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Proceedings National Academy of Sciences PROCEEDINGS OF THE NATIONAL ACADEMY OF SCIENCES OF THE UNITED STATES OF AMERICA Officers BRUCE ALBERTS, President of the JACK HALPERN, Vice President Academy PETER H. RAVEN, Home Secretary F. SHERWOOD ROWLAND, Foreign Secretary MILDRED S. DRESSELHAUS, Treasurer Editor-in-Chief NICHOLAS R. COZZARELLI Editorial Board PETER J. BICKEL JACK HALPERN PAUL R. SCHIMMEL of the MICHAEL T. CLEGG ERIC R. KANDEL STUART L. SCHREIBER Proceedings MARSHALL H. COHEN RICHARD A. LERNER CARLA J. SHATZ STANLEY N. COHEN HARVEY F. LODISH CHRISTOPHER A. SIMS MAX D. COOPER PHIL W. MAJERUS ALLAN C. SPRADLING JAMES E. DARNELL, JR. ARNO G. MOTULSKY LARRY R. SQUIRE IGOR B. DAWID RONALD L. PHILLIPS CHARLES F. STEVENS HERMAN N. EISEN TOM POLLARD JOANNE STUBBE RAYMOND L. ERIKSON STANLEY B. PRUSINER KARL K. TUREKIAN RONALD M. EVANS CHARLES RADDING IRVING L. WEISSMAN NINA FEDOROFF GIAN-CARLO ROTA SHERMAN M. WEISSMAN CHARLES FEFFERMAN DAVID D. SABATINI PETER G. WOLYNES JOSEPH L. GOLDSTEIN GOTTFRIED SCHATZ Publisher: KENNETH R. FULTON Managing Editor: FRANCES R. ZWANZIG Associate Editorial Manager: JOHN M. MALLOY Associate Manager for Production: JOANNE D'AMiCo Author/Member Support Coordinators: REID S. COMPTON, BARBARA A. BACON System Administrator: MARILYN J. MASON Manuscript Processor: JACQUELINE V. PERRY Secretary: BRENDA L. MCCOY Administrative/Systems Aide: DomTTE A. MAY Subscription Fulfillment: JULIA A. LiTTLE Office Assistant: CYNTHIA MATHEWS Correspondence: PROCEEDINGS OF THE NATIONAL ACADEMY OF SCIENCES, 2101 Constitution Avenue, NW, Washington, DC 20418 (via U.S. postal service) or 1010 Wisconsin Avenue, NW, Suite 530, Washington, DC 20007 (via courier service). Information for Contributors: See pp. i and ii (of this issue). Copyright: Volumes 90-93, copyright C) 1993-1996 by the National Academy of Sciences; Volumes 1-89, copyright as a collective work only with copyright to individual articles retained by the author(s). Requests for permission to reproduce all or parts of individual articles published in Volumes 1-89 should be addressed to the authors. Microforms of complete volumes are available to regular subscribers only and may be obtained from University Microfilms, Xerox Corporation, Ann Arbor, MI 48103. This journal is printed on acid-free paper effective with Volume 84, Issue 1. Subscriptions: All correspondence concerning subscriptions should be addressed to the Circulation Office of the PROCEEDINGS. Subscriptions are entered on a calendar year basis only. For 1996, subscription rates are as follows-in the United States: personal, $250; institutional, $595; elsewhere by surface mail: personal, $350; institutional, $695; elsewhere by Air Cargo at a surcharge of $208. Information regarding other air mail postage rates is available from the Circulation Office. 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PRINTED IN THE USA PROCEEDINGS OF THE NATIONAL ACADEMY OF SCIENCES OF THE UNITED STATES OF AMERICA (ISSN-0027-8424) is published biweekly by THE NATIONAL ACADEMY OF SCIENCES, 2101 Constitution Avenue, NW, Washington, DC 20418. 1996 by THE NATIONAL ACADEMY OF SCIENCES OF THE UNITED STATES OF AMERICA. POSTMASTER: Send address changes to: PROCEEDINGS OF THE NATIONAL ACADEMY OF SCIENCES OF THE UNITED STATES OF AMERICA, 2101 Constitution Avenue, NW, Washington, DC 20418. Coverphotograph: Hildreth Meiere's emblematic figure depicting Geology: One ofeight radialpanels on the dome ofthe Great Hall at the National Academy of Sciences. Proc. Natl. Acad. Sci. USA Vol. 93, June 1996 INFORMATION FOR CONTRIBUTORS (Revised May 1996) PURPOSE AND SCOPE the manuscript. All funding sources supporting the work must The PROCEEDINGS OF THE NATIONAL ACADEMY OF SCIENCES be acknowledged. USA publishes research reports, commentaries, reviews, collo- (iv) A completed form, assigning COPYRIGHT to the Na- quium papers, and actions of the Academy. In accordance with tional Academy of Sciences, must be returned to the PRO- the guiding principles established by George Ellery Hale in 1914, CEEDINGS office after the paper is accepted for publication. the PROCEEDINGS publishes brief first announcements of the (v) The Academy may distribute EMBARGOED copies of an Members' and Foreign Associates' (hereafter referred to as the article to the press prior to publication. Members) more important contributions to research and ofwork (vi) Manuscripts that report research involving HUMAN that appears to a Member to be of particular importance. SUBJECTS or the use of materials of human origin must be RESEARCH REPORTS describe the results of original experi- accompanied by institutional authorization. mental or theoretical research of exceptional importance. The (vii) Submission of manuscripts on research using RECOM- papers should be of the highest scientific quality and should be BINANT DNA implies that physical and biological containment intelligible to a broad scientific audience. conformed to National Institutes of Health guidelines or those COMMENTARIES call attention to papers of particular note of a corresponding agency. and are written at the invitation of the Editorial Board. (viii) Submission of manuscripts on research using UNIQUE REVIEws are usually solicited by the Editorial Board. MATERIALS (e.g., cloned DNAs; antibodies; bacterial, animal, COLLOQUIUM PAPERS are reports of symposia held under or plant cells; viruses; and computer programs) implies that the Academy auspices. authors will make them available to qualified researchers for noncommercial use. EDITORIAL POLICIES (ix) DATA BASES: Accession codes must be supplied for Submission and Review publication. Nucleic Acid Sequences: Authors must submit these data to Research Reports and Reviews may be submitted to the GenBank/EMBL/DNA Data Bank of Japan, and the acces- PROCEEDINGS through any of the following three tracks: sion number of the sequence must be supplied for publication. I. A Member of the Academy may "communicate" manu- Crystallographic Studies of Proteins and Other Biopolymers scripts for non-members, provided that the communicating Except Oligonucleotides: Authors must submit these data to the Member ensures that the paper undergoes thorough peer Protein Data Bank (PDB; Brookhaven National Laboratory). review prior to submission to the PROCEEDINGS. 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