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Biochemistry Biochemistry ( I S S N 0006-2960) is published biweekly by the American Chemical Society at 1155 16th Street, N . W . , Washington, D.C. 20036. Second-class postage paid at Washington, D . C , and additional mailing offices. P O S T M A S T E R : Send address changes to Biochemistry, Membership and Subscription Services, P.O. Box 3337, Columbus, Ohio 43210. American Chemical Society 1155 16th Street, N . W . Washington, D.C. 20036 (202) 872-4600 [TDD (202) 872-8733]

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Membership and Subscription Services American Chemical Society P.O. Box 3337, Columbus, Ohio 43210 (614) 421-3776 Journals Department American Chemical Society 2540 Olentangy River Road P.O. Box 3330, Columbus, Ohio 43210 (614) 421-3600, ext 3171 ( T E L E X 6842086) © Copyright 1986 by the American Chem­ ical Society. Copyright permission: An indi­ vidual may make a single reprographic copy of an article in this publication for personal use. Reprographic copying beyond that permitted by Section 107 or 108 of the U.S. Copyright Law is allowed, provided that the appropriate per-copy fee is paid through the Copyright Clearance Center, Inc., 27 Congress St., Salem, Massachusetts 01970. For reprint permission, please write to the Copyright Administrator, Books and Journals Division, at the ACS Washington address. Editorial

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Instructions to authors and copyright status form are printed in the first issue of each vol­ ume. Please conform to these instructions when submitting manuscripts. Manuscripts for publication should be sub­ mitted to the Editor, Hans Neurath, Depart­ ment of Biochemistry, University of Washing­ ton, Seattle, Washington 98195. Correspond­ ence regarding accepted papers and proofs should be directed to the Journals Department at the address above. Page charges of $30.00 per page may be paid for papers published in this journal. Payment does not affect accept­ ance or scheduling of papers. Bulk reprints of individual articles are available. For informa­ tion write to Distribution Office at the ACS Washington address. The American Chemical Society and its Ed­ itors assume no responsibility for the statements and opinions advanced by contributors. Registered names and trademarks, etc., used in this publication, even without specific indi­ cation thereof, are not to be considered unpro­ tected by law. Subscription and Business Information 1986 subscription prices including postage. All Other Europe Countries Air Service Air Service Included Mexico Included

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Biochemistry EDITOR

HANS NEURATH Department of Biochemistry University of Washington Seattle, Washington 98195 (206) 543-1690

ASSOCIATE EDITORS E A R L W.

DAVIE

University of Washington EDMOND H. FISCHER

University of Washington M I L T O N P. G O R D O N

University of Washington W I L L I A M W.

PARSON

University of Washington

EDITORIAL ADVISORY BOARD ROBERT H. ABELES

CLAUDE HÉLÈNE

Brandeis University

DEMETRIOS PAPAHADJOPOULOS

University of California, San Francisco

DAVID S. AULD

Museum National d'Histoire Naturelle, Paris

Harvard Medical School

JOHN W. B. HERSHEY

Massachusetts Institute of Technology

ROBERT L. BALDWIN

University of California, Davis

DAVID C. PHILLIPS

Stanford University

PETER C. HINKLE

University of Oxford

STEVEN J. BENKOVIC

Cornell University, Ithaca

JAMES H. PRESTEGARD

The Pennsylvania State University

TONY E. HUGLI

Yale University

GUNTER BLOBEL

Scripps Clinic & Research Foundation

PETER REICHARD

The Rockefeller University

HANS JÖRNVALL

Karolinska Institute, Stockholm

AKSEL A. BOTHNER-BY

Karolinska Institute, Stockholm

JAMES F. RIORDAN

Carnegie-Mellon University

H. RONALD KABACK

Harvard Medical School

BILL R. BRINKLEY

Roche Institute of Molecular Biology

RANDY W. SCHEKMAN

Baylor College of Medicine

PAUL KAESBERG

University of California, Berkeley

ERNESTO CARAFOLI

University of Wisconsin

HAROLD A. SCHERAGA

ΕΤΗ, Zurich

NEVILLE R. KALLENBACH

Cornell University, Ithaca GÜNTHER SCHÜTZ

MARY LOU PARDUE

MINOR J. COON

University of Pennsylvania

The University of Michigan

GEORGES KOEHLER

RAYMOND A. DWEK

Basel Institute of Immunology, Basel

German Cancer Research Center, Heidelberg

University of Oxford

JOSEPH KRAUT

EDWARD M. SCOLNICK

WERNER W.

University of California at San Diego

Merck Sharp & Dohme

German Cancer Research Center, Heidelberg

ROBERT J. LEFKOWITZ

G. GRAHAM SHIPLEY

Duke University

Boston University

CARL FRIEDEN

HENRY METZGER

ANTERO G. S O

IRVING H. GOLDBERG

NIADDK, National Institutes of Health

CHARLES C. SWEELEY

Harvard Medical School

PAUL L. MODRICH

Michigan State University

RICHARD W. HANSON

Duke University

EDWIN W. TAYLOR

Case Western Reserve University

ELIZABETH NEUFELD

University of Chicago

WILLIAM F. HARRINGTON

University of California, Los Angeles

Johns Hopkins University

BERT L. VALLEE

JAMES OFENGAND

Harvard Medical School

ROBERT HASELKORN

Roche Institute of Molecular Biology

KENSAL Ε. VAN HOLDE

FRANKE

Washington University

University of Miami

University of Chicago

Oregon State University

OSAMU HAYAISHI

SALIH J. WAKIL

Osaka Medical College

Baylor College of Medicine