Nucleic Acids Research

Executive editors

R.T.Walker, Birmingham, UK B.E.Griffin, London, UK P.Pearson, Baltimore, MD, USA R.J.Roberts, Cold Spring Harbor, R.B.Hallick, Tucson, AZ, USA J.M.Rosen, Houston, TX, USA NY, USA J.E.Hearst, Berkeley, CA, USA I.C .Eperon, Leicester, UK N.C.Martin, Louisville, KY, USA M.J.Gait, Cambridge, UK W.R.McClure, Pittsburgh, PA, USA Editorial board

M.Beato, Marburg, FRG W.T.Garrard, Dallas, 7X, USA S.E.V.Phillips, Leeds, UK A.J.Berk, Los Angeles, CA, USA F.Giannelli, London, UK R.H.Reeder, Seattle, WA, USA A.P.Bird, Edinburgh, UK P.J.Hagerman, Denver, CO, USA C.W.Schmid, Davis, CA, USA J.L.Bos, Utrecht, The Netherlands C.Helene, Paris, France Y.Shimura, Kyoto, Japan H.Buc, Paris, France A.J.Jeffreys, Leicester, UK R.Simpson, Bethesda, MD, USA K.Calame, New York, NY, USA C.J.Leaver, Oxford, UK D.Soll, New Haven, CT, USA N.Chua, New York, NY, USA H.Lehrach, London, UK B. Sollner-Webb, Baltimore, MD, USA D.A.Clayton, Stanford, CA, USA D.M.J.Lilley, Dundee, UK S.Spadari, Pavia, Italy A.Das, Farmington, CT, USA T.Lindahl, London, UK R.H.Symons, Adelaide, Australia M.L. DePamphilis, Nutley, NJ, USA S.M.Linn, Berkeley, CA, USA H.F.Tabak, Amsterdam, The M.P.Deutscher, Farmington, CT, USA J.T.Lis, Ithaca, NY, USA Netherlands J.E.Donelson, Iowa City, IA, USA R.McMacken, Baltimore, MD, USA S.Tilghman, Princeton, NJ, USA F.Eckstein, Gottingen, FRG D.A.Melton, Cambridge, MA, USA I.Tinoco, Jr, Berkeley, CA, USA J.D.Engel, Evanston, IL, USA J.R.Nevins, Durham, NC, USA Y.Wang, Shanghai, China P.T.Englund, Baltimore, MD, USA S.Nishimura, Tokyo, Japan D.J.Weatherall, Oxford, UK D.J.Finnegan, Edinburgh, UK E.Ohtsuka, Sapporo, Japan A.M.Weiner, New Haven, CT, USA H.Fukuhara, Orsay, France M.Olson, St Louis, MO, USA G.Zon, Foster City, CA, USA D.Gallwitz, Gottingen, FRG L.Philipson, Heidelberg, FRG

Editorial and Production

Joy Walker, UK Editorial Office Susan Jefferies, Production Editor Carol Cook, US Editorial Office Joyce Connolly, Production Assistant Sarah Brennan, Production Assistant

0IRL atPRESS OXFORD UNIVERSITY PRESS Oxford New York Tokyo Subscriptions

An annual subscription to Nucleic Acids Research comprises 24 issues plus one © IRL Press at Oxford University Press 1991. All rights reserved; no part of supplement, with an Annual Author and Subject Index. Subscriptions are entered this publication may be reproduced, stored in a retrieval system, or transmitted on a calendar year basis only. Price includes handling and delivery, by air beyond in any form or by any means, electronic, mechanical, photocopying, recording, Europe. or otherwise without either the prior written permission of the Publishers, or a licence permitting restricted copying issued in the UK by the Copyright Licensing Annual subscription rate (Volume 20, 1992): Agency Ltd, 90 Tottenham Court Road, London WIP 9HE, or in the USA by UK and Europe £625, Rest of World $1050. Back volume prices and subscription the Copyright Clearance Center, 27 Congress Street, Salem, MA 01970. rates for individuals are also available on request. Back volumes of this journal are available in hardcover from the publisher and Orders. Orders and payments from, or on behalf of, subscribers in the various in 16 mm microfilm, 35 mm microfilm and 105 microfiche from University geographical areas shown below should be sent to the Press office indicated. Microfilms International, 300 North Zeeb Road, Ann Arbor, MI 48106-1346, USA. Copies of articles published are also available from UMI. The Americas: Oxford University Press, PO Box Q, McLean, VA 22101, USA. Published by IRL Press at Oxford University Press, Eynsham, Oxford, UK. Japan: payment in Yen to sole agents (details on request), USACO Corporation, Tsutsumi Building, 13-12 Shimbashi 1-chome, Minato-ku, Tokyo 105, Japan. Second class postage paid at Jamaica, NY 11431, USA. Rest of the World: Oxford University Press, Pinkhill House, Southfield Road, POSTMASTER: send address changes to Nucleic Acids Research, Publications Eynsham, Oxford OX8 1JJ, UK. Expediting Inc., 200 Meacham Avenue, Elmont, NY 11003. Tel: (+44865 or 0865) 882283; telex: OXPRES 837330; Fax: (+44865 or 0865) 882890. Advertising. To advertise in Nucleic Acids Research contact Oxford University Press (US office) in the Americas or Oxford University Press (UK office) in the Disc conversion/typeset and printed by Information Press Ltd, Oxford, UK on Rest of the World (see addresses above). Permanent Paper. Author index 5811

