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reused as long as the same liquid is being pipetted.

*U.S. Patent No. 4406170 i ~ r ~ L n ~ f F

B R ] N K M A N N Quality products for research and control.

Reader Service No. 781 BRK-5452B

GENES

DEVELOPMENT

VOLUME 6 NUMBER 7 PAGES 1131-1356 JULY 1992

EDITORIAL BOARD

J. Adams (Melbourne, Australia) M. Ashburner (Cambridge, UK) J. Beckwith (Boston, USA) T. Cech (Boulder, USA) P. Chambon (Strasbourg, France) N.-H. Chua (New York, USA) E. Coen (Norwich, UK) J. Coffin (Boston, USA) S. Courtneidge (Heidelberg, FRG) E. De Robertis (Los Angeles, USA) R. Evans (La Jolla, USA) G. Fink (Cambridge, USA) P. Goodfellow (London, UK) S. Gottesman (Bethesda, USA) T. Graf (Heidelberg, FRG) C. Gross (Madison, USA) F. Grosveld (London, UK) M. Groudine (Seattle, USA) L. Guarente (Cambridge, USA) C. Guthrie (San Francisco, USA) W. Herr (Cold Spring Harbor, USA) J. Hodgkin (Cambridge, UK) B. Hogan (Nashville, USA) R. Horvitz (Cambridge, USA)

P. Ingham (Oxford, UK) N. Jones (London, UK) R. Losick (Cambridge, USA) J. Manley (New York, USA) D. McClay (Durham, USA} W. McGinnis (New Haven, USA) S. McKnight (Baltimore, USA) A. McMahon (Nutley, USA) P. Nurse (Oxford, UK) C. Nfisslein-Volhard (Tubingen, FRG) R. Palmiter (Seattle, USA) G. Rubin (Berkeley, USA) U. Schibler (Geneva, Switzerland) M. Scott (Stanford, USA) D. Solter (Freiburg, FRG) J. Steitz (New Haven, USA) J. Strathern (Frederick, USA) T. Taniguchi (Osaka, Japan) S. Ti lghman (Princeton, USA) R. Tjian (Berkeley, USA) H. Varmus (San Francisco, USA) E. Wagner (Vienna, Austria) V. Walbot (Stanford, USA) M. Wigler (Cold Spring Harbor, USA)

Editors T. Grodzicker (Cold Spring Harbor) N. Hastie (Edinburgh)

Managing Editor J. Cuddihy (Cold Spring Harbor)

Editorial~Production N. Dumser, Technical Editor C. Kuret, Production Editor L. Hearfield, Editorial Secretary

EDITORIAL OFFICES

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CENES & D~WLOPMENT {ISSN 0890-9369 is published monthly for $360 {institutional), $105 {individual making personal payment), $138 (Gene- tical Society of Great Britain members) by Cold Spring Harbor Labora- tory Press, Bungtown Road, Cold Spring Harbor, New York 11724, in association with the Genetical Society of Great Britain. Second-class postage is paid at Cold Spring Harbor and additional mailing offices. POSTMASTER: Send address changes to Cold Spring Harbor Laboratory Press, POB 100, 1 Bungtown Rd., Cold Spring Harbor, New York 11724- 2203. Subscription Price Orders may be sent to Cold Spring Harbor Labora- tory Press, Fulfillment Department, 10 Skyline Drive, Plainview, NY 11803-9729. Telephone: Continental US except NY State, 1-800-843- 4388; all other locations, 516-394-1930/1931/1932. FAX 516-349-1946. Volume 6, 1992, $360, U.S. institutional; $435, R.O.W. institutional. Personal subscription rate: $105, U.S.; $180, R.O.W. Price includes sur- face postage for U.S. and airlift for R.O.W. Genetical Society members, $138.00. All subscriptions are entered for the calendar year and must be prepaid. Personal subscriptions must be prepaid by personal check, credit card, or money order. All checks must be for US dollars and drawn on a US bank. Genetical Society members may also subscribe by check, payable to the Genetical Society, for £ 8 0 {includes airlift). Send to: Dr.

