/5 3 s.sz O OJ c D. 3 CA "7 "*" "" _2 C3 Ofi il "3 8 H ou 'S C/l *O SS ^ P-'O;?;^-^^ ss=ss (N Tt ri !n 5 ON oo OX c r3-f~- sO f^ir~-sor'--"/~i/^Ov i/~i r**j \o O O oo in ^> t 00 oo ON 5 o^ csr~- so r^-rj-^Ov^Tr-ov - ioooor~- r*~)O r-j r-- o r- oo r^-i oo mTfi/->i/^ Sv o' oo TI-' NO OX in ox m' in \f\ -t ON ^f i/~i . ^~ i r | ^ -i (N r\ ^D O ox n ^ Ox 1^ i^ -^ c ^1 ON' n o> ri ON' r- o' Ov ON NO ON < T3 *** -J c tf> *C 3 Q c o ri - 0. ro ~ ON O Z -2 ^J ri O O r i r CC m j^^ Tf P Q- ON (N NO **H c ' oo' ' ^ ^ Ox' c/3 O fN r I ^ u D 60 Qi G o O g; ~ ri ? m ^ r- vo r- oo Ox r] >j tj _' ^ +-> ON m o oo_ I 5s D rr "' *~ -rl \ T^-' ' ON' f^ ^ Ox ON^ Ox' O ~7 *~ e \ ~ oo ri w~) ON in . CQ ^ o <2 3 8 m "* 3- '* 'O Ox_ **- c/) "c ^~ a: fxj r i J m c c3 r~- r- &i ' Z H 2 ~ C Q .5 u 00 r?: C ~ rN Ox O U "2 '1 I 1 o oo in NO ON 1^ O 00 Ox o , "O c X t c g ^ J H4 (U Q. QQ U 1 ON' ' ~ t^ oo ON oo NO r ri O D u E c <*) 5 g 8 <*> 3 w > L"J= u 7 w > ~ * i fy* * ^ -rt f" 1 ^D *^~i OO sO ~" 'T s . OO i ^ Ox OO r- ^ 5 w M> So S ?! SS^o o >c P * NO NO OO >/"i OV fS S ^ A3 'C O* oo' so' r [ r-* so' trT ' r [ O r- ON' ' ON to s s 1 (/I I-n r! ^ S 5 Ox r i rxi 1 CL, ^ c v5 cd c fe 1 | ^ nrestricted Oo' O' Ov oo' rn Ov oo' Tf i^i c tn Ox r~l C' C (N oo ON' ** O cd a. H _j 60 T3 c CO 5> o u n w o t -> 21 II M Is llill lllilF 2<^ o-QO O c c l oi 'a -o J< C ! HZ a E U.H 10 Notes (o Financial Statements 1 1 Marine Biological Laboratory Notes to Financial Statements A. Pnrpi isc ( if the Labi >ruti iry The purpose of Marine Biological Laboratory (the "Laboratory") is to establish and maintain a laboratory or station for scientific study and investigations, and a school for instruction in biology and natural history. B. Significant accounting policies: Basis of presentation -fund accounting In order to ensure observance of limitations and restrictions placed on the use of resources available to the Laboratory, the accounts of the Laboratory' are maintained in accordance with the principles of fund accounting. This is the procedure by which resources are classified into separate funds in accordance with specified activities or objectives. Separate accounts are maintained for each fund; however, in the accompanying financial statements, funds that have similar characteristics have been combined into fund groups. Accordingly, all financial transactions have been recorded and reported by fund group. Externally restricted funds may only be utilized in accordance with the purposes established by the donor or grantor of such funds. However, the Laboratory retains full control over the utilization of unrestricted funds. Restricted gifts, grants, and other restricted resources are accounted for in the appropriate restricted funds. Restricted current funds are reported as revenue as the related costs are incurred (see Note K). Endowment funds are subject to restrictions requiring that the principal be invested in perpetuity with income available for use for restricted or unrestricted purposes by the Laboratory. Quasi-endowment funds have been established by the Laboratory for the same purposes as endowment funds: however, the principal of these funds may be expended for various restricted and unrestricted purposes. Fixed assets Fixed assets are recorded at cost. Depreciation is computed using the straight-line method over estimated useful lives of fixed assets. Contracts and grants Revenues associated with contracts and grants are recognized in the statement of support, revenues, expenses and changes in fund balances as the related costs are incurred (see Note K). The Laboratory reimbursement of indirect costs relating to government contracts and grants is based on negotiated indirect cost rates with adjustments for actual indirect costs in future years. Any over or underrecovery of indirect costs is recognized through future adjustments of indirect cost rates. Investments purchased by the Laboratory are carried at market value. Money market securities are earned at cost which approximates market value. Investments donated to the Laboratory' are carried at fair market value at the date of the gift. For determination of gain or loss upon disposal of investments, cost is determined based on the average cost method. The Laboratory is the beneficiary of certain endowment investments reported in the financial statements which are held in trust by others. Every ten years the Laboratory's status as beneficiary of these funds is reviewed to determine that the Laboratory's use of these funds is in accordance with the intent of the funds. The market values of these investments are $4, 125.093 and $4.039,803 at December 31. 1990, and 1989, respectively. Investment income ami distribution The Laboratory follows the accrual basis of accounting except that investment income is recorded on a cash basis. The difference between such basis and the accrual basis does not have a material effect on the determination of investment income earned on a year-to-year basis. Investment income includes income from the investments of specific funds and from the pooled investment account. Income from the pooled investment account is distributed to the participating funds on the market value unit basis (Note L). Annuities payable Amounts due to donors in connection with gift annuities is determined based on remainder value calculations which generally assure a rate of return at 10%, maximum payout terms of eighteen years, and interest payout rate of 8%. C. Land, buildings and equipment: The following is a summary of the unrestricted plant fund assets: 1990 1989 Land $ 689,660 $ 689,660 Buildings 16,955,015 16,926,715 Equipment 2,819.202 2,672,838 Construction in progress 1.969.713 580.598 22.433,590 20,869,811 Less accumulated depreciation (9.459.548) (8.858.047) $12,974,042 $12,011,764 12 Annual Report D. Retirement plan The Laboratory participates in the defined contribution pension program of the Teachers Insurance and Annuity Association College Retirement Equities Fund. Contributions amounted to $451,665 in 1990 and $393,422 in 1989. E. Restricted pledges and grants. As of December 31. 1990. the Laboratory reported active pledge and grant commitments outstanding of $492,939 (unaudited) to be received. The restricted pledges are not included in the financial statements since it is not practicable to estimate the net realizable value of such pledges. Restricted pledges of $467,337, and $14,151 and $1 1,451 are scheduled to be paid in 1991, 1992, and 1993, respectively. F. Intcrfiind borrowings: Current fund interfund balances at December 31 are as follows: Due to restricted endowment fund Due to restricted quasi-endowment funds 1990 $(4,750) (1.650) $(6.4001 1 989 $(2,190) (200) $(2,390) G. Mortgage and notes payable: Long-term debt at December 31, 1990 amounted to $1,265,000. The aggregate amount of redemption due for each of the next five fiscal years is as follows: 1991 1992 1993 1994 1995 Thereafter Less current portion Amount $ 65.000 60.000 60.000 60,000 60,000 960.000 1,265,000 65.000 $1.200.000 In 1989. the Laboratory issued $1,330,000 Massachusetts Industrial Finance Authority (MIFA) Series 1989 Bonds, which pay varying annual interest rates and mature on October 31, 201 1. The Series 1989 bonds are collateralized by a first mortgage on certain Laboratory property. The interest rate is adjustable and was 7.25% and 6.5% at December 31, 1990 and 1989. In compliance with the Laboratory's MIFA bond indenture, deposits with Shawmut Bank N.A., as trustee, represent investments in the debt service reserve fund of $143.006 in 1990 and $133.000 in 1989. H. Investments: The following is a summary of the cost and market value of investments at December 31. 1990 and 1989 and the related investment income and distribution of investment income for the years ended December 31. 1990 and 1989. Cost Market Investment Income Endowment ami Quasi-Endowmenl U.S. Government securities Corporate fixed income Common stocks Money market securities Real estate Total Less custodian and management fees Total Restricted Current Fund Certificates of deposits Money market securities Total Total investments 1990 1989 $ 1.934,834 6.663,376 4,284,165 590,735 343.247 $ 2.595.407 5.900,736 3,392,001 595,467 345,749 13,816.357 12,829,360 13.816,357 12,829.360 502,360 1.500.000 490,263 965.000 2,002,360 1.455.263 $15,818,717 $14.284.623 1990 1989 $ 1.957.512 6,809,264 6,346,778 590.735 343,247 $ 2.607.537 6.032.642 5,901,724 593.544 345.749 16,047.536 15,481.196 16.047,536 15,481,196 502,360 1.500.000 490,263 965,000 2.002.360 1,455,263 $18.049.896 $16.936.459 1990 $ 214.300 506,490 234,690 121,787 1,077,267 (55.592) 1.021.675 20.312 99.367 119.679 $1.141,354 1989 $ 1 34,394 363,439 196,452 87,994 782,279 (49.318) 732.961 34,785 I 75.205 209,990 $942,951 Notes to Financial Statements 13 I. Gift support for instruction: Current year unrestricted gifts includes $500,000 of gifts for the support of the Laboratory's instruction program available for indirect costs attributable to the instruction program. J. Tax-exempt status: The Laboratory is exempt from federal income tax under Section 501(c)3 of the Internal Revenue Code. K. Restricted current funds deterred support: The Laboratory defers recognition of revenue on current restricted funds until the related costs are incurred. Amounts received in excess of expenses are recorded as deferred support. The following summanzes the activity in the deferred support account for 1990 and 1989, respectively: 1990 Balance at beginning of year Additions: Gifts, endowment income and grants received Unrealized gains Realized gain Deductions: Funds expended under gifts and grants Transfers Balance at end of vear $4,450,222 8,499,360 19,868 1,649 7,977,187 219.294 $4,774.618 $2.951,662 9,382,212 30,672 7,542.909 371.415 $4,450.222 L. Accounting for pooled investments: The major portion of investment assets is pooled for investment purposes with each participating fund subscribing to, or disposing of. units at market value at the beginning of the current quarter. The unit participation of the funds at December 31, 1990 and 1989, respectively, is as follows: Endowment and similar funds: Quasi-unrestricted Quasi-restricted Restricted endowment 1990 3,909 7,436 39.401 50,746 1989 3.887 7.416 36.488 47.791 Pooled investment activity on a per-unit basis was as follows: Unit value at beginning of year Unit value at end of year Increase (decrease) in realized and unreal- ized appreciation Net income earned on pooled investments Total return on pooled investments $109.57 108.90 (.67) 5.99 $ 5.32 $100.00 109.57 9.57 5.73 $ 15.30 Investment income is distributed to individual funds as earned. SCHEDULE I MARINE BIOLOGICAL LABORATORY STATEMENT OF SUPPORT, REVENUES, EXPENSES AND CHANGES IN FUND BALANCES CURRENT FUNDS for the year ended December 31,1 990 SUPPORT AND REVENUES: Grant reimbursement of direct costs Recovery of indirect costs related to research and instruction programs Tuition Support activities: Dormitories Dining hall Library Biological Bulletin Research services Marine resources Investment income Gifts Change in deferred support Miscellaneous revenue Total support and revenues EXPENSES: Instruction Research Scholarships, fellowships, and stipends Support activities: Dormitories Dining hall Library Biological Bulletin Research services Marine resources Administration Sponsored projects administration Plant operations Other Total expenses Excess (deficit) of support and revenues over expenses Unrealized gain on investments Realized gain on investments Total gain on investments TRANSFERS AMONG FUNDS: Debt service Acquisition of fixed assets To unrestricted plant fund Housing transfer To support operations Instruction Capitalize ecosystems income Other Total transfers among funds Net change in fund balance Fund balances, beginning of year Fund balances, end of year Operating Fund $3.136.240 273.475 172,085 491,642 149,555 466,931 4.689.928 634,428 634.428 114.516 5.438,872 Auxiliary Enterprises Fund 579,898 141.118 658.115 443.676 1.867.738 278,221 1 .644.460 5.613.226 (174.3541 (21,653) 2.898 1 90,000 (5,045) 6.642 172.842 (1.512) 21.030 19,518 906,292 802,447 1,708,739 1,708,739 638,531 690,644 111,803 1.440.978 267.761 (65,000) (199.863) (2.898) (267,761) Total Current I 'nres-tricted Fund $3.136.240 906.292 802,447 273.475 172,085 491,642 149,555 466.931 6.398.667 634,428 634,428 114.516 7.147,611 638,531 690,644 579,898 141.118 658.1 15 443,676 1.979.541 278,221 1.644.460 7.054.204 93,407 (65,000) (21,653) (196,965) 190.000 (5.045) 3.744 (94.919) 11.512) 21.030 $ 19.518 Current Restricted Fund $4,872,529 484,749 236.366 674.423 6.268.067 1,979,547 (324.396) 1.655.151 224.740 8.147,958 1,370,282 5,228,030 356,253 172,398 347.996 6.791 35,014 1,009 432.408 7.950.181 197.777 19,868 1.649 21,517 (144,881) 135.209 5,045 (215,764) 1.097 (219.294) Total $4,872,529 3,136,240 4S4.749 906,292 802.447 273.475 172.085 728,008 149,555 1.141.354 12.666.734 2.613,975 (324,396) 339.256 15,295,569 1,370.282 5,228.030 356,253 638,531 690,644 752,296 141,118 1.006.1 1 1 450.467 2,014,555 278.221 1.645,469 432.408 15.004.385 291.184 19,868 1.649 21,517 (65,000) (144,881) (21,653) (196,965) 325.209 (215.764) 4.841 (314.213) (1.512) 21.030 $ 19,518 MARINE BIOLOGICAL LABORATORY STATEMENT OF SUPPORT, REVENUES, EXPENSES AND CHANGES IN FUND BALANCES ENDOWMENT FUNDS for the year ended December 31,1 990 SCHEDULE II Restricted Unrestricted Qiiasi- Endowmenl Endowment. Income for Unrestricted Purposes Endowment. Income for Restricted Purposes Quasi- Endowment Total Restricted SUPPORT AND REVENUES: Gifts Total support and revenues Excess of support and revenues over expenses Realized gains on investments Unrealized gain (lossl on investments Total gain (loss) on investments TRANSFERS AMONG FUNDS: Capitalize ecosystems income Endowment transfers Other transfers Total transfers among funds Net change in fund balances Fund balances, beginning of year Fund balances, end of vear $ 312.522 $ 2,450 $ 114,972 $ 314.972 312.522 2.450 314.972 314,972 312.522 2.450 314,972 314,972 9,488 $ 165,441 121,543 194.377 481,361 490,849 (11.451) (110.737) (113,465) (204.872) (429.074) (440,525) (1.963) 54.704 8.078 (10,495) 52,287 50.324 215.764 215.764 215.764 (190,000) (135,209) (135.209) (325.209) 2.260 1.921 (2.158) 12.277 12.040 14.300 (187,740) 1.921 (2.158) 92.832 92.595 (95.145) (189.703) 56.625 318.442 84,787 459,854 270.151 426.982 3.234.878 4.804.006 4.480.887 12.519.771 12.946.753 $237.279 $3.291.503 $5.122.448 $4,565.674 $12,979,625 $13,216,904 15 SCHEDULE III MARINE BIOLOGICAL LABORATORY STATEMENT OF SUPPORT, REVENUES, EXPENSES AND CHANGES IN FUND BALANCES PLANT FUNDS for the year ended December 31, 1990 SUPPORT AND REVENUES: Grant for capital additions Total support and revenues EXPENSES: Depreciation Plant operations Other Total expenses Excess (deficit) of support and revenues over expenses TRANSFERS AMONG FUNDS: Debt service Acquisition of fixed assets Transfers to unrestricted plant fund Housing transfers Other transfers Total transfers among funds Net change in fund balances Fund balances, beginning ol year Fund balances, end of year 601,501 8.500 610.001 (610.001) 65.000 180,662 245.662 (364.339) 10.272.954 $ 9,908,615 Unrestricted Repairs and Replacement Reserve $111,922 111.922 (II 1,922) (14.128) 199,863 227,892 413.627 301,705 148,820 Total Unrestricted Restricted Total SI. 429.322 $ 1.429,322 1,429.322 1,429,322 601.501 601,501 111.922 111.922 8.500 8.500 721.923 721,923 (721,923) 1,429,322 707,399 65,000 65.000 180,662 180,662 (14.128) (14.128) 199.863 199.863 227.892 (249.931) (22.039) 659.289 (249,931) 409,358 (62,634) 1.179,391 1.1 16.757 10.421.774 790.322 11.212.096 $450.525 $10,359,140 $1.969,713 $12,328.853 16 LI L LIE Report of the Librarian Library directors In 1990. Jane Fessenden, who had been the librarian and worked at the MBL for 29 years, retired. Jane made the Library a productive home for its users, improved the collection whenever financially possible, and lead the Library through innumerable changes. During her tenure, the Library was converted from the Dewey Decimal System to Library of Congress, the first "stack move" took place (after three years of planning), the reading rooms were redone, the Rare Books Room was created, the Copy Service Center was developed in its present location, staff was increased, and new technology was initiated. She will be best remembered, however, for her unique ability to provide the services and support most desired by our users. In December. Dr. David Stonehill accepted the position of Director of the MBL/WHOI Library and Scientific Information Research Center. Dr. Stonehill managed computer facilities on various NASA projects and directed computing services at academic institutions. From 1988 until his move to MBL, he was Director of Information Resources Managements for the Executive offices of the President of the United States. Under Dr. Stonehill, the Library plans to launch a Scientific Information Research Center, and we look forward to his leadership as we meld the traditional with the new methods of scientific information management. New age information delivery Using the report created by the Hughes Committee, the Library immediately began to implement some of its recommendations. A network manager was hired, and the network design was completed. What followed was a frenzy of equipment ordering and the installation of cable and wires to extend the network out of the library and into classrooms and labs in the Lillie and Loeb buildings and the Ecosystems Center. Because of the ease of communicating over the network, some of the traditional methods of the reference librarians moved into a new era during 1990. One advance. Current Contents loaded onto the librarians' computers, allowed us to send weekly up-dated bibliographies to requesters over the MBLnet. The Library's catalog has been converted to electronic records, and all the books in the library were barcoded as part of the automation of library systems. The CLAMS network of libraries came on-line in 1990, and scientists in their laboratories, who are on the Woods Hole network, can connect to the catalog and see whether the library holds the book they are seeking. Journal rates increase Our Library has studied the effect of rising book and subscriptions costs beginning with the comprehensive Rockefeller Journal Use study. In 1990, the close to 30% projected journal price increase reduced our options and resulted in the cancellation of 338 current subscriptions. Our User Panel worked throughout the summer with lists of journals targeted for cancellation, using criteria established by the Joint Users Committee, to insure the best possible preservation of our collection and services. This was not the only journal crisis of 1990 the space allocation for housing the journals had reached saturation. For over a year, the creation of a design to preserve our stack space has been developed and refined and the periodical collection will be moved during the beginning of 1991. The logical placement of the 17 18 Annual Report collections according to the alphabet will be maintained, albeit into two places. The A-Z arrangement will be in the front part of the stacks for currently received journals and also in the back wing for the pre-1976 journals. In an attempt to recover some of the escalating costs incurred in the Library, we have tried to create a model for users who are not directly associated with the contributing Woods Hole institutions. We have also changed our interlibrary loan policies and have increased resource sharing with a number of institutions, i.e.. Brandeis, University of Massachusetts Medical School, Wesleyan, University of Rhode Island and the Pell Laboratory. Escalating costs and U. S. postage rates drove the increase in interlibrary loan charges, and at the International Association of Marine Science Libraries and Information Centers' conference in Seattle, Washington, we were challenged, without ill will, but with some concern from the international community, about our rate increases. A solution to this problem may involve sharing the responsibility for serials collection development at the national and international level during the 1990s. Preservation Two documents that have been of historic and sentimental value to the scientists, staff, and visitors of the MBL were appraised and declared endangered. Study Nature Not Books the black charcoal, handwritten sign of Louis Agassiz had adhered to its backing and tears, holes, stains, rust, scratches, and smudges were evident everywhere. The same was true ot the two-page Last one to Go message to American soldiers by Katsuma Dan. Both of these documents were sent to the Northeast Document Conservation Center for treatment and have been returned to the library for display. Cathy Norton Acting Librarian Educational Programs Summer Courses Biology of Parasitism (June 10 to August 10) Co-Directors John Donelson, University of Iowa College of Medicine Carole Long, Hahnemann Medical College Faculty Steven Anderson, University of Iowa John Boothroyd. Stanford University Ted Bianco. Imperial College of Science & Technology, UK Patrick Farley, Hahnemann Medical College Steven Hajduk, University of Alabama, Birmingham Peter Ham, Liverpool School of Tropical Medicine, UK Michael E. Harris, University of Alabama, Birmingham Mary Alice Hartman, University of Kentucky Kwang S. Kim, University of Iowa Peter Kima, Hahnemann Medical College Yien Ming Kuo, Imperial College, UK Rick Martin. University of Iowa David Moser, University of Iowa Elonne Petrin, University of Cincinnati David Russell, New York University David Sachs, NIH Judy Sakanari, University of California, San Francisco Sam Turco. University of Kentucky Lecturers Nina Agabian, University of California, San Francisco Steven Beverley, Harvard Medical School Kent Campbell, Centers for Disease Control Dickson Despommier, Columbia University Paul Englund, Johns Hopkins School of Medicine Don Harn, Harvard University Stephanie James, NIH Keith Joiner, Yale University School of Medicine Patricia J. Johnson, UCLA Don Krogstad, Washington University, St. Louis Ira Mellman, Yale University School of Medicine George Nelson, Liverpool School, UK Ruth Nussenzweig, New York University Victor Nussenzweig, New York University Richard Olds, Brown University Bill Petri, University of Virginia Robert Sauer, Massachusetts Institute of Technology Alan Sher. NIH Irwin Sherman, University of California, Riverside Larry Simpson, UCLA Mitch Sogin, MBL Rick Tarleton. University of Georgia Merv Turner, Merck Sharp & Dohme Research Laboratory C. C. Wang, University of California, San Francisco Leon Weiss, University of Pennsylvania Don Wiley, Harvard University Dyann Wirth, Harvard University Students Wanida Asawanahasakda, Mahidol University, Thailand Fernanda R. Gadelha, University of Illinois Eileen S. Gruszynski, University of California, Los Angeles Michael J. Howard, Vanderbilt University Christopher A. Hunter, University of Glasgow, Scotland Gregory J. Jennings, Tulane University Christopher L. Leptak, Yale University Congjun Li, Worcester Foundation for Experimental Biology 19 20 Annual Report Leo X. Liu, Beth Israel Hospital/Harvard University James J. McCoy, University of Virginia Gloria I. Palma, Univ. del Valle. Colombia Laura J. Rocco, Johns Hopkins University Nicola N. Schweitzer, Imperial College of Science, UK Frank Seeber, Institiit fur Tropenhygeine, Germany Philippe G. Vandekerckhove, University of Leuven, Belgium Gayl Wall, University of Dundee, Scotland Embryology: Cell Differentiation and Gene Expression in Early Development (June 21 to July 30) Directors Eric Davidson, California Institute of Technology J. Richard Whitaker, MBL (Assistant Director) Faculty Michael Akam, University of Cambridge, UK Amy Bejsovic, University of Cambridge, UK Marianne Bronner-Fraser, University of California, Irvine Scott Fraser, University of California, Irvine Katherine Harding, Columbia University Janet Heasman, University of Cambridge, UK Linda Huffer, MBL Wendy Katz, California Institute of Technology Robert Kingsbury, Carnegie Institution, Baltimore Thomas Lallier, University of California, Irvine Robert Leclerc, University of Maryland Michael Levine, Columbia University David McClay, Duke University Steven McKnight. Carnegie Institution of Washington Robert Nickells, California Institute of Technology Jerome Regier, University of Maryland John Shuman, Carnegie Institution. Baltimore Paul Sternberg, California Institute of Technology Nicholas Torpey, University of Cambridge, UK Kellie Whittaker, California Institute of Technology Christopher Wylie, University of Cambridge, UK Students Kamran Ahmad, University of Utah Michael J. Bank, Columbia University Peter B. Bokor, Rockefeller University James B. Castelli-Gair, University of Madrid, Spain Robert A. Cornell. University of Washington Maria G. Di Bernardo, Italian National Research Council, Italv Win J. Dictus, University of Utrecht, The Netherlands Anne D. Donaldson, MRC Laboratory, UK Bruce W. Draper, University of Washington Silvia B. Frenk, King's College, Cambridge, UK Kareen M. Guida, University of Paris, France Nan Ho, University of California, Berkeley Jon F. Kayyem, California Institute of Technology Dangeruta Kersulyte, Acadamy of Sciences of Lithuania, USSR Daniel S. Kessler, Rockefeller University Mary Ellen Lane, Columbia University Thierry Lepage, University of Nice, France Donal T. Manahan. University of Southern California, Los Angeles Jeffrey R. Miller, Duke University Anne Marie Murphy, Johns Hopkins University Christof Niehrs, European Molecular Biology Lab., Germany Mary E. Pownall, University of Virginia Inge J. Van Wijk, Max-Planck Institute, Germany Tongweng Wang, University of Florida Marine Ecology: Concepts, Techniques and Applications of Molecular Probes (June 17 to July 28) Director J. Woodland Hastings. Harvard University Faculty Cheryl Booth, Falmouth, MA Ann Bucklin, Marine Biological Laboratory Thomas T. Chen, Center of Marine Biotechnology, University of Maryland Clara Cheng, University of Maryland Toby Cole, Hopkins Marine Station, Stanford University Lynna Hereford, Hopkins Marine Station, Stanford University Chun-Mean Lin, University of Maryland Kenneth Nealson, Great Lakes Research Center, LIniversity of Wisconsin, Milwaukee Dennis Powers, Hopkins Marine Station, Stanford University T. Roenneberg, University of Munich, Germany Simona Sorger, Hopkins Marine Station, Stanford University Keno Truper, Bonn, Germany Barbara Wimpee, Great Lakes Research Center, University of Wisconsin, Milwaukee Charles Wimpee, University of Wisconsin, Milwaukee Educational Programs 21 Lecturers David Caron Colleen Cavanaugh Penny Chisholm Ed De Long Paul Dunlap Brian Fry Linda Goff John Kessler Lynn Margulis James McCarthy Dan Morse Rob Olsen Hans Paerl Jack Palmer Ned Ruby G. Savior Ann Sesholz Bob Simon Mitch Sogin Felix Strumwasser John Waterbury Students Abdiel J. Alvarez, University of Puerto Rico Brian J. Binder, Massachusetts Institute of Technology Alice F. Brown, Brown University Ka Hou Chu, Chinese University of Hong Kong, Hong Kong Peter J. Edmunds, Northeastern University Oivind Enger, University of Bergen, Norway Jonathan B. Geller, University of Oregon Gregory J. Hinkle, University of Massachusetts, Amherst Robert E. Hodson, University of Georgia Eric R. Holm, Duke University Jerilyn Jewett-Smith, Whitman College Lisa M. Kann, University of Rhode Island James S. Maki, Harvard University Kirk D. Malloy, University of Delaware Adam G. Marsh. University of New Hampshire Tracie-Lynn Nadeau, University of Wisconsin Martin Polz, University of Vienna, Austria Michael C. Schmale, University of Miami Robin M. Schneider, University of Southern Louisiana Lynda P. Shapiro, Bigelow Laboratory for Ocean Sciences Robert L. Sinsabaugh, Clarkson University Erika Stephens. Harvard University Stephen C. Tsoi, University of Hong Kong, Hong Kong Karl E. Wommack, University of Maryland Microbiology: Molecular Aspects of Cellular Diversity (June 10 to July 26) Co-Directors Martin Dworkin, University of Minnesota John Breznak, Michigan State University Faculty Richard Behmlander, University of Minnesota Pamela Contag, University of Minnesota Christiane Dahl, University of Bonn, Germany Deborah Eastman. University of Minnesota Andrew M. Kropinski, Queen's University, Canada Hans Truper, University of Bonn, Germany Stefan Wagener, Michigan State University Lecturers Paul Dunlap, Woods Hole Oceanographic Institution Holger Jannasch, Woods Hole Oceanographic Institution J. Waterbury. Woods Hole Oceanographic Institution Students Diane K. Arwood, University of Southern Mississippi Joanna S. Brooke, University of Western Ontario, Canada Joseph P. Calabrese, West Virginia University Neena Din, University of British Columbia, Canada Amis Druka, Latvian State University, USSR Olivia T. Harriott, University of Connecticut Robert Huber, University of Regensburg, Germany Jennifer B. Klenz, University of Saskatchewan, Canada Judith A. Koskella, New York University Bridget E. Laue, University of Colorado Jared R. Leadbetter, Goucher College Timothy C. Lilburn, University of British Columbia, Canada Shi Liu, University of Oklahoma Lynn V. Mendelman, Harvard Medical School Elizabeth J. Orle, Colorado State University Mechthild Pohlschroder, University of Massachusetts, Amherst Frank J. Slack, Tufts University Barth F. Smets, University of Illinois Claire S. Ting, Cornell University Mary A. Wyka, Merck & Co., Inc. Neural Systems & Behavior (June 10 to August 1) Co-Directors Ronald Calabrese, Emory University Martha Constantine-Paton, Yale University 22 Annual Report Faculty Arthur Arnold, University of California, Los Angeles Alexander Borst, Max-Planck-Institut filr Biologisch Kybernetick, Germany John Byrne, University of Texas Medical School Thomas Carew, Yale University Leonard deary. University of Texas Medical School at Houston Robin Cloues, Harvard University Michael Davis, Yale University Robert M. Douglas, University of British Columbia, Canada Lise Eliot, Center for Neurobiology & Behavior Russell Fernald, University of Oregon Leslie Fischer, Columbia University William Frost, University of Texas Medical School Cole Gilbert, University of Arizona Dennis Gorlick, Columbia University Jon Hayashi, Arizona Research Laboratory Sally Hoskins, City College of CUNY John Koester, New York State Psychiatric Institute Richard B. Levine, University of Arizona Yurin Levy, Brandeis University Margaret Livingstone, Harvard Medical School Anne Lohoff, Columbia University Eduardo R. Macagno, Columbia University Emilie Marcus, Yale University Eve Marder, Brandeis University Michael P. Nusbaum, University of Alabama, Birmingham Mu-Ming Poo, Columbia University David J. Sandstrom, University of California Patricia Steen, Yale University Nacita Tabti, Columbia University Janis C. Weeks, University of Oregon Angela Wenning, Universitat Konstanz, Germany Michael Nitabach, Columbia University Lecturers Catherine Carr, University of Rochester Joe Martinez, Jr., University of California Students Robert A. Berkowitz, Washington LJniversity James P. Burke, University of Alabama, Birmingham Belinda S. Chang, Harvard University Miles G. Cunningham, Massachusetts Institute of Technology Graeme W. Davis, University of Massachusetts, Amherst Mayra Garcia-Ruiz, University of Nat. Autonoma, Mexico John F. Hamilton, Meharry Medical College, Nashville Tamara L. Harris, University of Ottawa, Canada Valerie L. Kilman, University of Illinois Barlett W. Mel, California Institute of Technology Brett D. Mensch, Baylor College of Medicine Alison R. Mercer, University of Otago, New Zealand Edward P. Monaghan, University of California, Berkeley Tanja Quenzer, Max-Planck-Institut, Germany Adina L. Roskies, University of California, San Diego Hyunjune S. Seung, Harvard University Deana L. Shackelford, University of Oklahoma Petra Skiebe, Universitat Hamburg, Germany John E. Spiro, University of California, San Diego Douglas Syme, University of California, Irvine Neurobiology (June 10 to August 11) Co-Directors Leonard Kaczmarek, Yale University Irwin Levitan, Brandeis University Christopher Miller, Brandeis University Faculty Cecilia Armstrong, University of Pennsylvania Gary Banker, LIniversity of Virginia Synnove Beckh, Max-Planck-Institiit fur Biophysikalische Chemie, Germany Andrew Czernik, Rockefeller University Jan De Weer, Duke University Judith Drazba, NINDS/NIH Keith Elmslie. Case Western Reserve University Stuart Firestein, Yale