journal of bacteriology · journalofbacteriology volume 171 * january 1989 * number1 samuelkaplan,...

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JOURNAL OF BACTERIOLOGY VOLUME 171 * JANUARY 1989 * NUMBER 1 Samuel Kaplan, Editor in Chief (1992) University of Illinois, Urbana Terrance J. Beveridge, Editor (1992) University of Guelph, Guelph, Canada James D. Friesen, Editor (1992) Hospital for Sick Children, Toronto, Canada June J. Lascelles, Editor (1989) University of California, Los Angeles Richard M. Losick, Editor (1993) Harvard University, Cambridge, Mass. L. Nicholas Ornston, Editor (1992) Yale University, New Haven, Conn. Robert H. Rownd, Editor (1990) Northwestern Medical School, Chicago, Ill. Kenneth N. Timmis, Editor (1992) GBF, Braunschweig, Federal Republic of Germany Graham C. Walker, Editor (1990) Massachusetts Institute of Technology, Cambridge, Mass. Robert A. Weisberg, Editor (1990) National Institute of Child Health and Human Development, Bethesda, Md. EDITORIAL BOARD Sankar Adhya (1990) Stuart J. Austin (1990) Frederick M. Ausubel (1989) Barbara Bachmann (1990) Manfred E. Bayer (1991) Margret H. Bayer (1989) Claire M. Berg (1989) Robert W. Bernlohr (1991) Mervyn Bibb (1991) Donald A. Bryant (1991) Richard Calendar (1991) Joseph M. Calvo (1990) A. M. Chakrabarty (1989) Mick Chandler (1990) Keith F. Chater (1991) Terrence G. Cooper (1990) Donald Court (1991) John E. Cronan, Jr. (1989) Jorge H. Crosa (1991) Victor de Lorenzo (1991) Bruce Demple (1991) Walter B. Dempsey (1989) Gary Ditta (1990) Timothy Donohue (1990) Ron J. Doyle (1991) David A. Dubnau (1989) S. Dusko Ehrlich (1991) Alan D. Elbein (1989) Bert Ely (1991) Wolfgang Epstein (1990) James G. Ferry (1989) David H. Figurski (1990) Timothy J. Foster (1989) Robert T. Fraley (1991) Michael Fried (1991) David I. Friedman (1989) Robert Gennis (1991) Costa Georgopolous (1990) David Gibson (1991) Jane Gibson (1991) Larry Gold (1991) Robert D. Goldman (1991) Susan Gottesman (1989) E. Peter Greenberg (1991) Nigel'Grindley (1990) Carol Gross (1990) Robert P. Gunsalus (1990) Scott R. Hagedorn (1991) R. E. W. Hancock (1990) Richard S. Hanson (1991) Shige Harayama (1990) Robert Haselkorn (1990) Gerald L. Hazelbauer (1990) George Hegeman (1991) Dennis Henner (1991) James Hopper (1991) Martha M. Howe (1990) Barbara Iglewski (1991) Karin Ihler (1990) Edward E., Ishiguro (1991) A. W. B. Johnston (1989) Clarence Kado (1991) David E. Kennell (1991) Wil N. Konings (1990) Dennis J. Kopecko (1990) Christopher Korch (1991) Susan F. Koval (1991) Viji Krishnapillai (1991) Terry Ann Krulwich (1990) Sidney Kushner (1991) Stuart B. Levy (1991) Mary E. Lidstrom (1990) Lasse Lindahl (1990) Jack London (1990) Sharon Long (1989) Paul S. Lovett (1990) Paul W. Ludden (1990) Ben J. J. Lugtenberg (1989) Robert Macnab (1991) Abdul Matin (1990) Philip Matsumura (1989) Larry McKay (1990) John Mekalanos (1991) S. Mizushima (1991) Edward A. Morgan (1990) Gisela Mosig (1991) Hiroshi Nikaido (1989) Staffan Normark (1991) Dennis Ohman (1991) William J. Paranchych (1991) John S. Parkinson (1990) Allen T. Phillips (1991) Anthony Pugsley (1991) Linda Randall (1990) Charles 0. Rock (1990) Barry P. Rosen (1989) Lucia B. Rothman-Denes (1989) Rudiger Schmitt (1989) June R. Scott (1990) Peter Setlow (1990) Howard A. Shuman (1991) Thomas J. Silhavy (1990) Gerald Smith (1991) Issar Smith (1990) Catherine Squires (1990) Gary Stacey (1991) Valley Stewart (1991) Anne 0. Summers (1990) Robert Switzer (1990) Godfried D. Vogels (1990) Judy D. Wall (1990) Barry Wanner (1990) Bernard Weisblum (1989) William B. Whitman (1991) Peter A. Williams (1989) Malcolm Winkler (1991) David Womble (1989) David R. Woods (1989) Henry C. Wu (1990) Duane C. Yoch (1989) Ryland Young (1990) Howard Zalkin (1991) David Zusman (1991) Helen R. Whiteley, Chairman, Publications Board Kirk Jensen, Director of Publications Sara Joslyn, Production Editor The Journal of Bacteriology (ISSN 0021-9193), a publication of the American Society for Microbiology, 1913 I St., N.W., Washington, DC 20006-5107, is devoted to the advancement and dissemination of fundamental knowledge concerning bacteria and other microorganisms. Instructions to authors are published in the January issue each year; reprints are available from the editors and the Publications Department. The Journal is published monthly, one volume per year. The nonmember subscription price is $340 per year; single copies are $30. The member subscription price is $41 (foreign, $71 [air drop shipping]) per year; single copies are $8. Correspondence relating to subscriptions, reprints, defective copies, availability of back issues, lost or late proofs, disposition of submitted manuscripts, and general editorial matters should be directed to the ASM Publications Department, 1913 I St., N.W., Washington, DC 20006-5107 (phone: 202 833-9680). Claims for missing issues from residents of the United States, Canada, and Mexico must be submitted within 3 months after publication of the issues; residents of all other countries must submit claims within 6 months of publication of the issues. Claims for issues missing because of failure to report an address change or for issues "missing from files" will not be allowed. Second-class postage paid at Washington, DC 20006, and at additional mailing offices. POSTMASTER: Send address changes to Journal of Bacteriology, ASM, 1913 I St., N.W., Washington, DC 20006-5107. Made in the United States of America. Printed on acid-free paper. Copyright ©) 1989, American Society for Microbiology. la: mI L1 9W J Ek Alfl A P 3t. All 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 pay the stated per-copy fee through the Copyright Clearance Center Inc., 21 Congress St., Salem, MA 01970, for copying beyond that permitted by Sections 107 and 108 of the U.S. Copyright Law. 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. Linda M. Illig, Managing Editor, Journals

