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JOURNAL OF VIROLOGYVOLUME 65 * JANUARY 1991 * NUMBER 1

Arnold J. Levine, Editor in Chief(1994)

Princeton UniversityPrinceton, N.J.Joan S. Brugge, Editor (1994)University ofPennsylvaniaPhiladelphia, Pa.Bernard N. Fields, Editor (1993)Harvard Medical SchoolBoston, Mass.

Stephen P. Goff, Editor (1994)Columbia UniversityNew York, N.Y.Peter M. Howley, Editor (1993)National Cancer InstituteBethesda, Md.Robert A. Lamb, Editor (1992)Northwestern UniversityEvanston, Ill.

Michael B. A. Oldstone, Editor (1993)Scripps Clinic & ResearchFoundation

La Jolla, Calif.Carol Prives, Editor (1991)Columbia UniversityNew York, N.Y.Thomas E. Shenk, Editor (1994)Princeton UniversityPrinceton, N.J.

EDITORIAL BOARDRafi Ahmed (1991)James Alwine (1991)Amiya K. Banerjee (1993)Kenneth I. Berns (1991)Joseph B. Bolen (1991)Thomas J. Braciale (1991)Dalius J. Briedis (1991)Thomas R. Broker (1992)Michael J. Buchmeier (1992)Robert Callahan (1991)Barrie J. Carter (1993)Mark Challberg (1992)Irvin Sy Chen (1993)Bruce Chesebro (1991)John M. Coffin (1992)Gary H. Cohen (1991)Charles N. Cole (1992)Peter L. Collins (1991)Richard A. Compans (1993)Richard C. Condit (1992)Neal Copeland (1992)Richard J. Courtney (1992)Clyde S. Crumpacker II (1991)Bryan R. Cullen (1993)Thomas Curran (1991)Samuel Dales (1991)Ronald C. Desrosiers (1993)Daniel DiMaio (1991)Walter Doerfler (1992)Elvera Ehrenfeld (1992)Robert N. Eisenman (1991)John Elder (1993)Suzanne U. Emerson (1992)Lynn W. Enquist (1991)Mary K. Estes (1992)Ellen Fanning (1991)Bernard Fleckenstein (1993)

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David M. Livingston (1991)Douglas R. Lowy (1992)Malcolm A. Martin (1992)Lois K. Miler (1991)Edward S. Mocarski (1993)Peter Model (1992)Bernard Moss (1992)Richard W. Moyer (1992)James Mullins (1993)Brian R. Murphy (1991)Nicholas A. Muzyczka (1993)Opendra Narayan (1991)Neal Nathanson (1993)Joseph R. Nevins (1991)Dennis O'Callaghan (1992)Peter Palese (1991)Enzo Paoletti (1991)Takis Papas (1993)J. Thomas Parsons (1992)George N. Pavlakis (1992)Ulf G. Pettersson (1992)James Pipas (1993)Lewis I. Pizer (1993)Nancy Raab-Traub (1993)Arnold B. Rabson (1993)Vincent Racaniello (1993)R. Frank Ramig (1991)Dan S. Ray (1992)Alan Rein (1993)Solon L. Rhode (1992)Harriet Robinson (1991)William S. Robinson (1992)Bernard Roizman (1991)John K. Rose (1991)Craig Rosen (1993)Naomi Rosenberg (1992)

Roland R. Rueckert (1991)Norman P. Salzman (1993)Charles E. Samuel (1992)Priscilla A. Schaffer (1993)Sondra Schlesinger (1992)Robert J. Schneider (1991)Manfred H. Schubert (1993)Bart Sefton (1991)Bert L. Semler (1992)Saul J. Silverstein (1991)Anna Marie Skalka (1993)Nahum Sonenberg (1993)Mark F. Stinski (1992)James Strauss (1993)Bill Sugden (1993)Jesse Summers (1992)Ron Swanstrom (1993)Peter J. Tattersal (1992)John M. Taylor (1993)Howard M. Temin (1991)Mary J. Tevethia (1993)David A. Thorley-Lawson (1991)Paula Traktman (1992)Inder Verma (1992)Luis V. Villarreal (1991)Peter K. Vogt (1993)Edward K. Wagner (1992)Robert G. Webster (1993)Gail Wertz (1991)Reed B. Wickner (1991)Eckard Wimmer (1991)Owen N. Witte (1992)Flossie Wong-Staal (1993)Moshe Yaniv (1992)Charles Hamish Young (1992)Harald zur Hausen (1992)

Barbara H. Iglewski, Chairman, Publications BoardJudith Nedrow, Production Editor

Linda M. Illig, Director, JournalsKenneth April, Assistant Production Editor

The Journal of Virology (ISSN 0022-538X), a publication of the American Society for Microbiology, 1325 Massachusetts Ave., N.W.,Washington, DC 20005-4171, is devoted to the dissemination of fundamental knowledge concerning viruses of bacteria, plants, and animals.Investigators are invited to submit reports of original research in all areas of basic virology, including biochemistry, biophysics, genetics,immunology, morphology, physiology, and pathogenesis and immunity. Instructions to authors are published in the January issue each year;reprints are available from the editors and the Journals Division. The Journal is issued monthly, one volume per year. The nonmembersubscription prices are $360 (U.S. and Canada) and $410 (foreign; air drop shipping) per year; single copies are $40. The member subscriptionprices are $49 (U.S. and Canada) and $98 (foreign; air drop shipping) per year; single copies are $10. Correspondence relating to subscriptions,reprints, defective copies, availability of back issues, lost or late proofs, disposition of submitted manuscripts, and general editorial mattersshould be directed to the ASM Journals Division, 1325 Massachusetts Ave., N.W., Washington, DC 20005-4171 (phone: 202 737-3600).Claims for missing issues from residents of the United States, Canada, and Mexico must be submitted within 3 months after publication ofthe issues; residents of all other countries must submit claims within 6 months of publication of the issues. Claims for issues missing becauseof failure to report an address change or for issues "missing from files" will not be allowed.Second-class postage paid at Washington, DC 20005, and at additional mailing offices.POSTMASTER: Send address changes to Journal of Virology, ASM, 1325 Massachusetts Ave., N.W., Washington, DC 20005-4171.Made in the U.S.A. Printed on acid-free paper.Copyright © 1991, American Society for Microbiology. El {tfl;A,J E MIf-, 1XI4tcAll Rights Reserved.The code at the top of the first page of an article in this journal indicates the copyright owner's consent that copies of the arti-cle may be made for personal use or for personal use of specific clients. This consent is given on the condition, however, that the copier paythe stated per-copy fee through the Copyright Clearance Center, 21 Congress St., Salem, MA 01970, for copying beyond that permitted bySections 107 and 108 of the U.S. Copyright Law. This consent does not extend to other kinds of copying, such as copying for generaldistribution, for advertising or promotional purposes, for creating new collective works, or for resale.