University School of Medicine Paul Forscher, Yale University Robert French, University of Calgary, Canada Sara Garber, University of Alabama, Birmingham Allison Hall, Case Western Reserve University Richard Horn. Roche Institute for Molecular Biology Richard Huganir, HHMI, Johns Hopkins Medical School Stephen Jones, Case Western Reserve University Richard Kramer, Columbia University Kyu-Ho Lee, Johns Hopkins Medical School Andrew Matus, Friedrich Meischer Institute, Switzerland Robert Miller, Case Western Reserve University Angus Nairn, Rockefeller University Randall Reed, HHMI, Johns Hopkins Medical School Thomas Reese, NINDS/NIH Talvinder Sihra, Rockefeller University Carolyn Smith, NINDS/NIH Walter Stuhmer, Max-Planck-Institiit fur Biophysikolische Chemie, Germany Educational Programs 23 Students Ricardo C. Araneda, Albert Einstein Medical School Sylwester Chyb, Wesleyan University Dan H. Cox, Tufts University Stuart D. Critz, University of Texas Medical School Peter F. Drain, MIT Kathryn J. Edson, University of Minnesota Julie A. Haack, University of Utah Lise R. Heginbotham, Harvard University Marc A. Post, University of Michigan Haohua Qian, University of Illinois Hanno M. Roder. Massachusetts Institute of Technology Maria A. Sosa, University of Florida Physiology: Cell and Molecular Biology (June 10 to July 21) Director Thomas D. Pollard. Johns Hopkins University Faculty Steven Almo, Johns Hopkins Medical School Kerry Bloom, University of North Carolina William Busa, Johns Hopkins University Antony Galione, Johns Hopkins University Neal R. Gliksman, University of North Carolina Robert Jensen. Johns Hopkins School of Medicine Margaret Kenna, University of North Carolina John Maslanski. Johns Hopkins University Jonathan McMenamin-Balano, University of Massachusetts, Boston Robert Palazzo, Marine Biological Laboratory Katherine Pollard, Johns Hopkins Medical School Ted Salmon, University of North Carolina Sue Schmidt, Glyndon, MD John Simon, University of North Carolina John Sinard, Johns Hopkins Medical School Tammy Smith, University of North Carolina Cynthia V. Stauffacher, Purdue University Murray Stewart, Medical Research Council, UK Elaine Yeh, University of North Carolina Students Robert L. Bacallao, University of California, Los Angeles Sandra A. Brockman, Carnegie Mellon University Thomas O. Carpenter, Yale University Isabelle A. Carre, SUNY, Stony Brook Joseph A. Cerro, Columbia University Marc D. Coltrera, University of Washington Tod A. Critchlow, Scripps Institute of Oceanography Spencer J. Danto, Cornell Medical College Michele I. Flatters, Tufts University Holly V. Goodson, Stanford University Supriya Jayadev, Duke University John R. Jordan, University of Utah David L. Keefe, Yale University Karen L. King, Florida State University Qingwen Li, University of Kansas Helen McNeill, University of Pennsylvania Michael E. Mendelsohn, Harvard/Brigham & Women's Hospital Christa S. Merzdorf, Harvard University Robert Mirro, University of Tennessee Karen M. Page, Dartmouth College Alice P. Pentland, Washington University, St. Louis Zhican Qu, Johns Hopkins University Joe W. Ramos, University of Virginia Jean F. Regal, University of Minnesota Eric A. Shelden, University of Massachusetts, Amherst Charles B. Shuster, Tufts University Thomas W. Smith, Brigham & Women's/Harvard Medical School Robin L. Stears, SUNY, Stony Brook Salme Taagepera, University of Virginia Charlotte M. Vines, Harvard Medical School Yingjian Wang, University of Miami Christiane Wiese, Johns Hopkins Medical School Elizabeth L. Winter, City College of New York Vicki L. Wolff, Brandeis University Qi Yang, University of Connecticut Guangwen Zhou, Oregon State University Short Courses Analytical and Quantitative Light Microscopy in Biology, Medicine, and Materials Science (May 10 to 18) Co-Directors Edward D. Salmon, University of North Carolina Greenfield Sluder, Worcester Foundation for Experimental Biology David E. Wolf, Worcester Foundation for Experimental Biology Faculty and Course Assistants Brad Amos, MRC, Cambridge, UK Orit Baha, Worcester Foundation for Experimental Biology Steven M. Block, Rowland Institute for Science Richard Cardullo, Worcester Foundation for Experimental Biology Walter Carrington, University of Massachusetts Medical School 24 Annual Report Gordon Ellis, University of Pennsylvania Fred Fay, University of Massachusetts Medical School JeffGelles, Brandeis University Richard Haugland, Molecular Probes, Eugene, OR Linda Huffer. MBL Shinya Inoue. MBL Anthony Moss, Worcester Foundation for Experimental Biology Rudolf Oldenbourg, MBL Stephen Parsons, University of North Carolina Robert V. Skibbens, University of North Carolina Kenneth R. Spring, NIH D. Lansing Taylor, Carnegie-Mellon University Richard Walker, University of North Carolina Students Julia Barsony, NIDDK/NIH Harold G. Bohlen, Indiana University Medical School Daniel J. Brat, Mayo Graduate School Barry J. Burbach, SUNY, Stony Brook John P. Caufield, Harvard Medical School Wendy Cheng, International Paper Company, Tuxedo, NY Matthew H. Chestnut, The Procter and Gamble Co., Cincinnati, OH Diana M. Cordova, Alcon Laboratories, Inc., Ft. Worth. TX Sascha Dho, The Hospital for Sick Children, Toronto, Canada Ulrich Dirnagl, Forschungslabor, Germany Cynthia J. Forehand, LIniversity of Vermont John J. Freeman. Monsanto Co., St. Louis, MO Jill Gemmill, University of Alabama, Birmingham John A. Hammer. Ill, NHLBI/NIH Ray S. Hartman, Children's Hospital of Los Angeles Donald T. Haynie, The Johns Hopkins University Walter J. Koroshetz, Massachusetts General Hospital Dan Green (left) demonstrates a ratio imaging package from Universal Imaging during an AQLM course session. Julia M. Lash, Indiana University School of Medicine Michael I. Lethem, University of North Carolina Steve Paddock, University of Wisconsin Shoshana Paglin, Boston University School of Medicine James R. Sellers. NHLBI/NIH Anna Spudich, Stanford University Fei Wang, Syracuse University Measurement and Control of Chemical Stimuli (April 25 to 30) Director Greg A. Gerhardt, University of Colorado Faculty and Course Assistants Barry W. Ache, C.V. Whitney Laboratory, University of Florida Jelle Atema, Boston University Marine Program, MBL Scott Brock, University of Colorado Marilyne Friedemann, University of Colorado John S. Kauer, Tufts New England Medical Center Stuart Firestein, Yale University School of Medicine Paul Moore. Boston University Marine Program, MBL Mike Palmer, University of Colorado Michael Parrish, University of Colorado Wayne L. Silver, Wake Forest University Students Carol E. Diebel, SUNY Health Science Center Richard L. Doty, Hospital of the University of Pennsylvania Heather L. Eisthen, Indiana University Tim Granata, Southeastern Massachusetts University Kristina-Viveka Hillegaart, Astra Research Centre, Sweden Herman K. Lehman, University of Arizona Celia Marrase. Boston University Marine Program, MBL Pricilla E. Purnick, Columbia University John R. Welborn, University of Southern California Methods in Computational Neuroscience (August 5 to September 1) Co-Directors James M. Bower, California Institute of Technology Christof Koch. California Institute of Technology Educational Programs 25 Lecturers and Instructors Paul Adams, SUNY, Stony Brook Edward Adelson, Massachusetts Institute of Technology Daniel Alkon, N1H Richard Anderson, Massachusetts Institute of Technology David Beeman, University of Colorado, Boulder Avis H. Cohen, Cornell University Norberto Grzywacz, Massachusetts Institute of Technology Nancy Kopell, Boston University Rudolfo Llinas, NYU Medical Center Kevan Martin, MRC, Oxford, UK Michael Mascagni, Washington, DC Kenneth Miller, University of California, San Francisco Mark Nelson, California Institute of Technology John Rinzel, NIH David Rumelhart, Stanford University Sylvia Ryckebusch, California Institute of Technology Terrence Sejnowski, Salk Institute Allen I. Selverston, University of California, San Diego John Uhley, California Institute of Technology David Van Essen, California Institute of Technology Lucia Vaina, Boston University Matthew Wilson, California Institute of Technology Students Aric Agmon, University of California, Irvine Hagai Agmon, Hebrew University of Jerusalem, Israel Dale Anders, University of California, San Diego Evyatar Av-Ron, The Weizmann Institute of Science, Israel Ellen Barton, University of Pennsylvania Eyal Bartfeld, Rockefeller University David Berkowicz, Yale University School of Medicine Neil J. Berman, Oxford University, UK Peter J. Braam, University of Utah Dennis Bray, Colorado State University Trevor Darrell, Massachusetts Institute of Technology Gyongyi Gaal, University of Pennsylvania Kurt Haas, Albert Einstein College of Medicine Dirk Kautz, University of Oregon Markus Lappe, NIH Sean Marrett, Montreal Neurological Institute, Canada Douglas Morton, Case Western Reserve University Dietmar Rapf, MPI fur biologische Kybernetik, Germany Walter Schneider, University of Pittsburgh Nelson Spruston, Baylor College of Medicine Christoph Staub, Brain Research Institute, Switzerland Fan-Gang Zeng, Syracuse University Molecular Evolution (August 19 to 31) Director Mitchell L. Sogin, MBL Faculty Michael Clegg, University of California, Riverside Daniel B. Davison, University of Houston W. Ford Doolittle, Dalhousie University, Canada Robert Dorit, Harvard University John W. Drake, National Institute of Environmental Health Sciences Joseph Felsenstein, University of Washington Walter M. Fitch, University of California, Irvine Barry G. Hall, University of Rochester Andrew Knoll, Botanical Museum of Harvard University David R. Maddison, Harvard University Peter Maloney, The Johns Hopkins Medical School Roger Milkman, The University of Iowa Gary Olsen, University of Illinois Norman R. Pace, Indiana University David Joseph Patterson, University of Bristol, UK Margaret Riley, University of Massachusetts, Amherst David A. Shub, SUNY, Albany Steve Smith, Harvard University Temple F. Smith, Dana Farber Cancer Institute Mark Wheelis, University of California, Davis Allan C. Wilson, University of California, Berkeley Elizabeth A. Zimmer, Smithsonian Institution Students Alexandra L. Basolo, University of Texas, Austin Heidi G. Blair, SUNY, Albany Judith Anne Blake, Smithsonian Institution Laura Blinderman, San Diego State University David J. Bogler, University of Texas, Austin Diane M. Bridge, Yale University Henner Brinkmann, Technische Universitat, Germany James R. Brown, Simon Eraser University, Canada Bruce E. Byers, University of Massachusetts, Amherst Robert E. Calhoon, Queens College 26 Annual Report Bruce J. Cochrane, University of South Florida Rafael O. De Sa, University of Texas, Austin Marcela Descalzi, University of Houston Philippe Djian, Harvard Medical School James W. Edwards, Salem College Bruce A. Eichhorst, University of North Dakota William Fischer, Los Alamos National Laboratory Dean A. Glawe, University of Illinois Robert M. Gould, New York State Institute for Basic Research Sharon P. Gowan, Harvard University Herbaria Lawrence I. Grossman, Wayne State University Radhey S. Gupta, McMaster University, Canada Raymond W. Holton, University of Tennessee Bradford E. James, Dartmouth College Matthew D. Kane, University of Illinois Jessica C. Kissinger, Indiana University Paul J. Kores, Tulane University Maja Kricker, National Institute of Environmental Health Sciences Uri Ladror, Chicago Medical School Laura Landweber, Harvard University Bang-Ning Lee, University of Texas, Houston Heikki Lehvaslaiho, University of Helsinki, Finland Wayne S. Leibel, Lafayette College Enrique P. Lessa, University of California, Berkeley Louise Ann Lewis, Ohio State University Paul O. Lewis, Ohio State University John M. Logsdon, Jr., Indiana University Kenneth L. McNally, University of California Linda K. Medlin, University of Bristol, UK Thomas Mills, University of Houston Judith Mongold, University of Massachusetts, Amherst Barbara Moore, Massachusetts Institute of Technology Rana Muzaffar. Florida State University Lena M. Nielsen, University of Lund, Sweden Elizabeth A. Oakenfull, MRC Haematology Unit, Oxford, UK Yves P. Quentin, Los Alamos National Laboratory James Ramsay, University of Utah Judith M. Rhymer, Smithsonian Institution Kenneth A. Rice, Harvard University Bernardo Rudy, NYU Medical Center Frank G. Salinas, University of Houston Leo C. Schalkwyk, Dalhousie University, Canada Michael Schloemann, Yale University Jeffrey D. Silberman, University of Miami Rama S. Singh, McMaster University, Canada Elizabeth M. Snella, Indiana University Elizabeth T. Snow, NYU Medical Center Peter L. Starkweather, University of Nevada Jeffrey L. Stein, University of California, San Diego Youngbae Suh, Louisiana State University Thomas R. Sutler, CUT, Research Triangle Park, NC Cheryl L. Tarr, University of North Dakota Richard N. Williams, Boise State University Charles G. Wray, Yale University Optical Microscopy & Imaging in the Biomedical Sciences (October 6 to 12) Co-Directors Nina Stromgren Allen, Wake Forest University Colin S. Izzard, SUNY, Albany Faculty and Course Associates Steven M. Block, Rowland Institute for Science Joseph De Pasquale, SUNY, Albany Alec Harootunian, HHMI, University of California, San Diego Fredric S. Fay, University of Massachusetts Medical Center Kenneth A. Jacobson, University of North Carolina John M. Murray, University of Pennsylvania Kenneth Orndorff, Dartmouth College Julia S. Sizemore, Wake Forest University Stephen J. Smith, Stanford University School of Medicine Kenneth R. Spring, NHLBI/NIH Anna Spudich. Stanford University School of Medicine Roger Tsien, HHMI, University of California, San Diego Lecturers Jan Hinsch, Leica, Inc., Rockleigh, NJ Shinya Inoue, MBL Ernst Keller, Carl Zeiss, Inc., Thornwood, NY Rudolf Oldenbourg, MBL Students Lawrence W. Argenbright, Boehringer Ingelheim Pharmaceuticals, Inc., Ridgefield, CT Elikplimi K. Asem, Purdue University Thomas G. Burke, City of Hope National Medical Center Dean Cole, Los Alamos National Laboratory R. Ford Denison, USDA Camille DiLullo, University of Pennsylvania Harold F. Dvorak, Beth Israel Hospital David H. Eidelman, McGill University, Canada T. R. Gowrishankar, University of Chicago John W. Hanrahan, McGill University, Canada Christopher M. Kenyon, Centre Hospitalier Thoracique de Montreal, Canada Educational Programs 27 Jeff Lansman, University of California, San Francisco Stephen Lin, Harvard Medical School William W. Mantulin, University of Illinois, Urbana Champaign Jose A. Mari Mutt, University of Puerto Rico Jane E. Minturn, Yale University School of Medicine Spering A. Scott, Purdue University Jacqueline Sterner. University of Rochester Scott J. Sternberg, Colorado State University Xiao-Ying Tien, Case Western Reserve University Susan M. Wall, NIH David C. Zawieja, Texas A&M University Pathogenesis of Neuroimmunologic Diseases (August 19 to 31) Co-Directors J. Murdoch Ritchie, Yale University School of Medicine Byron H. Waksman, Foundation for Microbiology Faculty Vahi E. Amassian, SUNY Health Science Center Barry G. W. Arnason, University of Chicago Joel A. Black, Yale University Pietro DeCamilli, Yale University Medical School Judah A. Denburg, McMaster Medical Center, Canada Marc A. Dichter, University of Pennsylvania Charles A. Dinarello, Tufts University Medical School Diane Griffin, The Johns Hopkins University Stephen L. Hauser, Massachusetts General Hospital Henry Khachaturian, NIMH/NIH Norman Latov, College of Physicians and Surgeons of Columbia University Carl M. Leventhal, NINDS/NIH W. Ian Lipkin, University of California. Irvine Cathy G. McAllister, University of Pittsburgh Dale E. McFarlin, NINDS/NIH John Newsom-Davis, Oxford University, UK Robert B. Nussenblatt, NEI/NIH Nathanial G. Pitts, National Science Foundation Jerome B. Posner, Memorial Sloan-Kettering Cancer Center Donald L. Price, Johns Hopkins Hospital Richard W. Price, University of Minnesota Health Center James W. Prichard, Yale University Medical School Cedric S. Raine, Albert Einstein College of Medicine Anthony T. Reder, University of Chicago David M. Regan, York University, Canada Stephen C. Reingold, National Multiple Sclerosis Society Benjamin F. Roy, Georgetown University School of Medicine Clifford B. Saper, University of Chicago Randolph B. Schiffer, Strong Memorial Hospital Eli E. Sercarz, University of California, Los Angeles Moon L. Shin, University of Maryland, Baltimore Michael E. Shy, Thomas Jefferson University Hospital Lawrence Steinman, Stanford Medical Center Stephen G. Waxman, Yale University Medical School Howard L. Weiner, Brigham and Women's Hospital Jerry Wolinsky, University of Texas Health Science Center at Houston Students Anat Achiron. Beilinson Medical Center, Israel Peter-Brian Andersson. Sir William Dunn School of Pathology, UK Jody L. Baron, Yale University School of Medicine Bruce F. Bebo, Jr., Texas A&M University John R. Bethea, University of Alabama, Birmingham Helen Clare Bodmer, Institut de Chimie Biologique, France Arlene R. Collins, SUNY, Buffalo Steven W. Dow, Colorado State University Lorise C. Gahring, Research Institute of Scripps Clinic Maureen N. Gannon, Rockefeller University Claude Paul Genain, University of Kentucky Koenraad Gijbels, University of Leuven, Belgium Jonathan D. Glass, Johns Hopkins Hospital John J. Hemperly, Becton Dickinson Research Center Nancy A. Johnson, Washington University Medical School Abraham Kessler, Weizmann Institute of Science, Israel Judith Luber-Narod, University of Massachusetts Medical School Mary Ann McKee, Columbia Presbyterian Hospital Rune Midgard, Molde County Hospital, Norway Nicholas T. Potter, University of Connecticut School of Medicine Reijo Salonen, University of Turku, Finland Jun-ichi Satoh, University of British Columbia, Canada Tiziana Savio, Institute Superiore di Sanita. Italy Daryth D. Stallone, University of Pennsylvania School of Medicine Sharon A. Stranford. Hahnemann University Ursula I. Wesselmann, Northwestern University Jurgen Zielasek, Diabetes Research Institute, Germany Summer Research Programs Principal Investigators Alkon, Daniel, NIH, LMCN Armstrong, Clay M., University of Pennsylvania Armstrong, Peter, University of California Augustine, George J., University of Southern California Baker, Robert, New York University Medical Center Barlow, Robert, Syracuse University Barry, Daniel, University of Michigan Medical Center Barry, Susan R., University of Michigan Bass, Andrew H., Cornell University Bates, William R., Carleton University, Canada Bearer, Elaine L., University of California Beauge, Luis. Institute M. y M. Ferreyra, Argentina Begenisich, Ted, University of Rochester Medical Center Bennett, Michael V. L., Albert Einstein College of Medicine Bezanilla, Francisco, University of California, Los Angeles Bloom, George S., University of Texas Southwestern Medical Center Bodznick, David, Wesleyan University Borgese, Thomas A., Lehman College, CUNY Boron, Walter F., Yale University School of Medicine Borst, David W., Illinois State University Boyer, Barbara C, Union College Brady, Scott, University of Texas Southwestern Medical Center Brehm, Paul, SUNY, Stony Brook Brown, Joel E., Washington University School of Medicine Bryant, Shirley, University of Cincinnati Bug, William J., Columbia University Burdick, Carolyn J., Brooklyn College Burger, Max M., Friedrich Miescher-Institute, Switzerland Callaway, Joseph C., University of North Carolina Cariello, Lucio, Stazione Zoologica, Italy Chang, Donald C., Baylor College of Medicine Chappell, Richard L., Hunter College, CUNY Charlton. Milton, University of Toronto, Canada Charmantier, Guy, Universite des Sciences et Techniq. du Languedoc, France Charmantier-Daures, Mireille, Universite des Sciences et Techniq. du Languedoc, France Clay, John, NIH, NINDS Cohen, Lawrence B., Yale University School of Medicine Cohen, William D., Hunter College, CUNY Collin, Shaun P., University of California, San Diego Cooperstein, Sherwin J., University of Connecticut Health Center Cuppoletti, John, University of Cincinnati D'Alessio, Giuseppe, University of Naples, Italy D'Avanzo, Charlotte, Hampshire College De Weer, Paul, Washington University School of Medicine Deitmer, Joachim W., University of Kaiserslautern, Germany Dunlap, Kathleen, Tufts Medical School Eckberg, William R., Howard University Ehrlich, Barbara E., University of Connecticut Health Center Feinman, Richard, SUNY Health Science Center Ferguson, Donald, University of Cincinnati Fieber, Lynne A., University of Miami Fink. Rachel, Mount Holyoke College Fishman, Harvey M., University of Texas Medical Branch Gadsby, David C., Rockefeller University Gainer, Harold, NIH, NINDS Garrick, Rita Anne, New Jersey Medical School and Fordham University Giuditta. Antonio, University of Naples, Italy 28 Principal Investigators 29 Glynn, Paul, Hunter College Goldman, Robert D., Northwestern University Medical School Gonzalez-Serratos, Hugo, University of Maryland School of Medicine Griff, Edwin, University of Cincinnati Haimo, Leah T., University of California Hardin, John A., Yale University Heiny, Judith, University of Cincinnati Hernandez-Cruz, Arturo, Roche Institute of Molecular Biology Highstein, Stephen M., Washington University Holman, Molly A., The Whitney Laboratory, University of Florida Hoskin, Francis C.G.. Illinois Institute of Technology Hoy, Ronald R., Cornell University Hummon. William, Ohio University Ilan. Joseph, Case Western Reserve University Ilan, Judith, Case Western Reserve University Ip. Wallace, University of Cincinnati Johnston, Daniel, Baylor College of Medicine Josephson, Robert K., University of California Kaminer, Benjamin, Boston University School of Medicine Kaneshiro, Edna S., University of Cincinnati Kaplan, Ehud, Rockefeller University Kaplan, Ilene M., Union College Karp, Richard, University of Cincinnati Kremer, James N., University of Southern California Kriebel, Mahlon, SUNY Health Science Center, Syracuse Langford, George, University of North Carolina, Chapel Hill Laufer, Hans, University of Connecticut Lee, Youngsook, Harvard University Lehman, Michael, University of Cincinnati Levin, Jack, University of California School of Medicine, San Francisco Levine, Robert Paul, Washington University School of Medicine Lian, Jane, University of Massachusetts Medical School Linck, Richard W., University of Minnesota Lipicky, Raymond John, U. S. Food and Drug Administration Lisman, John, Brandeis University Llinas, Rodolfo R., New York University Medical Center Loewenstein, Werner R.. University of Miami School of Medicine Lohmann, Kenneth J., University of Washington Malchow. Robert Paul, University of Illinois College of Medicine Malinowska, D. H., University of Cincinnati Martinez, Joe L., Jr., University of California Matteson, Donald R., University of Maryland Meinertzhagen, Ian A., Dalhousie University, Canada Metuzals, Janis, University of Ottawa, Canada Mitchison, Timothy, University of California, San Francisco Moreno, Alonso P., Albert Einstein College of Medicine Nasi, Enrico, Boston University School of Medicine Nelson, Leonard, Medical College of Ohio Noe, Bryan D., Emory University School of Medicine Northcutt, R. Glenn, University of California, San Diego Obaid, Ana Lia, University of Pennsylvania School of Medicine Pant, Harish, NIH, NINDS Pappas. George D., University of Illinois, Chicago Parysek. Linda, University of Cincinnati Peckol, Paulette, Smith College Pierce, Sidney K., University of Maryland, College Park Pierson, Beverly K., University of Puget Sound RarTerty. Nancy S., Northwestern University Rakowski, Robert F., UHS/The Chicago Medical School Render, JoAnn, Hamilton College Ripps, Harris, University of Illinois College of Medicine Rose, Birgit, University of Miami School of Medicine Ross. William N., New York Medical College Ruderman, Joan V., Harvard Medical School Russell, John M.. University of Texas Medical Branch, Galveston Ryan, Una S., Washington University Salzberg, Brian M., University of Pennsylvania School of Medicine Segal, Sheldon, Rockefeller Foundation 30 Annual Report Sheetz, Michael P., Washington University Medical School Silver, Robert B., Cornell University Sloboda, Roger D., Dartmouth College Speksnijder, Johanna E., Utrecht University, The Netherlands Sperelakis, Nicholas, University of Cincinnati Spiegel, Evelyn, Dartmouth College Spiegel, Melvin, Dartmouth College Spray, David C, Albert Einstein College of Medicine Stein, Gary, University of Massachusetts Medical School Steinacker, Antoinette, Washington University Stracher, Alfred, SUNY Health Science Center, Brooklyn Stuart, Ann E., University of North Carolina Swandulla, Dieter, Max-Planck-Institut, Germany Swenson, {Catherine I., Harvard Medical School Tanguy, Joelle, Northwestern University Telzer, Bruce R., Pomona College Terada, Hirotoshi, Hamamatsu Photonics, K. K., Japan Treistman, Steven N., Worcester Foundation for Experimental Biology Trinkaus, John Philip, Yale University Troll, Walter, New York University Medical Center Tucker, Edward B., Baruch College Tykocinski. Mark L., Case Western Reserve University Tytell, Michael, Bowman Gray School of Medicine of Wake Forest University Vaisberg, Eugeni A., Protein Research Institute, USSR Vallee, Richard, Worcester Foundation for Experimental Biology Van Egeraat, Jan M., Vanderbilt University Vogel, Steven S., NIH, NIDDK Weiss, Dieter G., Zoology Institute, Germany Welsford, Jan G.. University of Pennsylvania School of Medicine Wolszon, Laura R., Columbia University Worden, Mary Kate, Harvard Medical School Yeh, Jay Z., Northwestern University Zigman, Seymour, University of Rochester School of Medicine and Dentistry Zottoli, Steven J., Williams College Zukin, R. Suzanne, Albert Einstein College of Medicine Summer investigators (left to right) David Borst, Guy Charmantier, Mireille Charmantier-Daures, Hans Laufer, and Brian Tsukimura Other Research Personnel Akeson, R. A., University of Cincinnati Altamirano, Anibal A.. University of Texas Medical Branch Andrews. S. Brian, NIH. NINDS Arnold, John, University of Hawaii Bamrungphol, Wattana. University of Pennsylvania Bargeron. Mary, Syracuse University Batten, Bruce E., Dublin, OH Bechtold-Imhof, Ruth, Boston University Medical Center Belmont, Lisa, University of California, San Francisco Bernal-Martinez, Juan, University of Connecticut Health Center Bersoff, Rochelle. Washington University School of Medicine Bezprozvanny, Ilya, University of Connecticut Health Center Blanchard, Charles, Cambridge, MA Bloom, Jonathan, Northwestern University Bloom, Theodora, Harvard Medical School Blubaugh, Diane, University of Puget Sound Bollner, Tomas, Dalhousie University, Canada Braithwaite, Scott, SUNY, Stony Brook Breitweiser, Gerda E., Johns Hopkins University School of Medicine Brezina, Vladimir, University of Connecticut Health Center Brient, Heather L., Ohio University Brodwick, M., University of Texas Medical Branch Brooks, Brian P., University of Pennsylvania Other Research Personnel 31 Buelow, Neal, Syracuse University Buerkett, Christopher G.. Illinois State University Burkart. Werner, Paul Scherrer Institut, Switzerland Campos de Carvalho, Antonio, Federal University of Rio De Janeiro, Brazil Capano. Carla Perrone. Stazione Zoologica, Italy Caputo, Carlo, IVIC-Inst. Venezolano de Inv. Ciento, Venezuela Carey, John P., University of Washington Caviedes. Pablo, NIH, NIDDK Chow, Robert H., University of Pennsylvania Cohen, Avrum, Yale University School of Medicine Collin, Carlos, NIH, LMCN Correa, Ana Maria, University of California, Los Angeles Cota, Gabriel, Centre de Investigacion del IPN, Mexico Couch, Ernest, Texas Christian University Davis. Marion, Yale University School of Medicine Demers. David, University of California, Riverside Dermietzel. Rolf. University of Regensburg, Germany Derrick, Brian, University of California, Berkeley Dessev, George N., Northwestern University Medical School DiPolo, Reinaldo, IVIC, Venezuela Diebel, Carol, SUNY Health Science Center at Brooklyn Dohrmann, Cord, Harvard Medical School Drazba, Judith, NIH, NINDS Evans, Wayne A., Ohio University Faeta, Hillary H.. Ohio University Falk, Chun Xiao, Yale University Medical School Floyd, Carl C., NIH, NINDS Flucher, Bernhard E., NIH, NINDS Forman, Robin, Medical College of Virginia Franzini-Armstrong, Clara, University of Pennsylvania Fujiki, Hirota, National Cancer Center Research Institute, Japan Gao, Qian, University of Pennsylvania School of Medicine Gavi, Benny, SUNY, Binghamton Genao. Ivan, Lehman College Gerosa, Daniela, Friedrich Miescher Institut, Switzerland Gill-Kumar, Pritam, U. S. Food and Drug Administration Goldman, Anne E., Northwestern University Medical School Gomez, Maria, Boston University Gomez-Laganas, Froylan M., Centra de Investigacion del IPN, Mexico Goodwill, Ken Gould, Robert, New York State Institute for Basic Research Grant, Philip, NIH, NINDS Grassi. Daniel. Ft. Lauderdale. FL Hammar. Kasia, NIH Hamosh, Leora Y., Johns Hopkins School of Medicine Haneji, Tatsuji, Chiba University School of Medicine, Japan Hernandez, Michael R., University of Texas Medical Branch Herzog, Eric, Syracuse University Hessinger. David, Loma Linda University Hogan, Emilia, Yale University School of Medicine Homola, Ellen. University of Connecticut Huang, Jack, NIH, NIDDK Huddie, Patrick L., NIH, NINDS Huerta. Patricio, Brandeis University Jaffe, David, Baylor College of Medicine Johnston, Jennifer, Dartmouth College Jordan, John R., University of Utah Kadam, P. A., The Population Council Kahana, Alon, Brandeis University Kenner. Glenda S., Ohio University Key, Brian, Children's Hospital Medical Center, Cincinnati Khanna, Anu, Lehman College Kim, Nam Hew, Albert Einstein College of Medicine Kingston, Samuel, New York KJein, Kathryn, Emory University School of Medicine Knudsen, Knud D.. U. S. Food and Drug Administration Koide, Samuel S., Population Council Komura, Hitoshi, University of Pennsylvania School of Medicine Konnerth, Arthur, Max-Planck-Institiit fur Biophysikalische Chemie, Germany Kowtha, Vijayanand C., NIH Kozlowski, David, University of Connecticut Krishna, Gobal, NIH Kronidou, Nafsika, Dartmouth College Kuhns, William, Hospital for Sick Children, Canada Kumar, Sanjay S., University of Pennsylvania Landau, Matthew, Stockton State College Larsen, Mark, University of Puget Sound Lasser-Ross, Nechama, New York Medical College Latorre, Ramon, Universidad de Chile, Chile Lederhendler, I. Izja, NIH, NINDS 32 Annual Report Leidigh. Christopher, Brown University Leonard II, Edward E., University of Pittsburgh Levitan, Herbert, University of Maryland Lin, Jen-Wei, New York University Medical Center Locke, Rachel, Washington University Lowe, Kris, New College Luca, Frank, Harvard Medical School Martin, Melissa, Illinois State University McDonald, John K., Emory University School of Medicine Menichini, Enrico, University of Naples, Italy Miledi, Ricardo, University of California Milgram, Sharon L., Emory University School of Medicine Mimori, Tsuneyo, Keio University. Japan Misevic, Gradimir. University Hospital of Basel, Switzerland Moogan, Teresa, Hunter College Moreira, Jorge E., NIH, NINDS Moshiach, Simon, NIH Murray, Sandra A., University of Pittsburgh Niclas, Joshua, University of California, San Francisco Nitabach, Michael, Massachusetts Institute of Technology Norgren. Robert, University of Cincinnati Olds, James, NIH, LMCN Sagi, Amir, Hebrew University, Israel Sakakibara, Manabu, Toyohashi, Japan Sala, Salvador, University of Maryland, Baltimore Sanchez, Ivelisse, Hunter College Sanchez-Andres, Juan V., NIH Sato, Eimei, Kyoto University, Japan Schiminovich, Samuel, Englewood, NJ Seemes, Eliana, University of Sao Paulo, Brazil Sheetz, Jennifer, Woods Hole, MA Sheller, Rebecca, University of Texas, Austin Shibuya, Ellen, Harvard Medical School Sivaramakrishnan, Shobhana, University of Southern California Sosnicki, Andrzej A., University of Pennsylvania Spires, Sherrill, University of Rochester Steffen, Walter, University of Minnesota Stewart, Patricia, University of Rochester Stokes, Darrell R., Emory University Stout, Matthew P., SUNY, BufTalo Sugimori, Mutsuyuki, New York University Medical Center Sweet, Hyla C, Union College Syme, Douglas, University of California Tewari, Kirti, University of Texas Medical Branch Todaro, M. Antonio D., University of Modena, Italy Tsukimura. Brian, Illinois State University Turner, Robert, Lehman College Papaconstandinov, Eleni, University and Cantonal Hospital, Switzerland Pardo, Alex, Hampshire College Parsey, Ramin. University of Maryland Parsons, Stephen, University of