Abe,T. 5797 Dunn,R.J. 5673 Kobayashi,Y. 5792 Adra,C.N. 5755 Ecker,D.J. 5743 Koczan,D. 5591 Akhtar,S. 5543,5551 Endo,T. 5537 Kopun,M. 5725 Alloing,G. 5525 Evans,G.A. 5796 Kreklywich,C.N. 5794 Alvarrez,V. 5806 Fang,G. 5515 Krumlauf,R. 5497 Arinami,T. 5801 Farrar,G.L. 5800 Kuhlemeier,C. 5491 Arnold,H.H. 5645 Flavell,A.J. 5533 Kumar-Singh,R. 5800 Atkins,J.F. 5607 Fodde,R. 5805 Lai,K.-M. 5763 Auld,V.J. 5673 Fortini,P. 5569 Landfear,S. 5787 Azorin,F. 5639 Fox,R. 5795 Lane,S. 5793 Backendorf,C. 5633 Franze,A. 5561 Lavedan,C. 5804 Bagley,R. 5802 Fregeau,C.J. 5583 Le Blancq,S.M. 5790 Basu,S. 5551 Freier,S.M. 5743 Leclair,B. 5755 Beggs,J.D. 5483 Froguel,P. 5799 Lench,N.J. 5796 Belt,P.B.G.M. 5633 Fuchs,P.P. 5603 Lesage,S. 5799 Bentley,D.R. 5803 Gardner,E. 5795 Leupin,W. 5507 Bernard,L.E. 5794 Gecz,J. 5806 Liu,Q. 5703 Bernardi,G. 5561 Gesteland,R.F. 5607 L6pez-Larrea,C. 5806 Best-Belpomme,M. 5689 Geyer,C.K. 5799 Lovell-Badge,R. 5497 Bignami,M. 5569 Ghahraman,P. 5788 Lucas,A.M. 5802 Bleackley,R.C. 5583 Gonzalez,C. 5743 Ma,C. 5749 Bobrow,M. 5803 Griffiths,D.E. 5791 Macaya,G. 5561 Bonfanti,M. 5739 Grimaldi,G. 5661 MacDonald,M.H. 5575 Braun,T. 5645 Gupta,S.K. 5673 Manzini,G. 5625 Braus,H. 5785 Hagamuchi,H. 5801 Maraia,R.J. 5695 Breakfield,X.O. 5798 Hanson,S. 5787 Marino,M. 5661 Breukel,C. 5804,5805 Hardesty,B. 5749 Martinez-Balbas,M.A. 5639 Brieley,C. 5533 Hayakawa,H. 5597 Martinez-Naves,E. 5806 Broggini,M. 5739 Herman,B. 5543 Marzluff,W.F. 5653 Broom,A.D. 5719 Hickson,I.D. 5519 Matassi,G. 5561 Bryant,L.A. 5533 Hill,J. 5791 Mathew,C.G. 5803 Byrne,J. 5795 Hobom,G. 5591 Matsui,Y. 5763 Callen,D.F. 5793 Hoeijmakers,J.H.J. 5633 Matsuura,S. 5792 Carpenter,S. 5796 Hofmann,S. 5491 McBumey,M.W. 5755 Carrera,P. 5639 Hogan,B.L.M. 5763 McElligott,D.L. 5796 Cech,T.R. 5515 Hoke,G.D. 5743 Mejean,V. 5525 Cera,C. 5707 Holman,K. 5793 Melis,R. 5561 Chan,S.-K. 5681 Hopwood,J.J. 5796 Mentink,M.M. 5800 Chazin,W.J. 5507 Horn,G.T. 5798 Merril,C.R. 5797 Chen,D. 5703 Hotaling,E. 5713 Miyamura,T. 5792 Chen,L.Z. 5793 Humphries,P. 5800 Moerman,D.G. 5669 Chen,X. 5703 Hunt,A.G. 5575 Mogen,B.D. 5575 Chen,Y. 5703 Hutchison III,C.A. 5731 Morris,C.P. 5796 Chollet,A. 5507 Iba,H. 5537 Motomiya,M. 5797 Christopherson,M.S. 5719 Ishizawa,M. 5792 Mulligan,L.M. 5795 Claverys,J.-P. 5525 Jardine,K. 5755 Nakamura,Y. 5804 Cohen,D. 5799 Jones,D.G.L. 5785 Napierski,I. 5725 Condron,B.G. 5607 Jordan,A.S. 5800 Nelson,P.V. 5796 Content,J. 5789 Jost,J.-P. 5771 Nishina,H. 5537 D'Incalci,M. 5739 Juliano,R.L. 5543,5551 Nowell,P.C. 5799 Davies,C.J. 5731 Junien,C. 5804 Ochmachi,T. 5797 de Mendonca Lima,L. 5789 Karran,P. 5569 Odijk,H. 5633 Deaven,L. 5804 Kase,R.S. 5790 Oetting,W.S. 5800 Degrave,W. 5789 Kennedy,J.L. 5801 Okuno,H. 5537 Devedjian,J.-C. 5525 Khan,P.M. 5805 Oliva,R. 5786 Dogliotti,E. 5569 Kidd,K.K. 5801 Owttrim,G.W. 5491 Donald,K.A.I.G. 5791 Kiff,J.E. 5669 Ozelius,L. 5798 Dowdy,S.F. 5763 Kikuchi,H. 5797 Paillard,S. 5619 Draper,K. 5743 King,M.W. 5777 Paker,N. 5795 Driver,V,B, 5743 King,R.A. 5800 Palumbo,M. 5707 Dubois,J. 5802 Kitayama,K. 5801 Pandey,N.B. 5653 Instructions to Authors