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This book will very effectively update interestedscientists and students on the current status ofresearch into ADP-ribosylating toxins and re-lated topics and will point the way for futureadvances.

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v .

Author IndexAdams, Sally E., 450Adlish, John D., 373Akgun, Ercan, 382Amanuma, Hiroshi, 132Anderson, Carl W., 326Anderson, Mark G., 51Antoni, Beth Ann, 396Aronoff, Rachel, 71Aurigemma, Rosemarie E., 23

Babiuk, L. A., 263Bachmayer, Helmut, 81Barbosa, Miguel S., 292Benjamin, Thomas L., 335,

342, 350Bennett, Robert P., 272Bergeron, Dominique, 7Bergeron, Louise, 281, 532Berrie, Eleanor L., 450Billaud, Marc, 415Bohnlein, Ernst, 81Bohnlein, Sabine, 81Bornkamm, Georg W., 415Bradley, Johnathan, 326Brian, David A., 320Britt, William J., 138

Calderone, Albert, 335Calender, Alain, 415Campos, M., 263Carter, Barrie J., 396Caton, Andrew J., 364Chejanovsky, Nor, 396Chen, Irvin S. Y., 546Cheng, Seng H., 170Chesebro, Bruce, 308Cheynier, Remi, 225Chittenden, Thomas, 514Clements, Janice E., 51Cordier, Martine, 415Courtney, Richard J., 520Craven, Rebecca C., 272Crittenden, L. B., 313Crook, Tim, 505

Daar, Eric S., 489Dawe, Clyde J., 335, 342de Goede, Ruud E. Y., 356Delassus, Sylvie, 225DeLuca, Neal A., 299Demers, G. William, 473de Wolf, Frank, 356Dillon, Patrick J., 445Doerfier, Walter, 89Dorfman, Tatyana, 281Dormont, D., 541Drlica, Karl, 551Dubensky, Thomas W., 342

Earl, Patricia L., 31Eick, Dirk, 415Ekstijn, G. L., 98Espino, Pearl C., 170Esteban, Mariano, 494, 499

Fay, Jaydie, 138Federspiel, M. J., 313Fields, Bernard N., 123Finberg, Robert W., 220Fincham, Valerie J., 461Fisher, Chris, 505

Fitzpatrick, D. R., 263Flamand, Anne, 123Freed, Eric O., 190Frese, Knut, 457Freund, Robert, 335, 342, 350Fujimura, Tsutomu, 155

Gagner, Jean-Pierre, 123Galloway, Denise A., 473Garcea, Robert L., 350Garcfa, J. A., 1Garoff, Henrik, 147Gerhard, Walter, 364Gong, Shiaoching, 499Graham, Kathryn, 61Gras, G. S., 541Green, Patrick L., 546Greene, Warner C., 405Grez, Manuel, 382Griffiths, Joanne C., 450Grundstrom, Thomas, 42Guan, Nin, 483

Haberman, Ann M., 364Hakura, Akira, 479Halbert, Christine L., 473Hallberg, Bengt, 42Hanly, Sarah M., 405Happ, Brigitte, 89Harber, James J., 326Harland, R., 263Harmsen, M., 98Harris, Stephen J., 450Harvey, Robert, 170Haseltine, William, 281, 532Hauber, Joachim, 81Helenius, Ari, 232Hemming, Val G., 103Ho, David D., 489Hofer, Lotte, 81Hofmann, Martin A., 320Holdsworth, Linda N., 450Hom, Richard C., 220Hong, Tao, 551Hughes, S. H., 313Hunt, Arthur G., 511Hunter, Eric, 272

Icho, Tateo, 155Ikawa, Yoji, 132Inoue, Hirokazu, 479Ishimoto, Akinori, 526

Jackson, Alan C., 537Jasin, Maria, 440Jhagjhoor-Singh, S. S., 98Jong, Song-Muh J., 180

Kabat, David, 464Kaufman, Peter A., 405Kawaoka, Yoshihiro, 389Kim, Jerome H., 405Kingsman, Alan J., 450Kingsman, Susan M., 450Kniess, Norbert, 138Koenig, Scott, 31Kootstra, Neeltje A., 356Kowalski, Mark, 281Kozak, Christine A., 7, 464Kozak, Susan L., 464

Kraaijeveld, C. A., 98Krikorian, Charles R., 112Kulkosky, Joseph, 213

Lahijani, Roya S., 373Lai, Chingfeng, 499Lain, S., 1Landau, Nathaniel R., 162Laurence, Jeffrey, 213Laux, Gerhard, 415Lenoir, Gilbert M., 415Lever, Andrew, 532Levine, Arnold J., 514Li, Jian, 89Li, Xi Ling, 489LiUjestrom, Peter, 147Linial, Maxine, 71Littman, Dan R., 162, 440Liu, Fenyong, 206Lowy, Douglas R., 292