North Carolina, Chapel Hill Perozo, Eduardo, University of California, Los Angeles Piccoli, Renata, University of Naples, Italy Plant. Charles P., Tufts University Powers, Maureen K., Vanderbilt University Pumplin, David W., University of Maryland, Baltimore Quigley, James P., SUNY, Stony Brook Rafferty, Keen A., University of Illinois Rasgado-Flores, Hector, University of Health Sciences/ Chicago Medical School Reese, Thomas, NIH, NINDS Rodriguez, Katrin, University of Illinois Romero, Adarli, Washington University Roth, William W., Emory University School of Medicine Russell, Joshua C., University of Texas Medical Branch Ueno, Hiroshi, Rockefeller University Vargas, Fernando, U. S. Food and Drug Administration Vogel, Jackie M., Illinois State University Wache, Susanne C., University of Connecticut Wadsworth, Patricia, University of Massachusetts Watson, Win, University of New Hampshire Wells, Dan, Lexington, MA Wiercinski, Floyd J., Northeastern Illinois University Wu, Jian-Young, Yale University School of Medicine Zakevicius, Jane M., University of Illinois College of Medicine Zecevic, Dejan, University of Belgrade, Yugoslavia Zheng, Qiang, Baylor College of Medicine Zhi-quo, Liang, Population Council Zigman, Bunnie Rose, University of Rochester School of Medicine and Dentistry Zipser, David, E. Lansing, MI Library Readers 33 Library Readers: General Adler, Elizabeth, MBL Apter, Nathanial, Nova University Baccetti, Baccio, University of Siena, Italy Barrett, Dennis, University of Denver Bartolucci, Simonetta, Naples, Italy Burdick, Jonathan, MBL Bursztajn, S., Baylor College of Medicine Carriere, Rita M., Downstate Medical Center Child, Frank M., Trinity College Chinard, Francis P., New Jersey Medical School Cobb, Jewel Plummer, California State University, Fullerton Cohen, Leonard, American Health Foundation DeSimone, Douglas W., University of Virginia Health Science Center DeToledo-Morrell, Leyla, Rush Medical Center Dowling, John, Harvard University Duncan, Thomas K., Nichols College Edds, Kenneth T., SUNY, Buffalo Eisen, Herman, Massachusetts Institute of Technology Farmanfarmaian, A., Rutgers University Fox, Tom, Harvard Medical School Frenkel, Krystyna, NYU Medical Center Friedler, Gladys, Boston University School of Medicine German, James L., The New York Blood Center Gilbert, Daniel L., NIH Goldstein, Moise H., John Hopkins University Goodman. Dewitt S.. Columbia University Gormley, Gerard, MBL Guttenplan, Joseph, NYU Dental Center Hill, Richard, Michigan State University Humphreys, Tom, University of Hawaii International Wildlife, MBL Kaltenbach, Jane, Mount Holyoke College Karlin, Arthur, Columbia University Kelly, Robert. U.I.C. College of Medicine Kisten & Babitsky, MBL Klemow, Kenneth M.. Wilkes University Kremer, James N., University of South California Laderman, Aimlee D., Yale University Lee, John J., City College of CUNY Levitz, Mortimer, NYU Medical Center Marine Research, MBL May, Ronald, MBL Mitchell, Ralph, Harvard University Mooseker, Mark S., Yale University Musacchia, X. J., University of Louisville Olins, Ada L., University of Tennessee, Oak Ridge Olins, Donald, University of Tennessee, Oak Ridge Oschman, James L., MBL Passano, Leonard, University of Wisconsin Peisach, Jack, Albert Einstein College of Medicine Pollen, Daniel, University of Massachusetts Medical School Prosser, C. Ladd. University of Illinois Prusch, Robert, Gonzaga University Robinson, Denis, MBL Romagnani, Sergio, Universita di Firenze, Italy Rossi, Mose, Naples, Italy Rourke, John, MBL Russell-Hunter, W. D., Syracuse University Sanger, Jean, University of Pennsylvania Sanger, Joseph W., University of Pennsylvania Schippers. Jay M., WAFRA, New York, NY Shriftman, Mollie-Starr, North Nassau Health Center Singh, Ajai Pratap, Bareilly College Solomon, Dennis, MBL Stein, Leonard, Health Sciences Center SUNY Stephens, Philip J., Villanova University Stephenson, William K., Earlham College Sweet, Frederick, Washington University School of Medicine Szent-Gyorgyi, Andrew, Brandeis University Szulman, Aron, MaGee Womens Hospital Trager. William, The Rockefeller University Van Holde, Kensal E., Oregon State University Vaina, Lucia, Boston University Wagner, Robert R., University of Virginia Wallace, Robert W.. MBL Warren, Leonard, Wistar Institute Wilber, Charles G., Colorado State University Wimpee. Charles, MBL Wittenberg, Jonathan, Albert Einstein College of Medicine Wolken, Jerome, Carnegie Mellon University Worgul, Basil, Columbia University Worgul, Kathleen, MBL Young, Wise, NYU Medical Center Young, Lily, NYU Medical Center 34 Annual Report Library Readers: Desks Anderson, Everett, Harvard Medical School Avioli, Louis V., Jewish Hospital, St. Louis Boyer, John F., Union College Candelas, Graciela C., University of Puerto Rico Chaet, Alfred B., University of W. Florida Clark, Arnold, MBL Cohen, Seymour, MBL Collier, Marjorie M., St. Peters College Copeland, Eugene, MBL Corliss, Bruce, Duke University Crews, David, University of Texas, Austin Czinn, Steven J., RB&C Hospital, Cleveland Festoff, Barry, VA Medical Center, Kansas City Fussell, Catherine, University of Pennsylvania Gibson, Kevin, University of Pittsburgh Gross, Paul R., University of Virginia Grossman, Albert, NYU Gruner, John, NYU Medical Center Haubrich, Robert, Denison University Herskovitz, Theodore, Fordham University Inoue, Sadayuki, McGill University, Canada Katz, George M., Merck, Sharp & Dohme King, Kenneth, Childrens Hospital Krane, Stephen, Mass. General Kravitz, Edward, Harvard Medical School Leighton, Joseph, Peralta Cancer Research Lorand, Laszlo, Northwestern University Malbon, Craig C., SUNY Mauzarall, David, Rockefeller University Mizell, Merle, Tulane University Morrell, Frank, St. Lukes Medical Center, Chicago Narahashi, Toshio, Northwestern University Medical School Nickerson, Peter A., SUNY, Buffalo Paton, David, MBL Person, Philip, VA Medical Center, Brooklyn Robinson, Denis, MBL Roth, Jay, University of Connecticut Roth, Lorraine, MBL Shanklin, Douglas, University of Tennessee Shepard, Frank, Woods Hole Database Shepro, David, Boston University Sonnenblick, B. P.. Rutgers University Spector, Abraham. Columbia University Speck, William, Case Western Reserve Stuart, Ann, University of North Carolina Sundquist, Eric, USGS, Woods Hole Sydlik, Mary Anne, Eastern Michigan University Tweedel, Kenyon, University of Notre Dame Webb, Marguerite, MBL Wittenberg, Beatrice, Albert Einstein College of Medicine Yow, Frank, Kenyon College Library Readers: Rooms Hines, Michael, Duke University Medical School Moore, John W., Duke University Medical School Rabinowitz, Michael. Harvard Medical School Reynolds, George, Princeton University Weidner, Earl, Louisiana State University Weissman, Gerald, NYU Medical Center Zweig, Ronald, MBL Domestic Institutions Represented Alabama, University of Birmingham Albert Einstein College of Medicine American Bionetics, Inc. American Psychological Association Ames Laboratory Analytical Luminescence Laboratory Applied Biosystems Arizona Research Laboratory Arizona, University of Arizona, University of. School of Medicine Aspen Research Institute Atlantex & Zieler Instrument Corporation Axon Instruments, Inc. Bareilly College Baruch College of CUNY Baylor College of Medicine Beckman Instruments, Inc. Bethesda Research Labs Bio-Rad Laboratories Bodega Marine Station Boston University Boston University School of Medicine Bowling Green State University Bowman Gray School of Medicine Brandeis University Brigham & Women's Hospital Brinkmann Instruments. Inc.. Brooklyn College of CUNY Brown University Institutions Represented Bryn Mawr College Bunion Instrument Company, Inc. California Institute of Technology California, University of, Berkeley California, University of, Davis California, University of, Irvine California, University of, Los Angeles California, University of. Riverside California, University of. San Diego California, University of, San Francisco California, University of, Santa Cruz Cambridge Instruments Cambridge Technology Carnegie Institute of Washington Carnegie-Mellon University Case Western Reserve University Center for Agricultural Biotechnology Center for Marine Biotechnology Center for Microbial Ecology, Michigan State University Chicago, University of Children's Hospital Medical Center Ciba Corning Diagnostics Corp. Cincinnati, University of City College of New York City University of New York (CUNY) Clark University Colgate University Colorado College Colorado, University of. Boulder Columbia University Columbia University College of Physicians and Surgeons Connecticut, University of Connecticut, University of. Health Center Cornell University Coy Laboratory Products Crimson Camera Technical Sales CSPI, Inc. Dage MTI. Inc. 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School of Medicine Michigan State University Michigan, University of, Ann Arbor Minnesota, University of Molecular Probes Monsanto Company Mount Holyoke College Murray State University National Institutes of Health/NINDS National Institutes of Health/NIEHS National Institutes of Health/NIDDK National Jewish Center for Immunology & Respiratory Medicine National Science Foundation Naval Medical Research Institute Neslab Instruments, Inc. New Brunswick Scientific Company, Inc. New College of the University of South Florida New England Medical Center New Hampshire, University of New Jersey Medical School New York Medical College New York University Medical Center 36 Annual Report Nikon, Inc. North Carolina, University of. 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Williams College Wisconsin, University of Wistar Institute Woods Hole Oceanographic Institution Worcester Foundation for Experimental Biology Yale University Yale University School of Medicine Zeiss, Carl, Inc. Foreign Institutions Represented Academy of Sciences, USSR Basel, University of, Switzerland Belgrade, University of, Yugoslavia Bergen, University of, Norway Bielefeld, University of, Germany Biozentrum, Basel. Switzerland Bonn, University of, Germany British Museum of Natural History, UK Calgary, University of, Canada Cambridge, University of, UK Carlton LIniversity, Canada Catania. University of, Italy Centro de Investigacion del IPN, Mexico Chiba University, Japan Chiba University School of Medicine, Japan Chile, University of. C.E.C.S., Chile Institutions Represented 37 CINEVESTAV-IPN, Mexico Cologne, University of, Germany Dalhousie University, Canada Dalhousie University Medical School, Canada Ecole normale superieure, France Federal University of Rio de Janeiro, Brazil Freie Universitat Berlin, Germany Friedrich Miesener Institut. Switzerland Gottingen, University of, Germany Hamamatsu Photonics, K. K.. Japan Hebrew University, Israel Hong Kong, University of. Hong Kong Imperial College of Science & Technology, UK Institute for Biological Research. Yugoslavia Instituto di Biologia dello Sviluppo, CNS, Italy Instituto M. y. M. Ferreyra, Argentina I.V.I.C., Venezuela Kaiserslautern, University of, Germany Katholieke Universiteit Leuven, Belgium Keio University, Japan Kobe University, Japan Koln, University of, Germany Konstanz, University of, Germany Kyoto University, Japan Life Sciences Institute, Israel London School of Hygiene & Tropical Medicine, UK London, University of, Egham, UK Max-Planck-Institut fur Biophysikalische Chemie, Germany Max-Planck-Institiit, Gottingen, Germany McGill University, Canada Medical Research Council, UK Milan, University of, Italy Modena, University of, Italy Montreal. University of, Canada Naples, University of, Italy Nice, University of, France Nottingham, University of, UK Osnabriick, University of, Germany Ottawa, University of, Canada Paris, University of, France Paul Scherrer Institute, Switzerland Philipps-Universitat, Marburg, Germany Philipps-Universitat, Germany Protein Research Institute, LISSR Queens College, UK Queens University, Canada Queensland, University of, Australia Regensburg, University of, Germany Sao Paulo, University of, Brazil Saskatchewan. University of, Canada St. Andrews University, Scotland, UK Seoul National University, Korea Siena, University of. Italy Simon Bolivar, University of, Venezuela State University of Utrecht, The Netherlands Stazione Zoologica, Italy Stockholm, University of, Sweden Swiss Federal Institute, Switzerland Swiss Federal Institute of Technology, Switzerland Toronto, University of, Canada Tubingen, University of, Germany Universita di Firenze, Italy Universita di Palermo, Italy Universite des Sciences et Techniq. du Languedoc, France University College, UK University Hospital of Basel, Switzerland Utrecht University, The Netherlands Vienna, University of, Austria Vrije Universiteit Brussel, Belgium Wolfson College, UK Z.L.F. Kantonsspital, Basel, Switzerland Zoologie Institut, Germany Year-Round Research Programs Boston University Marine Program Atema, Jelle (Professor of Biology, Program Director) Humes. Arthur G. (Professor of Biology Emeritus) Strickler, J. Rudi (Professor of Biology, Program Director) Tamm, Sidney L. (Professor of Biology) Valiela, Ivan (Professor of Biology) I '/siting tacultv and investigators Ache. Barry (C. V. Whitney Lab, St. Augustine. Florida) Bamhach, Richard (Virginia Polytech and State University) Bloomer, Sherman (Boston University) Borisy, Gary (University ot Wisconsin, Madison) Caballero. Pascual (University of Las Palmas. Gran Canaria) Chang, Donald (Baylor College of Medicine. Texas) D'Avanzo. Charlene (Hampshire College) Dionne. Vincent (University of California) Gerhardt, Greg (University of Colorado) Good. Michael (Lltrecht University, The Netherlands) Hinga. Kenneth (LJniversity of Rhode Island) Josephson, Robert (University of California, Irvine) Kauer. John (Tufts University) Kaufman, Les (New England Aquarium) Kremer, James (University of Southern California) Linck, Richard W. (University of Minnesota. St. Paul) Marrase, Celia (University of Barcelona, Spain) Meinertzhagen, Ian (Dalhousie University, Canada) Nakamura. Shogo (Toyama University, Japan) Patterson. David (University of Bristol. UK) Peckol, Paulette (Smith College) Perez Castillo. Fernando (CIQRO. Mexico) Rhoads, Donald (Adjunct Professor of Geology) Rietsma. Carol (SUNY, New Paltz) Rosenbaum. Joel (Yale University) Sardet. Christian (Villefranche-sur-Mer. France) Sardu. Rafael (Centre d'Estudio Avancats de Blanes, Spain) Singarajah, K. V. (University of Brazil. Brazil) Steffan, Walter (University of Minnesota, St. Paul) Speksnijder. Annelies (Utrecht University, The Netherlands) Sulak. Ken (Huntsman Marine Science Centre, Canada) Terasaki, Mark (NIH) Research stall Foreman, Kenneth (Research Associate) Gerardo. Hortense (Research Associate) Tamm, Signhild (Senior Research Associate) Voigt. Rainer (Research Associate) Teaching assistants and stall Alber, Merryl (Course Assistant) Coughlin, David (Course Assistant) Cowan, Diane (Course Assistant) Gomez, George (Course Coordinator) Hahn, Dorothy (Senior Administrative Secretary) Hersh. Douglas (Course Assistant) LaMontagne. Michael (Course Assistant) Mosiach. Simon (Course Assistant) Mulsow. Sandor (Course Assistant) Paullin. Susanne (Assistant to the Director) Scholz, Nathaniel (Course Assistant) Sunlcy, Madeline (Administrative Manager) Varela, Diana (Course Assistant) Graduate students Alber, Merryl Banta. Gary Brewer, Matthew Bohachevsky, Boris Bryden, Cynthia Coughlin. David Cowan, Diane Elskus. Adria Gallager. Scott Gomez. George Hersh. Douglas Hwang, Jiang-shiou Karavanich. Christy Katz, Andrea Year-Round Research Programs 39 Kennedy, Blain LaMontagne, Michael Lavalli. Kan Lindstrom, Daniel Mazel. Charles Merrill, Carl Moore. Paul Mosiach, Simon Mulsow, Sandor Portnoy. John Scholz. Nathaniel Strubel, liana Tamse Armando Trager, Geoffrey Usup, Gires Varela, Diana Weinstein, Diana White, David Undergraduate student* I-'all 1990 Annis, Eric Boulanger, Lisa Brand. Michelle (University of California. San Diego) Carrama. Carolyn Christine. Stephanie Collumb, Chris (Trinity University) Christman, Laurie Cruse, Jennifer Deiner, Michael (Tulane) DeSantis, Michael Dougall, David (University of Pittsburg, Johnstown) Fernandez, Cecilia Forrest. Davina Fox, Pamela Gibson. Mark (Wesleyan) Goodbred, Steven Guertin, Laura (Bucknell) Harris. Matthew Haven. Michael Hover, Eric (Lawrence University) Kinkade, Christopher Kozlowski, Wendy MacDonald, Robin McKee, Nancy (Middlebury) McNeil. Sean Polsky, Matthew Russell, John (Franklin & Marshall) Seton-Haris, Genevieve Shaz, David Smith. Douglas Smith. Kirsten Spradlin, Trevor Staunton, Edward Steenburgh. Eric Stousland. Brett (Lawrence University) Tello. Christine Valdes. Hugo Warner, Jacqueline BUMP graduate student Diane Cowan explains the finer points of the lobster to BUMP undergrads. Youngberg, Jill Ziemba, Robert Visiting graduate .stndentx Fall 1991) ll'HMS Burner Michael (Lawrence University) Jones, Fynn (Vanderbilt) Klaper. Rebecca (University of Illinois) Maselli. Andrew (Knox College) Naessig, Tricia (Augusta College) Theis, Lisa (Lawrence University) Summer undergraduate interns Altes, Hester Bergles, Dwight Butler, Nina (Westown School. PA) Casterline, Jennifer Call, Christopher Hoagland. Todd (Bucknell) Lacomis, Lynne (SUNY. Binghamton) Pfeifer, Shaili Pardo. Alex (Five Colleges) Reischauer, Alyssa (USC) Sanders, Sophie (Dalton School) Short, Graham Szulgit, Greg Fall undergraduate intern Waggett, Caryl (Brown) 40 Annual Report Laboratory ofJelle Atema Organisms use chemical signals as their main channel of information about the environment. These signals are transported in the marine environment by turbulent currents, viscous flow, and molecular diffusion. Receptor cells extract signals through various filtering processes. Currently, the lobster with its exquisite sense of taste and smell, is our major model to study the signal filtering capabilities of the whole animal and its narrowly tuned receptor cells. Research focuses on amino acids (food signals) and pheromones (courtship and dominance) neurophysiology of receptor cells, behavior guided or modulated by chemical signals, and computational models of odor plumes and neural filters. Laboratory of Arthur G. Humes Research interests include systematics, development, host specificity, and geographical distribution of copepods associated with marine invertebrates. Current research is on taxonomic studies of copepods from invertebrates in the tropical Indo-Pacific area, and poecilostomatoid and siphonostomatoid copepods from deep-sea hydrothermal vents and cold seeps. Laboratory of Ivan Valiela Our major research activity involves the Waquoit Bay Land Margin Ecosystems Research Project. This work examines how human activity in coastal watersheds (including landscape use and urbanization) increases nutrient loading to groundwater and streams. Nutrients in groundwater are transported to the sea, and, after biogeochemical transformation, enter coastal waters. There, increased nutrients bring about a series of changes. The Waquoit Bay LMER is designed to help us to understand and model the coupling of land use and consequences to receiving waters, and to study the processes involved. A second long-term research topic is the structure and function of salt marsh ecosystems, including the processes of predation, herbivory. decomposition, and nutrient cycles. The Ecosystems Center The Center was established in 1975 to promote research and education in ecosystems ecology. Eleven senior scientific staff and forty research assistants and support staff study the terrestrial and aquatic ecology of a wide variety of ecosystems ranging from northern Europe (trace gas emission from acid- rain affected forests) to the Alaskan Arctic (long-term studies of the controls of tundra, lake, and stream biota) to the Harvard Forest (long-term studies of the effects of disturbance in forest ecosystems) to Buzzards Bay (controls of anaerobic decomposition). Many projects, such as those dealing with sulfur transformations in lakes and nitrogen cycling in the forest floor, investigate the movements of nutrients and make use of the Center's mass spectrometry laboratory (directed by Brian Fry) to measure the stable isotopes of carbon, nitrogen, and sulfur. The research results are applied wherever possible to questions of the successful management of the natural resources of the earth. In addition, the ecological expertise of the staff is made available to public affairs groups and government agencies who deal with such problems as acid rain, ground water contamination, and possible carbon dioxide-caused climate change. Opportunities are available for postdoctoral fellows. Staff and consultants Hobbie, John E., Co-Director Melillo, Jerry M., Co-Director Banta, Gary Bauman, Carolyn Berger, Laurel Bowles, Francis Bowles, Margaret Castro, Nancy Cochran, Wendy Davis, Sarah Deegan. Linda Dornblaser. Mark Downs, Martha Fafinski, Stephen Fry. Brian Garritt, Robert Giblin, Anne Gregg, Tim Griffin, Elisabeth Helfrich, John Hopkinson, Charles Hullar, Meredith Jesse. Martha Jordan, Marilyn Kicklighter, David Kracko, Karen Laundre, James Michmerhuizen, Catherine Miliefsky, Michelle Moller, Bernard Nadelhoffer, Knute Padien. Daniel Pallant. Julie Parmentier. Nancy Peterson. Bruce Raich, James Rastetter, Edward Regan, Kathleen Ricca, Andrea Russell, Ann Saupe. Susan Schwamb, Carol Schwarzman, Elisabeth Semino. Suzanne Shaver, Gaius Shulman, Laura Steudler, Paul Tholke, Kristin Tucker. Jane Bowden. Richard Peterjohn. William Castro. Mark Ryan. Michael Klmg, George Wainright. Sam McKane, Robert I 'iMiinx scholars Joyce, Linda, U.S.D.A. Forest Service McGuire. David, U.S.D.A. Forest Service Neill. Christopher. University of Massachusetts, Amherst Laboratory for Marine Animal Health The laboratory provides diagnostic, consultative research, and educational services to the institutions and scientists of the Woods Hole community concerned with marine animal health. Diseases of wild, captive, and cultured animals are investigated. Staff Abt, Donald A., Director and The Robert R. Marshak Term Professor of Aquatic Animal Medicine and Pathology, School of Veterinary Medicine, University of Pennsylvania Year-Round Research Programs 41 A view of Alaska's Brooks Range form the Ecosystems Center's Toolik Lake research station. Photo by Mark Dornblaser. Bullis, Robert A., Research Associate in Microbiology, University of Pennsylvania Leibovitz, Louis, Director Emeritus McCafferty, Michelle, Histology Technician, University of Pennsylvania Moniz, Priscilla C., Secretary Smolowitz, Roxanna. Research Associate in Pathology, University of Pennsylvania Wadman. Elizabeth A., Microbiology Technician, University of Pennsylvania Laboratory of Aquatic Biomedicine This laboratory investigates leukemias of soft shell clams. Monoclonal antibodies developed by this laboratory and techniques in molecular biology are used to investigate the differences between normal and leukemic cells and their ontogeny. The impact of pollutants on leukemogenesis is currently being studied with an emphasis on regional superfund sites. Staff Reinisch, Carol L., Investigator, MBL, and Chairperson Department of Comparative Medicine, Tufts University School of Veterinary Medicine Miosky, Donna, Laboratory Technician White. Marja. Postdoctoral Fellow Laboratory of Cell Biochemistry This laboratory studies developmental, metabolic, and environmental influences on the genetic regulation of cellular enzymes. Current emphasis is on the gene products involved in hepatic heme biosynthesis and utilization in marine fish. These processes are responsive to hormonal and nutritional signals as well as to environmental pollutants and carcinogens. This work is being conducted with fish liver in vivti, with primary cultures of normal hepatocytes. and with cultured hepatoma cells isolated from a fish tumor. Gene activity is quantitated with cDNA probes, and the relevant genes are being cloned in bacteria to define better the actions of chemical inducers. Other research is concerned with translocation of proteins between various subcellular compartments both in fish hepatocytes and in invertebrate eggs before and after fertilization. Staff Cornell, Neal W., Senior Scientist Bruning, Grace, Research Assistant Foley. Kathleen, Research Assistant I'tviting Fox, T. O., Harvard Medical Center 42 Annual Report Laboratory of Developmental Genetics This research group studies the early gene control of cellular differentiation pathways (cell lineage determination) in the embryos of tunicates and other marine invertebrate species. Staff Whittaker, J. Richard, Senior Scientist Crowther, Robert, Research Assistant Loescher, Jane L., Research Assistant Meedel, Thomas H.. Assistant Scientist I 'isitint( investigators Collier, J. R., Brooklyn College Lee, James J., Columbia University, College of Physicians & Surgeons Laboratory* of Judith P. Grassle Studies on the population genetics and ecology of marine invertebrates living in disturbed environments, especially of sibling species in the genus Capilclla (Polychaeta). Staff Grassle, Judith P., Senior Scientist Feinsilber, Sigalit, Research Assistant Mills, Susan W.. Research Assistant Laboratory ofHaryln O. Halvorson This laboratory is interested in the molecular process of sporulation and germination in members of the genus Bacillus. Our earlier work has involved characterization of the ger.l gene in Bacillus subtilis, and determination of the germination requirements of marine endospore-forming bacteria. Over the past year, we have isolated a large number of sporeformers from various marine environments like deep sea cores and sediments. Our intention is to characterize these bacteria at the molecular level, with emphasis on genes associated with sporulation and germination. Protocols based on DNA fingerprinting and quantitative hybridizations have been developed to differentiate these bacteria from one another, as well from terrestrial sporeformers. The hybridization data has shown that the bacterial isolates are not closely related to one another. Numerical taxonomic methods are also being used to cluster the various isolates. The physiologically interesting sporeformers will also be characterized by physical mapping using rare-cutting restriction endonucleases. Staff Halvorson. Harlyn O., Principal Investigator Chikarmane, Hemant, Assistant Scientist Glick, Beatriz. Research Assistant Pratt. Sara. Research Assistant VanLoov. Lori. Research Assistant I isiting investigators Anderson, Porter, University of Rochester Keynan, Alex, Hebrew University, Jerusalem, Israel Kornberg, Hans, Christ's College, Cambridge. UK. Vincent, Walter, University of Delaware Yashphe. Jacob, Hebrew University, Jerusalem, Israel Laboratory of Shiny a Inoue Mechanism of mitosis and related motility. Development of high resolution 3-D video microscope systems. High resolution polarized light microscopy of muscle fibrils. Physical origin of edge birefringence and image formation in the polarized light microscope. Staff Inoue, Shinya, Distinguished Scientist Knudson, Robert. Instrument Development Engineer Oldenbourg, Rudolf. Visiting Assistant Scientist Stukey, Jetly, Research Assistant Szent-Gyorgyi, David. Research Assistant Taracka, Richard. Instrument Maker Woodward. Bertha M., Laboratory Manager I 'isiting investigators Inoue, Theodore, Universal Imaging Corporation, West Chester, Pennsylvania Silver, Robert B., Cornell University. Ithaca, New York Stemmer, Andreas, Lhiiversity of Basel, Switzerland Laboratory of Alan M. Ku^irian The research explores the functional morphology and ultrastructure of various organ systems in opisthobranch mollusks. The program includes mariculture of the nudibranch, Hermissenda crassicornis, with emphasis on developing reliable culture methods for rearing and maintaining the animal as a research resource. Studies include optimization of adult and larval nutrition, control of facultative pathogens and disease, and development of morphologic criteria for staging larvae and juveniles. These morphologic studies stress the ontogeny of neural and sensory structures. Concurrent with these studies is the development of a new technique to obtain and reconstruct serial block face images (SBFI) of epoxy embedded tissue actually sectioned inside an SEM by an in situ miniature ultramicrotome. Additional collaborative research includes histochemical investigations on strontium's role in initiating calcification in molluscan embryos (shell and statoliths). as well as immunocytochemical labelling of cell-surface and secretory product antigens of neurosecretory neurons in the eye of . l/'/r.v/fl. Staff Alan M. Kuzirian. Assistant Scientist Catherine T. Tamse, Research Assistant Year-Round Research Programs 43 Dr. Alan Kuzirian and Catherine Tamse in the laboratory. Photo by George Liles. Laboratory of Molecular Evolution The major research effort of this laboratory is the structure analysis of ribosomal RNA. Similarities between small subunit ribosomal RNA sequences are used to infer the evolutionary history of eukaryotic microorganisms and to design molecular probes for studies in marine ecology. Staff Sogin, Mitchell L., Director Bhattacharya. Debashish, Postdoctoral Fellow Bibeau, Claude, Research Technician Bucklin, Ann, Visiting Scientist Stickel, Shawn, Research Technician Wainwright, Patricia, Postdoctoral Fellow Laboratory of Neuroendocrinology This laboratory studies the molecular and cellular bases of two neural programs that regulate different important behaviors in the model mollusk Aplysia. Research is conducted on the mechanisms of the neuronal circadian oscillators located in the eyes. These circadian oscillators drive the circadian activity rhythm ot the animal, which is concerned with the daily timing of food gathering and of prolonged rest. Additional research is conducted on a group of neuroendocrine cells that produce a peptide, "egg-laying hormone," that initiates egg laying and associated behaviors. The laboratory is interested in how the three-dimensional shape of this peptide hormone allows a highly specific interaction with its receptor and the intracellular processes that are triggered by it. In another project, the laboratory has discovered and is continuing research on a novel second messenger enzyme, an NADase, in the oocytes of Aplysia. that generates cyclic ADPR, a Ca :+ -mobilizing product. Staff Strumwasser, Felix, Director Cox, Rachel L., Senior Research Assistant Glick, David, Senior Postdoctoral Fellow Hellmich, Mark, Postdoctoral Fellow Rainville, Carol, Laboratory Assistant Vogel, Jackie, Research Assistant Laboratory of Robert E. Palazzo This laboratory studies the biochemical regulation of cellular events during meiosis and mitosis. An integral part of the research effort is the design of reconstitution systems that faithfully execute cell cycle dependent events under defined conditions. Current cell biological, immunochemical, biochemical, and microscopic methodologies are employed. Using marine eggs as a material source, assays have been developed that allow the study of germinal vesicle breakdown (GVBD), aster formation, and reactivation of isolated mitotic apparatus in vitro. Current focus of the laboratory is on the identification of cell cycle dependent regulatory events with major emphasis on protein phosphorylation and other post- translational modifications. The ultimate goal is the identification of key enzymes and target substrates that are involved in the regulation of cell division and are highly conserved during evolution. Laboratory of Monica Riley Research in this laboratory focuses on the molecular evolution and gene expression in the bacterium Escherichia coli. In a collaborative effort, a database containing information on the intermediary metabolism and biochemical pathways of E. coli is being developed. When completed, this database is expected to contain information on each metabolic reaction, the enzyme, the reactants, products, cofactors, activators, inhibitors, kinetics, equilibrium constants, binding constants, etc. Related research is on the evolution of the K. coli DNA and organization of the genes in the chromosome. Comparative nucleotide and amino acid sequence data provide information on the evolutionary relationships of E. coli genes to homologous genes in related bacteria. Laboratory of Sensory Physiology Since 1973, the laboratory has conducted research on various aspects of vision. Current studies focus on photoreceptor cells, on their light-absorbing pigments, and on their biochemical reactions initiated by light stimulation. Microspectrophotometric and biochemical techniques are used to study the receptors of both vertebrates (amphibia, fish, and mammals) and invertebrates (horseshoe crab and squid). 