Please read these instructions carefully and follow them strictly to ensure that Computer papers will only be considered if they describe new applications or the review and publication of your paper is as efficient as possible. Please note novel algorithms relevant to nucleic acid sequence detenmination or analysis. Small that on submission all manuscripts must be accompanied by a manuscript submittal improvements in existing programs of variations of well-established algorithms form, a copy of which can be found at the end of these instructions. will generally not be suitable. Manuscripts dealing with computer analysis of already published sequences SCOPE OF THE JOURNAL are acceptable only if new and interesting findings emerge. Nucleic Acids research provides rapid publication for papers on physical, chemical, For The Record biochemical and biological aspects of nucleic acids, their constituents and proteins Several areas of research are suitable for presentation in this section. with which they interact. In addition, a section entitled Nucleotide Sequence Update (1) Sequence data: This is limited to the description of sequences of genes whose publishes recent database entries available on the EMBL File Server. products are relevant to nucleic acid metabolism and interactions. Structure RNAs and their genes are appropriate. The sequence must be complete and code for ORGANIZATION OF THE JOURNAL an identified product. Flanking regions should be included in the data deposited where available. The Papers are Journal will not print the actual sequence but it must have listed under: Surveys & Summaries, Molecular Biology, Enzymology, been submitted to the Data Library as indicated by an accession number. Papers Chemistry, Genome Structure and Mapping, For The Record and New Methods. must contain information about the sequence in addition to the annotation that Surveys & Summaries (-8-10 printed pages) has been submitted to the sequence database. Typically that information might This section is intended to provide a format for brief reviews. The section heading include a discussion of the broader significance of the sequence or features is deliberately chosen so that material not normally acceptable in a formal review contained within it. Such information might have been gained either by article can be presented. experimentation or by computer analysis. A copy of the sequence deposited with the Data Library should be available for the referees and in the case of DNA Full-length papers (8 printed pages) sequences should be accompanied by a letter indicating that both strands have This section includes papers placed in the Molecular Biology, Enzymology, been sequenced with appropriate overlapping sequence runs. Sequences of Chemistry, Genome Structure and Mapping and New Methods headings. pseudogenes are not acceptable. For The Record (1 printed page) Sequences not accompanied by a Data Library accession number will be returned to the authors. This section contains data not appropriate for publication as a full paper but which Protein and are of use to other (2) nucleic acid sequence motifs: New motifs that represent signals scientists. It is not intended for preliminary publication of in DNA or RNA will be considered. Modifications to previously described motifs data which will eventually be published elsewhere or for information that would must include a be better included as parts of full-length papers. demonstration of increased utility. Protein sequence motifs are Publication of papers in this section limited to those present in the proteins relevant to nucleic acid metabolism and takes place within 6 weeks. interactions. (3) Methods: This includes useful improvements on existing methods. However, CRITERIA FOR ACCEPTANCE authors are advised that, due to intense competition for publication in this section, Full-length papers only those manuscripts reporting developments of the highest originality and The Journal can only publish a fraction of the manuscripts it receives. For this usefulness will be published and trivial or obvious variations on established methods reason, the Editors insist that all manuscripts considered for publication must are unacceptable. In all cases the method described should be in sufficient detail present some novel development and meet the criteria of originality, timeliness, for it to be reproduced. Supplementary data may be supplied at submission to significance and scientific excellence. Technical progress has greatly increased indicate that the method has been fully tested. Descriptions of new vectors are the ease of obtaining new sequence information-the Editors will consider only not acceptable. those manuscripts in which new sequences