Macaulay, Colin, 61Mach, Michael, 138Marshall, John, 170Martin, Kelsey, 232Martin, M. T., 1McFadden, Grant, 61McKeating, Jane A., 489Merrill, Janet, 170Miedema, Frank, 356Miller, Irving L., 396Miller, Paul, 213Miller, Ruth, 16Miyasaka, Michio, 479Moore, John P., 450Morrison, Lynda A., 123Moss, Bernard, 31Moudgil, Tarsem, 489Mukaigawa, Jun, 245Mullaney, Steven, 220Murphy, Ellen, 551Murphy, John F., 511Myers, Dawn J., 190

Nayak, Debi P., 245Nelbock, Peter, 445Nishi, Masahiro, 132North, Thomas W., 308

Oh, Sang-Jin, 514Ojakian, George K., 494Oosterlaken, T. A. M., 98Ornelles, David A., 424Otteson, Elmer W., 373

Page, Kathleen A., 162, 440Palladino, Giuseppe, 364Paque, Ronald E., 16Parker, M. D., 263Paul, Ralph, 464Pawlita, Michael, 468Pedersen, Niels C., 308Perkins, Ann, 445Pfleiderer, Michael, 254Pinter, Abraham, 551Pirker, F. Peter, 81Poliquin, Laurent, 7Porter, David D., 103Porter, Helen G., 103Poznansky, Mark, 532

Prince, Gregory A., 103Provencher, L. P., 313

Rabson, Arnold B., 396Rassart, Eric, 7Read, G. Sullivan, 112Remington, Kathryn Martin,

308Rhee, Sung, 272Rhoads, Robert E., 511Riechmann, J. L., 1Rimsky, Laurence T., 405Risser, Rex, 190Robinson, James E., 489Rodriguez, Dolores, 494Rodriguez, Juan-Ramon, 494Roizman, Bernard, 206Rosen, Craig A., 445Routledge, Edward, 254Ruprecht, Ruth M., 220Ryan-Poirier, Kathleen A.,

389Rychlik, Wojciech, 511

Sahli, Roland, 350Scherle, Peggy A., 364Schilken, Dorothea, 457Schiller, John T., 292Schreiber, Martha, 61Schuetze, Scott, 464Schuitemaker, Hanneke, 356Schwyzer, Martin, 195Sczakiel, Georg, 468Senior, John M., 450Sethna, Phiroze B., 320Shaw, John G., 511Shenk, Thomas, 424Shepard, Alyssa A., 299Siddell, Stuart G., 254Sikder, Santosh K., 213Smith, Alan E., 170Smith, Ralph E., 23Snippe, H., 98Sodroski, Joseph, 281, 532Sommer, Marvin, 520St. Jeor, Stephen C., 373Stauber, Roland, 254Stitz, Lothar, 457Stuart, David, 61Suentzenich, Karl-Otto, 415Sun, Nai-Chau, 489

Takami, Yasunari, 479Tanaka, Harutaka, 526Taylor, Alexander H., 364Tersmette, Matthijs, 356Thornell, Anders, 42Torgersen, Jofrid L., 23Trempe, James P., 396Ts'o, Paul 0. P., 213

van den Hurk, J. V., 263van Drunen Littel-van den

Hurk, S., 263Vass, William C., 292Vogt, Bernd, 195Vousden, Karen H., 505

Wain-Hobson, Simon, 225Wang, Lu-Hai, 180

i

ii AUTHOR INDEX

Watanabe, Naoko, 132Wehrly, Kathy, 308Wickner, Reed B., 155Widner, William R., 155

Wills, John W., 272Wimmer, Eckard, 326Wirth, Urs V., 195Wyke, John A., 461

Xie, Yiming, 546

Yanagawa, Shin-Ichi, 526Yates, John L., 483

Zamb, T. J., 263Ziegler, Marion, 382Zimber-Strobl, Ursula, 415Zimmermann, Klaus, 81

J. VIROL.

JOURNAL OF VIROLOGY, Jan. 1991

JOURNAL OF VIROLOGY

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Page ProofsThe printer sends page proofs, the copy-edited

manuscript, and a page charge/reprint order form tothe author. As soon as the page proofs are corrected(within 48 h), they should be mailed to the ASMJournals Division.The proof stage is not the time to make extensive

corrections, additions, or deletions. Important newinformation that has become available between accep-tance of the manuscript and receipt of the proofs maybe inserted as an Addendum in Proof with the permis-sion of the editor. If references to unpublished data orpersonal communications are added, include writtenassurance that permission to cite them has beengranted. Limit changes to correction of spelling errors,incorrect data, grammatical errors, and updated infor-mation for "submitted" and "in press" references.

Questions about late proofs and problems in theproofs should be directed to the ASM Journals Divi-sion, telephone (202) 737-3600.Reprints

Reprints (in multiples of 100) may be purchased bycontributors. An order form that includes a tableshowing the cost of reprints is sent with each proof.ORGANIZATION AND FORMAT

Regular PapersRegular full-length papers should include the ele-

ments described in this section.

Title. Each manuscript should present the results ofan independent, cohesive study; thus, numbered se-ries titles are not allowed. Avoid the main title/subtitlearrangement, complete sentences, and unnecessaryarticles. On the title page, include the title, runningtitle (not to exceed 54 characters and spaces), name ofeach author, address(es) of the institution(s) at whichthe work was performed, each author's affiliation, anda footnote indicating the present address of any authorno longer at the institution where the work was per-formed. Place an asterisk after the name of the authorto whom inquiries regarding the paper should bedirected, and give that author's telephone number.