44 Annual Report Staff Harosi, Ferenc, Director, Associate Scientist, MBL. and Boston University School of Medicine Szuts, Ete, Associate Scientist, MBL, and Boston University School of Medicine I 'isiting investigators Evans, Barbara I., University of Oregon, Eugene Hawryshyn, Craig W., University of Victoria, Canada Lall, Abner B., Johns Hopkins University Laboratory ofOsamu Shimomura Biochemical studies of the various types of bioluminescent systems. Preparation of the improved forms of aequorin for measuring intracellular free calcium. Staff Shimomura, Osamu, Senior Scientist, MBL, and Boston University School of Medicine Shimomura, Akemi, Research Assistant I'isiting investigator Nakamura, Hideshi, Harvard University Laboratory of Raquel Sussman We investigate the molecular mechanism of DNA damage- inducible functions in E culi. Present studies deal with novel genes that affect radiation-induced mutagenesis and analysis of RecA functions. Staff Sussman, Raquel, Associate Scientist Hemant Chikarmane, Postdoctoral Research Associate Dudley. Karen, Research Assistant National Vibrating Probe Facility We are exploring the roles of ionic currents, gradients, and waves in controlling development. We focus on controls of pattern and controls by calcium ions. Staff Jaffe, Lionel, Senior Scientist and Facility Director Kuhtreiber, Willem, Physiologist McLaughlin, Jane, Research Assistant Miller, Andrew, Research Associate Sanger, Richard, Technician Shipley, Alan, Technician I 'isiling investigators Allen, Nina, Wake Forest College Bates, William, Carleton University, Canada Buonano, Mark. Massachusetts Institute of Technology Chrystal, Jane, University of Sidney, Australia Cornwall, Carter, Boston University Danilchik. Michael, Wesleyan University Gillot, Isabelle. Station Zoologique. Villefranche-sur-mer, France Isaacs, Hugh, Brookhaven National Laboratories Kinnamon, Sue, Colorado State University Koshian, Leon, Cornell University Lucas, William, University of California, Davis McConnaughey, Ted, Marine Biological Laboratory O'Donnell, Michael, McMaster University, Canada Ripps, Harris, University of Illinois College of Medicine Rubinacci, Allessandro, University of Milan, Italy Sardet, Christian, Station Zoologique, Villefranche-sur-mer, France Saunders. Mary Jane, University of South Florida, Tampa Schiefelbein, John, University of Michigan Shapiro. James, University of Chicago Smith, Peter, Cambridge University, UK Speksnijder, Johanna, University of Utrecht. The Netherlands Other Year-Round Investigators and Staff Stephens, Raymond E., Principal Investigator Szent-Gyorgy, Gwen, Research Assistant Warren, Lisa, Research Assistant Honors Friday Evening Lectures Fellowships Dennis Powers, Stanford University. Hopkins Marine Station, 29 June "Adapting to a Changing Environment: Genetic and Physiological Mechanisms" Joan Ruderman, Harvard Medical School, 6 July "Controlling Cell Division" James Spudich, Stanford University School of Medicine, 13 July "Manifestation of a Molecular Motor: From Muscle to Amoebae" Ricardo Miledi, University of California, 19, 20 August (Forbes Lectures) "How to Study the Brain Using Frog Oocytes" James Tiedje, Michigan State University, 27 July "Destruction ofPCBs and Other Pollutants by a New Class of Anaerobic Sediment Microorganisms" Ruth Sager, Dana-Farber Cancer Institute, 3 August "Tumor Represser Genes" A. James Hudspeth, University of Texas Southwestern Medical Center, 10 August "How the Ear's Works IVork: Mechanoelectrical Transduction, Frequency Tuning, and Synoptic Transmission by Hair Cells of the Internal Ear" Phil Leder, Harvard Medical School, 17 August "Limb Deformity: A Pleiotropic Mutation Governing Embryonic Pattern Formation in the Mouse" Brian Fry, MBL Ecosystems Center, 24 August "Consumers and Carbon Isotopes: Good Chemistry in the Sea" Thoru Pederson, Worcester Foundation for Experimental Biology, 31 August "Between Nucleus and Cytoplasm: The Baroque Process of Making mRNA as Studied from the Perspective of Cell Biology" Robert Day Allen Fellowship Eugeni A. Vaisberg, Protein Research Institute, USSR Frederick B. Bang Fellowship Fund Elaine Bearer, University of California, San Francisco Jean and Ratsuma Dan Fellowship Douglas E. Koshland (financial support by Japanese Dan Fellowship), Carnegie Institution of Washington M. G. F. Fuortes Fellowship Joseph Charles Callaway, University of North Carolina, Chapel Hill Stephen W. Kuffler Fellowship Joachim W. Deitmer, University of Kaiserslauten, Zoologie, Germany Dieter Swandulla, Max-Planck-Institiit, Germany Hayden-Baille Fellowship Neena Din, University of British Columbia, Canada Anne Donaldson, MRC Laboratory of Molecular Biology, UK Faith Miller Fellowship Elaine L. Bearer, University of California, San Francisco Youngsook Lee, Harvard University 45 46 Annual Report Nikon Fellowship Timothy Mitchison. University of California, San Francisco Science Writing Fellowships Robin Henig. Freelance Bruce Jacobs. Freelance Bill Krasean, Kalamaioo Gazelle June Kinoshita, Freelance Kerning Kuo, Voice of America Beth Livermore, Freelance Michael Skoler, National Public Radio Pamela Weintraub, Omni Rick Weiss, Science News H. Burr Steinbach Memorial Fellowship Youngsook Lee, Harvard University Steps Toward Independence MBL Summer Fellowship Elaine Bearer, University of California, San Francisco Robert Paul Malchow, University of Illinois JoAnn Render, Hamilton College Katherine Swenson, Harvard Medical School Johanna E. Speksnijder, Utrecht University, The Netherlands Eugeni Vaisberg, Protein Research Institute, USSR Scholarships APA Fellow Maria A. Sosa, University of Florida John F. Hamilton, Meharry Medical College ARCS Foundation Fellow Daniel S. Kessler, Rockefeller University ASCB Fellows Abdiel J. Alvarez, University of Puerto Rico Robert L. Bacallao, University of California, Los Angeles Miles G. Cunningham. Massachusetts Institute of Technology Charlotte M. Vines, Harvard Medical School Biology Club of CUNY Elizabeth L. Winter, City College of New York Father Arsenius Boyer Scholarship Fund Haohua Qian. University of Illinois C. Lalor Burdick Scholarship Ann Marie Murphy, Johns Hopkins University Gary N. Calkins Memorial Scholarship Hanno M. Roder, Massachusetts Institute of Technology Frances S. Claff Memorial Scholarship Ka Hou Chu, Chinese University of Hong Kong, Hong Kong Edwin Grant Conklin Memorial Scholarship Sylwester Chyb, Wesleyan University Lucretia Crocker Endowment Fund Silvia E. Frenk, King's College, Cambridge. UK Bernard Davis Scholarship Diane K. Arwood, University of Southern Mississippi Joanna S. Brooke, University of Western Ontario, Canada Joseph P. Calabrese, West Virginia University Judith A. Koskella, New York University Bridget E. Laue, University of Colorado Jared R. Leadbetter, Goucher College Shi Liu, University of Oklahoma Lynn V. Mendelman, Harvard Medical School Elizabeth J. Ode, Colorado State University Mechthild Pohlschroder, University of Massachusetts, Amherst William F. and Irene Diller Scholarship Fund Sylwester Chyb, Wesleyan University Caswell Grave Scholarship Ricardo Araneda. Albert Einstein College of Medicine Sylwester Chyb, Wesleyan University Kareen M. Guida. University of Paris, France Aline D. Gross Scholarship Anne Marie Murphy. Johns Hopkins University Arthur Klorfein Fund Scholarship Dangeruta Kersulyte, Academy of Science, Lithuania, USSR Honors 47 Thierry Lepage, University of Nice, France Kareen M. Guida. University of Paris, France Edwin S. Linton Memorial Scholarship Michael C. Schmale, University of Miami S. O. Mast Founders Scholarship Peter J. Edmunds. Northeastern University Ka Hou Chu, Chinese University of Hong Kong, Hong Kong Center for Microbial Ecology, Michigan State Scholarship Olivia T. Harriott, University of Connecticut Alberto Monroy Fellow Maria G. DiBernardo, Institute di Biologia dello Sviluppo, CNS, Italy James S. Mountain Memorial Fund Scholarship Karen M. Page, Dartmouth College Eric A. Shelden, University of Massachusetts, Amherst Salme Taagepera, University of Virginia Sandra A. Brockman, Carnegie-Mellon University Tod A. Critchlow, Scripps Institute of Oceanography Planetary Biology Internship Jennifer E. Klenz, University of Saskatchewan. Canada William Townsend Porter Foundation Fellowship Abdiel J. Alvarez, University of Puerto Rico Robert L. Bacallao, University of California, Los Angeles Miles G. Cunningham, Massachusetts Institute of Technology Domingo T. Rivera, Worcester Foundation for Experimental Biology Charlotte M. Vines, Harvard Medical School Ebenezer Yamoah, University of Alberta, Canada Herbert \V. Rand Scholarship Brian J. Binder, Massachusetts Institute of Technology Sandra A. Brockman, Carnegie-Mellon University Isabelle Carre, SUNY, Stony Brook Joseph Cerro, Columbia University Ka Hou Chu, Chinese University of Hong Kong, Hong Kong Tod A. Critchlow, Scripps Institute of Oceanography Michele I. Flatters, Tufts University Holly Y. Goodson, Stanford University Jerilyn Jewett-Smith, Whitman College John R. Jordan, University of Utah David L. Keefe, Yale University Karen L. King, Florida State University Qingwen Li, University of Kansas Helen McNeill, University of Pennsylvania Christa S. Merzdorf. Harvard University Robert Mirro, University of Tennessee Qi Yang, University of Connecticut Society for General Physiologists Scholarships Robert A. Berkowitz. Washington University Daniel H. Cox, Tufts University Holly Y. Goodson, Stanford University Daniel S. Kessler, Rockefeller University Marjorie W. Stetten Fund Haohua Qian. University of Illinois Surdna Foundation Scholarship Peter J. Edmunds, Northeastern University Oivind Enger, University of Bergen, Norway Robert E. Hodson, University of Georgia James S. Maki, Harvard University Martin Polz. University of Vienna. Austria Stephen C. Tsoi, University of Hong Kong, Hong Kong William Morton Wheeler Family Founders' Scholarship Hanno M. Roder, Massachusetts Institute of Technology Awards Lewis Thomas Award Natalie Angier, The New York Times MBL Tour Guides Award for Outstanding Science Presentation to the General Public Nancy Rafferty, Northwestern University Seymour Zigman, University of Rochester School of Medicine Board of Trustees and Committees Corporation Officers and Trustees Ex officio Honorary Chairman of the Board of Trustees, Denis M. Robinson. Key Biscayne. FL Chairman of the Board of Trustees, Prosser Gifford. Washington. DC President of the Corporation and Director. Harlyn O. Halvorson. Marine Biological Laboratory. Woods Hole, MA Treasurer. Robert D. Manz. Helmer & Associates. Waltham. MA Clerk of the Corporation, Kathleen Dunlap, Tufts University School of Medicine. Boston, MA Class of 1994 Trustees Robert D. Goldman, Northwestern University, Chicago, IL Rodolfo R. Llinas. New York University Medical Center, New York. NY Thomas D. Pollard, Johns Hopkins University, Baltimore. MD Joan V. Ruderman, Harvard University School of Medicine, Boston, MA Joseph Sanger, University of Pennsylvania School of Medicine. Philadelphia, PA Ann Stuart, Marine Biological Laboratory. Woods Hole, MA Class of 1994 Trustees-at- Large Frederick Bay. Gaston Snow Beekman & Bogue, New York. NY Mary-Ellen Cunningham. Grosse Pointe Farms, MI Robert W. Pierce, Boca Grande, FL Irving Rabb, University Place at Harvard Square, Cambridge. MA Class of 1993 Garland E. Allen, Washington University, St. Louis. MO Jelle Atema, Manne Biological Laboratory, Woods Hole, MA William L. Brown, Weston, MA Alexander W. Clowes, University of Washington School of Medicine. Seattle, WA Barbara Ehrlich, University of Connecticut. Farmington, CT Edward A. Kravitz, Harvard Medical School. Boston. MA Robert E. Mainer, The Boston Company, Boston, MA Jerry M. Melillo. Marine Biological Laboratory, Woods Hole, MA Roger D. Sloboda. Dartmouth College, Hanover, NH Class of 1992 Norman Bernstein, Bernstein Management, Inc., Washington, DC Ellen R. Grass, Grass Foundation, Quincy, MA Neil Jacobs, Hale & Dorr, Boston. MA Sir Hans Kornberg, Christ's College, Cambridge, UK George Langford, University of North Carolina, Chapel Hill. NC Jack Levin, V.A. Medical Center. San Francisco, CA Evelyn Spiegel, Dartmouth College, Hanover, NH Andrew G. Szent-Gyorgyi. Brandeis University, Waltham, MA Kensal E. VanHolde, Oregon State University, Corvallis. OR Stanley W. Watson. Associates of Cape Cod. Inc.. Falmouth, MA Class of 1991 Robert B. Barlow Jr.. Syracuse University, Syracuse, NY Dieter Blennemann, Carl Zeiss, Inc., Thornwood, NY James M. Clark, Palm Beach, FL Wensley G. Haydon-Baillie, Porton, Int., London. UK Laszlo Lorand, Northwestern University, Evanston. IL Lionel I. Rebhun. University of Virginia, Charlottesville, VA Carol L. Reinisch, Tufts University School of Veterinary Medicine. Boston, MA Brian M. Salzberg, University of Pennsylvania. Philadelphia, PA Sheldon J. Segal, The Rockefeller Foundation. New York, NY Emeriti John B. Buck, NIH, Bethesda. MD Seymour S. Cohen. Woods Hole, MA Arthur L. Colwin, Key Biscayne, FL Laura Hunter Colwin, Key Biscayne. FL D. Eugene Copeland. MBL, Woods Hole, MA Sears Crowell. Indiana University, Bloomington, IN Alexander T. Daignault, Boston, MA William T. Golden, New York, NY Teru Hayashi, Woods Hole, MA Ruth Hubbard. Cambridge, MA Lewis Kleinholz, Reed College. Portland. OR Maurice E. Krahl, Tucson, AZ Charles B. Metz, Miami, FL Keith R. Porter, University of Pennsylvania, Philadelphia, PA C. Ladd Prosser, University of Illinois. Urbana. IL S. Meryl Rose. E. Falmouth, MA W. D. Russell-Hunter, Syracuse University, Syracuse, NY John Saunders, Jr., Waquoit, MA Mary Sears, Woods Hole. MA Homer P. Smith, Woods Hole, MA W. Randolph Taylor, Ann Arbor, MI D. Thomas Tngg, Wellesley, MA Walter S. Vincent, Woods Hole, MA George Wald, Cambridge, MA 48 Trustees and Committees 49 Executive Committee of the Board of Trustees Prosser Gifford*. Chairman Robert B. Barlow Jr.. 1991 Ray L. Epstein* Robert D. Goldman, 1994 Harlyn O. Halvorson* Robert E. Mainer. 1993 Robert D. Manz* Jerry M. Melillo. 1992 Irving W. Rabb, 1991 Sheldon J. Segal. 1992 D. Thomas Tngg, 1990 Trustee Committees 1990 Audit Robert Mainer. Chairman Ray L. Epstein* Robert D. Manz* Sheldon J. Segal Andrew G. Szent-Gyorgyi D. Thomas Tngg Kensal E. Van Holde Stanley W. Watson Compensation Prosser Gifford, Chairman Robert E. Mainer Robert D. Manz Irving W. Rabb D. Thomas Tngg Investment D. Thomas Trigg, Chairman William L. Brown Ray L. Epstein* William T. Golden Maunce Lazarus Werner R. Loewenstein Robert D. Manz* Irving W. Rabb W. Nicholas Thorndike Standing Committees for the Year 1990 Buildings & Grounds Institutional Animal Care and Use Craig Dorman. WHOI Ray Epstein Kenyon Tweedell. Chairman Leslie D. Garrick*. Chairman George Grice, WHOI Alfred Chaet Robert B. Bullis John W. Speer* Lawrence B. Cohen Alfred Chaet Gary Walker. WHOI Richard Cutler* Ray L. Epstein* Alan Fein Ferenc Harosi Edward Jaskun Andrew Mattox* Library Joint Users Donald B. Lehy* Thomas Meedel Philip Person Evelyn Spiegel Fellowships Thoru Pederson, Chairman Martha Constantme-Paton Ray L. Epstein* Leslie D. Garrick* Ann Giblin George M. Langford Eduardo Macagno Carol L. Reinisch J. Richard Whittaker Housing, Food Service, and Child Care Thomas Reese. Chairman Jelle Atema Andrew Bass Susan Barry Donald Chang Richard Cutler* Stephen Highstein LouAnn King* ex officio Institutional Biosafety Raquel Sussman. Chairman Paul De Weer Paul Englund Harlyn O. Halvorson* Paul Lee Donald B. Lehy* Joseph Martyna Andrew Mattox* Alfred W. Senft Instruction Roger Sloboda. Chairman Ray L. Epstein* Rachel Fink Leslie D. Garrick* Leah Haimo Ron Hoy Hans Laufer Joan Ruderman Robert Silver Ray Stephens Library Joint Management Harlyn O. Halvorson*, Chairman Garland Allen Garland Allen. Chairman Henry J. B. Dick. WHOI A. Farmanfarmaian Lionel Jafle Catherine Norton* John Teal. WHOI Geoff Thomson, WHOI Page Valentine, USGS Carole Winn*. WHOI Marine Resources Robert Goldman. Chairman Donald Abt William Cohen Richard Cutler* Toshio Narahashi George Pappas Roger Sloboda Melvin Spiegel Antoinette Steinacher John Valois* Radiation Safety Ete Z. Szutz, Chairman David W. Borst Richard L. Chappell Sherwin J. Cooperstein Louis M. Ken- Andrew Mattox* Robert Rakowski Walter Vincent 50 Annual Report Research Services Robert Palazzo Robert Silver Paul Steadier Steven Treistman Brian Noe, Chairman , van Valie|a Peter Armstrong Richard Va ,, ee Robert B. Barlow jr. Research Space Richard Cutler* Safety Barbara Ehrhch Jose P h San 8 er - Chairman Ken Forman ' 3au ' ^ e Wecr J nn Hobbie, Chairman Ehud Kaplan Ra y L - Epstein* D. Eugene Copeland Samuel S Koide Leslie D. Garnck* Richard Cutler* Aimlee Laderman John Hildebrand Edward Enos* Jack Levin David Landowne Louis Kerr Andrew Mattox* Hans Laufer Alan Kuzirian Eduardo Macagno Donald B. Lehy* Jerry M. Melillo Andrew Mattox* *c\v afficio Joan V. Ruderman Paul Steudler Laboratory Support Staff* Biological Bulletin Clapp. Pamela L., Managing Editor Puckett. Kathryn Stone, Beth Ready Computer Services Tollios, Constantine D.. Manager Schmidt. Valerie Controller's Office Speer. John W.. Controller Accounting Services Binda. Ellen F. Campbell. Ruth B. Davis. Doris C. Ghetti. Pamela M, Gilmore. Mary F. Hobbs. Roger W.. Jr. Hough, Rose A. Oliver. Elizabeth Poravas. Maria Riley, Janis E. Chein Room Chisholm. Caroline G. Miller. Lisa A. Sadowski. Edward A. Purchasing Hall. Lionel E.. Jr. Schorer. Timothy M. Copy Service Center Mountlbrd. Rebecca J.. Supervisor Jackson. Jacquelyn F. Ridley, Sherie Including persons who joined or left I he staff during 1990. Development Office Avers, Donald E.. Director Berthel, Dorothy Lessard, Kelley J. Thimas, Lisa M. Director 's Office Halvorson. Harlyn O., President and Director Burrhus, I. Elaine Catania. Didia Epstein, Ray L. Kinneally. Kathleen R. Power. Linda M. Watkins, Joan E. Gray Museum Backus, Richard H., Curator Armstrong, Ellen P. Montiero. Eva Housing King. LouAnn D.. Conference Center and Housing Manager Barnes. Susan M. Farrell, Bermee R. Johnson, Frances N. Krajewski. Viola I. Kuil, Elisabeth Mancevice, Denise M. McNamara, Noreen Sadovsky, Sebastian Telephone Office Baker, Ida M. Geggatt. Agnes L. Ridley. Alberta W. Human Resources Goux, Susan P., Manager Donovan. Murcia H. Library Norton, Catherine N., Acting Librarian Ashmore, Judith A. Costa, Marguerite E. Fessenden. Jane Fisher, Susan Goux, Randal Huguenin. Sanders Keenan, Patrick M. Mirra, Anthon> J. Mountford. Rebecca J. Nelson. Heidi Pratson, Patricia G. Showalter, Christine M. Tamm, Ingrid deVeer. Joseph M. Wnght. Rosemary MBL Associates Liaison Scanlon, Deborah Public Information Office Liles, George W., Jr., Director Anderson, Judith L. Kaye-Peterson, Amy Stone, Beth Ready Safety Services Mattox, Andrew H.. Safety Officer Apparatus Barnes. Franklin D. Haskins, William A. Martin. Lowell V. Nichols, Francis H., Jr. Shipping and Receiving Geggatt. Richard E. Illgen, Robert F. Monteiro, Dana 52 Annual Report Services, Projects, and Facilities Cutler. Richard D.. Manager Enos, Joyce B. Kurland. Charles J. Buildings and Grounds Lehy, Donald B., Superintendent Baldic. David P. Blumsack. Jeffrey J. Blunt, Hugh F. Bourgoin, Lee E. Carini, Robert J. Fish, David L., Jr. Fuglister, Charles K. Gonsalves, Walter W., Jr. Hathaway, Peter Jones, Leeland Justason. C. Scott Lochhead, William M. Lunn. Alan G. MacLeod. John B Me Adams. Herbert M., Ill Mills, Stephen A. Olive, Charles W.. Jr. Rattacasa. Frank D. Schoepf. Claude deVeer, Robert L. Weeks, Gordon W. Electron Microscopy Lab Kerr. Louis M. Housekeeping Services Jerry Phillabaum, General Supervisor Allen, Wayne D. Anderson, Lewis B. Boucher, Richard L. Collins, Paul J Conlin, Henry P. Gibbons. Roberto G. Krajewski, Chester J. Lynch, Henry L. Instrument Development I. til' Robert Knudson Machine Shop Sylvia, Frank E. Marine Resources Center Valois, John J.. Manager Enos. Edward G.. Jr.. Superintendent Fisher, Harry T.. Jr. Hanley. Janice S. Moniz, Pnscilla C. Monteiro. Dana Revellese, Christopher Sullivan, Daniel A. Tassinari, Eugene Torres, Sophie J. Photolab Colder, Linda M. Colder, Robert J. Rugh, Douglas E. Sponsored Programs Garnck, Leslie D.. Administrator Dwane. Florence Huffer. Linda Lynch, Kathleen F. Animal Care Facility Hanley. Janice S. Shephard, Jennifer Summer Support Stall' Albrecht, Helen Amon, Tyler C. Anderson, Penny Ashmore. Lynne E. Avers, Andrew D. Bolton. Hugh Burke, Sean Cadwalader. George, Jr. Campbell, Robert E. Capobianco, James A. Child, Malcolm S. Cishek. Dawn Clinard, Nathan Cullen, Timothy Cutler, Laura Demir, Oktay Dias, John. Ill Donovan, Erin Donovan, Jason P. Ford, John. Ill Grassle, J. Thomas Halpm. Michelle Hamilton, Elizabeth R. Hibhitt, Catherine Hrynyshyn, James Hullum, Rebekah Illgen, Robert C. Kaplan. Liat Kinneally, Kara J. Langton, Lori Leatherbee. Amy Marini. Michael F. Montroll, Charles Northern, Marc D. Peal, Jennifer Peal, Richard W. Philbm, Linda M. Phillips, Daniel Piazza. Lucia Remsen, Andrew S. Roderick, Paul Rosenkranz, Margalil St. Jean, Jeannette Scherer, Aimee Sheetz, Jonathan Sheffield. James Shepherd. Jennifer Shock, Duane Snyder. Matthew Snyder, Rebecca Sofferman. Rebecca Swope. John G. Tilghman. Alison Varao. John Vardac. Michael Wetzel, Ernest D. Wilkes, Jennifer Members of the Corporation' Life Members Abbott, Marie, c/o Vaughn Abbott. Flyer Rd., East Hartland, CT 06027 Beams, Harold W., Department of Biology, University of Iowa, Iowa City, IA 52242 Bernheimer, Alan \V., Department of Microbiology, New York University Medical Center, 550 First Ave., New York, NY 10016 Bertholf, Lloyd M., Westminster Village #2114, 2025 E. Lincoln St., Bloomington. IL 61701 Bodian, David, 4100 North Charles St., #913, Baltimore, MD 21218 Bridgman, A. Josephine, 715 Kirk Rd., Decatur, GA 30030 Buck, John B., NIH, Laboratory of Physical Biology. Room 1 12. Building 6 Bethesda, MD 20892 Burbanck, Madeline P., Box 15134, Atlanta, GA 30333 Burbanck, William D., Box 15134. Atlanta. GA 30333 Carpenter, Russell L., 60-H Lake St., Winchester, MA 01890 Chase, Aurin, Department of Biology, Guyot Hall, Princeton University, Princeton, NJ 08544 (resigned) Clark, Arnold M., 53 Wilson Rd.. Woods Hole, MA 02543 Cohen, Seymour S., 10 Carrot Hill Rd., Woods Hole, MA 02543-1206 Colwin, Arthur, 320 Woodcrest Rd., Key Biscayne, FL 33149 Colwin, Laura Hunter, 320 Woodcrest, Key Biscayne, FL 33149 Copeland, D. E., 41 Fern Lane. Woods Hole. MA 02543 Costello, Helen M., Carolina Meadows. Villa 137. Chapel Hill, NC 27514 Crouse, Helen, Address unknown Kailla, Patricia M., 2149 Loblolly Lane, Johns Island. SC 29455 Ferguson, James K. \V.. 56 Clarkehaven St., Thornhill, Ontario L4J 2B4 Canada Fries, Erik F., 41 High Street, Woods Hole. MA 02543 (deceased) Goldman, David, 63 Loop Rd., Falmouth, MA 02540 Graham, Herbert, 36 Wilson Rd.. Woods Hole, MA 02543 Green, James W., 409 Grant Ave., Highland Park, NJ 08904 Grosch, Daniel S., 1222 Duplin Road. Raleigh, NC 27607 Hamburger, Viktor, Department of Biology, Washington LIniversity. St. Louis, MO 63 130 Hamilton, Howard L., Department of Biology, University of Virginia. 238 Gilmer Hall. Charlottesville, VA 22901 Harding, Clifford V., Jr., Wayne State University School of Medicine. Department of Ophthalmology, Detroit, MI 48201 Haschemeyer, Audrey E. V., 2 1 Glendon Road, Woods Hole, MA 02543 Hauschka, Theodore S., FD1, Box 781, Damariscotta. ME 04543 Hisaw, F. L., 5925 SW Plymouth Drive, Corvallis, OR 97330 * Including action of the 1990 Annual Meeting. Hollaender, Alexander, Council for Research Planning, 1717 Massachusetts Ave. NW. Washington, DC 20036 (deceased) 1 1 ul>l>:irn. William D., Department of Zoology. Ohio University, Athens. OH 45701 Humphreys, Susie H., Research & Development. Kraft. Inc., 801 Waukegan Rd., Glenview, IL 60025 Humphreys, Tom D., L'niversity of Hawaii. PBRC, 41 Ahui St., Honolulu, HI 96813 Hunter, Robert D., Department of Biological Sciences, Oakland University, Rochester. MI 48309-4401 Hunter, W. Bruce, Box 321. Lincoln Center, MA 01773 Hurwitz, Charles, Basic Science Research Lab, Veterans Administration Hospital, Albany, NY 12208 Hurwitz, Jerard, Sloan Kettenng Institute for Cancer Research, 1275 York Avenue. New York. NY 1 1021 Huxley, Hugh E., Department of Biology, Rosenstiel Center, Brandeis University, Waltham, MA 02154 Hynes, Thomas J., Jr., Meredith and Grew, Inc., 160 Federal Street, Boston. MA 021 10-1701 Han, Joseph, Department of Developmental Genetics and Anatomy, Case Western Reserve University School of Medicine, Cleveland, OH 44 106 Ingoglia, Nicholas, Department of Physiology, New Jersey Medical School, 100 Bergen St., Newark, NJ 07103 Inoue, Saduyki, Department of Anatomy, McGill University Cancer Centre, 3640 University St.. Montreal, PQ, H3A 2B2, Canada Inoue, Shinya, Marine Biological Laboratory, Woods Hole, MA 02543 Isselbacher, Kurt J., Massachusetts General Hospital, 32 Fruit Street, Boston, MA 02114 Issidorides, Marietta, R., Department of Psychiatry, University of Athens, Moms Petraki 8, Athens, 140 Greece Izzard, Colin S., Department of Biological Sciences, SUNY, 1400 Washington Ave., Albany, NY 12222 Jacobs, Neil, Hale & Dorr, 60 State St.. Boston. MA 02019 Jacobson, Antone G., Department of Zoology, University of Texas, Austin, TX 78712 Jaffe, Lionel, Marine Biological Laboratory, Woods Hole. MA 02543 Jannasch, Holger W., Department of Biology, Woods Hole Oceanographic Institution, Woods Hole. MA 02543 Jeffery, William R., Bodega Marine Laboratory. Box 247, Bodega Bay, CA 94923 Jones, Meredith L., Division of Worms, Museum of Natural History, Smithsonian Institution. Washington, DC 20560 Josephson, Robert K., Department of Biological Sciences, University of California, Irvine, CA 92717 Members of the Corporation 59 Kabat, E. A., Department of Microbiology, College of Physicians and Surgeons, Columbia University, 630 West 168th St., New York, NY 10032 Kaley, Gabor, Department of Physiology, Basic Sciences Building, New York Medical College. Valhalla, NY 10595 kaltenbach, Jane, Department of Biological Sciences, Mount Holyoke College. South Hadley, MA 01075 kaminer. Benjamin, Department of Physiology. School of Medicine, Boston University, 80 East Concord St., Boston. MA 021 IS Kane, Robert E., PBRC. University of Hawaii. 41 Ahui St., Honolulu. HI 96813 kaneshiro, Edna S., Department of Biological Sciences. University of Cincinnati. Cincinnati. OH 45221 kao, Chien-yuan, Department of Pharmacology, Box 29, SUNY, Downstate Medical Center. 450 Clarkson Avenue. Brooklyn. NY 11203 kaplan, Ehud, Department of Biophysics. The Rockefeller University, 1230 York Ave., New York, NY 10024 karakashian, Stephen J., Apt. 16-F. 165 West 91st St., New York. NY 10024 km INI. Arthur, Department of Biochemistry and Neurology, Columbia University. 630 West 168th St.. New York, NY 10032 Katz, George M., Fundamental and Experimental Research Labs. Merck Sharp and Dohme. P. O. Box 2000. Rahway. NJ 07065 kelly, Robert E., Department of Anatomy. College of Medicine. University of Illinois. P. O. Box 6998. Chicago, IL 60680 kemp, Norman E., Department of Biology, University of Michigan, Ann Arbor, MI 48109 kendall, John P., Faneuil Hall Associates, 176 Federal Street, 2nd Floor, Boston, MA 02110 kendall, Richard E., Commissioner of Environmental Management, 100 Cambridge Street, Room 1905, Boston, MA 02202 kerr, Louis M., Marine Biological Laboratory, Woods Hole, MA 02543 keynan, Alexander, Hebrew University. Jerusalem, Israel kiehart, Daniel P., Department of Cellular and Developmental Biology, Harvard University. 16 Divinity Street, Cambridge, MA 02138 kirk, Mark D., Division of Biological Sciences. University of Missouri, Columbia, MO 652 I 1 klein, Morton, Department of Microbiology, Temple University Medical School, Philadelphia, PA 19103 klotz, Irving M., Department of Chemistry, Northwestern University, Evanston. IL 60201 knudson, Robert A., Marine Biological Laboratory. Woods Hole, MA 02543 koide, Samuel S., Population Council, The Rockefeller University, 1230 York Avenue, New York, NY 10021 kornberg. Sir Hans, The Master's Lodge, Christ's College. Cambridge CB2 3BU. England. UK kosower, Edward M., Ramat-Aviv. Tel Aviv, 69978 Israel krahl, M. E., 2783 W. Casas Circle. Tucson, AZ 85741 krane, Stephen M., Arthritis Unit, Massachusetts General Hospital, Fruit Street, Boston, MA 021 14 krauss, Robert, FASEB, 9650 Rockville Pike, Bethesda. MD 20814 kravitz, Edward A., Department of Neurobiology. Harvard Medical School, 220 Longwood Ave., Boston, MA 02 1 1 5 kriebel, Mahlon E., Department of Physiology, SUNY Health Science Center, Syracuse, NY 13210 kristan, William B., Jr., Department of Biology B-022, University of California San Diego, La Jolla, CA 92093 kropinski, Andrew M. B., Department of Microbiology/Immunology, Queen's University. Kingston. Ontario K7L 3N6, Canada kuhns, William ,1., Hospital for Sick Children, Department of Biochemistry Research, Toronto, Ontario M5G 1X8, Canada kuhtruiber, Willem M., Marine Biological Laboratory, Woods Hole. MA 02543 kusano, kiyoshi, NIH. Bldg. 36. Room 4D-20, Bethesda. MD 20892 kuzirian, Alan M., Marine Biological Laboratory. Woods Hole. MA 02543 Laderman. Aimlee, Yale University, New Haven, CT 06520 LaMarche, Paul H., Eastern Maine Medical Center. 489 State St., Bangor. ME 04401 I ainlis. Dennis M. D., Department of Developmental Genetics and Anatomy, Case Western Reserve University School of Medicine, Cleveland. OH 44106 Landowne, David, Department of Physiology. P. O. Box 016430, University of Miami School ot Medicine. Miami. FL 33101 Langford, George M., Department of Physiology. CB7545 University of North Carolina School of Medicine. Chapel Hill. NC 27599-7545 Laster, Leonard, University of Massachusetts Medical School. 55 Lake Avenue. North, Worcester. MA 01655 Laufer, Hans. Department of Biological Science, Molecular and Cell Biology, Group U-125, University of Connecticut, Storrs, CT 06268 Lazarow, Paul B., Department of Cell Biology and Anatomy, Mount Sinai Medical School. Box 1007, 5th Avenue & 100th Street, New York, NY 10021 Lazarus, Maurice, Federated Department Stores, Inc.. Sears Cresent, City Hall Plaza. Boston. MA 02108 Leadbetter, Edward R., Department of Molecular and Cell Biology, U-131, University of Connecticut, Storrs, CT 06268 Lederbcrg, Joshua, The Rockefeller LIniversity, 1230 York Ave.. New York. NY 10021 Lee, John J., Department of Biology, City College of CLINY, Convent Ave. and 138th St.. New York, NY 10031 Lehy, Donald B., Marine Biological Laboratory, Woods Hole, MA 02543 Leibovitz, Louis, 3 Kettle Hole Road, Woods Hole, MA 02543 Leighton, Joseph, 2324 Lakeshore Avenue. #2, Oakland, CA 94606 Leighton, Stephen, NIH, Bldg. 13 3WI3, Bethesda. MD 20892 Leinwand, Leslie Ann, Department of Microbiology and Immunology. Albert Einstein College of Medicine, 1 300 Morris Park Ave., Bronx, NY 10461 Lerman, Sidney, Eye Research Lab. Room 41, New York Medical College, 100 Grasslands Ave., Valhalla. NY 10595 Lerner, Aaron B., Yale University, School of Medicine, New Haven, CT06510 Lester, Henry A., 156-29 California Institute of Technology, 156-29, Pasadena, CA 91 125 Levin, Jack, Veterans Administration Medical Center, I 13 A. 4150 Clement St., San Francisco, CA 94 1 2 1 Levinthal, Cyrus, Department of Biological Sciences, Columbia University. Broadway and 1 16th Street. New York. NY 10026 (deceased) Levitan, Herbert, Department of Zoology, University of Maryland, College Park, MD 20742 Levitan, Irwin B., Department of Biochemistry, Brandeis University, Waltham, MA 02254 Linck, Richard \\ '., Department of Anatomy, Jackson Hall, University of Minnesota, 321 Church Street. S. E., Minneapolis. MN 55455 Lipicky, Raymond J., Department of Cardio-Renal/Drug Prod. Div., FDA, Rm. 16B-45, 5600 Fishers Lane, Rockville, MD 20857 Lisman, John E., Department of Biology, Brandeis LIniversity, Waltham, MA 02254 Liuzzi, Anthony, 55 Fay Rd., Box 184. Woods Hole. MA 02543 60 Annual Report Llinas. Rodolfo R., Department of Physiology and Biophysics, New York University Medical Center. 