are accompanied by complementary (4) NMR assignment data. data with relevance to genomic organization, transcription, RNA processing, ACCESSION NUMBERS & ATOMIC COORDINATES expression and genetic analysis; the reported sequence must shed significant new light on basic questions of structural or functional interest. Sequences of cDNAs Authors should be aware that all new sequence information, including that which or genes whose products are not relevant to nucleic acids are not acceptable. extends a previously determined sequence already present in the database (and Manuscripts dealing with comparative sequence analyses will be considered if which already has an accession number) must be submitted to the Data Library the genes under study are of relevance to nucleic acid metabolism. Sequences for a new accession number. This number must be provided when submitting of genes for well-studied RNAs (as well as the RNA sequences themselves) are the manuscript. For details see Volume 20 (1). not acceptable unless of unusual interest. Also, it is unnecessary and usually The Journal now requires that atomic coordinates for crystal structures should undesirable to print all the sequence. Provided that the sequence is available for be deposited with a database prior to manuscript submission. Appropriate databases the referees and has been deposited in the databank, only those parts relevant are as follows. Nucleosides, nucleotides and other small molecules: Cambridge to the discussion of the paper will be printed. Crystallographic Data Centre (CCDC). Proteins, polypeptides etc.: Protein Data Methods papers are considered only if an outstanding and potentially very useful Bank (PDB). oligonucleotides: CCDC or PDB. For details on deposition of data advance is clearly documented. Otherwise, the Editors prefer that such information see Volume 19 (13). be included in the 'Methods' section of a paper presenting new data of interest. Such new methods can be highlighted and listed separately in the Table of Contents JOURNAL POLICIES (see Vol. 19, issue no. 18). Except in extraordinary cases, manuscripts presenting It is our continued policy that all strains, clones, cell lines, hybridomas and construction of cloning vehicles or clone banks, preparation of monoclonal computer programs that are described in publication in the Journal should be made antibodies, identification of isoschizomer restriction enzymes or those describing immediately available to any qualified investigator upon request. Authors submitting routine syntheses of oligonucleotides are not published. manuscripts to Nucleic Acids Research must agree to adhere to this policy. The Genome mapping: for prokaryotic genomes only complete chromosome maps Editors are prepared to deny further publication rights in the Journal to authors will be considered. For higher organisms, unless extraordinary circumstances unwilling to abide by this principle. prevail, genome structure and mapping papers should introduce new technical Submission of a paper implies that it reports unpublished work and that it is advances or should describe a complete intron-exon structure for a gene relevant not under consideration elsewhere. The signature of the corresponding author to nucleic acid metabolism and interactions. is on behalf of all authors and assumes that they are in complete agreement with Chromosome assignment papers where only cytological data are prsented are the contents of the paper and are prepared to abide by the policies of the Journal. not acceptable. likewise, simple NMR assignment papers are not acceptable. Only If previously published tables, illustrations, or more than 200 words of text those that present new information not previously available from other methods are to be included, then the copyright holder's permission must be obtained, copies are considered. Manuscripts dealing with small molecule-nucleic acid interactions of any such permission letters should be enclosed with the paper. It is the are considered only if they present substantial new information relevant to nucleic responsibility ofthe authors to obtain authorzation for all personal communications. acid structure. Appropriate letters should accompany manuscripts. PREPARATION OF MANUSCRIPTS * Save the text as a word-processor file. If you have the facility to save the text as an ASCII file, make a copy ofthe file(s) in this format. This will speed Nucleic Acids Research is a fully typeset American Quarto size Journal. For full- up manual typesetting in the event that the word-processor file is not length papers, rapid publication will be maintained by using electronic typesetting from author-supplied floppy disk. However, three hard copies should still