Abstract. Limit the abstract to 250 words or fewerand concisely summarize the basic content of thepaper without presenting extensive experimental de-tails. Avoid abbreviations and do not include dia-grams. When it is essential to include a reference, usethe References citation but omit the article title. Be-cause the abstract will be published separately byabstracting services, it must be complete and under-standable without reference to the text.

. .


Introduction. The introduction should supply suffi-cient background information to allow the reader tounderstand and evaluate the results of the presentstudy without referring to previous publications on thetopic. The introduction should also provide a rationalefor the present study. Choose references carefully toprovide the most salient background rather than anexhaustive review of the topic.

Materials and Methods. The Materials and Methodssection should include sufficient technical informationto allow the experiments to be repeated. When cen-trifugation conditions are critical, give enough infor-mation to enable another investigator to repeat theprocedure: make of centrifuge, model of rotor, tem-perature, time at maximum speed, and centrifugalforce (x g rather than revolutions per minute). Forcommonly used materials and methods (e.g., mediaand protein determinations), a simple reference issufficient. If several alternative methods are com-monly used, it is helpful to identify the method brieflyas well as to cite the reference. For example, it ispreferable to state "cells were broken by ultrasonictreatment as previously described (9)" rather than tostate "cells were broken as previously described (9)."The reader should be allowed to assess the methodwithout constant reference to previous publications.Describe new methods completely and give sources ofunusual chemicals, equipment, or microbial strains.When large numbers of microbial strains or mutantsare used in a study, include tables identifying thesources and properties of the strains, mutants, bacte-riophages, plasmids, etc.A method, strain, etc., used in only one of several

experiments reported in the paper may be described inthe Results section or very briefly (one or two sen-tences) in a table footnote or figure legend.

Results. In the Results section, include the rationaleor design of the experiments as well as the results;reserve extensive interpretation of the results for theDiscussion section. Present the results as concisely aspossible in one of the following: text, table(s), orfigure(s). Data in tables (e.g., cpm of radioactivity)should not contain more significant figures than theprecision of the measurement allows. Illustrations(particularly photomicrographs and electron micro-graphs) should be limited to those that are absolutelynecessary to show the experimental findings. Numberfigures and tables in the order in which they are citedin the text, and be sure to cite all figures and tables.

Discussion. The Discussion should provide an inter-pretation of the results in relation to previously pub-lished work and to the experimental system at handand should not contain extensive repetition of theResults section or reiteration of the introduction. Inshort papers, the Results and Discussion sections maybe combined.

Acknowledgments. Acknowledgments of financialassistance and of personal assistance should be givenin separate paragraphs. The usual format for acknowl-edgment of grant support is as follows: "This workwas supported by Public Health Service grantCA-01234 from the National Cancer Institute."

Appendixes. Appendixes, which contain supplemen-tary material to aid the reader, are permitted. Titles,authors, and References sections that are distinct fromthose of the primary article are not allowed. If it is notfeasible to list the author(s) of the appendix in thebyline or the Acknowledgment section of the primaryarticle, rewrite the appendix so that it can be consid-ered for publication as an independent article, eitherfull-length or Note style. Equations, tables, and figuresshould be labeled with the letter "A" preceding thenumeral to distinguish them from those cited in themain body of the text.

References. The References section must include allrelevant sources, and all listed references must becited in the text. Arrange the citations in alphabeticalorder by first author and number consecutively. Abbre-viate journal names according to Serial Sourcesfor theBIOSIS Data Base (BioSciences Information Service,1990). Cite each listed reference by number in the text.Follow the styles shown in the examples below.

1. Arens, M., and T. Yamashita. 1978. In vitro terminationof adenovirus DNA synthesis by a soluble replicationcomplex. J. Virol. 25:698-702.

2. Berry, L. J., R. N. Moore, K. J. Goodrum, and R. E.Crouch, Jr. 1977. Cellular requirements for enzymeinhibition by endotoxin in mice, p. 321-325. In D.Schlessinger (ed.), Microbiology-1977. American Soci-ety for Microbiology, Washington, D.C.

3. Cox, C. S., B. R. Brown, and J. C. Smith. J. Gen.Genet., in press.*

4. Dhople, A., I. Ortega, and C. Berauer. 1989. Effect ofoxygen on in vitro growth of Mycobacterium leprae,abstr. U-82, p. 168. Abstr. 89th Annu. Meet. Am. Soc.Microbiol. 1989.

5. Fitzgerald, G., and D. Shaw. In A. E. Waters (ed.),Clinical microbiology, in press. EFH Publishing Co.,Boston.

6. Gill, T. J., III. 1976. Principles of radioimmunoassay, p.169-171. In N. R. Rose and H. Friedman (ed.), Manualof clinical immunology. American Society for Microbi-ology, Washington, D.C.

7. Gustlethwaite, F. P. 1985. Letter. Lancet ii:327.8. Jacoby, J., R. Grimm, J. Bostic, V. Dean, and G. Starke.

Submitted for publication.9. Jensen, C., and D. S. Schumacher. Unpublished data.

10. Jones, A. (Yale University). 1990. Personal communica-tion.

11. Leadbetter, E. R. 1974. Order II. Cytophagales nomennovum, p. 99. In R. E. Buchanan and N. E. Gibbons(ed.), Bergey's manual of determinative bacteriology,8th ed. The Williams & Wilkins Co., Baltimore.

12. Miller, J. H. 1972. Experiments in molecular genetics, p.352-355. Cold Spring Harbor Laboratory, Cold SpringHarbor, N.Y.

IV


13. Powers, R. D., W. M. Dotson, Jr., and F. G. Hayden.1982. Program Abstr. 22nd Intersci. Conf. Antimicrob.Agents Chemother., abstr. 448.