550 First Ave., New York, NY 10016 Loew, Franklin M., Tufts University School of Veterinary Medicine, 200 Westboro Rd.. N. Grafton. MA 01536 Loewenstein, Birgit R., Department of Physiology and Biophysics, R- 430. University of Miami School of Medicine, Miami, FL 33101 Loewenstein, Werner R., Department of Physiology and Biophysics, University of Miami. P. O. Box 016430. Miami, FL 33101 London, Irving M., Massachusetts Institute of Technology, E-25-551. Cambridge, MA 02 139 Longo, Frank J., Department of Anatomy, University of Iowa, Iowa City. IA 52442 Lorand, Laszlo, Department of Biochemistry and Molecular Biology, Northwestern University, 2153 Sheridan Road. Evanston, IL 60208 Luckenbill-Edds, Louise, 155 Columbia Ave., Athens, OH 45701 Luria, Salvador E., 48 Peacock Farm Rd., Lexington, MA 02173 (deceased) Macagno, Lduardo R., 1003B Fairchild, Department of Biosciences, Columbia University. New York. NY 10027 MacNichol, E. F., Jr., Department of Physiology. Boston University School of Medicine, 80 E. Concord St., Boston, MA 021 18 Maglott-Duffield, Donna R., American Type Culture Collection, 12301 Parklawn Drive, Rockville, MD 20852-1776 Maienschein, Jane Ann, Department of Philosophy. Arizona State University, Tempe, AZ 85287-2004 Mainer, Robert, The Boston Company, One Boston Place. OBP-15-D, Boston. MA 02 168 Malbon, Craig Curtis, Department of Pharmacology. Health Sciences Center, SUNY. Stony Brook, NY 1 1744-8651 Manalis, Richard S., Department of Biological Sciences, Indiana University Purdue University at Fort Wayne, 2101 Coliseum Blvd.. E. Fort Wayne. IN 46805 Mangum, Charlotte P., Department of Biology, College of William and Mary. Williamsburg, VA 23185 Manz, Robert D., Helmer and Associates, Suite 1310. 950 Winter St., Walthan, MA 02 154 Margulis, Lynn, Botany Department, University of Massachusetts. Mornll Science Center. Amherst. MA 01003 Marinucci, Andrew C, 102 Nancy Dnve, Mercerville, NJ 08619 Marsh, Julian B., Department of Biochemistry and Physiology. Medical College of Pennsylvania, 3300 Henry Ave., Philadelphia. PA 19129 Martin, Lowell V'., Marine Biological Laboratory, Woods Hole, MA 02543 Martinez-Palomo, Adolfo, Seccion de Patologia Experimental, Cinvesav-ipn, 07000 Mexico. D.F. A. P.. 140740, Mexico Maser, Morton, Woods Hole Education Assoc., P. O. Box EM. Woods Hole. MA 02543 Mastroianni, Luigi, Jr., Department of Obstetrics and Gynecology, Hospital of the University of Pennsylvania, 106 Dulles, 3400 Spruce Street, Philadelphia. PA 19104-4283 Matteson. Donald R., Department of Biophysics, University of Man-land School of Medicine, 660 Redwood Street, Baltimore, MD 21201 Mautner, Henry G., Department of Biochemistry, Tufts University School of Medicine, 136 Harrison Ave., Boston, MA 021 1 1 Mauzerall. David. The Rockefeller University, 1230 York Ave., New York, NY 10021 McCann, Frances, Department of Physiology, Dartmouth Medical School, Hanover, NH 03755 McLaughlin, Jane A., Marine Biological Laboratory, Woods Hole, MA 02543 McMahon, Robert F., Department of Biology. Box 19498. University of Texas. Arlington. TX 76019 Meedel, Thomas, Marine Biological Laboratory. Woods Hole, MA 02543 Meinert/hagen, Ian A., Department of Psychology, Life Sciences Center. Dalhousie University, Halifax, Nova Scotia B3H 451. Canada Meiss, Dennis E., 462 Soland Avenue, Hayward, CA 94541 Melillo, Jerry A., Ecosystems Center, Marine Biological Laboratory, Woods Hole, MA 02543 Mellon, DeForest, Jr., Department of Biology. Gilmer Hall. University of Virginia, Charlottesville, VA 22903 Mellon, Richard P., P. O. Box 187. Laughlintown. PA 15655 Metu/.als, Janis, Department of Pathology. University of Ottawa, Ottawa, Ontario, K1H 8M5 Canada Metz, Charles B., 7220 SW 124th St.. Miami. FL 33156 Milkman, Roger, Department of Biology. University of Iowa, Iowa City, IA 52242 Mills, Robert, 10315 44th Avenue. W 12 H Street. Bradenton. FL 33507-1535 Mitchell. Ralph, DAS. Harvard University, 29 Oxford Street. Cambridge. MA 02 1 38 Miyamoto, David M., Department of Biology. Drew University, Madison, NJ 07940 Mizell, Merle, Laboratory of Tumor Cell Biology, Tulane University, New Orleans, LA 70118 Moore, John W., Department of Neurobiology, Box 3209, Duke University Medical Center, Durham, NC 27710 Moore, Lee E., Department of Physiology and Biophysics, University of Texas Medical Branch, Galveston, TX 77550 Morin, James G., Department of Biology, University of California, Los Angeles, CA 90024 Murrell, Frank, Department of Neurological Science. Rush Medical Center. 1753 W. Congress Parkway, Chicago, IL 60612 Morse, M. Patricia, Marine Science Center, Northeastern University, Nahant, MA 01 908 Morse, Robert W., Box 574. N. Falmouth. MA 02556 Morse, Stephen Scott, The Rockefeller University, 1230 York Ave., Box 2. New York, NY 10021-6399 Mote, Michael I., Department of Biology. Temple University. Philadelphia. PA 19122 Mountain, Isabel, Vinson Hall #112, 6251 Old Dominion Drive, McLean, VA 22101 Muller, Kenneth J., Department of Physiology and Biophysics. University of Miami School of Medicine. Miami. FL 33101 Murray, Sandra Ann, Department of Neurology, Anatomy and Cell Science, University of Pittsburgh School of Medicine, Pittsburgh, PA 15261 Musacchia, Xavier J., Department of Physiology and Biophysics, University of Louisville School of Medicine, Louisville, KY 40292 Nabrit, S. M., 686 Beckwith St., SW, Atlanta. GA 30314 Nadelhoffer, knute. Marine Biological Laboratory. Woods Hole. MA 02543 Naka, Ken-ichi, 2-9-2 Tatsumi Higashi. Okazaki, Japan 444 Nakajima, Shigehiro, Department of Pharmacology and Cell Biology, University of Illinois College of Medicine at Chicago. 808 S. Wolcott Street, Chicago. IL 60612 Nakajima, Yasuko, Department of Anatomy and Cell Biology. University of Illinois College of Medicine at Chicago. M/C 512, Chicago, IL60612 Narahashi, Toshio, Department of Pharmacology, Northwestern University Medical School, 303 East Chicago Ave., Chicago. IL 60611 Members of the Corporation 61 Nasatir, Maimon, Department of Biology. University of Toledo, Toledo. OH 43606 Nelson, Leonard, Department of Physiology. CS10008, Medical College of Ohio. Toledo, OH 43699 Nelson, Margaret C., Section of Neurohiology and Behavior. Cornell University. Ithaca. NY 14850 Nicholls, John G., Biocenter. Klingelbergstrasse 70. Basel 4056. Switzerland Nickerson, Peter A., Department of Pathology, SUNY, Buffalo. NY 14214 Nicosia, Santo V., Department of Pathology. University of South Florida. College of Medicine. Box 11. 12901 North 30th St., Tampa. FL 33612 Noe, Bryan D., Department of Anatomy and Cell Biology, Emory University School of Medicine, Atlanta, GA 30322 Northcutt, R. Glenn, Department of Neuroscience, A-001. Scripps Institution of Oceanography, La Jolla, CA 92093-0201 Norton, Catherine N., Marine Biological Laboratory. Woods Hole. MA 02543 Obaid, Ana Lia, Department of Physiology and Pharmacy, University of Pennsylvania School of Medicine, B-400 Richards Bldg. Philadelphia. PA 19104-6085 Oertel, Donata, Department of Neurophysiology. University of Wisconsin. 281 Medical Science Bldg.. Madison. WI 53706 O'Herron, Jonathan, 45 Swifts Lane. Darien. CT 06820 Ohki, Shinpei, Department of Biophysical Sciences, SUNY at Buffalo, 224 Can Hall, Buffalo, NY 14214 Olins, Ada L., University of Tennessee-Oak Ridge. Graduate School of Biomedical Sciences, Biology Division ORNL, P. O. Box 2009, Oak Ridge. TN 37831-8077 Olins, Donald E., University of Tennessee-Oak Ridge, Graduate School of Biomedical Sciences. Biology Division ORNL, P. O. Box 2009, Oak Ridge, TN 37831-8077 O'Melia. Anne F., 16 Evergreen Lane, Chappaqua, New York 10514 Oschman, James L., 31 Whittier Street, Dover, NH 03820 Palazzo, Robert E., Marine Biological Laboratory. Woods Hole. MA 02543 Palmer, John D., Department of Zoology, University of Massachusetts. Amherst, MA 01002 Palti, Yoram, Rappaport Institution, Technion, POB 9697, Haifa, 31096 Israel Pant, Harish C, NINCDS/NIH, Laboratory of Neurochemistry. Bldg. 36. Room 4D-20. Bethesda. MD 20892 Pappas, George D., Department of Anatomy, College of Medicine, University of Illinois, 808 South Wolcott St.. Chicago, IL 60612 Pardee, Arthur B., Department of Pharmacology. Harvard Medical School. Boston. MA 02 1 1 5 Pardy, Roosevelt L., School of Life Sciences. University of Nebraska. Lincoln, NE 68588 Parmentier, James L., Becton Dickinson Research Center. P. O. Box 12016, Research Triangle Park, NC 27709 Passano, Leonard M., Department of Zoology. Birge Hall, University of Wisconsin, Madison, WI 53706 Pearlman, Alan L., Department of Physiology, School of Medicine. Washington University, St. Louis. MO 631 10 Pederson, Thoru, Worcester Foundation for Experimental Biology, Shrewsbury, MA 01 545 Perkins, C. D., 400 Hilltop Terrace, Alexandria. VA 22301 Person, Philip, Research Testing Labs. Inc., 167 E. 2nd St.. Huntington Station, NY 1 1746 Peterson, Bruce J., Ecosystems Center, Marine Biological Laboratory, Woods Hole, MA 02543 Pethig, Ronald, School of Electronic Engineering Science, University College of N. Wales. Dean St., Bangor, Gwynedd. LL57 IUT, UK Pfohl. Ronald J., Department of Zoology. Miami University, Oxford. OH 45056 Pierce, Robert \V., 4851 Shore Lane. P. O. Box 1404, Boca Grande, FL 33921 Pierce, Sidney K., Jr., Department of Zoology, University of Maryland, College Park, MD 20742 Poindexter, Jeanne S., Science Division, Long Island University. Brooklyn Campus, Brooklyn, NY 1 1201 Pollard. Harvey B., NIH, NIDDK.D. Bldg. 8, Rm. 401, Bethesda, MD 20892 Pollard, Thomas D., Department ot Cell Biology and Anatomy. Johns Hopkins University, 725 North Wolfe St., Baltimore, MD 21205 Poole, Alan F., P. O. Box 533, Woods Hole. MA 02543 Porter, Beverly H., 13617 Glenoble Drive, Rockville. MD 20853 Porter, Keith R., Department of Biology, Leidy Laboratories, Rm. 303, University of Pennsylvania. Philadelphia. PA 19104-6018 Porter, Mary E., Department of Cell Biology and Neurology, Llniversity of Minnesota, 4-147 Jackson Hall, Minneapolis, MN 55455 Potter. David, Department of Neurobiology, Harvard Medical School, Longwood Avenue, Boston, MA 021 15 Potts, William T., Department of Biology, University of Lancaster. Lancaster. England. UK Powers, Maureen K., Department of Psychology, Vanderbilt Llniversity. Nashville. TN 37240 Pratt. Melanie M., Department of Anatomy and Cell Biology, University of Miami School of Medicine (R124), P. O. Box 016960, Miami, FL 33101 Prendergast, Robert A., Wilmer Institute, Johns Hopkins Hospital, 601 N. Broadway, Baltimore, MD 21205 Presley, Phillip H., Carl Zeiss. Inc., I Zeiss Drive. Thornwood. NY 10594 Price. Carl A., Waksman Institute of Microbiology, Rutgers University, P. O. Box 759, Piscataway, NJ 08854 Prior. David J., Department of Biological Sciences. NAU Box 5640, Northern Arizona University, Flagstaff, AZ 8601 1 Prusch, Robert D., Department of Life Sciences, Gonzaga University, Spokane, WA 99258 Purves, Dale, Department of Neurobiology. Duke University Medical School, Box 3209, Durham, NC 27710 Quigley, James, Department of Pathology. SUNY Health Science Center. BHS Tower 9, Rm. 140. Stony Brook, NY 1 1794 Rabb, Irving V\ ., 1010 Memonal Drive. Cambridge, MA 02138 Rabin, Harvey, DuPont Co., CRD. Exp. Station 328/358, Wilmington. DE 19880 Rabinowitz, Michael B., Marine Biological Laboratory. Woods Hole. MA 02543 Rafferty, Nancy S., Department of Anatomy. Northwestern University Medical School, 303 E. Chicago Avenue, Chicago, IL 60611 Rakowski, Robert K., Department of Physiology and Biophysics. UHS/The Chicago Medical School, 3333 Greenbay Rd.. N. Chicago, IL 60064 Ramon, Fidel, Dept. de Fisiologia y Bionsca, Centre de Investigacion y de Estudius Avanzados del ipn, Apurtado Postal 14-740. D.F. 07000, Mexico 62 Annual Report Rabb, Irving VV., 1010 Memorial Drive. Cambridge, MA 02138 Rabin, Harvey, DuPont Co., CRD, Exp. Station 328/358, Wilmington, DE 19880 Rabinowitz, Michael B., Marine Biological Laboratory. Woods Hole. MA 02543 Rafferty, Nancy S., Department of Anatomy, Northwestern University Medical School, 303 E. Chicago Avenue. Chicago, IL 60611 Rakowski, Robert F., Department of Physiology and Biophysics, UHS/The Chicago Medical School, 3333 Greenbay Rd., N. Chicago, IL 60064 Ramon, Fidel, Dept. de Fisiologia y Bionsca. Centra de Investigacion y de Estudius Avanzados del ipn. Apurtado Postal 14-740, D.F. 07000, Mexico Ranzi, Silvio, Sez Zoologia Sc Nat, Via Coloria 26, 120133. Milano. Italy Rastetter, Edward B., Ecosystems Center. Marine Biological Laboratory, Woods Hole, MA 02543 Ratner, Sarah, Department of Biochemistry, Public Health Research Institute. 455 First Ave., New York. NY 100 Id Rebhun, Lionel I., Department of Biology, Gilmer Hall. L'niversity of Virginia, Charlottesville, VA 22901 Reddan, John R., Department of Biological Sciences, Oakland University, Rochester, MI 48309-4401 Reese, Barbara F., NINCDS/NIH, Bldg. 36, Room 3B26, 9000 Rockville Pike. Bethesda, MD 20892 Reese, Thomas S., NINCDS/NIH, Bldg. 36. Room 2A27. 9000 Rockville Pike. Bethesda. MD 20892 Reiner, John M., 1 1 1 Emerson St.. Apt. 623. Denver, CO 80218 Reinisch, Carol L., Tufts University School of Veterinary Medicine, 136 Harrison Avenue, Boston, MA 021 I I Reynolds, George T., Department of Physics, Jadwin Hall, Princeton University. Princeton, NJ 08544 Rice, Robert V.. 30 Burnham Dr., Falmouth. MA 02540 Rich, Alexander, Department of Biology, Massachusetts Institute of Technology, Cambridge, MA 02 1 39 Rickles. Frederick R., Department of Medicine. Division of Hematology-Oncology, University of Connecticut Health Center, Farmmgton, CT 06032 Ripps, Harris, Department of Ophthalmology, University of Illinois College of Medicine, 1855 W. Taylor Street, Chicago, IL 6061 1 Robinson, Denis M., 200 Ocean Lane Drive #908. Key Biscayne, FL 33149 Rosenbaum, Joel L., Department of Biology, Kline Biology Tower. Yale University, New Haven, CT 06520 Rosenberg, Philip, School of Pharmacy. Division of Pharmacology, University of Connecticut. Storrs. CT 06268 Rosenbluth, Jack, Department of Physiology. New York University School of Medicine. 550 First Ave., New York, NY 10016 Rosenbluth, Raja, Department of Biological Sciences, Simon Fraser University. Burnaby, BC, V5A 1S6, Canada Roslansky, John, Box 208, Woods Hole, MA 02543 Roslansky, Priscilla F., Box 208, Woods Hole, MA 02543 Ross, William N., Department of Physiology. New York Medical College, Valhalla, NY 10595 Roth, Jay S., 18 Millneld Street, P. O. Box 285, Woods Hole. MA 02543 Rowland. Lewis P., Neurological Institute. 710 West 168th St., New York, NY 10032 Ruderman, Joan V'., Department of Anatomy and Cell Biology, Harvard University School of Medicine, Boston. MA 021 15 Rushforth, Norman B., Department of Biology, Case Western Reserve University, Cleveland, OH 44106 Russell-Hunter, \V. D., Department of Biology. Lyman Hall 012, Syracuse University, Syracuse. NY 13244 Saffo, Mary Beth, Institute of Marine Sciences. 272 Applied Sciences. University of California, Santa Cruz. CA 95064 Sager, Ruth, Dana Farber Cancer Institute, 44 Binney St., Boston. MA 021 15 Salama, Guy, Department of Physiology, University of Pittsburgh, Pittsburgh. PA 15261 Salmon, Edward D., Department of Biology. Wilson Hall, CB3280, University of North Carolina, Chapel Hill, NC 27599 Salzberg, Brian M., Department of Physiology, University of Pennsylvania. B-400 Richards Bldg.. Philadelphia. PA 19104-6085 Sanborn, Richard C., 1 1 Oak Ridge Road. Teaticket, MA 02536 Sanger, Jean M., Department of Anatomy. School of Medicine. University of Pennsylvania, 36th and Hamilton Walk, Philadelphia. PA 19174 Sanger, Joseph, Department of Anatomy, School of Medicine, University of Pennsylvania. 36th and Hamilton Walk, Philadelphia, PA 19174 Sato, Hidemi, Faculty of Social Science, Nagano University, Shiminogo, Ueda, Nagano 386- 1 2, Japan Sattelle, David B., AFRC Llnit-Department of Zoology, University of Cambridge. Downing St.. Cambridge CB2 3EJ, England, UK Saunders, John \\ ., Jr.. P. O. Box 381. Waquoit Station. Waquoit. MA 02536 Saz, Arthur K., Department of Immunology. Georgetown University Medical School, Washington. DC 20007 Schachman, Howard K., Department of Molecular Biology, University of California, Berkeley. CA 94720 Schalten, Gerald P., Integrated Microscopy Facility for Biomedical Research. University of Wisconsin. 1 1 17 W. Johnson St.. Madison, WI 53706 Schatten, Heide, Department of Zoology, University of Wisconsin. Madison, WI 53706 Schiff, Jerome A., Institute for Photobiology of Cells and Organelles, Brandeis University, Waltham, MA 02254 Schmeer, Arline C., Mercenene Cancer Research Institute, Hospital of Saint Raphael, New Haven. CT 0651 1 Schmidek, Henry H., Henry Ford Neurosurgical Institute. Henry Ford Hospital. Detroit. MI 48202 Schnapp, Bruce J., Department of Physiology, Boston University Medical School, 80 East Concord Street. Boston. MA 02 1 1 8 Schneider, E. Gayle, Department of Obstetrics and Gynecology. Yale University School of Medicine. 333 Cedar St.. New Haven, CT 06510 Schneiderman, Howard A., Monsanto Company. 800 North Lindbergh Blvd., D1W. St. Louis, MO 63166 (deceased) Schuel, Herbert, Department of Anatomical Sciences, SUNY, Buffalo, Buffalo. NY 14214 Schuetz, Allen W., School of Hygiene and Public Health, Johns Hopkins University, Baltimore, MD 21205 Schwartz. James H., Center for Neurobiology and Behavior. New York State Psychiatric Institute Research Annex. 722 W. 168th St., 7th Floor. New York, NY 10032 Scofield, Virginia Lee, Department of Microbiology and Immunology, UCLA School of Medicine, Los Angeles, CA 90024 Scott, Allan C., 1 Nudd St., Waterville, ME 04901 Sears, Mary, P. O. Box 152. Woods Hole, MA 02543 Segal, Sheldon J., Population Division, The Rockefeller Foundation, 1 133 Avenue of the Americas, New York. NY 10036 Selman, Kelly, Department of Anatomy. College of Medicine. University of Honda. Gainesville. FL 32601 Members of the Corporation 63 Shanklin, Douglas R., Department of Pathology, Room 584, University of Tennessee College of Medicine, 800 Madison Avenue, Memphis, TN 38163 Shapiro, Herbert, 6025 North 13th St., Philadelphia, PA 19141 Shaver, Gaius R., Ecosystems Center, Marine Biological Laboratory, Woods Hole, MA 02543 Shaver, John R., 18 Las Parras, Cayey, PR 00633 Sheetz, Michael P., Department of Cell Biology and Physiology. Washington LIniversity Medical School. 606 S. Euclid Ave., St. Louis, MO 63 110 Shepard, David C, P. O. Box 44, Woods Hole, MA 02543 Shepro, David, Department of Microvascular Research, Boston University, 5 Cummington St.. Boston. MA 02215 Sher, F. Alan, Immunology and Cell Biology Section. NIAID/NIH. Laboratory of Parasitic Disease, Building 5, Room 1 1 4, Bethesda, MD 20892 Sheridan, William F., Biology Department, LIniversity of North Dakota, Box 8238, University Station, Grand Forks, ND 58202- 8238 Sherman, I. W., Department of Biology, LIniversity of California. Riverside, CA 92521 Shimomura, Osamu, Marine Biological Laboratory, Woods Hole, MA 02543 Siegel, Irwin M., Department of Ophthalmology, New York University Medical Center, 550 First Avenue, New York. NY 10016 Siegelman, Harold W., Department of Biology. Brookhaven National Laboratory, Upton, NY 1 1973 Silver, Robert B., Department of Physiology. Cornell University, 822 Veterinary Research Tower, Ithaca, NY 14853-6401 Sjodin, Raymond A., Department of Biophysics, University of Maryland, Baltimore. MD 21201 Skinner, Dorothy M., Oak Ridge National Laboratory, P. O. Box 2009, Biology Division, Oak Ridge. TN 37831 Sloboda, Roger D., Department of Biological Sciences. Dartmouth College. Hanover, NH 03755 Sluder, Greenfield, Worcester Foundation for Experimental Biology, 222 Maple Ave.. Shrewsbury. MA 1 545 Smith, Michael A., J 1 Sinabung, Buntu #7. Semarang, Java, Indonesia Smith, Ralph I., Department of Zoology. LJniversity of California, Berkeley. CA 94720 Sogin, Mitchell, Marine Biological Laboratory. Woods Hole, MA 02543 Sorenson, Martha M., Depto de Bioquimica-RFRJ. Centro de Ciencias da Saude-I. C. B., Cidade Universitaria-Fundad, Rio de Janeiro, Brasil 21.910 Speck, William T., Department of Pediatrics. Case Western Reserve University, Cleveland, OH 44106 Spector, Abraham, College of Physicians and Surgeons, Columbia University, 630 West 168th Street, New York, NY 10032 Speer, John W., Marine Biological Laboratory, Woods Hole, MA 02543 Sperelakis, Nicholas, Department of Physiology & Biophysics, University of Cincinnati, Cincinnati, OH 45267-0576 Spiegel, Evelyn, Department of Biological Sciences, Dartmouth College, Hanover, NH 03755 Spiegel, Melvin, Department of Biological Sciences. Dartmouth College, Hanover, NH 03755 Spray, David C., Albert Einstein College of Medicine, Department of Neurosciences. 1300 Morris Park Avenue, Bronx, NY 10461 Steele, John Hyslop, Woods Hole Oceanographic Institution, Woods Hole, MA 02543 Steinacker, Antoinette, Dept. of Otolaryngology, Washington University. School of Medicine, Box 8115. 4566 Scott Avenue, St. Louis, MO 63 110 Steinberg, Malcolm. Department of Biology, Princeton University, Princeton, NJ 08540 Stephens, Grover C., Department of Ecol. and Evol. Biology, School of Biological Sciences. University of California. Irvine, CA 92717 Stephens, Raymond E., Marine Biological Laboratory. Woods Hole, MA 02543 Stetten, DeWitt, Jr., NIH, Bldg. 16, Room 1 18, Bethesda, MD 20892 (deceased) Stetten, Jane Lazarow, 4701 Willard Ave., Chevy Chase, MD 20815 Steudler, Paul A., Ecosystems Center, Marine Biological Laboratory. Woods Hole, MA 02543 Stokes, Darrell R., Department of Biology. Emory University, Atlanta, GA 30322 Stommel, Elijah W., Marine Biological Laboratory, Woods Hole, MA 02543 Stracher, Alfred, Department of Biochemistry. SUNY Health Science Center, 450 Clarkson Ave.. Brooklyn, NY 1 1203 Strehler, Bernard L., 2235 25th St., #217. San Pedro. CA 90732 Strickler, J. Rudi, Center for Great Lakes Studies. 600 East Greenfield Ave., Milwaukee, WI 53204 Strumasser, Felix, Marine Biological Laboratory, Woods Hole, MA 02543 Stuart, Ann E., Department of Physiology, Medical Sciences Research Wing 206H. University of North Carolina, Chapel Hill, NC 27599- 7545 Sugimori, Mutsuyuki, Department of Physiology and Biophysics, New York University Medical Center, 550 First Avenue, New York, NY 10016 Summers, William C., Huxley College of Environmental Studies, Western Washington University, Bellingham, WA 98225 Suprenant, Kathy A., Department of Physiology and Cell Biology, 4010 Haworth Hall. University of Kansas, Lawrence, KS 66045 Sussman, Maurice, 72 Carey Lane, Falmouth. MA 02540 Sussman, Raquel B., Marine Biological Laboratory, Woods Hole, MA 02543 Sweet, Frederick, Box 8064, Washington University School of Medicine. 499 South Euclid, St. Louis. MO 631 10 Sydlik, Mary Anne, Department of Biology, Eastern Michigan University, Ypsilanti, MI 48197 Szent-Gyorgyi, Andrew, Department of Biology. Brandeis University, Bassine 244. 415 South Street. Waltham, MA 02254 Szuts, Ete Z., Laboratory of Sensory Physiology, Marine Biological Laboratory, Woods Hole. MA 02543 Tabares, Lucia, AVDA. Department of Physiology, Sanchez, Pizjuan 4, 411009 Seville, Spain Tamm, Sidney L., Boston University Marine Program. Marine Biological Laboratory, Woods Hole. MA 02543 Tanzer, Marvin L., Department of Oral Biology, Medical School, University of Connecticut, Farmington, CT 06032 Tasaki, Ichiji, Laboratory of Neurobiology, NIMH/NIH, Bldg. 36, Rm. 2B-16. Bethesda, MD 20892 Taylor, Douglass L., Center for Fluorescence Research, Carnegie Mellon University, 440 Fifth Avenue. Pittsburgh, PA 15213 Teal, John M., Department of Biology, Woods Hole Oceanographic Institution, Woods Hole, MA 02543 Telfer, \\ illiam H., Department of Biology, University of Pennsylvania, Philadelphia. PA 19104 Telzer, Bruce, Department of Biology, Pomona College, Claremont, CA 9 1 7 1 I 64 Annual Report Thorndike, W. Nicholas, Wellington Management Company. 28 State St., Boston. MA 02 109 Trager, William, Rockefeller University, 1230 York Ave., New York, NY 10021 Travis, D. M., Veterans Administration Medical Center, 2101 Elm Street, Fargo, ND 58102 Treistman, Steven N., Worcester Foundation for Experimental Biology, 222 Maple Avenue, Shrewsbury, MA 01545 Trigg, D. Thomas, 1 25 Grove St.. Wellesley, MA 02 1 8 1 Trinkaus, J. Philip, Department of Biology. Box 6666, Yale University, New Haven, CT 065 1 1 Troll, Walter, Department of Environmental Medicine, College of Medicine. New York University. New York, NY 10016 Troxler, Robert F., Department of Biochemistry, School of Medicine, Boston University. 80 East Concord St., Boston. MA 02 1 1 8 Tucker, Edward B., Department of Natural Sciences. Baruch College, CUNY, 17 Lexington Ave., New York, NY 10010 Turner, Ruth D., Mollusk Department, Museum of Comparative Zoology. Harvard University. Cambridge. MA 02138 Tvteedell, Kenyon S., Department of Biological Sciences, University of Notre Dame, Notre Dame, IN 46656 Tytell. Michael. Department of Anatomy, Bowman Gray School of Medicine, Wake Forest University. Winston-Salem. NC 27103 Ueno, Hiroshi, Department of Biochemistry. The Rockefeller University. 1230 York Ave.. New York. NY 10021 Valiela, Ivan, Boston University Marine Program, Marine Biological Laboratory, Woods Hole, MA 02543 Vallee, Richard, Cell Biology Group. Worcester Foundation for Experimental Biology, Shrewsbury, MA 01545 Valois, John, Marine Biological Laboratory, Woods Hole, MA 02543 Van Holde, kensal. Department of Biochemistry and Biophysics. Oregon State University. Corvallis, OR 97331-6503 Vincent, Walter S., 16 F. R. Lillie Road, Woods Hole, MA 02543 Vogel, Steven S., LBM, NIDDK/NIH, Bldg. 10, Rm. 9B04. Bethesda. MD 20894 Waksman, Byron, Foundation for Microbiology. 300 East 54th St.. New York, NY 10022 Wall, Betty, 9 George St.. Woods Hole. MA 02543 Wallace, Robin A., Whitney Laboratory, 9505 Ocean Shore Blvd., St. Augustine, FL 32086 Wang, Ching Chung, Department of Pharmaceutical Chemistry, University of California, San Francisco, CA 94143 Warner, Robert C., Department of Molecular Biology and Biochemistry. University of California, Irvine, CA 92717 Warren, Kenneth S., Maxwell Communications Corp., 866 Third Avenue. New York. NY 10022 Warren, Leonard, Wistar Institute, 36th and Spruce Streets, Philadelphia, PA 19104 \\aterbury, John B., Department of Biology. Woods Hole Oceanographtc Institution, Woods Hole, MA 02543 Watson, Stanley, Associates of Cape Cod, Inc., P. O. Box 224, Woods Hole, MA 02543 Associate Members Waxman, Stephen G., Department of Neurology, LCI 708. Yale School of Medicine. 333 Cedar Street. New Haven. CT 06510 Webb, H. Marguerite, Marine Biological Laboratory. Woods Hole. MA 02543 Weber, Annemarie, Department of Biochemistry and Biophysics, School of Medicine. University of Pennsylvania. Philadelphia. PA 19066 Weidner, Earl, Department of Zoology and Physiology, Louisiana State University, Baton Rouge, LA 70803 Weiss, Dieter G., Institut fur Zoologie. Technische Universitat Munchen. 8046 Garching, FRG Weiss, Leon P., Department of Animal Biology, School of Veterinary Medicine, University of Pennsylvania, Philadelphia, PA 19104 Weiss, Paul A., Address unknown Weissmann, Gerald, New York University Medical Center. 550 First Avenue. New York, NY 10016 Werman, Robert, Neurobiology Unit, The Hebrew University, Jerusalem, Israel Westerfield, R. Monte, The Institute of Neuroscience. LIniversity of Oregon, Eugene, OR 97403 V\ hittaker, J. Richard, Marine Biological Laboratory, Woods Hole, MA 02543 Wigley, Roland L., 35 Wilson Road, Woods Hole, MA 02543 Wilson, Darcy B., Medical Biology Institute. I 1077 North Torrey Pines Road, La Jolla. CA 92037 Wilson, T. Hastings. Department of Physiology, Harvard Medical School, Boston, MA 02 1 1 5 \\ilkovsky, Paul, Department of Ophthalmology, New York University Medical Center, 550 First Ave., New York, NY 10016 Wittenberg, Jonathan B., Department of Physiology and Biophysics, Albert Einstein College, 1300 Morris Park Ave.. Bronx, NY 01461 Wolfe, Ralph, Department of Microbiology, 131 Burrill Hall, University of Illinois, Urbana, IL 61801 \\olken, Jerome J.. Department of Biological Sciences. Carnegie Mellon University, 440 Fifth Ave.. Pittsburgh, PA 15213 \\orgul, Basil V., Department of Ophthalmology, Columbia University, 630 West 168th St., New York, NY 10032 W'u, Chau Hsiung, Department of Pharmacology. Northwestern LIniversity Medical School, Chicago, IL 6061 1 Wyttenbach, Charles R., Department of Physiology and Cell Biology. University of Kansas, Lawrence, KS 66045 Yeh, Jay Z., Department of Pharmacology. Northwestern University Medical School. Chicago. IL 6061 1 Zigman, Seymour, School of Medicine and Dentistry. LIniversity of Rochester. 260 Cnttenden Blvd., Rochester, NY 14620 Zigmond, Richard E., Center for Neurosciences, School of Medicine, Case Western Reserve University, Cleveland. OH 44106 Zimmerberg, Joshua J., NIH, Bldg. 12A. Room 2007, Bethesda, MD 20892 Zottoli, Steven J., Department of Biology. Williams College, Williamstown, MA 01267 Zucker, Robert S., Neurobiology Division, Department of Molecular and Cellular Biology, University of California. Berkeley, CA 94720 Zukin, Ruth Suzanne, Department of Neuroscience, Albert Einstein College of Medicine, 1300 Morris Park Ave., Bronx, NY 10461 Ackroyd, Dr. Frederick W. Adams, Dr. Paul Adelberg, Mrs. Edward A. Ahearn. 