translatable. be * Label the disk with the first-named author and complete the manuscript supplied-one for the Editor and two for the referees-but they need not be camera- submission form. ready. For The Record papers will be typeset from a hard copy using a document reader. * When sending the disk, ensure that it is adequately protected to avoid damage by bending or X-ray inspection. Full-length papers For The Record submission Length: Papers should not usually exceed 8 printed pages. The typical number These papers will be typeset manually from the hard copy and proofs sent to of words on a text-only printed page is 1240. In calculating the final length of the corresponding author by fax. For The Record manuscripts are restricted in the manuscript please allow a 1/2-page for each figure or table. length to 1 printed page. This corresponds to approximately 1000 words without Sections of the manuscript: The abstract should be a single paragraph and not any allowance for figures or tables. Supplementary data should be supplied at exceeding 200 words. The references should be numbered in order of appearance submission where appropriate. and must be listed numerically. The citation ofjournals (abbreviated in the style Proofs of Chemical Abstracts), books and multi-author books and sequence data not published and only available from the databank entry should conform with the In the interest of speed, no proofs will be sent to authors submitting manuscripts following examples on disk. For papers that are manually typeset, proofs will be sent by fax and should be returned within 24 hours. 1. Holdsworth,M.J., Bird,C.R., Ray,J., Schuch,W. and Grierson,D. (1987) Nucleic Acids Res., 15, 7312-739. SUBMISSION TO THE EDITORS 2. Huynh,T.V., Young,R.A. and Davies,R.W. (1988) In Glover,D.M. (ed.), Three hard copies of the manuscript, including one original set of figures and DNA cloning-A Practical Approach. IRL Press, Oxford, Vol. I, pp. 49-78. two others of good definition suitable for the referees, accompanied by a disk 3. Maniatis,T., Fritsch,E.F. and Sambrook,J. (1982) Molecular Cloning: A if appropriate, and a completed manuscript submittal form (including telephone Laboratory Manual. Cold Spring Harbor University Press, Cold Spring and fax numbers and email address) should be submitted as follows: Harbor. 4. Muller,S.J. and Caradonna,S. (unpublished). X52486. From the Americas, Full-length papers and For The Record to: The Editor, Nucleic Acids Research, Oxford University Press, PO Box Q, McLean References of the type Smith et al. (1989) must not be used. VA 22101, USA. 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However, all sequence figures must be typed to the full width Birmingham B15 2TT, UK. of the page. Any lettering should be approximately 2 mm in height. if submitting Tel: (+4421 or 021) 476 1688 Fax: (+4421 or 021) 477 5376 figures that require reduction, please ensure that the lettering will be clearly legible For The Record papers to: after reduction to final size. Photographs should be provided as high-quality glossy The Editor, Nucleic Acids Research, Oxford University Press, Pinkhill House, prints in order to withstand the inevitable loss of contrast and detail inherent in Eynsham, Oxford OX8 1JJ, UK the printing process. Line drawings should be provided as good-quality hard copies Tel: (+44865 or 0865) 882283 Fax: (+44865 or 0865) 882890 suitable for reproduction as submitted. No additional artwork, redrawing or typesetting will be done by the Publishers. 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April 12-15, 1992 SHEFFIELD UNIVERSITY, UK

Following the highly succesful First Meeting in 1989, NACON-II will have Four Themes for Recognition Studies: Nucleic Acid:Nucleic Acid DNA Protein:DNA Protein Nucleic Acid:Drug RNA:Protein The meeting will be fully residential in Tapton Hall, Sheffield University and Registration will be restricted to 160 participants. Plenary Lectures will be given by: Richard Brimacombe Thomas Lindahl Simon Phillips Fritz Eckstein Steven Neidle Wojciech Stec Claude Helene Eiko Ohtsuka Larry Wakelin Angus Lamond Art Pardi Kurt Wuthrich David Lilley Dinshaw Patel

The Sixth Gobind Khorana Lecture will be delivered by Bob Wells Presentations of Short Lectures and Posters are invited. Please submit an Abstract by December 31, 1991 for consideration. A limited number of bursaries will be available for younger research workers.

Further details, Registration and Abstract forms are available from: Professor Mike Blackburn, Department of Chemistry, Sheffield University Sheffield, S3 7HF, UK Tel: +44 (0)742 768555 x4474, Fax: +44 (0)742 738673 Please mark all correspondence NACON-II REGISTRATION DEADLINE: JANUARY 31, 1992