14. Sigma Chemical Co. 1989. Sigma manual. Sigma Chem-ical Co., St. Louis, Mo.

15. Smith, J. C. April 1970. U.S. patent 484,363,770.16. Smyth, D. R. 1972. Ph.D. thesis. University of Califor-

nia, Los Angeles.17. Yagupsky, P., and M. A. Menegus. 1989. Intraluminal

colonization as a source of catheter-related infection.Antimicrob. Agents Chemother. 33:2025. (Letter.)

* Note that an "in press" reference to an ASMpublication should state the control number (e.g., JVI576-91) or the name of the publication if it is a book.Notes

Submit Notes in the same way as full-length papers.They receive the same review, they are not publishedmore rapidly than full-length papers, and they are notconsidered preliminary communications. The Noteformat is intended for the presentation of brief obser-vations that do not warrant full-length papers.Each Note must have an abstract of no more than 50

words. Do not use section headings in the body of theNote; report methods, results, and discussion in asingle section. Paragraph lead-ins are permissible. Thetext should be kept to a minimum and if possibleshould not exceed 1,000 words; the number of figuresand tables should also be kept to a minimum. Materialsand methods should be described in the text, not infigure legends or table footnotes. Present acknowledg-ments as in full-length papers, but do not use aheading. The References section is identical to that offull-length papers.Minireviews

Minireviews are brief summaries (limit of 4 printedpages) of developments in fast-moving areas. Theymust be based on published articles; they may addressany subject within the scope of JVI. Minireviews maybe either solicited or proffered by authors respondingto a recognized need. Irrespective of origin, minire-views are subject to editorial review.Letters to the Editor

Letters to the Editor must include data to supportthe writer's argument and are intended only for com-ments on articles published previously in the journal.They may be no more than 500 words long. Send lettersto the Journals Division. They will be processed andsent to the editor who handled the article in question.If the editor believes that publication is warranted, hewill solicit a reply from the author of the article andmake a recommendation to the editor in chief. Finalapproval for publication rests with the editor in chief.All letters intended for publication must be typeddouble spaced.ErrataThe Erratum section provides a means of correcting

errors (e.g., typographical) in published articles.

Changes in data and the addition of new material arenot permitted. Send errata directly to the JournalsDivision.

Author's CorrectionsThe Author's Correction section provides a means

of adding citations that were overlooked in a publishedarticle. The author who failed to cite a reference andthe author whose paper was not cited must agree tosuch a publication; the editor, editor in chief, andchairman of the Publications Board will not be in-volved. Letters from both authors must accompanythe author's correction sent to the Journals Division.

DisclaimersStatements disclaiming governmental or any other

type of endorsement or approval will be deleted by theJournals Division.

ILLUSTRATIONS AND TABLESThe figure number and authors' names should be

written on all figures, either in the margin or on theback (marked lightly with a soft pencil). For micro-graphs especially, the top should be indicated as well.Do not clasp figures to each other or to the manu-

script with paper clips. Insert small figures in anenvelope.

Continuous-Tone and Composite PhotographsWhen submitting continuous-tone photographs

(e.g., polyacrylamide gels), keep in mind the journalpage width: 5'/16 inches for a single column and 67/8inches for a double column (maximum). Include onlythe significant portion of an illustration. Photos mustbe of sufficient contrast to withstand the inevitable lossof contrast and detail inherent in the printing process.Submit one photograph of each continuous-tone figurefor each copy of the manuscript; photocopies are notacceptable. If possible, the figures submitted should bethe size they will appear when published so that noreduction is needed. If they must be reduced, makesure that all elements, including labeling, can with-stand reduction and remain legible.

If a figure is a composite of a continuous-tonephotograph and a drawing or labeling, the originalcomposite must be provided for the printer (i.e., not aphotograph of the composite). This original, labeled"6printer's copy," may be sent with the modifiedmanuscript to the editor.

Electron and light micrographs must be direct cop-ies of the original negative. Indicate the magnificationwith a scale marker on each micrograph.

Color PhotographsColor photographs are discouraged. However, if

they are necessary, include an extra copy at the timeof manuscript submission so that a cost estimate forprinting may be obtained. The cost of printing colorphotographs must be borne by the author.

v


DrawingsSubmit graphs, charts, sequences, complicated

chemical or mathematical formulas, diagrams, andother drawings as glossy photographs made fromfinished drawings not requiring additional artwork or

typesetting. Computer-generated graphics producedon high-quality laser printers are also usually accept-able. No part of the graph or drawing should behandwritten. Both axes of graphs must be labeled.Most graphs will be reduced to one-column width (35/16inches, and all elements in the drawing should be largeenough to withstand this reduction. Avoid heavy let-ters, which tend to close up when reduced, andunusual symbols, which the printer may not be able toreproduce in the legend.

In figure ordinate and abscissa scales (as well as

table column headings), avoid ambiguous use of num-bers with exponents. Usually, it is preferable to use

the International System of Units (,u for 10-6, m forl0o-, k for 103, M for 106, etc.). A complete listing ofSI symbols can be found in the International Union ofPure and Applied Chemistry (IUPAC) "Manual ofSymbols and Terminology for Physicochemical Quan-tities and Units" (Pure Appl. Chem. 21:3-44, 1970).Thus, a representation of 20,000 cpm on a figureordinate is to be made by the number 20, accompaniedby the label kcpm.When powers of 10 must be used, the Journal

requires that the exponent power be associated withthe number shown. In representing 20,000 cells per ml,the numeral on the ordinate would be "2" and thelabel would be "i04 cells per ml" (not "cells per ml x

10-4'"). Likewise, an enzyme activity of 0.06 U/mlwould be shown as 6, accompanied by the label 10-2U/ml. This is equivalent to the label "hundredths ofunits per ml" or the preferable "cU/ml" for the SIconvention. The preferred designation would be 60mU/ml (milliunits per ml).