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Mizell, Dr. and Mrs. Merle Monroy. Mrs. Alberto Montgomery, Dr. and Mrs. Charles H. Montgomery, Mrs. Raymond B. Moore, Drs. John and Betty Morgan, Miss Amy Morse, Mrs. Charles L.. Jr. Morse, Dr. M. Patricia Moul. Mrs. Edwin T. Munson, Mr. William Murchelano, Dr. Robert Murray, Mr. David M. Myles-Tochko, Drs. Christina J. and John Nace, Dr. and Mrs. Paul Nace, Mr. Paul F., Jr. Naugle, Mr. John E. Neall, Mr. William G. Nelson. Dr. and Mrs. Leonard Nelson, Dr. Pamela Newton. Mr. William E. Nickerson. Mr. and Mrs. Frank L. Norman, Mr. and Mrs. Andrew E. Norman Foundation Norris, Mr. and Mrs. Barry Norris, Mr. and Mrs. John A. Norris, Mr. William Norton. Mrs. Thomas J. O'Connell, Dr. and Mrs. Clifford Offenback, Dr. Jack A. O'Herron, Mr. and Mrs. Jonathan Olszowka, Dr. Janice S. O'Neil, Mr. and Mrs. Barry T. O'Rand, Mr. and Mrs. Michael O'Sullivan, Dr. Renee Bennett Ott, Drs. Philip and Karen Palmer, Mr. and Mrs. David Pappas, Dr. and Mrs. George D. Park, Mr. and Mrs. Malcolm S. Parmenter, Dr. Charles Parmenter, Miss Carolyn L. Pearce, Dr. John B. Pearson, Mrs. Oscar H. Peltz. Mr. and Mrs. William L. Pendergast, Mrs. Claudia Pendleton, Dr. and Mrs. Murray E. Peri, Mr. and Mrs. John B Perkins, Mr. and Mrs. Courtland D. Person, Dr. and Mrs. Philip Peters, Mr. and Mrs. Frederick S. Peterson, Mr. and Mrs. E. Gunnar Peterson, Mr. and Mrs. E. Joel Peterson, Mr. Raymond W. Petty, Mr. Richard F. Pteiffer, Mr. and Mrs. John Plough. Ms. Frances Plough. Mr. and Mrs. George H. Plough, Mrs. Harold H. Pointe. Mr. Albert Pointe, Mr. Charles Porter, Dr. and Mrs. Keith R. Pothier, Dr. and Mrs. Aubrey Press, Dr. Frank Proskauer, Mr. Joseph H. Proskauer, Mr. Richard Prosser, Dr. and Mrs. C. Ladd Psaledakis, Mr. Nicholas Psychoyos, Dr. Alexandre Putnam, Mr. and Mrs. Allan Ray Putnam, Mr. and Mrs. William A., Ill Rankin, Mrs. John Raphael, Ms. Ellen S. Raymond, Dr. Samuel Reese. Miss Bonnie Regis. Ms. A. Kathy Reingold, Mr. Stephen C. Reynolds, Mrs. George Reynolds, Dr. John L. Reynolds. Mr. and Mrs. Robert M. Rezmkoff, Mrs. Paul Ricca. Dr. and Mrs. Renato A. Righter, Mr. and Mrs. Harold Riina, Mr. John R. Riley. Dr. Monica Robb, Mrs. Alison A. Roberts, Miss Jean Roberts, Mr. Mervin F. Robertson, Mrs. C. W. Robinson. Dr. Denis M. Robinson, Mr. John G. Robinson, Mr. and Mrs. Marius A. Root. Mrs. Walter S. Rosenthal. Miss Hilde Members of the Corporation 67 Roslansky, Drs. John and Priscilla Ross, Dr. and Mrs. Donald Ross, Dr. Robert Ross. Dr. Virginia Roth, Dr. Stephen Rowan. Mr. Edward Rowe, Dr. Don Rowe, Mrs. William S. Rugh, Mrs. Roberts Ryder, Mr. Francis C. Sager. Dr. Ruth Sanidas. Dr. and Mrs. Dennis J. Sardinha, Mr. George J. Saunders, Dr. and Mrs. John W. Saunders, Mrs. Lawrence Saunders, Lawrence, Fund Sawyer. Mr. and Mrs. John E. Saz. Mrs. Ruth L. Schlesinger. Dr. and Mrs. R. Walter Schwamb. Mr. and Mrs. Peter Schwartz, Dr. Lawrence Scott, Mrs. George T. Scott, Mr. and Mrs. Norman E. Sears. Mr. Clayton C. Sears. Mr. and Mrs. Harold B. Sears, Mr. Harold H. Seaver, Mr. George Seder, Mr. John Segal, Dr. and Mrs. Sheldon J. Selby, Dr. Cecily Senft. Dr. and Mrs. Alfred Shanklm. Dr. D. R. Shapiro, Mr. and Mrs. Howard Shapley, Dr. Robert Sharp. Mr. and Mrs. Robert W. Shemin, Dr. and Mrs. David Shepro, Dr. and Mrs. David Sherblom. Dr. James P. Sichel, Dr. Enid Siegel, Mr. and Mrs. Alvin Simmons. Mr. Tim Simon, Mr. and Mrs. Stephen Singer, Mr. and Mrs. Daniel M. Smith, Drs. Frederick E. and Marguerite A. Smith. Mr. and Mrs. Homer P. Smith. Mr. Van Dorn C. Snyder, Mr. Robert M. Solomon, Dr. and Mrs. A. K. Sonnenblick, Mrs. Perle Specht, Mr. and Mrs. Heinz Speck, Dr. William T. Spiegel. Drs. and Mrs. Melvin and Evelyn Spotte, Mr. Stephen Steele, Mrs. John H. Steele, Dr. Robert E. Stein, Mr. Ronald Steinbach. Mrs. H. Burr Stetson. Mrs. Thomas J. Stetten, Dr. Gail Stetten. Dr. and Mrs. H. DeWitt. Jr. Stracher. Dr. Dorothy Sudduth. Dr. William Swain, Mr. Albert H. Sussman. Dr. and Mrs. Maurice Swanson. Mrs. Carl P. Swift, Mr. and Mrs. E. Kent Swope. Mrs. Gerard, Jr. Swope, Mr. and Mrs. Gerard L Szent-Gyo'rgyi, Dr. Andrew Taber, Mr. George H. Talamas-Rohana, Dr. Eduardo Taylor, Mr. James K. Taylor. Dr. and Mrs. W. Randolph Tietje, Mr. Emil D., Jr. Timmins. Mrs. William Tochko, Dr. John S. Todd, Mr. and Mrs. Gordon F. Tolkan. Mr. and Mrs. Norman N. Trager, Mrs. William Trigg, Mr. and Mrs. D. Thomas Troll. Dr. and Mrs. Walter Trousof. Miss Natalie Tucker, Miss Ruth Tully. Mr. and Mrs. Gordon F. Ulbnch, Mr. and Mrs. Volker Valois, Mr. and Mrs. John Vancouver Public Aquarium Van Buren, Mrs. Harold Van Holde, Mrs. Kensal E. Veeder, Mrs. Ronald A. Veeder, Ms. Susan Vincent. Mr. and Mrs. Samuel W. Vincent, Dr. Walter S. Vonderhaar, Dr. William Wagner, Mr. Mark Waksman, Dr. and Mrs. Byron H. Ward, Dr. Robert T. Ware. Mr. and Mrs. J. Lindsay Warren, Dr. Henry B. Warren, Dr. and Mrs. Leonard Watt, Mr. and Mrs. John B. Weeks, Mr. and Mrs. John T. Weift'enbach, Dr. and Mrs. George Weinstein. Miss Nancy B. Weisberg, Mr. and Mrs. Alfred M. Weissmann, Dr. and Mrs. Gerald Wheeler, Dr. and Mrs. Paul S. Wheeler, Dr. William M. Whitehead. Mrs. Fred Whitney. Mr. and Mrs. Geoffrey G., Jr. Wichterman, Dr. and Mrs. Ralph Wickersham, Mr. and Mrs. A. A. Tilney Wiese, Dr. Konrad Wilber. Mrs. Clare M. Wilhelm. Dr. Hazel S. Willis. Mr. Herbert F. Wilson, Mr. and Mrs. Leslie J. Wilson, Mr. and Mrs. T. Hastings Winn. Dr. William M. Winsten, Dr. Jay A. Witting, Miss Joyce Woitkoski, Miss Nancy Wolfinsohn, Mrs. Wolfe Woodwell, Dr. and Mrs. George M. Wrigley, Mrs. Roland Yntema, Mrs. Chester L. Young, Miss Nina L. Zinn, Dr. and Mrs. Donald J. Zipf. Dr. Elizabeth Gift Shop Volunteers Marian Adelberg Michael Goldring Bertha Person Louise Atkins Rose Grant Julia Rankin Barbara Atwood Martha Griffin Virginia Reynolds Gloria Borgese Edith Grosch Lilyan Saunders Jennie Brown Jean Halvorson Elsie Scott Elisabeth Buck Helen Hodosh Deborah Senft Patricia Case Adele Hoskins Charlotte Shemin Summers Case Pauline Hyde Manlyn Shepro Shirley Chaet Sona Jones Cynthia Smith Julia Child Sally Karush Marguerite Smith Vera Clark Ruth Ann Laster Louise Specht Margaret Clowes Barbara Little Judith Stetson Villa Crowell Sarah Loessel Dorothy Stracher Elizabeth Daignault Winnie Mackey Mary Ulbrich Janet Daniels Constance Martyna Barbara van Holde Alma Ebert Florence Mixer Barbara Whitehead Margaret German Lorraine Mizell Clare Wilber Rebeckah Glazebrook Phyllis Meyers MEL Tour Guides Teni Hayashi Betsy Bang Sally Loessel Lola Robinson John Buck Isabel Mountain Donald Zinn Sears Crowell Julie Rankin Margery Zinn Certificate of Organization Articles of Amendment Bylaws of the MBL Certificate of Organization (On File in the Office of the Secretary of the Commonwealth) No. 3170 We. Alpheus Hyatt. President. William Stanford Stevens. Treasurer, and William T. Sedgwick. Edward G. Gardiner. Susan Mims and Charles Sedgwick Mmot being a majority of the Trustees of the Marine Biological Laboratory in compliance with the requirements of the fourth section of chapter one hundred and fifteen of the Public Statutes do hereby certify that the following is a true copy of the agreement of association to constitute said Corporation, with the names of the subscribers thereto: We. whose names are hereto subscribed, do, by this agreement, associate ourselves with the intention to constitute a Corporation according to the provisions of the one hundred and fifteenth chapter of the Public Statutes of the Commonwealth of Massachusetts, and the Acts in amendment thereof and in addition thereto. The name by which the Corporation shall be known is THE MARINE BIOLOGICAL LABORATORY The purpose for which the Corporation is constituted is to establish and maintain a laboratory or station for scientific study and investigations, and a school for instruction in biology and natural history. The place within which the Corporation is established or located is the city of Boston within said Commonwealth. The amount of its capital stock is none. In Witness Whereof, we have hereunto set our hands, this twenty seventh day of February in the year eighteen hundred and eighty-eight, Alpheus Hyatt, Samuel Mills. William T. Sedgwick, Edward G. Gardiner. Charles Sedgwick Minot. William G. Farlow, William Stanford Stevens, Anna D. Phillips, Susan Mims, B. H. Van Vleck. That the first meeting of the subscribers to said agreement was held on the thirteenth day of March in the year eighteen hundred and eighty-eight. In Witness Whereof, we have hereunto signed our names, this thirteenth day of March in the year eighteen hundred and eighty-eight. Alpheus Hyatt. President. William Stanford Stevens, Treasurer, Edward G. Gardiner. William T. Sedgwick. Susan Mims. Charles Sedgwick Minot. (Approved on March 20. 1888 as follows: / hereby a-mfr that it appears upon an examination of the within written certificate and the records of the corporation duly submitted to my inspection, that the re- quirements of sections one, two and three of chapter one hundred and fifteen, and sections eighteen, twenty and twenty-one of chapter one hundred and six, of the Public Statutes, have been complied with and 1 hereby approve said certificate this twentieth day of March A.D. eighteen hundred and eighty-eight. Charles Endicott Commissioner of Corporations) Articles of Amendment (On File in the Office of the Secretary of the Commonweallh) We, James D. Ebert. President, and David Shepro. Clerk of the Marine Biological Laboratory, located at Woods Hole. Massachusetts 02543, do hereby certify that the following amendment to the Articles of Organization of the Corporation was duly adopted at a meeting held on August 15, 1975, as adjourned to August 29. 1975, by vote of 444 members, being at least two-thirds of its members legally qualified to vote in the meeting of the corporation: Voted: That the Certificate of Organization of this corporation be and it hereby is amended by the addition of the following provisions: "No Officer, Trustee or Corporate Member of the corporation shall be personally liable for the payment or satisfaction of any obligation or liabilities incurred as a result of, or otherwise in connection with, any commitments, agree- ments, activities or affairs of the corporation. "Except as otherwise specifically provided by the Bylaws of the corporation, meetings of the Corporate Members of the corporation may be held anywhere in the United States. "The Trustees of the corporation may make, amend or repeal the Bylaws of the corporation in whole or in part, except with respect to any provisions thereof which shall by law, this Certificate or the bylaws of the corporation, require action by the Corporate Members." The foregoing amendment will become effective when these articles of amendment are filed in accordance with Chapter 180. Section 7 of the General Laws unless these articles specify, in accordance with the vote adopting the amendment, a later effective date not more than thirty days after such filing, in which event the amend- ment will become effective on such later date. In Witness whereof ami Under the Penalties of Perjury, we have hereto signed our names this 2nd day of September, in the year 1975. James D. Ebert, President: David Shepro, Clerk. (Approved on October 24, 1975, as follows: I hereby approve the within articles of amendment and. the filing fee in the amount of $10 having been paid, said articles are deemed to have been filed with me this 24th day of October. 1975. Paul Guzzi Secrelan of the Commonwealth) Bylaws of the Corporation of the Marine Biological Laboratory (Revised August 17, 1990) (These Brians hem' been e.\lensirely amended by the Board ol Trustees and are \ubieel lo approval br the Corporation II these amendments are not approved by the Corporation, the existing Bylaws mil then remain in place.) 68 Bylaws of the Corporation 69 ARTICLE I THE CORPORATION A Name ami Purpose. The name of the Corporation shall be The Marine Bio- logical Laboratory The Corporation's purpose shall be to establish and maintain i laboratory or station for scientific study and investigation and a school for in- struction in biology and natural history. B Sondtscntninalion The Corporation shall not discriminate on the basis of age, religion, color, race, national or ethnic origin, sex or sexual preference in its policies on employment and administration or in its educational and other programs. ARTICLE II MEMBERSHIP I Members. The Members of the Corporation ("Members") shall consist of persons elected by the Board of Trustees (the "Board"), upon such terms and conditions and in accordance with such procedures, not inconsistent with law or these Bylaws, as may be determined by the Board- At any regular or special meeting of the Board, the Board may elect new Members. Any Member may vote at any meeting of the Members either in person or by proxy executed no more than three months prior to the date of such meeting. Except as otherwise limited therein, proxies shall entitle the persons named therein to vote at any adjournment of such meeting, but shall not he valid after final adjournment of such meeting. Proxies need not be sealed or attested and proxy purported to be executed by or on behalf of a Member entitled to vote shall be deemed valid unless challenged at or poor to its exercise. Members shall serve until their death or resignation unless earlier removed with or without cause by the affirmative vote of two-thirds ot the Trustees then in office. Any Member who has retired from his or her home institution may. on written request to the Corporation, be designated a Life Member. Life Members shall not have the right to vote and shall not be assessed for dues. 6 Meetings The annual meeting of the Members shall be held on the Friday following the second Tuesday in August of each year, at the Laboratory of the Corporation in Woods Hole. Massachusetts, at 9:30 a.m. If no annual meeting is held in accordance with the foregoing provision, a special meeting may be held in lieu thereof with the same effect as the annual meeting, and in such case all references in these Bylaws, except in this Article II. B., to the annual meeting of the Members shall be deemed to refer to such special meeting. Members shall transact business as may properly come before the meeting. Special meetings of the Members may be called by the Chairman or the Trustees, and shall be called by the Clerk, or in the case of the death, absence, incapacity or refusal by the Clerk, by any other officer, upon written application of Members representing at least ten percent of the smallest quorum of Members required for a vote upon any matter at the annual meeting of the Members, to be held at such time and place as may be designated. The Chairman of the Board shall preside at all meetings of the Corporation. C Notice of Meetings. Notice of any annual meeting or special meeting of Members, if necessary, shall be given by the Clerk by mailing notice of the time and place and purpose of such meeting at least 1 5 days before such meeting to each Member at his or her address as shown on the records of the Corporation. D ll'u/ivr ol Notice Whenever notice of a meeting is required to be given a Member under any provision of the Articles or Organization or Bylaws of the Corporation, a written waiver thereof, executed before of after the Meeting by such Member, or his or her duly authorized attorney, shall be deemed equivalent to such notice. E Adjournments Any meeting ot the Members may be adjourned to any other time and place by the vote of a majority of those Members present or represented at the meeting, whether or not such Members constitute a quorum, or by any officer entitled to preside at or to act as Clerk of such meeting, if no Member is present or represented. It shall not be necessary to notify any Members of any adjournment unless no Member is present or represented at the meeting which is adjourned, in which case, notice of the adjournment shall be given in accordance with Article II E. Any business which could have been transacted at any meeting of the Members as originally called may be transacted at an adjournment thereof. ARTICLE III ASSOCIATES OF THE CORPORATION l\\oiwies ol the Corporation. The Associates of the Marine Biological Laboratory shall be an unincorporated group of persons (including associations and corporations) interested in the Laboratory and shall be organized and operated under the general supervision and authority of the Trustees. The Associates of the Marine Biological Laboratory shall have no voting nghts. ARTICLE IV BOARD OF TRUSTEES A Pon-ers The Board of Trustees shall have the control and management of the affairs of the Corporation. The Trustees shall annually elect a Chairman ot the Board who shall serve until his or her successor is selected and qualified. They shall annually elect a President of the Corporation who shall also be the Vice Chairman of the Board and Vice Chairman of meetings of the Corporation. They shall annually elect a Treasurer. They shall elect a Clerk, who shall be a resident of Massachusetts and shall serve a term of four years. Eligibility for re-election of the Clerk shall be in accordance with the content of this Article IV as applied to Corporate or Board Trustees. They shall elect Trustees-at-Large as specified in this Article IV. They shall appoint a Director of the Laboratory for a term not to exceed five years, provided the term shall not exceed one year, if the candidate has attained the age of 65 years prior to the dale of the appointment. They may choose such other officers and agents as they shall think best. They may fix the compensation of all officers and agents of the Corporation and may remove them at any time. They may fill vacancies occurring in any of the offices. The Board shall have the power to choose an Executive Committee from their own number as provided in Article V. and to delegate to such Committee such of their own powers as they may deem expedient in addition to those powers conferred by Article V. They shall, from time to time, elect Members to the Corporation upon such terms and conditions as they shall have determined, not inconsistent with law or these Bylaws. B Composition and Election. There shall be four groups of Trustees: ( 1 1 Trustees (the "Corporate Trustees") elected by the Members according to such procedures, not inconsistent with these Bylaws, as the Trustees shall have determined. Except as provided below, such Trustees shall be divided into four classes of six, one class to be elected each year to serve for a term of four years. Such classes shall be designated by the year of expiration of their respective terms. (2) Trustees ("Trustees-at-Large"). Nominees for Trustees-at-Large shall be introduced at the annual meeting of the Corporation for subsequent election by the Board according to such procedures, not inconsistent with these Bylaws, as the Trustees shall have determined. Such Trustees-at-Large shall be divided into four classes of four Trustees, one class to be elected each year to serve for a term ol lour years. Such classes shall be designated by the year of expiration of their respective terms. It is contemplated that, unless otherwise determined by the Trustees for good reason. Trustees-at-Large. shall be individuals who have not been considered for elections as Corporate Trustees. (3) Trustees e\ offiao shall be the Chairman, the President, the Treasurer, the Clerk and the Director of the Laboratory. (4) Trustees emeriti shall include any Member who has retired from his or her home institution and has requested to serve as a Trustee emeritus provided he or she has served at least two terms as a Trustee. A Trustee ex officio is eligible to serve as a Trustee ementus provided he or she has served as Trustee e\ officio for at least eight years. Trustees ex ofticio and emeriti shall have all the nghts of the Trustees, except that Trustees emeriti shall not have the right to vote. (5) The total number of Corporate Trustees and Trustees-at-Large elected in any year (excluding Trustees elected to fill vacancies which do not result from expiration of a term) shall not exceed ten. The number of Trustees-at-Large so elected shall not exceed four and, unless otherwise determined by vote of the Trustees, the number of Corporate Trustees so elected shall not exceed six. Corporate Trustees shall always constitute a majority on the Board of those elected or approved by the Members. (6) Newly elected Trustees shall take office at the February meeting of the Board, but may participate in discussions at intervening meetings following their election, without voting rights. (7) The Trustees and officers shall hold their respective offices until their suc- cessors are chosen and quahfied. C. Eligibility A Corporate Trustee or a Trustee-at-Large who has served an initial term of at least two years' duration shall be eligible for re-election to a second term, but shall be ineligible for re-election to any subsequent term until two years have elapsed after he/she has last served as a Trustee. D Removal. Any Trustee may be removed from office at any time without cause, by vote of a majority of the Members entitled to vote in the election of Trustees; or for cause, by vote of two-thirds of the Trustees then in office. A Trustee may be removed for cause only if notice of such action shall have been given to all of the Trustees or Members entitled to vote, as the case may be. prior to the meeting at which such action is to be taken and if the Trustee to be so removed shall have been given reasonable notice and opportunity to be heard before the body proposing to remove him or her. E I 'ticuncies Any vacancy in the Board, unless and until filled by the Members at any annual or special meeting of the Members, may be filled by vote of a majority of the remaining Trustees present at a meeting of Trustees at which a quorum is present or by appointment of all of the Trustees if less than a quorum shall remain in office. F. Meetings. The annual meeting of the Trustees shall be held promptly after the annual meeting of the Members at the Laboratory in Woods Hole, Massachusetts. Special meetings of the Trustees may be called by the Chairman, the President or 70 Annual Report by any seven Trustees to be held at such time and place as may be designated. The Chairman of the Board, when present, shall preside overall meetings of the Trustees. Wntten notice shall be sent to a Trustee's usual or last known place of residence at least two weeks before the meeting. Notice of a meeting need not be given to any Trustee if a written waiver of notice executed by such Trustee before or after the meeting is hied with the records of the meeting, or if such Trustee shall attend the meeting without protesting prior thereto or at its commencement the lack of notice given to him or her. G Quorum Twenty-five Members shall constitute a quorum at any meeting. Except as otherwise required by law or these Bylaws, the affirmative vole of a majority of the Members voting in person or by proxy at a meeting attended by a quorum (present in person or by proxy) shall consitute action on behalf of the Members. // Transfers of Interests in Land. There shall be no transfer of title or long-term lease of real property held by the Corporation without pnor approval of not less than two-thirds of the Trustees. Such real property transactions shall be finally acted upon at a meeting of the Board only if presented and discussed at a pnor meeting of the Board. Either meeting may be a special meeting and no less than four weeks shall elapse between the two meetings. Any property acquired by the Corporation after December I, 1989 may be sold with the pnor approval of not less than two-thirds of the Trustees (other than any Trustee or Trustees with a direct or indirect financial interest in the transaction being considered for approval) who are present at a regular or special meeting of the Board at which there is a quorum. ARTICLE V COMMITTEES A Executive Committee The Executive Committee is hereby designated to consist of not more than ten Trustees, including the t'.v n/licio Trustees (Chairman of the Board. President. Treasurer, and Director of the Laboratory); and six additional Trustees, two of whom shall be elected by the Board each year, to serve for a three- year term. Beginning with the Members elected for terms ending in 1990. one of the Trustees elected to serve on the Executive Committee shall be a Trustee-at-Large. Beginning with the Members elected for terms ending in 1991. the Trustees will elect, to the Executive Committee. Trustees to ensure that the Committee includes four Cor- porate Trustees and two Trustees-at-Large. The Chairman of the Board shall act as Chairman of the Executive Committee and the President as Vice Chairman. The Executive Committee shall meet at such times and places and upon such notice and appoint such subcommittees as the Committee shall determine. The Executive Committee shall ha\e and may exercise all the powers of the Board dunng the mter\als between meetings of the Board except those powers specifically withheld, from time to time, by vote of the Board or by law. The Executive Committee may also appoint such committees, including persons who are not Trustees, as it may. from time to time, approve to make recommendations with respect to matters to be acted upon by the Executive Committee or the Board. The Executive Committee shall keep appropnate minutes of its meetings, which shall be reported to the Board. Any actions taken by the Executive Committee shall also he reported to the Board. The elected Members of the Executive Committee shall constitute a Standing Committee for the Nomination of Officers, responsible for making nominations at each annual meeting of the Members and of the Board for candidates to rill each office as the respective terms of office expire (Chairman of the Board. President. Treasurer, Clerk and Director of the Laboratory). B Board Committee-. Henerallv The Board shall have the power, by vote of a majonty of the Trustees then in office, to elect an Investment Committee, a Nom- inating Committee and any other committee and, by like vote, to delegate thereto some or all of the powers of the Board except those which by law. the Articles of Organization or these Bylaws they are prohibited from delegating. The members of any such committee shall have such tenure and duties as the Trustees shall determine. The Investment Committee, which shall oversee the management of the Corporation's endowment funds and marketable securities shall include as t'.v n/fieio members, the Chairman of the Board, the Treasurer and the Chairman of the Audit Committee, together with such Trustees as may be required for not less than two-thirds of the Investment Committee to consist of Trustees. Except as otherwise provided by these Bylaws or determined by the Trustees, any such com- mittee may make rules for the conduct of its business, but, unless otherwise provided by the Trustees or in such rules, its business shall be conducted as nearly as possible in the same manner as is provided by these Bylaws lor the Trustees ( Iciion', Without a Meeting Any action required or permitted to be taken at any meeting of the Executive Committee or any other committee elected by the Trustees may be taken without a meeting if all Members of such committees consent to the action in wnting and such wntten consents are filed with the records of meetings. Members of the Executive Committee or any other committee elected by the Trustees may also participate in any meeting by means of a telephone con- ference call, or otherwise take action in such a manner as may. from time to time, be permitted by law. ARTICLE VI OFFICERS A Enumeraiiiin The officers of the Corporation shall consist of a President, a Treasurer and a Clerk, and such other officers having the powers of President. Treasurer and Clerk as the Board may determine, and a Director of the Laboratory. The Corporation may have such other officers and assistant officers as the Board may determine, including (without limitation) a Chairman of the Board, and one or more Vice-Presidents, Assistant Treasurers or Assistant Clerks. Any two or more offices may be held by the same person. An officer need not be a Member or Trustee of the Corporation. If required by the Trustees, any officer shall give the Corporation a bond lor the faithful performance of his or her duties in such amount and with such surety or sureties as shall be satisfactory to the Trustees. B Tenure Except as otherwise provided by law. by the Articles of Organization or by these Bylaws, the President. Treasurer, and all other officers shall hold office until the first meeting of the Board following the annual meeting of Members and thereafter, until his or her successor is chosen and qualified. C. Resignation Any officer may resign by delivenng his or her wntten resignation to the Corporation at its principal office or to the President or Clerk and such resignation shall be effective upon receipt unless it is specified to be effective at some other time or upon the happening of some other event. D Removal The Board may remove any officer with or without cause by a vote of a majonty of the entire number of Trustees then in office, at a meeting of the Board called for that purpose and for which notice of the purpose thereof has been given, provided that an officer may be removed for cause only after having an opportunity to be heard by the Board at a meeting of the Board at which a quorum is personally present and voting. E lummy A vacancy in any office may be filled for the unexpired balance of the term by vote of a majonty of the Trustees present at any meeting of Trustees at which a quorum is present or by written consent of all of the Trustees if less than a quorum of Trustees shall remain in office. F. Dinrlor T he Director shall be the chief operating officer and. unless otherwise voted by the Trustees, the chief executive officer of the Corporation. The Director shall, subject to the direction of the Trustees, have general supervision of the Lab- oratory and control of the business of the Corporation. At the annual meeting, the Director shall submit a report of the operations of the Corporation for such year and a statement of its affairs, and shall, from time to time, report to the Board all matters within his or her knowledge which the interests of the Corporation may require to be brought to its notice. G. Deputy Direetor. The Deputy Director, if any. or if there shall be more than one. the Deputy Directors in the order determined by the Trustees, shall, in the absence or disability of the Director, perform the duties and exercise the powers of the Director and shall perform such other duties and shall have such other powers as the Trustees may. from time to time, prescribe. // r<0.005]. This bar graph is based on 15 no- stimulus control experiments and 30 single-arm experiments, 15 each for "nonlayer. no eggs" and "egg layer, eggs"; in the single-arm experi- ments, animals were choosing between a stimulus in one arm and no stimulus in the other. without egg cordons [X : (2) = 34. 12; P < 0.005], dem- onstrating that egg-laying animals are attractive. Because the "egg-laying animal" stimulus has two components the egg layer and its egg cordon subse- quent experiments focused on their relative contributions to pheromonal attraction. More animals were attracted to egg layers without egg cordons than had been attracted to nonlayers without egg cordons, and fewer made no choice (Fig. 3); the difference in response patterns was statistically significant [X 2 (2) = 15.38; P < 0.005], dem- onstrating that the egg layer is a source of aggregation pheromones. Identical results were obtained when a non- layer with a recently deposited egg cordon was used as the stimulus (Fig. 3), indicating that the egg cordon is also SEXUAL PHEROMONES IN APLYSIA 85 0} a co n 3 15 10 Positive No choice Negative Nonlayer Egg layer Nonlayer Egg layer No eggs No eggs Eggs Eggs Stimulus Figure 3. Egg layers are attractive to Aplysia braxiliana: a larger number of animals was attracted to an egg layer without eggs than was attracted to a nonlayer without eggs, and the patterns of responses to the two stimuli are significantly different [X 2 (2) = 15.38; P < 0.005]. Egg cordons are also attractive to A. brasiliana: a larger number of animals was attracted to a nonlayer with eggs than was attracted to a nonlayer without eggs, and the response patterns are significantly different [X : (2) = 15.38; P < 0.005]. The effects of egg layers and egg cordons are not additive at the concentrations tested: the response patterns for a nonlayer with eggs and an egg layer without eggs are identical, and do not differ significantly from the pattern obtained for an egg layer with eggs [X 2 (2) = 2.83; 0.25 < P < 0.50]. This bar graph is based on 60 single-arm experiments, 1 5 per stimulus; in each experiment, animals were choosing between a stimulus in one arm and no stimulus in the other. a source of pheromonal activity. Neither pattern differed significantly from that obtained for an egg-laying animal with an egg cordon [X 2 (2) = 2.83; 0. 10 < P < 0.25], dem- onstrating that the effects of the layer-derived and cordon- derived factors are not additive at the concentrations tested. Subsequent experiments examined whether animal- derived factors are required for the attractiveness of the egg cordon and whether the attraction is visually mediated. Two series of experiments were performed. In the first, an egg cordon without any animal served as the stimulus. The level of attraction and pattern of responses were identical to those obtained using an egg layer and its cor- don as the stimulus (Figs. 3, 4), demonstrating that egg cordons are sufficient to attract conspecifics. In the second series of experiments, a "sham" cordon (a tangled mass of silastic tubing; vol = 2 ml) served as the stimulus. The level of attraction and pattern of responses differed sig- nificantly from those observed with the egg cordon [X 2 (2) = 1 5.27; P < 0.005], but did not differ from those observed in no-stimulus control experiments [Fig. 4; X 2 (2) = 1.70; 0.25 < P < 0.50]. The differential responses to egg cordons and sham cordons suggests that the attraction is chemi- cally rather than visually mediated. The results of these two series of experiments, in conjunction with those re- ported above, demonstrate that both egg layers and egg cordons are sufficient to attract conspecifics, but that nei- ther is uniquely required. As a first step toward identifying tissue sources of the cordon-derived aggregation pheromones, acidic extracts of the atrial gland (equivalent to 50% of the material in one gland) were assayed for the ability to increase the attractiveness of a nonlaying animal when placed in the surrounding ASW. A higher level of attraction to nonlay- ing animals and lower level of no-choice responses was observed when the extract was present (Fig. 5); the dif- 0) a E c ffl 4) n 15 10 Positive No choice Negative No animal No animal No animal No eggs Sham eggs Eggs Stimulus Figure 4. Animals are not required for an egg cordon to be attractive to Aplysia brasiliana: a larger number of animals was attracted to an egg cordon without an animal than was attracted to a sham cordon without an animal, and the response pattern was significantly different [x : (2) = 15.27; P < 0.005]. The sham cordon was a tangled mass of silastic tubing, 2 ml in volume. The attractiveness of an egg cordon is not visually mediated: the pattern of responses to a sham cordon without an animal did not differ significantly from the no-stimulus control (no animals, no eggs) (X-(2) = 1.70; 0.25 < P < 0.50]. (Note: for the no- stimulus control, movement to the left was defined as a positive response and movement to the right a negative response.) This bar graph is based on 15 no-stimulus control experiments and 30 single-arm experiments, 1 5 each for "no animal, sham eggs" and "no animal, eggs"; in the single- arm experiments, animals were choosing between a stimulus in one arm and no stimulus in the other. 