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Claims for missing issues must be received within 4 months of issue date. Change of address Please enclose recent mailing label with address change; allow 4 weeks. Advertising To advertise in Genes & Development, contact Nancy Kuhle, Advertising Manager, Cold Spring Harbor Laboratory Press, Cold Spring Harbor, New York 11724-2203; telephone 516-367-8351. Photo Copy Authorization to photocopy items for internal or personal use, or the internal or personal use of specific clients, is granted by Cold Spring Harbor Laboratory Press for libraries and other users registered with the Copyright Clearance Center {CCC) Transactional Reporting Service, provided that the base fee of $3.00 per copy is paid directly to CCC, 21 Congress St., Salem, MA 01970 (0890-9369/92 $3.00 + 0.). This consent does not extend to other kinds of copying, such as copying for general distribution for advertising or promotional purposes, for creating new collective works, or for resale. Copyright © 1992 by Cold Spring Harbor Laboratory Press

Contents GENES & DEVELOPMENT July 1992

Research papers

Production of human papillomavirus and modulation of the infectious program in epithelial raft cultures Sheila C. Dollard, Jeffrey L. Wilson, Lisa M. Demeter, William Bonnez, Richard C. Reichman, Thomas R. Broker, and Louise T. Chow

1131

Wild-type p53 mediates positive regulation of gene expression through a specific DNA sequence element Gerard P. Zambetti, Jill Bargonetti, Kristen Walker, Carol Prives, and Arnold J. Levine

1143

The human heat shock protein hsp70 interacts with HSF, the transcription factor that regulates heat shock gene expression Klara Abravaya, Michael P. Myers, Shawn P. Murphy, and Richard I. Morimoto

1153

DnaK and DnaJ heat shock proteins participate in protein export in Escherichia coli Jadwiga Wild, Elliot Altman, Takashi Yura, and Carol A. Gross

1165

Isolation and characterization of RATI: an essential gene of Saccharomyces cerevisiae required for the efficient nucleocytoplasmic trafficking of mRNA David C. Amberg, Alan L. Goldstein, and Charles N. Cole

1173

Gene expression during preimplantation mouse development Jay L. Rothstein, Dabney Johnson, Julie A. DeLoia, Jacek Skowronski, Davor Solter, and Barbara Knowles

1190

Isolation of novel murine maternal mRNAs regulated by cytoplasmic polyadenylation Fernando J. Sall~s, Andrew L. Darrow, Marcia L. O'Connell, and Sidney Strickland

1202

Replication of distinct scrapie prion isolates is region specific in brains of transgenic mice and hamsters Rolf Hecker, Albert Taraboulos, Michael Scott, Keh-Ming Pan, Shu-Lian Yang, Marilyn Torchia, Klaus Jendroska, Stephen J. DeArmond, and Stanley B. Prusiner

1213

Translational activation of the non-AUG-initiated c-myc 1 protein at high cell densities due to methionine deprivation Stephen R. Hann, Karen Sloan-Brown, and Gerald D. Spotts

1229

Analysis of the functional role of the Polycomb chromo domain in Drosophila melanogaster Sabine Messmer, Axel Franke, and Renato Paro

1241

The Drosophila cellularization gene hullo produces a blastoderm-specific transcript whose levels respond to the nucleocytoplasmic ratio Lesilee Simpson Rose and Eric Wieschaus

1255

Trans and cis requirements for intron mobility in a prokaryotic system Jonathan Clyman and Marlene Belfort

1269

Signal transduction in Saccharomyces cerevisiae requires tyrosine and threonine phosphorylation of FUS3 and KSS1 Anton Gartner, Kim Nasmyth, and Gustav Ammerer

1280

Constitutive mutants of the protein kinase STE11 activate the yeast pheromone response pathway in the absence of the G protein Brian J. Stevenson, Nelson Rhodes, Beverly Errede, and George 1 =. Sprague Jr.

1293

Order of action of components in the yeast pheromone response pathway revealed with a dominant allele of the STEll kinase and the multiple phosphorylation of the STE7 kinase Bradley R. Cairns, Sandra W. Ramer, and Roger D. Kornberg

1305

SPT3 interacts with TFIID to allow normal transcription in Saccharomyces cerevisiae David M. Eisenmann, Karen M. Arndt, Stephanie L. Ricupero, John W. Rooney, and Fred Winston

1319

Topoisomerases and yeast rRNA transcription: negative supercoiling stimulates initiation and topoisomerase activity is required for elongation Michael C. Schultz, Steven J. Brill, Qida Ju, Rolf Sternglanz, and Ronald H. Reeder

1332

The RNA polymerase II ternary complex cleaves the nascent transcript in a 3' --, 5' direction in the presence of elongation factor SII Michael G. Izban and Donal S. Luse

1342

Cover Computer color enhancement of HPV virus particles in the nucleus of the mature, productive keratinocyte. (For details, Dollard et al., p. 1131.1