Presentation of Nucleic Acid SequencesNucleic acid sequences of limited length which are

the primary subject of a study may be presentedfreestyle in the most effective format. Longer nucleicacid sequences must be presented in the followingformat to conserve space. Submit the sequence as

camera-ready copy of dimensions 8Y2" x 11" (orslightly less) in standard (portrait) orientation. Printthe sequence in lines of 100 bases, each in a nonpro-

portional (monospace) font which is easily legiblewhen published at 100 bases/6 inches. Uppercase andlowercase letters may be used to designate the exon/intron structure, transcribed regions, etc., if the low-ercase letters remain legible at 100 bases/6 inches.Number the sequence line by line; place numerals,representing the first base of each line, to the left of thelines. Minimize spacing between adjacent lines ofsequence, leaving room only for annotation of thesequence. Annotation may include boldface, underlin-ing, brackets, boxes, etc. Encoded amino acid se-

quences may be presented, if necessary, immediately

above the first nucleotide of each codon, using thesingle-letter amino acid symbols. Comparisons of mul-tiple nucleic acid sequences should conform as nearlyas possible to the same format.

Figure LegendsLegends should provide enough information so that

the figure is understandable without frequent referenceto the text. However, detailed experimental methodsmust be described in the Materials and Methodssection, not in a figure legend. A method that is uniqueto one of several experiments may be reported in alegend only if the discussion is very brief (one ortwo sentences). Define all symbols and abbreviationsused in the figure that have not been defined else-where.

TablesType each table on a separate page. Arrange the

data so that columns of like material read down, notacross. The headings should be sufficiently clear sothat the meaning of the data will be understandablewithout reference to the text. See the Abbreviationssection of these instructions for those that should beused in tables. Explanatory footnotes are acceptable,but more extensive table "legends" are not. Footnotesshould not include detailed descriptions of the exper-iment. Tables must include enough information towarrant table format; those with fewer than six piecesof data will be incorporated into the text by the copyeditor. A well-constructed table is shown below:

TABLE 1. Distribution of protein and ATPase in fractions ofdialyzed membranesa

ATPaseMembranes Fraction

from: U/mg of Total Uprotein

Control Depleted membrane 0.036 2.3Concentrated supernatant 0.134 4.82

El treated Depleted membrane 0.034 1.98Concentrated supernatant 0.11 4.6

a Specific activities of ATPase of nondepleted membranes from control andtreated bacteria were 0.21 and 0.20, respectively.

Tables that can be photographically reproduced forpublication without further typesetting or artwork arereferred to as "camera ready." They should not behand lettered and must be carefully prepared to con-form with the style of the journal. The advantage ofsubmitting camera-ready copy is that the material willappear exactly as envisioned by the author, and nosecond proofreading is necessary. This is particularlyadvantageous when there are long, complicated tablesand when the division of material and spacing areimportant.

Vi


NOMENCLATURE

Chemical and Biochemical NomenclatureThe recognized authority for the names of chemical

compounds is Chemical Abstracts (Chemical Ab-stracts Service, Ohio State University, Columbus) andits indexes. The Merck Index (llth ed., 1989; Merck &Co., Inc., Rahway, N.J.) is also an excellent source.For biochemical terminology, including abbreviationsand symbols, consult the following: Biochemical No-menclature and Related Documents (1978; reprintedfor The Biochemical Society, London), Instructions toAuthors of the Journal ofiBiological Chemistry and theArchives of Biochemistry and Biophysics (first issuesof each year), and the Handbook ofBiochemistry andMolecular Biology (G. D. Fasman, ed., CRC Press,Inc., 3rd ed., 1976).Do not express molecular weight in daltons; molec-

ular weight is a unitless ratio. Molecular mass isexpressed in daltons.For enzymes, use the recommended (trivial) name

assigned by the Nomenclature Committee of the In-ternational Union of Biochemistry as described inEnzyme Nomenclature (Academic Press, Inc., 1984).If a nonrecommended name is used, place the proper(trivial) name in parentheses at first use in the abstractand text. Use the EC number when one has beenassigned, and express enzyme activity either in katals(preferred) or in the older system of micromoles perminute.

Nomenclature of VirusesNames used for viruses should be those approved

by the International Committee on Taxonomy of Vi-ruses (ICTV) and published in the 4th Report of theICTV: Classification and Nomenclature of Viruses(Intervirology 17:23-199, 1982). If desired, synonymsmay be added parenthetically when the name is firstmentioned. Approved generic (or group) and familynames may also be used.

Nomenclature of BacteriaBinary names, consisting of a generic name and a

specific epithet (e.g., Escherichia coli), should be usedfor all bacteria. Names of higher categories may beused alone, but specific and subspecific epithets maynot. A specific epithet must be preceded by a genericname the first time it is used in a paper. Thereafter, thegeneric name should be abbreviated to the initialcapital letter (e.g., E. coli), provided there can be noconfusion with other genera used in the paper. Namesof all taxa (phyla, classes, orders, families, genera,species, subspecies) are printed in italics; strain des-ignations and numbers are not.

Genetic NomenclatureWhen appropriate for viral genetic systems, use the

recommendations of Demerec et al. (Genetics 54:61-76, 1966) as a guide.

(i) Phenotype designations must be employed whenmutant loci have not been identified or mapped. Phe-notype designations generally consist of three-lettersymbols; these are not italicized and the first letter ofthe symbol is capitalized. It is preferable to use romanor arabic numerals (instead of letters) to identify aseries of related phenotypes. Thus, a series of bacte-riocin-tolerant mutants might be designated Toll,Tolll, TollIl, etc., or a series of nucleic acid polymer-ase mutants might be designated Poll, Pol2, Pol3, etc.Wild-type characteristics can be designated as Tol+ orPol+ and, when necessary for clarity, negative super-scripts (Tol- Pol-) can be used to designate mutantcharacteristics. Superscript letters may be used tofurther delineate phenotypes (e.g., Strs for strep-tomycin sensitivity). Phenotype designations shouldbe defined.