86 S. D. PAINTER ET AL CO 'c a E z 15 10 Positive No choice Negative Nonlayer Nonlayer Nonlayer No eggs AQE Eggs Stimulus Figure 5. Secretory products of the Aply.iia californica atrial gland (or A. brasiliana AG-LE) may contribute to the attractiveness of an egg cordon. The number of animals attracted to a nonlayer was increased when an extract of the A. californica atrial gland (AGE) was placed in the surrounding ASW. The pattern of responses differed significantly from that observed for a nonlaying animal without eggs [X 2 (2) = 10.58: F < 0.01], but did not differ significantly from that for a nonlaying animal with eggs [ V(2) = 0.44; 0.90 < P < 0.95]. This bar graph is based on 45 single-arm experiments. 1 5 per stimulus: in each experiment, an- imals were choosing between a stimulus in one arm and no stimulus in the other. Terence in response patterns was significant [X 2 (2) = 10.58; P < 0.0 1 ]. Interestingly, the level of attraction and pattern of responses did not differ significantly from those ob- tained when a recently deposited conspecific egg cordon was placed in the same location [Fig. 5; X 2 (2) = 0.44; 0.75 < P < 0.90], suggesting that products of the A. californica atrial gland (or A. brasiliana AG-LE) might significantly contribute to the attractiveness of an egg cordon. Induction of mating activity Animals. A pool of 205 sexually mature Aplysia brasil- iana individuals was used in these studies. Small plastic fish tags ( 1 1 mm in diameter; Howitt Plastics, Molalla, Oregon) were sutured to the caudal region of the right parapodium so that individuals could be identified. Four criteria were used to select animals for each experiment: ( 1 ) the animal must not have laid eggs during the preced- ing 24 h; (2) the animal must not have participated in a behavioral experiment during the preceding 24 h; (3) the animal must not have been tested with the stimulus; and (4) all of the animals in an experiment must have been housed in the same aquarium. Once selected, the animals were randomly assigned to treatments. All experiments were begun between 9 and 10 am, because there is evi- dence of a circadian rhythm in Aplysia mating behavior (A.J'asciata. Susswein et al.. 1983, 1984). Relative contributions of the egg layer and egg cordon: experimental protocol and statistical analyses. Eight an- imals were used in each experiment (Fig. 6). One animal was injected with 0. 1 ml of atrial gland extract and placed in a 4-1 plastic beaker containing 3 1 of aerated non-con- ditioned ASW; this treatment induced egg laying, usually within 30 min, and the egg layer conditioned the ASW in the beaker. A second animal was handled and placed in an identical beaker; this treatment did not induce egg laying, but the nonlayer conditioned the ASW. When the injected animal finished laying eggs (70.3 4.1 min after injection; mean S.E.M.), it was removed, rinsed in fresh non-conditioned ASW and transferred to a third beaker; the handled animal was treated in the same way and transferred to a fourth beaker. Nontreated animals were then distributed among the four beakers so that each con- tained two animals. The resulting experimental conditions are: ( 1 ) two nonlayers in animal-conditioned ASW with an egg cordon; (2) two nonlayers in animal-conditioned ASW without an egg cordon; (3) one egg layer and one nonlayer in non-conditioned ASW without an egg cordon; and (4) two nonlayers in non-conditioned ASW without an egg cordon. The reproductive activity of each individual was as- sessed at 10-min intervals for 270 min. Three categories of mating activity were recognized (male, female, and si- multaneous hermaphrodite); egg-laying activity was also recorded, but courtship was not. For calculation purposes, animals that did not mate or lay eggs during the obser- vation period were assigned a 280-min latency for that activity. Although this approach underestimates the dif- ference in mean latency to mating between strongly pos- itive and control conditions, the effect is relatively small because at least 85% of the animals mated in every ex- perimental condition tested. Statistical significance was assessed by a one-way analysis of variance, followed by Duncan's multiple range test for pairwise comparisons. When time courses of mating activity were compared, statistical significance was assessed by X 2 analysis of in- dividual time points. Egg cordon volume was measured at the end of each experiment by ASW displacement in a graduated cylinder and averaged 1.7 ml (1.7 0.2 ml; mean S.E.M.). Twenty experiments were performed in this series. Results. Animal-conditioning the ASW with a nonlay- ing animal resulted in an increase in the percentage of SEXUAL PHEROMONES IN APLYSIA 87 Handle animal; inject atrial gland extract I Conditions ASW; lays eggs Transfer egg layer to Conditioned ASW Non-conditioned ASW with eggs without eggs I I Add 2 nonlayers Add 1 nonlayer Handle animal; no injection i Conditions ASW; does not lay eggs Transfer nonlayer to Conditioned ASW Non-conditioned ASW without eggs without eggs I Add 2 nonlayers Add 1 nonlayer 2 nonlayers in Egg layer, nonlayer in conditioned ASW non-conditioned ASW with eggs without eggs 2 nonlayers in conditioned ASW without eggs 2 nonlayers in non-conditioned ASW without eggs Figure 6. Flow diagram of the protocol used in the first series of mating experiments. One animal was injected with an extract of the atrial gland to induce egg laying and placed in a beaker containing aerated non-conditioned ASW; a second animal was handled and placed in a second beaker containing aerated non- conditioned ASW. When egg deposition was complete, the egg layer was transferred to a third beaker and the handled nonlayer to a fourth beaker. Additional nontreated animals were then distributed among the four beakers so that each contained a pair of Aplysia: the resulting experimental conditions are indicated at the bottom of the diagram. Mating and egg-laying behaviors were scored at 10-min intervals for 270 min: animals failing to exhibit a behavior during the observation period were assigned a 280-min latency for calculations. animals mating at early time periods relative to non-con- ditioned ASW controls (Fig. 7A); the difference was sta- tistically significant at three time periods 40, 50, and 60 min [X'( 1 ) > 3.84 for each; P < 0.05]. The mean latency to mating was also reduced (Fig. 7B), but did not differ significantly from that of the non-conditioned ASW con- trols (P = 0.29; one-way analysis of variance). It is im- portant to note that these effects, although small, were consistently observed when the ASW was animal-condi- tioned. Comparable results were obtained in an indepen- dent series of experiments performed in our laboratory (A. R. Gustavson, unpubl. data). The studies used a dif- ferent pool of A. brasiliana and animal-conditioned the ASW for 60 rather than 70 min, but produced quantita- tively similar responses. A higher percentage of animals mated at early time periods relative to the non-conditioned ASW controls [X 2 (l) > 3.84 at 30, 40, 70, 80. 90, 100, and 1 10 min; P < 0.05 for each]; the mean latency to mating was reduced, but the change was not statistically significant (P = 0.28; one-way analysis of variance). The consistency of these two sets of results suggests that ani- mal-derived factors induce mating, but that their activity or concentration is relatively low under the conditions tested. The idea that animal-derived factors induce mating in Aplysia is consistent with a recent report in the literature that the amount of time that A. fasciata spend mating is a function of the number of animals available as copu- latory partners (Ziv el al, 1989) and thus, presumably, a function of the concentration of animal-derived factors in the ASW. Similar, but quantitatively larger, effects were observed when the ASW was animal-conditioned by an egg layer and contained an egg cordon (Fig. 7 A. B). The percentage of animals mating at early time periods was increased relative to non-conditioned ASW controls, and the difference was statistically significant for every observation period from 10 through 1 10 min [X 2 (l) > 3.84 for each; P < 0.05]. The mean latency to mating was significantly reduced (P = 0.002; one-way analysis of variance). Assuming that animal-con- ditioning the ASW with an egg layer is comparable to ani- mal-conditioning with a nonlayer (see below), these results demonstrate that cordon-derived factors induce mating, and suggest that the effects of the animal-derived and cordon- derived factors may be additive. S. D. PAINTER ET AL E c a o 01 5 Q. 100 - 80 40 20 Nonlayer, No eggs Conditioned Nonlayer, No eggs Non-conditioned 40 80 120 160 200 240 280 Time (mini 150 _ 125 1 CD 100 75 50 25 Nonlayer Nonlayer Nonlayer No eggs No eggs Eggs Non-cond. Conditioned Conditioned Stimulus Figure 7. Both animal-derived and cordon-derived factors induce mating activity in Aplysia brasiliana. (A) The percentage of animals mating at early time periods was increased by animal-conditioning the ASW with a nonlaying animal; the difference was significant at 40, 50, and 60 mm [X 2 ( 1 ) > 3.84; P < 0.05]. The percentage was further increased by animal-conditioning the ASW with an egg-laying animal and leaving the egg cordon in the ASW; the percentages were significantly higher than those obtained in non-conditioned ASW without an egg cordon at every observation period from 10 through 1 10 min (X'U) > 3.84 for each; P < 0.05]. The experimental protocol is shown in Figure 6 (n = 20 experiments). Because Aplysia tend to mate in bouts lasting approximately 60 min and the bouts are often separated by periods during which no mating occurs (Leonard and Lukowiak, 1987), it is not possible to read a mean latency to mating directly from this graph. (B) The latency to mating (mean S.E.M.) was reduced by animal-conditioning the ASW with a nonlaying animal, but the difference was not statistically significant (P = 0.29; one-way analysis of variance). The latency was further reduced in animal-conditioned ASW with an egg cordon, and differed significantly from that obtained for non-conditioned ASW without an egg cordon (P < 0.002; one-way analysis of variance). Animals did not distinguish between recent egg layers and nonlayers in non-conditioned ASW without an egg cordon. There were no significant differences in the time courses of mating activity (Fig. 8A), in the latencies to mating (Fig. 8B), or in the sexual role first assumed by the animals (Table I). These results suggest that there is not a prolonged change in the motivational state of the egg layer (i.e.. an increase in receptivity to courtship) that persists in the absence of an egg cordon. More importantly, they suggest that the egg cordon, rather than the egg layer, may be primarily responsible for the relatively short la- tencies to mating observed when animals are actively lay- ing eggs. The experiments did not address the question of whether specific layer-derived or animal-derived factors are required for the induction of mating by egg cordons, however, and this issue was examined in the next series of experiments. Induction of mating by egg cordons in non-conditioned ASW ': experimental protocol and statistical analyses. Five animals, selected as described above, were used in each experiment (Fig. 9). One was injected with atrial gland extract and placed in a beaker to lay eggs. When deposition was complete, the egg cordon was removed, quickly rinsed, and transferred to a second beaker containing non- conditioned ASW; a pair of animals was then placed in this beaker and another pair placed in a third beaker that contained only non-conditioned ASW. The resulting ex- perimental conditions are: (1) two nonlayers in non-con- ditioned ASW with an egg cordon; and (2) two nonlayers in non-conditioned ASW without an egg cordon. Repro- ductive behavior was assessed for each animal at 10-min intervals for 270 min and analyzed as in the preceding experiments. Egg volume was measured after each ex- periment and averaged 2.0 ml (2.0 0.3 ml; mean S.E.M. ). Fifteen experiments were performed. Results. Placing a recently deposited egg cordon in the non-conditioned ASW surrounding two nonlaying ani- mals significantly increased the percentage of animals mating in 9 of the first 14 observation periods [Fig. 10A; X : ( 1 ) > 3.84 at 20-40 min, 70 min, and 100-140 min: P < 0.05 for each], and significantly reduced their mean latency to mating relative to the control group (Fig. 10B; P < 0.01; one-way analysis of variance). These results demonstrate that cordon-derived factors alone are suffi- cient to induce mating. Induction of egg-laying activity Egg deposition was also monitored in the experiments described above. Neither animal-derived nor cordon-de- rived factors significantly affected the percentage of ani- mals laying eggs (Table II), and the low percentages made calculations of mean latency to deposition meaningless. Because Aplysia brasiliana lays eggs more frequently when SEXUAL PHEROMONES IN APLYSIA 89 100 Egg layer, No eggs Non-conditioned Nonlayer, No eggs Non-conditioned 40 80 120 160 200 240 280 Time (min) 150 125 c E o> 100 75 50 25 B Nonlayer No eggs Non-cond. Egg layer No eggs Non-cond. Stimulus Figure 8. Aplysia brasiliana does not distinguish between recent egg layers and nonlayers in non-conditioned ASW without an egg cordon. (A) Recent egg layers and nonlayers mated at the same frequency in non-conditioned ASW without an egg cordon. The experimental protocol is shown in Figure 6 (n = 20 experiments). (B) There is no difference in mean latency to mating between recent egg layers and nonlayers in non- conditioned ASW without an egg cordon. caged alone rather than in pairs (Blankenship et a!., 1983), the experiments were repeated with one animal in each beaker rather than two. Single-animal experiments: protocol. Four test animals, selected as described in the mating studies, were used in each experiment. One was placed in each of four beakers and egg-laying activity assessed at 10-min intervals for 270 min. The ASW in each beaker contained a different combination of animal-derived and cordon-derived fac- tors: ( 1 ) non-conditioned ASW without an egg cordon (negative control); (2) non-conditioned ASW with an egg cordon (cordon-derived factors only); (3) animal-condi- tioned ASW without an egg cordon (animal-derived fac- tors only); and (4) animal-conditioned ASW with an egg cordon (both animal-derived and cordon-derived factors). The conditions in each beaker were established as de- scribed in the section on mating (see Figs. 6 and 9). The volumes of the stimulus egg cordons were measured at the end of every experiment and averaged 3.2 ml (3.2 0.6 ml; mean S.E.M.). In five experiments, all animals that were not induced to lay eggs were injected with atrial gland extract to verify that they were physiologically com- petent to do so; all laid egg cordons in response to the injection, demonstrating that the experimental conditions were not interfering with the activity. Results. Neither animal-derived nor cordon-derived factors had a significant effect on egg deposition (Ta- ble II). Discussion Pheromonal attraction These studies have shown that Aplysia brasiliana is not attracted to nonlaying animals in the absence of an egg cordon. The results contrast with those of Lederhendler and colleagues (1977), which showed that Aplysia dac- tylomela is attracted to nonlaying conspecifics and that the magnitude of the attraction increases as the number of stimulus animals increases. We do not know whether the difference reflects species differences or whether it re- sults from differences in experimental design (e.g., from differences in concentration produced by using 5-min rather than 60-min conditioning periods), and we have not examined the possibility that a group of nonlaying A. brasiliana would be attractive. We have, however, tested A. californica in T-maze experiments and have found that A. californica, like A. brasiliana, is not attracted to non- laying conspecifics under these conditions (S. D. Painter, unpubl. data). These results are consistent with earlier studies by Audesirk (1977), which showed that A. cali- fornica is not attracted to nonlaying animals in Y-maze experiments. Although there is electrophysiological evi- dence that A. californica detects the odors of nonlaying conspecifics (Audesirk and Audesirk, 1977; Chase, 1979), there is no evidence to date that the electrophysiological response is to species-specific odors (Chase, 1979) and no behavioral evidence that the odors are attractive. Aplysia brasiliana is attracted to egg-laying animals with egg cordons. The ability to distinguish egg-laying animals with egg cordons from nonlaying animals without egg cordons was previously described in burrowing studies in this species (Aspey and Blankenship, 1976). In those studies, when a nonlaying animal was introduced into an aquarium containing a burrowed conspecific, the intro- duced animal burrowed; when an egg-laying animal was 90 S. D. PAINTER ET AL Table I The effects of recent egg deposition on sexual role in Aplysia brasiliana Conditions t & of animals first mating as % Not mating Animal Female Male Hermaphrodite Conditioned ASW. egg cordon present" Non-conditioned Egg layer 90 10 ASW, no egg cordon* Non-conditioned Egg layer 45 30 1 5 10 ASW, no egg cordon* Nonlayer 40 40 10 10 " From Painter et al ( 1989). Some egg layers were actively laying eggs when the second animal was introduced into the chamber. The ASW was animal-conditioned by the egg layer, n = 20. * In each case, the experimental animal was introduced into a new chamber at the beginning of the observation period and the ASW was not animal-conditioned. Egg layers were animals that had just completed egg deposition; nonlayers were those that had not laid within the preceding 24 h. The activity pattern of an egg layer in non-conditioned ASW without an egg cordon differed significantly from that of an egg layer in animal- conditioned ASW with an egg cordon [X 2 (3) = 16.67; P < 0.005], but did not differ from that of a nonlayer in non-conditioned ASW without an egg cordon (X : = 1.11; 0.75 < P < 0.90). n = 20 for each. introduced, however, the burrowed animal emerged to mate with it. The distinction is also evident in mating experiments ( Painter t'//., 1989), which showed that egg- laying animals have significantly shorter latencies to mat- ing than do sexually mature but nonlaying animals. Other species of Aplysia also make the distinction. Copulatory Handle animal; inject atrial gland extract Conditions ASW; lays eggs Conditioned ASW with egg layer Transfer eggs to Non-conditioned ASW without eggs Non-conditioned ASW without eggs Add 2 nonlayers Add 2 nonlayers 2 nonlayers in non-conditioned ASW with eggs 2 nonlayers in non-conditioned ASW without eggs Figure 9. Flow diagram of the protocol followed in the second series of mating experiments. One animal was injected with an extract of the atnal gland to induce egg laying and placed in a beaker containing aerated non-conditioned ASW. When egg deposition was complete, the egg cordon was removed from the container, quickly rinsed, and transferred to a second beaker containing non-conditioned ASW. Two nontreated animals were then added to the beaker containing the egg cordon, and two others added to a third beaker containing only non-conditioned ASW. The resulting experimental conditions are indicated at the bottom of the diagram. Reproductive behaviors were scored at 10-min intervals for 270 min. SEXUAL PHEROMONES IN APLYSIA 91 100 * 80 OB c 60 ra CL 40 20 Nonlayer, Eggs Non-conditioned Nonlayer, No eggs Non- conditioned 40 80 120 160 200 240 280 Time (min) 200 150 100 50 Nonlayer No eggs Non-cond. Nonlayer Eggs Non-cond. Stimulus Figure 10. High concentrations of animal-denved factors are not required for the induction of mating activity by recently deposited egg cordons. (A) A higher percentage of nonlaying .l/Vrau brasiliana mated at early time periods when an egg cordon was placed in the surrounding non-conditioned ASW; the increase was statistically significant at 20, 30,40. 70, and 100-140 min [X 2 (l)> 3.84 in each case; P < 0.05]. The experimental protocol is shown in Figure 9 (n = 15 experiments). (B) Placing an egg cordon in the non-conditioned ASW surrounding two nonlaying animals significantly reduced their latency to mating relative to nonlaying animals without an egg cordon (P < 0.0 1 ; one-way analysis of variance). The experimental protocol is shown in Figure 9 (n = 15 expenmentsl. chains of A. californica, for example, are both more stable and more attractive when the first animal in the chain is laying eggs (Audesirk, 1977). The present studies have shown that both egg-layer- derived and cordon-derived factors contribute to the at- tractiveness of an egg-laying animal. Although there are two apparent sources of this pheromonal activity, it re- mains to be demonstrated whether the two sets of factors differ biochemically. There is extensive and prolonged contact between the egg layer and its cordon, both during and following oviposition, which would facilitate a transfer of activity between the two. The attractiveness of a nonlaying animal was increased by placing a recently deposited egg cordon in the same arm of the T-maze, providing the basis for a simple bioas- say system in which to identify potential tissue sources of the "cordon-derived" pheromonal activity. Extracts of the A. californica atrial gland were assayed in this system and also increased the attractiveness of a nonlaying animal, suggesting that secretory products of the A. californica atrial gland (or A. brasiliana AG-LE) may contribute to the cordon-derived activity. The extracts each contained 50% of the material in a single atrial gland and were as attractive as A. brasiliana egg cordons, suggesting that the A. brasiliana and A. californica aggregation pheromones may be chemically similar or identical. This issue is cur- rently being examined by HPLC analyses of egg cordon eluates, and by compositional and microsequence analyses of active fractions. It is worth noting that extracts of the A. californica atrial gland also induce copulatory activity in A. brasiliana (Painter et ai. 1989), and that field studies suggest that Aplysia aggregation pheromones may not be entirely species-specific (A. vaccaria, for example, has been observed in association with A. californica aggregations: Kupfermann and Carew, 1974). Mating activity The experiments showed that both animal-derived and cordon-derived factors reduce the latency to mating in Table II Neither animals nor egg cordon', stimulated egg-laying activity in Aplysia brasiliana % Animals laying eggs Conditions 2 Animals in chamber 1 Animal in chamber Non-conditioned ASW No egg cordon 0" 0" Egg cordon 6.7 6.7" Animal-conditioned ASW No egg cordon 7.5' O d Egg cordon 12.5' I0 d " n = 1 5; 30 possible egg-laying episodes. Not significantly different [X-(D = 2.14:0.10 < /><0.25]. * n = 15; 15 possible egg-laying episodes. Not significantly different [X 2 (D = 1.07:0.25 < P < 0.50]. ' n = 20; 40 possible egg-laying episodes. Not significantly different [X 2 (l) = 1.44:0.10 - Enzyme-Treated No Addition ' / i I 6 12 18 24 48 Duration Larval Exposure (hr) Figure 4. Inaetivation of the cell wall-associated inducer by treatment with digestive enzymes from abalone. Equivalent amounts of inducer were incubated in parallel ( I h, 28C) with or without exposure to en- zymes. Samples then were washed by ultrafiltration. Assays with the chromogenic substrate for sulfatase. p-nitrophenylsulfate. confirmed that all of the enzyme was removed from the paniculate samples by this washing procedure. The washed samples then were assayed in quadru- plicate for remaining morphogenic activity (in parallel with assays with no addition). After 24 h of exposure of the larvae to each of the three assay conditions, fresh samples of the inductive crustose coralline red alga Hydmlithon boergesenii (CCA) were added to duplicate samples of each assay type (open symbols), to assess the remaining responsiveness of the larvae. pet. Patella vulgata (Table II). A significant reduction in the dose-dependent activity of the cell wall-associated morphogen resulted from this enzyme treatment. When Table II Effect of sulfatase purified from Patella vulgata on cell wall-associated morphogen Inducer (equivalents)' Untreated Enzyme-treated Metamorphosis- 0+0 1 70 10 6 100 1 6 20 1 1 70 10 1 The decalcified morphogen was incubated with or without enzyme for 1 h at 28C. and subsequently washed free of enzyme, as described in Materials and Methods. One equivalent corresponds to 0.25 ml of the original decalcified morphogen. Portions of the two samples then were assayed, either singly or in pre-mixed combination, in the relative amounts shown. 2 Assays were conducted with 5 larvae/trial, and scored for metamor- phosis after 24 h: results = mean S.D.; n = 2 trials for each condition. 10 D. E. MORSE AND A. N. C. MORSE 100 ,- _ 80 - None Trps. Papn |3-Gal. c-Gal 0-Glu Hyal Bactri Haiiotis Patella Wu JOOu 200u 200u Sulfatase Figure 5. Effects of purified enzymes used as probes for structural determinants of the cell wall-associated inducer of larval metamorphosis. Trps. = trypsin; Papn. = papain; /J-Gal. = /i-galactosidase; -Gal. = - galactosidase; fi-Glu. = /j-glucuronidase; Hyal. = hyaluromdase; sulfatases purified from the bacterium Acrohuctcr acrnt;cne.t, the limpet Piilclln vitlgiild. and the abalone Haliolis craclierndii. were used in the amounts (enzyme units) indicated. Assays, after treatment and washing of the paniculate samples, were performed in duplicate; all other details as in Figure 4 and Materials and Methods. equivalent amounts of the enzyme-treated and the un- treated morphogen were mixed and assayed together, no reduction in the activity of the untreated morphogen was observed, thus confirming the conclusion that the mol- luscan enzymes inactivated the inducer, and not the larvae. Larvae exposed to molluscan digestive enzyme- treated inducer, and either subsequently (Fig. 4) or si- multaneously (Table II) exposed to untreated inducer, re- sponded normally and completed metamorphosis. The digestive enzyme preparation from Haiiotis that inactivated the coral morphogen (Fig. 4) is a relatively crude mixture containing high quantities of a sulfatase (Spaulding and Morse, 1991) and lower quantities of /3- glucuronidase and several other enzymes. When purified molluscan sulfatase and /i-glucuronidase were tested sep- arately, the sulfatase was a potent inactivator of the cell wall-associated morphogen, whereas jtf-glucuronidase had little if any significant activity (Fig. 5). The data show that sulfatases purified from Haliolis, Patella, and from a bac- terium all inactivate the morphogen, and that the effect of the Haiiotis sulfatase is concentration-dependent. (The slight inactivation caused by treatment with the /3-gluc- uronidase preparation is likely to reflect the activity of the small amount of sulfatase known to still contaminate this preparation.) In similar tests, five other purified enzymes were used to probe for essential features of the morphogen structure. Of these, only hyaluronidase, which cleaves sulfated poly- saccharide chains, reduced the activity of the insoluble, cell wall-associated morphogen (Fig. 5). The proteolytic enzymes trypsin and papain, and the exosaccharidases a- galactosidase and 0-galactosidase, were completely with- out effect. These results, and those presented above. strongly suggest that a sulfated polysaccharide is an es- sential component of the morphogen recognized by the Agaricia humilis larvae. Enzymes release and degrade soluble morphogen Four purified endopolysaccharidases, including agarase, endo-/i-galactosidase, lysozyme, and hyaluronidase, re- leased a soluble morphogen from the insoluble cell wall preparation (Fig. 6). The subsequent time-dependent de- cline in soluble morphogen activity seen with prolonged exposure to agarase suggests that this enzyme may con- tinue to attack the solubilized inducer; this suggestion is confirmed by experiments presented below. None of the other enzymes tested in the experiment shown in Figure 5 released any detectable soluble activity. The enzyme-solubilized morphogen was a potent in- ducer, causing the Agaricia humilis larvae to quickly at- tach either to the sides or bottoms of the clean polystyrene assay beakers, and to undergo rapid and normal meta- morphosis and post-metamorphic growth (cf. Fig. 10). These results, and those with the alkali-solubilized ma- terial discussed below, prove that the larvae do not require any specific tactile stimulus from the morphogen, but re- spond solely to its chemical structure. Experiments in which aliquots of the cell wall-associated morphogen were incubated in the presence and absence of purified enzymes, inside dialysis tubing, confirmed that agarase and other endosaccharidases continue to attack and subsequently degrade the soluble morphogenic mol- ecules they first release (as suggested by the data in Fig. 100 - Digestion (hr) Figure 6. Solubilization of morphogen by treatment with purified endopolysaccharidases. Agarase (squares): endo-/3-galactosidase (filled circles); hyaluronidase (upward triangles); lysozyme (downward triangles); no enzyme controls (open circles). Other procedures similar to experiment shown in Figure 5. Details are in Materials and Methods. CHARACTERIZATION OF CORAL MORPHOGEN 111 100 - _ 80 - + Agarase tn O 9- o E 3 QJ 60 - 0246 + Agarase, 1 sample) No Enzyme! No Inducer J "j 10 12 14 16 18 20 22 Digestion (hr) Figure 7. Enzymatic solubilization and further purification of small morphogen. The insoluble, cell wall-associated inducer of larval meta- morphosis was digested inside a dialysis tubing ( retention limit = 1 4,000 Da) with purified agarase. Small morphogens released from the insoluble inducer were allowed to accumulate in the external dialysate, which was removed and assayed, and replaced with fresh external medium, every two hours. An otherwise identical sample was digested in parallel with no change of dialysate for 22 h, after which the external medium was removed and assayed (dotted line). Other details as described in the text; all assays were performed in duplicate under standard conditions. Con- trols conducted in parallel included assays of dialysates changed every 2 h from a sample incubated with no enzyme, and larvae incubated with no additions; these gave 0% metamorphosis. Results show meta- morphosis induced by the dialysates. 