(ii) Genotype designations are similarly indicated bythree-letter locus symbols. These are lowercase italic(e.g., pol src). If several loci govern related functions,these are distinguished by italicized capital lettersfollowing the locus symbol.

(iii) Wild-type alleles are indicated with a super-script plus (ara+ his'). A superscript minus is not usedto indicate a mutant locus; thus one refers to an aramutant rather than an ara- strain.

(iv) The rules for genetic nomenclature of viruses(phages) differ from those of bacteria. As a generalrule, the entire description of a virus is italicized,including the designations am or sus (amber) and ts(temperature sensitive). Superscripts are employed toindicate hybrid genomes. Genetic symbols may beone, two, or three letters. For example, a mutantstrain of A might be designated as XcI857int2redll4susAll; this strain carries mutations in genes cI, int,and red and a suppressible (sus) mutation in gene A. Astrain designated Ximm21att434 would represent a hy-brid of phage A which carries the immunity region(imm) of phage 21 and the attachment (att) region ofphage 434. Host DNA insertions into viruses should bedelineated by square brackets, and the genetic sym-bols and designations for such inserted DNA shouldconform to those employed for the host genome.Genetic symbols for phage X can be found in Echolsand Murialdo (Microbiol. Rev. 42:577-591, 1978) andSzybalski and Szybalski (Gene 7:217-270, 1979).

(v) Nomenclature of restriction endonucleasesshould follow the recommendations of Roberts (p.757-768, in Bukhari et al., ed., DNA Insertion Ele-ments, Plasmids, and Episomes, Cold Spring HarborLaboratory, 1977).

"Mutant" vs. "mutation." Authors are reminded ofthe distinction between a mutation (an alteration of theprimary sequence of the genetic material) and a mu-tant (a strain carrying one or more mutations). Onemay speak about the mapping of a mutation, but onecannot map a mutant. Likewise, a mutant has nogenetic locus, only a phenotype.

.ii


ABBREVIATIONS AND CONVENTIONS

Patient IdentificationWhen isolates are derived from patients in clinical

studies, do not identify them by using the patients'initials, even as part of a strain designation. Changethe initials to numerals or use randomly chosen letters.Do not give hospital unit numbers; if a designation isneeded, use only the last two digits of the unit. (Note:Established designations of some viruses and celllines, although they consist of initials, are acceptable[e.g., JC virus, BK virus, HeLa cells].)

Verb TenseASM strongly recommends that for clarity you use

the past tense to narrate particular events in the past,including the procedures, observations, and data ofthe study that you are reporting. Use the present tensefor your own general conclusions, the conclusions ofprevious researchers, and generally accepted facts.Thus, most of the abstract, Materials and Methods,and Results sections will be in the past tense, and mostof the introduction and some of the Discussion will bein the present tense.Be aware that it may be necessary to vary the tense

in a single sentence. For example, it is correct to say"White (30) demonstrated that XYZ cells grow at pH6.8," "Figure 2 shows that ABC cells failed to grow atroom temperature," and "Air was removed from thechamber and the mice died, which proves that micerequire air." In reporting statistics and calculations, itis correct to say "The values for the ABC cells arestatistically significant, indicating that the drug inhib-ited...."For an in-depth discussion of tense in scientific

writing, see p. 158-160 in How to Write and Publish aScientific Paper, 3rd ed.

AbbreviationsGeneral. Abbreviations should be used as an aid to

the reader, rather than as a convenience to the author,and therefore their use should be limited. Abbrevia-tions other than those recommended by the IUPAC-IUB (Biochemical Nomenclature and Related Docu-ments, 1978) should be used only when a case can bemade for necessity, such as in tables and figures.

It is often possible to use pronouns or to paraphrasea long word after its first use (e.g., "the drug," "thesubstrate"). Standard chemical symbols and trivialnames or their symbols (folate, Ala, Leu, etc.) may beused for terms that appear in full in the neighboringtext.

It is strongly recommended that all abbreviationsexcept those listed below be introduced in the firstparagraph in Materials and Methods. Alternatively,define each abbreviation and introduce it in parenthe-ses the first time it is used; e.g., "cultures were grownin Eagle minimal essential medium (MEM)." Gener-ally, eliminate abbreviations that are not used at least

five times in the text (including tables and figurelegends).Not requiring introduction. In addition to abbrevia-

tions for Systeme International d'Unites (SI) units ofmeasurement, other common units (e.g., bp, kb, andDa), and chemical symbols for the elements, thefollowing should be used without definition in the title,abstract, text, figure legends, and tables: DNA (deoxy-ribonucleic acid); cDNA (complementary DNA);RNA (ribonucleic acid); cRNA (complementaryRNA); RNase (ribonuclease); DNase (deoxyribonu-clease); rRNA (ribosomal RNA); mRNA (messengerRNA); tRNA (transfer RNA); AMP, ADP, ATP,dAMP, ddATP, GTP, etc. (for the respective 5' phos-phates of adenosine and other nucleosides) (add2'-, 3'-, or 5'- when needed for contrast); ATPase,dGTPase, etc. (adenosine triphosphatase, deoxygua-nosine triphosphate, etc.); NAD (nicotinamide ade-nine dinucleotide); NAD+ (nicotinamide adenine dinu-cleotide, oxidized); NADH (nicotinamide adeninedinucleotide, reduced); NADP (nicotinamide adeninedinucleotide phosphate); NADPH (nicotinamide ade-nine dinucleotide phosphate, reduced); NADP+ (nic-otinamide adenine dinucleotide phosphate, oxidized);poly(A), poly(dT), etc. (polyadenylic acid, polydeoxy-thymidylic acid, etc.); oligo(dT), etc. (oligodeoxythy-midylic acid, etc.); Pi (orthophosphate); PP1 (pyro-phosphate); UV (ultraviolet); PFU (plaque-formingunits); CFU (colony-forming units); MIC (minimalinhibitory concentration); MBC (minimal bactericidalconcentration); Tris [tris(hydroxymethyl)amino-methane]; DEAE (diethylaminoethyl); A260 (absorb-ance at 260 nm); EDTA (ethylenediaminetetraaceticacid); and AIDS (acquired immunodeficiency [or im-mune deficiency] syndrome). Abbreviations for celllines (e.g., HeLa) also need not be defined.The following abbreviations should be used without

definition in tables:

amt (amount)approx (approximately)avg (average)concn (concentration)diam (diameter)expt (experiment)exptl (experimental)

ht (height)sp gr (specific gravity)tr (trace)vs (versus)wt (weight)

mo (month)mol wt (molecular weight)no. (number)prepn (preparation)SD (standard deviation)SE (standard error)SEM (standard error of the mean)

sp act (specific activity)temp (temperature)vol (volume)wk (week)yr (year)

Reporting Numerical Data

Standard metric units are used for reporting length,weight, and volume. For these units and for molarity,use the prefixes m, pR, n, and p for 10-3, 10-6, 1o-9,and 10-2, respectively. Likewise, use the prefixes cfor 102 and k for i03. Avoid compound prefixes such asm,u or [tR. Use ,ug/ml or ,ug/g in place of the ambiguous

. .i.


ppm. Units of temperature are presented as follows:37°C or 324 K.When fractions are used to express units such as

enzymatic activities, it is preferable to use wholeunits, such as g or min, in the denominator instead offractional or multiple units, such as ,ug or 10 min.For example, "pmol/min" would be preferable to"nmol/10 min," and ",umol/g" would be preferable to"nmol/,ug." It is also preferable that an unambig-uous form such as exponential notation be used; forexample, ",uwmol g-1 min-m" is preferable to "pRmol/g/min."See the CBE Style Manual, 5th ed., for more de-

tailed information about reporting numbers. Also con-tained in this source is information on the appropriateSI units for the reporting of illumination, energy,frequency, pressure, and other physical terms. Alwaysreport numerical data in the appropriate SI unit.

Isotopically Labeled CompoundsFor simple molecules, labeling is indicated in the

chemical formula (e.g., 14Co2, 3H20, H235S04).

Brackets are not used when the isotopic symbol isattached to the name of a compound that in its naturalstate does not contain the element (e.g., 32S-ATP) orto a word that is not a specific chemical name (e.g.,131I-labeled protein, "4C-amino acids, 3H-ligands, etc.).For specific chemicals, the symbol for the isotope

introduced is placed in square brackets directly pre-ceding the part of the name that describes the labeledentity. Note that configuration symbols and modifiersprecede the isotopic symbol. The following examplesillustrate correct usage:

["4C]urea[2,3-3H]serine[ey32P]ATPSV40 [32P]DNA

L-[methyl-"4C]methionine[a-14C]lysineUDP-[U-'4C]glucosefructose 1,6-[1-32P]bisphosphate

This journal follows the same conventions for iso-topic labeling as the Journal ofBiological Chemistry,and more detailed information can be found in theInstructions to Authors of that journal (first issue ofeach year).

IX

NOTICE TO AUTHORSEffective with the January 1991 issues, articles in all ASM journals will contain aReferences section instead of a Literature Cited section. Thus, items that were notpermitted in the Literature Cited section (unpublished data, personal communications,manuscripts in preparation [which must be changed to "unpublished data"], manu-scripts submitted, "in press" references, pamphlets, abstracts, patents, theses,dissertations, newsletters, letters to the editor, editorials, and material that has notbeen peer reviewed) must be placed in the References section. Samples of correctcitations are shown below.

The Journals Division will assume the responsibility of incorporating text referencesinto the new section for accepted manuscripts during most of 1991. Thereafter, authorswill be expected to submit their manuscripts with the correct format or be asked tocorrect them at the modification stage.

If you have any questions, please feel free to call the production editor or JenniferOwens, Manager, Editorial Style and Training.

1. Anagnostopoulos, C., and J. Spizizen. 1961. Requirements for transformation in Bacillus subtilis.J. Bacteriol. 81:741-746.

2. Berry, L. J., R. N. Moore, K. J. Goodrum, and R. E. Couch, Jr. 1977. Cellular requirements forenzyme inhibition by endotoxin in mice, p. 321-325. In D. Schlessinger (ed.), Microbiology-1977.American Society for Microbiology, Washington, D.C.

3. Cox, C. S., B. R. Brown, and J. C. Smith. J. Gen. Genet., in press.*4. Dhople, A., I. Ortega, and C. Berauer. 1989. Effect of oxygen on in vitro growth of Mycobacterium

leprae, abstr. U-82, p. 168. Abstr. 89th Annu. Meet. Am. Soc. Microbiol. 1989.5. Finegold, S. M., W. E. Shepherd, and E. H. Spaulding. 1977. Cumitech 5, Practical anaerobic

bacteriology. Coordinating ed., W. E. Shepherd. American Society for Microbiology, Washington,D.C.

6. Fitzgerald, G., and D. Shaw. In A. E. Waters (ed.), Clinical microbiology, in press. EFH PublishingCo., Boston.

7. Gustlethwaite, F. P. 1985. Letter. Lancet ii:327.8. Jacoby, J., R. Grimm, J. Bostic, V. Dean, and G. Starke. Submitted for publication.9. Jensen, C., and D. S. Schumacher. Unpublished data.

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* Note that an "in press" reference to an ASM publication should state the control number(e.g., AEM 576-91) or the name of the publication, if it is a book.