6). The external dialysates (including the permeant small morphogens released) were allowed to diffuse and accu- mulate outside of the membranes for a fixed interval, after which they were removed for assay, and replaced with fresh external liquid for the next interval of incubation. With purified agarase, and 2-h intervals for accumulation, removal, and replacement of the external dialysates. we observed a time-dependent increase and subsequent de- cline in the rate of appearance of dialyzable morphogenic activity with M < 14,000 Da (Fig. 7). However, far less of the total morphogenic activity could be detected in an otherwise identical incubation, in which only one sample of dialysate was collected and assayed after prolonged in- cubation (24 h). All of the dialysates were held at 28C until the end of the experiment (24 h) and assayed si- multaneously, to control for any time-dependent decay of activity. Because the difference between the two parallel incubations thus was in the limitation (2 h interval) or prolongation (24 h interval) of the opportunity for the small dialyzable morphogen to diffuse back and forth be- tween the outside and inside of the dialysis tubing, we conclude that prolonged exposure of released, dialyzable morphogen to the agarase resulted in degradation of the active morphogen. We therefore conclude that agarase, which cleaves sulfated polysaccharides, particularly at 0- 1,4-linked galactose residues, not only releases the active morphogen from a sulfated polysaccharide parent mole- cule, but degrades the morphogen itself, as well. Therefore, the structure of the morphogen would seem to contain a sulfated polysaccharide with galactose residues. In similar experiments, we compared dialysis mem- branes with calibrated porosites ofca. 10,000-14,000 Da and ca. 6.000-8,000 Da, and two other endosaccharidases in addition to agarase (Fig. 8). When lysozyme was used we again saw a time-dependent increase and subsequent decline in the rate of accumulation of dialyzable mor- phogen with M < 14,000 Da. There was a similar, but displaced, rise and fall in the accumulation of smaller morphogen (M < 8000 Da), suggesting that these smaller molecules may be derived, in part, by enzymatic cleavage of the larger dialyzable inducers. With the 8-h intervals 100 80 60 40 20 M< 14,000 Agarase M < 8,000 Lysozyme 0-8 8-16 16-24 Digestion (hr) Figure 8. Enzymatic solubilization and further purification of small morphogen through dialysis membranes of two different porosities. Ex- periment performed as in Figure 7, except that: agarase, lysozyme and endo-i-galactosidase were compared; the digestions were performed in parallel in dialysis tubings with retention limits of 14.000 Da (hatched bars) and 8,000 Da (solid bars); the external dialysates were changed and saved for assays every 8 h. 12 D. E. MORSE AND A. N. C. MORSE Table III Specificities and effects <>l'en:yinc probes ;<csenii was incubated with vigorous stirring at pH 12.0. At the times indicated, aliquots were withdrawn, neutralized to pH 8.2. filtered through nitrocellulose filters (0.2 ^m. 47 mm diam.), and the soluble fractions assayed (in duplicate 10 ml samples) under the standard conditions. Details are in Materials and Methods. Results shown are the average percentages of larvae induced to attach and metamorphose by the soluble fractions, S.D. Results of a parallel and otherwise identical incubation held continuously at pH 8.2 are included for comparison. polysaccharide (cleaved by hyaluronidase, agarase, and sulfatase), with multiple, substituted residues of N-ace- tylglucosamine (linkages cleaved by lysozyme and endo- itf-galactosidase) and multiple residues of galactose (link- ages cleaved by agarase and endo-/3-galactosidase). Protein (or peptide) does not appear to be an essential component, nor do several specific free or terminal sugars or saccharide units. Hydrolysis with dilute alkali releases small anionic morphogen containing sulfate The finding that specific endopolysaccharidases can re- lease a soluble morphogen from the insoluble cell wall- associated fraction suggested that non-enzymatic, partial hydrolysis under mild conditions might yield similar re- sults. While some desulfation also would occur, hydrolysis with sufficiently dilute alkali would be expected to yield oligosaccharide fragments with retention of at least some of the original sulfate groups (Percival and McDowell, 1967). As predicted, exposure of the decalcified, partic- ulate cell wall fraction to dilute alkali (pH 11-12) released a soluble, sulfate-containing morphogen. The increase in yield of soluble morphogenic activity as a function of the time of incubation at pH 12 is shown in Figure 9. The apparent decline in yield after 6 h suggests that the mor- phogen itself is slowly inactivated at pH 12. Exposure to strong acid (4 N HC1) or strong base (4 TV NaOH) causes more rapid inactivation. The morphogen released by mild alkaline hydrolysis is of relatively low molecular weight, strongly anionic, sul- fate-containing, and unstable. Estimates of the apparent molecular weight were made by three independent meth- ods: ultrafiltration through a calibrated membrane, di- alysis through a calibrated dialysis membrane, and Seph- adex gel filtration (Table IV). In ultrafiltration, all of the applied activity was recovered in the YM-5 membrane ultrafiltrates, consistent with M < 5000 Da. In dialysis through a calibrated membrane, all of the recovered ac- tivity (equal to that in the non-dialyzed control) was found in the first external dialysate, (see Materials and Methods), consistent with M < 2000 Da. These results were con- firmed by gel-filtration through Sephadex G-10, in which all of the applied activity was eluted at the end of the included volume, indicating M < 2000 Da. These findings all indicate, therefore, that the morphogen solubilized by alkaline hydrolysis may be as small as 2000 Da or less (Table IV). This low molecular weight material is strongly anionic; it binds tightly to DEAE Sephadex and to DEAE-nitro- cellulose (Table V). Because the material does not bind to Sephadex or nitrocellulose alone, without DEAE sub- stitution, the binding is most likely dependent on ionic interaction. This binding proves to be very strong; little activity could be removed from the DEAE Sephadex by elution with 0.4 M NaCl, and 75% of the initially applied activity was found still adsorbed to the resin (Table V). Similarly, 63% 13% of the activity applied to a DEAE- nitrocellulose filter was found still adsorbed to the filter, even after elution with 0.4 N HC1. That the morphogen was not eluted from DEAE by 0.4 TV HC1 indicates that its ionic binding was dependent on more strongly anionic groups than uronic acids or other simple carboxylates (pK ca. 2). These results are consistent with the conclusion, based on sulfatase sensitivity (cf. Figs. 4, 5; Table II). that the morphogen contains sulfate groups (see Discussion). The presence of sulfate esters in this solubilized mor- phogen was confirmed independently by turbidimetric analysis with barium after strong acid hydrolysis (method according to Beeley, 1985). The yields obtained indicate Table IV Estimation of molecular weight of the small morphogen solubilized by partial alkaline hydrolysis Method M Estimate (Da) Ultrafiltration Dialysis Sephadex gel filtration <5000 <2000 <2000 Details are as described in Materials and Methods. Independent ex- periments verified that there was no significant adsorption of the mor- phogen to the gel-filtration matrix or to the ultrafiltration or dialysis membranes. 114 D. E. MORSE AND A. N. C. MORSE Table V Adsorption of the small morphoxen. xolubilizeil by partial alkaline hydrolysis, to DEAE Recovery Adsorbant Eluant or fraction Units DEAE Sephadex Application 1,000 1 m.U Tris 1 mM Tns + 0.4 A/ NaCl 50 5 Remaining on DEAE 753 75.3 (Total) (80.3) DEAE Nitrocellulose Application 640 1 mM Tris 1 mM Tns + 0.4 A/ NaCl 0.4 N HCI Remaining on DEAE 400 80 63 13 (Total) (63 13) Details are as described in Materials and Methods. Morphogen re- covered in the high-salt eluate. and that remaining on the DEAE, induced larvae to attach firmly to the polystyrene assay beakers and metamorphose normally. Control experiments demonstrated that the DEAE adsorhants without the applied morphogen had no such activity. Additional controls showed that there is no binding of the soluble morphogen either to Seph- adex or nitrocellulose alone, without the DEAE ion-exchange groups present. the presence ofca. 8-14% (w/w) sulfate in the alkali-sol- ubilized morphogenic fraction. Agaricia humilis larvae exposed to the alkali-solubilized low molecular weight morphogen, either free in solution or bound to DEAE (Sephadex or nitrocellulose) showed rapid, normal, and complete metamorphosis. The activity of both the soluble and DEAE-bound morphogen was concentration- or dose-dependent. Activity, specificity, and stability oj the small soluble morphogen The small, dialyzable morphogens obtained in the ex- periments shown in Figures 6-9 induce rapid and normal larval settlement, attachment, complete metamorphosis, and normal post-metamorphic growth (Fig. 10). These processes induced by the small morphogenic molecules are indistinguishable from those induced by the native, intact nongeniculate coralline alga. The low molecular weight morphogens produced by enzymatic or alkaline hydrolysis of the paniculate cell wall fraction from Hydrolithon boergesenii (or associated microbial symbionts) induce metamorphosis of at least one other agariciid coral, in addition to Agaricia humilis. Larvae of the sympatric A. lenuifolia. which also are in- duced to settle and metamorphose by the intact Hydro- lithon boergesenii (A. Morse et ai, in prep.), are induced to metamorphose by the dialyzable morphogen released by agarase digestion of the cell wall fraction from this alga, with an efficiency comparable to that exhibited for the A. humilis larvae (Table VI). In both of these species, the coral larvae are induced to attach to the walls or bot- tom of the polystyrene containers, metamorphose com- pletely, and begin normal post-metamorphic growth in response only to the low molecular weight chemical mor- phogen. In contrast, larvae of the sympatric ahermatypic coral, Tubastraea aurea, which are not induced to me- tamorphose by Hydrolithon boergesenii. are not induced by the low molecular weight morphogen enzymatically released from the algal cell wall fraction (Table VI). This demonstrates that the morphogenic activity of this chem- ical is biologically specific for those larvae that respond to the intact alga. Whereas the decalcified, cell wall-associated, insoluble morphogen is relatively stable when frozen at - 10C, the small, dialyzable morphogen (M < 8000 Da) released from this parent material by endo-0-galactosidase is stable for only a few days at -10C. The lower molecular weight morphogen released by mild alkaline hydrolysis (M < 2000) proved to be even more unstable. Even when frozen at -10C, this material lost activity with a half- life that varied in different preparations between 24 and 72 h. Activities of known compounds We have tested a wide variety of sulfated and non-sul- fated polysaccharides and related polymers, including several substrates for the enzymes used in this study. These (all pre-adjusted to the pH of ambient seawater prior to testing) have included: agar; agarose; agaric acid; alginic acid; ascophyllan; X-, (-, and K-carrageenans; fucoidan, furcellaran; laminarin; chitin; di-N-acetylchitobiose; tri- N-acetylchitotriose; keratan sulfate; chondroitin sulfates A, B, and C; heparin; heparan sulfate; hyaluronic acid; asialofetuin; dextran sulfate; pentosan sulfate; polyvinyl sulfate; polyanethol sulfate; pectin; amylopectin; cellulose; cellulose sulfate; sulfoethylcellulose; and sulfopropyl- sepharose. Most of these proved inactive with Agaricia humilis larvae. Only the /,-carrageenan (a sulfated polymer rich in galactose), fucoidan (a sulfated polymer rich in fucose), and keratan sulfate (a sulfated glycosaminoglycan) induced metamorphosis, although this activity was weak and evident only at very high concentrations (>5 mg/ml). Discussion Recent field and laboratory studies have shown that recruitment of the shallow- water agariciid corals, Agaricia humilis and A. tenmfolia, is determined in part by larval recognition of a chemical inducer of substratum-specific settlement and metamorphosis (Morse et ai. 1988). This inducer is associated with specific nongeniculate coralline red algae. In the case of A. humilis, only certain coralline CHARACTERIZATION OF CORAL MORPHOGEN 115 Figure 10. Normal attachment and metamorphosis of Agaricia humilis larva induced by the soluble morphogen. Scanning electron micrographs of the endoskeleton of a post-metamorphic corallite attached to polystyrene, dorsolateral (a) and dorsal (b) views, and for comparison, a pre-metamorphic planula larva (c). The pronounced calcified septa and the attachment plaque are clearly visible in the 2-day post-metamorphic corallite. Diameter of the corallite ca. 1.5 0.1 mm; length of the planula ca. 0.7 0.1 mm. The soluble morphogen was obtained by digestion with endo-ff-galactosidase. followed by dialysis, as in Figure 8. Tissue- digestion, fixation, dehydration, critical-point drying, and electron microscopy were by standard procedures. red algae contain the inducing morphogen (Fig. 1: Table I: cf. Morse et ai. 1988; A. Morse and R. Steneck, in prep.). We have extended our previous finding that Agaricia humilis larvae maintain both the stringency of their de- pendence upon the alga-associated morphogen, and the specificity of this requirement, for at least 30 days follow- ing their release (Fig. 1 ). While most larvae probably settle and metamorphose in less than 30 days in the natural environment, this capacity to delay metamorphosis in the absence of the required morphogen can enhance both the dispersal of the larvae, and the substratum-specificity of the final distribution of recruits. Although much of scler- actinian recruitment may be locally seeded (Bak and En- gle, 1979; Rylaarsdam, 1983; Baggett and Bright, 1985; Sammarco and Andrews, 1988), the larvae of A. humilis (cf.Fig. 1 ; also Morse etal., 1988) and certain other species (cf. Harrigan, 1972; Richmond, 1981, 1985, 1987; Har- rison et ai. 1984; Scheltema, 1986; Morse et ai. 1988; Richmond and Hunter, 1990) are capable of distant dis- persal as well. Indeed, temporal pulses of scleractinian recruitment at certain sites on the Great Barrier Reef are dependent on the settlement of larvae produced at distant locations (Wallace, 1985; Babcock, 1988). In addition to the importance of hydrodynamic, topographic, and geo- graphic features for the control of larval dispersal and consequent recruitment (e.g.. Frith et ai. 1986; Rough- garden et ai, 1988; Sammarco and Andrews, 1988; Black, 1988), and the effects of predation, nutrition, competition, and other biotic factors, studies of A. humilis (Morse et ai. 1988; and those reported here) indicate that for some coral species, the stringency and specificity of the larval requirements for the induction of settlement and meta- morphosis also can be important in controlling the spatial distribution of recruits. A similar larval requirement for a substratum-specific biochemical inducer of settlement Table VI Induction oj metamorphosis of Agaricia humilis and A. tenuifolia /an'ae. and absence of induction o/'Tubastraea aurea, by the low molecular weight (LA/H'J morphogen Treatment Metamorphosis at 24 h (X S.D.) LMW morphogen Seawater control Agaricia humilis Agaricia tenuifolia Tubastraea aurea 95% 10% (n = 4) 0% 0% (n = 5) 87% 23% (n = 3) 0% 0% (n = 5) 0% 0% (n = 3) 0% 0% (n = 3) Soluble morphogen was prepared by agarase digestion of the particulate cell wall preparation of Hydrolithon boergesenii (agarase. 1 mg. 1000 units), and dialysis through a membrane retaining molecules with M > 14,000 Da, as described in Materials and Methods. Larvae of each of the three coral species were assayed for metamorphosis in response to the dialyzable morphogen and seawater controls in parallel: 5 larvae/trial: n = number of trials. 116 D. E. MORSE AND A. N. C. MORSE and metamorphosis, and larval recognition of the chem- ical inducer in the ocean environment, recently were shown to determine the fine-scale spatial distribution of recruitment of the polychaete, Phragmatopoma califomica (Jensen and Morse, 1990). Chemical nature ofihe inducer The inducer of Agaricia humilis settlement and meta- morphosis that is associated with specific nongeniculate coralline red algae is chemical in nature (Table I; cf. Morse el a/.. 1988). This morphogenic substance is insoluble in a wide range of solvents, apparently because it is associated with cell wall polysaccharides (with which the morphogen is partially purified; cf. Fig. 2). Solubilized fractions of the inducer can be generated, however, by hydrolysis with enzymes or mild alkali. The fact that these solubilized fractions are sufficient to trigger normal attachment and metamorphosis of A. humilis larvae on clean polystyrene surfaces (Fig. 10) proves that the requirement of the larvae for a morphogenetic inducer is satisfied by a chemical substance from the inductive algal substratum. Non-re- cruiting algal substrata do not yield this activity (Table I). These results thus confirm and extend the finding that larval recognition of the inductive but insoluble paniculate fraction, partially purified from the settlement-inducing algal surface, is dependent upon the integrity of a perio- date-sensitive (Fig. 3) and sulfatase-sensitive (Fig. 5; Table II) chemical structure. These sensitivities, the insolubility of the crude inducer, and its insensitivity to proteolytic enzymes, suggested that the inducer is associated with a sulfated polysaccharide. Consistent with this suggestion, we have found that the inducer can be solubilized by cleavage with purified en- doglycosidases that can act on sulfated polysaccharides (Figs. 6-8). As summarized in Table III. the cleavage by agarase and by endo-0-galactosidase indicates that the substrate polymer contains galactose units; cleavage by lysozyme and by endo-0-galactosidase indicates that this polymer also contains 0-1,4 linked N-acetylglucosamine or N-acetylglucosamine sulfate units (cf. Li el a/.. 1982; Kitamikadocfa/., 1982; Scudder et at.. 1983, 1984; Mor- rice et at.. 1983a, b; Usov and Ivanova, 1987). Because continued hydrolysis by each of these enzymes leads first to solubilization. and then to progressive inactivation of the inducer, the inductive moiety itself probably contains the above-mentioned sites of hydrolysis. The morpho- genetic inducer is thus associated with, and may itself contain, a sulfated glycosaminoglycan, i.e., a sulfated polysaccharide that includes multiple N-acetylglucos- amine and galactose units. The finding that partial alkaline hydrolysis liberates a sulfate-containing, strongly anionic, small morphogen is consistent with this conclusion. Re- cently, hydrolysis, ion-exchange HPLC, and sensitive de- tection of the resolved monosaccharides by pulsed am- perometry have independently confirmed that glucos- amine (derived from N-acetylglucosamine) and galactose are indeed principal components of this solubilized small morphogen (M. Hardy, Dionex Corp., pers. comm.). Sul- fated polysaccharides that contain amino sugars and are rich in galactose, similar to those in foliose red algae (cf, Percival and McDowell, 1967; Percival, 1978; Mc- Candless, 1981; Yaphe, 1984), previously were found in other coralline red algae (Turvey and Simpson, 1965). Keratan sulfate, which proved to be slightly active as a morphogen, has three characteristics that make it unique among the known sulfated glycosaminoglycans tested. In these same features, it also, to some degree, resembles the morphogen recognized by Agaricia humilis larvae: ( 1 ) it contains multiple galactose-/3-l,4-N-acetylglucosamine units (whereas heparin, heparan sulfate, and the three dif- ferent chondroitin sulfates all contain glucuronic acid or other uronic acid units in place of galactose); (2) it is readily cleaved by endo-0-galactosidase (Scudder el at.. 1983), whereas the other known compounds tested are not; and (3) the keratan sulfates from some sources cannot be eluted from anion exchangers at salt concentrations below 3-4 M (Roden et til., 1972; cf. our results with DEAF, Table V), whereas the other sulfated glycosami- noglycans are readily eluted at significantly lower con- centrations (Roden et at.. 1972). Very highly sulfated gal- actans from red algae also are not eluted from DEAF, or else are eluted only at high concentrations of urea or at high temperature (Yaphe, 1984). However, the weak morphogenetic activity evident at only high concentra- tions of keratan sulfate, carrageenan, and fucoidan indi- cates that although these substances may be somewhat similar to the natural inducer, they differ in structural features (possibly including positions of the sulfate groups or other linkages) from those of the natural morphogen. The morphogenetic activity of the natural inducer sol- ubilized from the cell wall fraction obtained from Hy- drolithon boergesenii or its associated microflora is bio- logically specific (Table VI). This molecule induces normal attachment and metamorphosis in larvae of two species of Agaricia that are also induced by the intact alga from which the morphogen was obtained. However, the mol- ecule fails to induce these reactions in larvae of the sym- patric Tubastraea aurea. which are not induced by the intact alga. The high specificity of the Agaricia humilis larvae for morphogens associated with only certain nongeniculate coralline red algae (Fig. 1; Table I; Morse et a/.. 1988; A. Morse and R. Steneck, in prep.) is apparently a reflection of the chemical specificity of the larval receptors for only some unique sulfated glycosaminoglycans. This suggestion is supported by the finding that a wide variety of synthetic natural sulfated polysaccharides. glycosaminoglycans, and CHARACTERIZATION OF CORAL MORPHOGEN 117 structurally related polymers have either little or no mor- phogenetic activity. A better understanding of the chem- ical basis for the high specificity of the larval receptors awaits further information about the stereochemistry and structure of the organic sulfate esters and other substitu- ents in the natural morphogen. Relation to other systems Sulfated glycosaminoglycans and other sulfated poly- saccharides have been widely implicated in other highly specific cell recognition phenomena that control differ- entiation, including inter-phyletic symbiosis, fertilization, aggregation of sponge cells, the "homing" of circulating mammalian lymphocytes into the lymph nodes, pattern formation in the developing nervous system, and the me- tastasis and invasiveness of tumors. The number of struc- tural permutations of such molecules, and hence the complexity of coding of cell recognition ideotypes, far ex- ceed those of the proteins, which are more fully under- stood (Drickamer, 1988; Sharon and Lis, 1989). Recent evidence has shown that the host specificity for nodule induction in leguminous plants by the nitrogen- fixing symbiotic bacterium, Rliiiobium meliloti, is deter- mined by recognition of a unique sulfated glycosamino- glycan signal (a tetrasaccharide containing one sulfate and one acylamino group) produced by the bacterium (Ler- ouge el al.. 1990). Recognition, by the host plant, of this inducing bacterial signal, and the resulting specificity of the inter-phyletic symbiosis, has long been thought to be governed by a class of receptors known as lectins, present on the plant root hair surfaces (Kijne et ai. 1989). Re- cently, this suggestion was strongly confirmed when the intergeneric transfer between plants of the DNA coding for one such root lectin resulted in the transfer of host specificity for nodulation induced by the bacterium (Diaz et ai. 1989). Recognition of specific sulfated polysaccha- rides at the surface of Strongylocentrotus pwpiiratus sea urchin eggs by conspecific sperm has been demonstrated to be essential for fertilization (Rossignol et al., 1984; DeAngelis and Glabe, 1987). In this reaction (DeAngelis and Glabe, 1987), and in the binding of heparin to anti- thrombin (Atha et ai, 1985), the locations of the sulfate esters on the sugars have been found to be of critical im- portance; in the latter case, removal of one specific sulfate reduced the affinity of specific binding by as much as 10,000-fold (Atha et ai, 1985). The lengths of the sulfated polysaccharides are also critical determinants of the strength (and hence, the specificity) of these binding re- actions (Hoylaerts et ai, 1984; DeAngelis and Glabe, 1987). Evidence also suggests that lectin-like recognition of cell surface sulfated glycosaminoglycans controls an essential phase in the species-specific reaggregation and subsequent differentiation of sponge cells (Henkart et ai. 1973; Turner and Burger, 1973; Jumblatt et ai, 1980; Conrad et ai, 1984; Diehl-Seifert et ai, 1985, 1989; Mar- goliash et ai, 1965; Coombe et ai, 1987; Coombe and Parish, 1988; Schroder et ai, 1988;Gramzowe/a/.. 1989). Recognition and binding of sulfated glycosaminoglycan moieties of cell surface proteoglycans also controls adhe- sion, tissue-specific differentiation and growth in a wide variety of mammalian and other higher systems (Edelman, 1985; Fransson, 1987; Ruoslahti, 1989). The structures and positions of the sulfate esters can be critically im- portant in the control of these functions as well (Fransson, 1987). Sulfated glycosaminoglycans and other sulfated polysaccharides bind to lectin-like receptors on the sur- faces of lymphocytes (Parish et ai, 1984; Parish and Snowden, 1985; Chong and Parish, 1985, 1986; Thurn and Underbill, 1986; Brenan and Parish, 1986; Brandley et ai. 1987). Recently, the genes coding for two distinct "homing receptors" from the surfaces of mammalian lymphocytes have been cloned and sequenced (Gallatin et ai, 1986; Yednock et ai, 1987a; Goldstein et ai, 1989; Holzmann et ai, 1989; Siegelman et ai. 1989; Stamen- kovic et ai, 1989; Stoolman, 1989). Lectin-like recogni- tion of specific sulfated or other anionic carbohydrates in the target lymphoid tissues, mediated by these receptors, is thought to direct the homing of specific subsets of lym- phocytes from the circulation to adhere to the blood vessel endothelia of their target lymphoid organs (lymph nodes. Peyer's patches, etc.), where the recruited lymphocytes then differentiate to produce antibodies (Brenan and Par- ish, 1986; Gallatin et ai, 1986; Yednock et ai, 1987a, b; Jalkanenc/a/.. 1988; Stoolman, 1989; Coombe and Rider, 1989). The parallels between this lymphocyte homing reaction and the settlement and metamorphosis ofAgaricia humilis larvae are potentially interesting. In both, substratum- specific "recruitment," attachment, and differentiation are apparently induced by recognition of a non-diffusing, substratum-specific sulfated polysaccharide. Our further observation that the partially purified sulfated polysac- charide that induces A. humilis larvae to attach and me- tamorphose also induces murine lymphocytes to undergo mitosis (Morse and Eardley, unpub. obs.), may therefore be worth further investigation. The specificity of this latter reaction is unclear, however, as a wide variety of sulfated polysaccharides and related polymers, including several carrageenans, fucoidan, and ascophylan, induce lympho- cyte mitosis. Whether the same subset of lymphocytes responds to each of these compounds has not yet been determined. In contrast, the A. humilis larvae respond only slightly and incompletely to carrageenan and fucoi- dan, and only at very high concentrations. Cell-surface recognition of polysaccharides and other complex carbohydrates in such non-immune systems is generally considered to be mediated by lectins, a broad 18 D. E. MORSE AND A. N. C. MORSE class of ubiquitous, carbohydrate-specific receptors that recently has been redefined (Barondes, 1988; Drickamer, 1988; Sharon and Lis, 1989). Mitchell and his colleagues first demonstrated that the settlement and metamorphosis of larvae of the polychaete, Janua brasiliensis, are me- diated by a lectin-like recognition of inductive exopoly- saccharides produced by specific bacteria (Kirchman el a/., 1982a, b; Mitchell and Kirchman, 1984; Maki and Mitchell, 1985, 1986). These authors first pointed out the similarities between this larval settlement reaction and other lectin-mediated recognition phenomena, including the root nodule-bacteria symbiosis discussed above. Wei- ner et al. (1985) and Bonar el al. ( 1986) also have shown that specific bacterial exopolysaccharides may play a role in the induction of settlement and metamorphosis of Crassostrea virginica and C. gigas oyster larvae, although induction in those systems is complex, and more than one class of compound is known to be involved (Coon et al., 1985; Coon and Bonar, 1987; Fitt el al., 1989; Bonar et al., 1990). The results reported here, demonstrating sensitivity of the Agaricia humilis morphogen to cleavage by endo-/3- galactosidase and agarase, (and resistance to the exogly- cosidic /3-galactosidase), indicate that the morphogen contains essential internal /3-galactoside units. This finding may be of particular interest in view of the suggested im- portance of j3-galactoside-specific lectins in controlling differentiation in higher animal systems (Barondes et al., 1988; Sharon and Lis. 1989). Remaining problems Two problems remaining are the determination of the complete chemical structure of the inducing molecule recognized by the Agaricia humilis larvae, and the un- equivocal identification of the biological source of this inducer. A number of compounds that induce metamor- phosis of various marine invertebrate larvae have been partially purified from the respective inductive substrata, and the structures of these compounds have been partially characterized. But we are aware of only two natural in- ductive molecules that have been completely character- ized. These include the algal molecules that induce meta- morphosis of the scallop, Pecten ma.\imits (Yvin et al., 1985), and the hydrozoan, Corync uchidai (Kato et al., 1975). Significantly, both of these are small molecules soluble in organic solvents, and thus amenable to gas- chromatography and mass spectroscopy. In a large num- ber of the other cases investigated, however, the native inducers have proved to be either water soluble (e.g., Highsmith, 1982; Hadfield and Scheuer, 1985; Burke, 1 986; Hadfield and Pennington, 1 990), or polymeric and insoluble (e.g., Jensen and Morse, 1984. 1990; Morse et al.. 1988). [We are not including in this discussion such molecules as potassium or calcium salts, fatty acids, cyclic nucleotides, or other widely active effectors of depolar- ization, protein phosphorylation, or signal transduction pathways: these all have been shown to induce meta- morphosis of larvae without species- or substratum-spec- ificity (eg., Baloun and Morse, 1984; Morse, 1985, 1990; Yool et al.. 1986; Pechenik and Heyman, 1987; Jensen and Morse, 1990; Jensen et al., 1990)]. The inducer of Agaricia humilis metamorphosis described here is in its native form associated with a substratum-specific insol- uble polymer. Partial hydrolysis with either enzymes or dilute alkali releases a smaller, water-soluble and strongly anionic inducer which is markedly unstable. This insta- bility has hindered analyses of the active morphogen. The use of highly purified enzymes, and employment of their specificities for selective cleavage, solubilization, and as probes of the structural determinants of morphogenetic activity, may prove widely useful in further studies of such otherwise intractable molecules. The inductive molecule that we have described is ob- tained from homogenates of the nongeniculate coralline red alga, Hydrolithon boergesenii. Larvae of Agaricia hu- milis are induced to metamorphose by contact with intact specimens of that alga, and recruits of the coral are found preferentially on that alga in the field (Morse et al., 1988; A. Morse and R. Steneck, in prep.). Three other species of anthozoan, including the scleractinian Agaricia ten- iiifo/ia (Morse et al., 1988), the temperate octocoral Al- cyonnim sidcriiim (Sebens, 1983a. b), and a tropical gor- gonian, Plexanra sp. (Lasker, 1990), also have been found to settle and metamorphose in response to crustose red algal surfaces. But in each of these cases, the inductive molecule could have been produced by bacteria or other microorganisms associated with the algal surfaces. Bacteria or bacterial films have been implicated in the control of larval settlement and metamorphosis in a few other cni- darians in which these processes are chemically induced. Larvae of the hydroid, Hydractinia echinata. settle and metamorphose in response to films of the bacterium, Al- teromonas sp., on shells inhabited by hermit crabs (Spin- dler and Miiller, 1972). Cassiopea andromeda (scypho- zoan) larvae also are induced to settle and metamorphose by bacteria, apparently in response to soluble peptides produced by the action of bacterial degradative enzymes (Fitt and Hofmann, 1985; Fitt et a/., 1987; Hofmann and Brand, 1987). Bacterial films have been widely implicated in the control of larval settlement and metamorphosis in many other kinds of invertebrates as well (Wilson, 1955; Cameron and Hinegardner, 1974; Brancato and Woola- cott, 1982; Kirchman et at.. 1982; Mitchell and Kirchman, 1984; Bonar et al. 1986; Maki and Mitchell. 1986; Fitt et al.. 1989; Maki et al., 1990). Moreover, as Maki et al. ( 1990) have suggested in the case of barnacle larvae, the larval response may depend in a complex way on the CHARACTERIZATION OF CORAL MORPHOGEN 119 interaction between bacteria and the surface on which they are attached. The structure of the morphogen recognized by Agaricia hunulis larvae, suggested by the results reported here to be a sulfated glycosaminoglycan, would be equally con- sistent with a molecule of the red algal cell wall and of a cell wall or other exopolymer produced by an associated bacterium (Percival and McDowell, 1967; Mackie and Preston, 1974; Sanford et a/., 1977; McCandless, 1981; Drews and Weckesser, 1982; Boyle and Reed, 1983). We have found, however, that the activity of crude homog- enates appears markedly enhanced following decalcifi- cation, consistent with the unmasking of constituents of the algal cell wall. Attempts to culture the algal cells ax- enically from isolated protoplasts, by the methods of Polne-Fuller and Gibor (1984) and Kloareg et al. (1989), and attempts to culture the alga-associated microbial symbionts, may help further resolve the source of the in- ducer. 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