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Page 1: JOURNAL OF BACTERIOLOGY · JOURNALOFBACTERIOLOGY VOLUME 171 * JANUARY 1989 * NUMBER1 SamuelKaplan, Editor in Chief(1992) University ofIllinois, Urbana TerranceJ. Beveridge, Editor(1992)

JOURNAL OF BACTERIOLOGYVOLUME 171 * JANUARY 1989 * NUMBER 1

Samuel Kaplan, Editor in Chief (1992)University of Illinois, UrbanaTerrance J. Beveridge, Editor (1992)University of Guelph, Guelph,Canada

James D. Friesen, Editor (1992)Hospital for Sick Children,

Toronto, CanadaJune J. Lascelles, Editor (1989)University of California, Los Angeles

Richard M. Losick, Editor (1993)Harvard University, Cambridge, Mass.L. Nicholas Ornston, Editor (1992)Yale University, New Haven, Conn.Robert H. Rownd, Editor (1990)Northwestern Medical School,

Chicago, Ill.

Kenneth N. Timmis, Editor (1992)GBF, Braunschweig,Federal Republic of Germany

Graham C. Walker, Editor (1990)Massachusetts Institute of

Technology, Cambridge, Mass.Robert A. Weisberg, Editor (1990)National Institute of Child

Health and HumanDevelopment, Bethesda, Md.

EDITORIAL BOARDSankar Adhya (1990)Stuart J. Austin (1990)Frederick M. Ausubel (1989)Barbara Bachmann (1990)Manfred E. Bayer (1991)Margret H. Bayer (1989)Claire M. Berg (1989)Robert W. Bernlohr (1991)Mervyn Bibb (1991)Donald A. Bryant (1991)Richard Calendar (1991)Joseph M. Calvo (1990)A. M. Chakrabarty (1989)Mick Chandler (1990)Keith F. Chater (1991)Terrence G. Cooper (1990)Donald Court (1991)John E. Cronan, Jr. (1989)Jorge H. Crosa (1991)Victor de Lorenzo (1991)Bruce Demple (1991)Walter B. Dempsey (1989)Gary Ditta (1990)Timothy Donohue (1990)Ron J. Doyle (1991)David A. Dubnau (1989)S. Dusko Ehrlich (1991)Alan D. Elbein (1989)Bert Ely (1991)Wolfgang Epstein (1990)James G. Ferry (1989)David H. Figurski (1990)

Timothy J. Foster (1989)Robert T. Fraley (1991)Michael Fried (1991)David I. Friedman (1989)Robert Gennis (1991)Costa Georgopolous (1990)David Gibson (1991)Jane Gibson (1991)Larry Gold (1991)Robert D. Goldman (1991)Susan Gottesman (1989)E. Peter Greenberg (1991)Nigel'Grindley (1990)Carol Gross (1990)Robert P. Gunsalus (1990)Scott R. Hagedorn (1991)R. E. W. Hancock (1990)Richard S. Hanson (1991)Shige Harayama (1990)Robert Haselkorn (1990)Gerald L. Hazelbauer

(1990)George Hegeman (1991)Dennis Henner (1991)James Hopper (1991)Martha M. Howe (1990)Barbara Iglewski (1991)Karin Ihler (1990)Edward E., Ishiguro (1991)A. W. B. Johnston (1989)Clarence Kado (1991)David E. Kennell (1991)

Wil N. Konings (1990)Dennis J. Kopecko (1990)Christopher Korch (1991)Susan F. Koval (1991)Viji Krishnapillai (1991)Terry Ann Krulwich (1990)Sidney Kushner (1991)Stuart B. Levy (1991)Mary E. Lidstrom (1990)Lasse Lindahl (1990)Jack London (1990)Sharon Long (1989)Paul S. Lovett (1990)Paul W. Ludden (1990)Ben J. J. Lugtenberg (1989)Robert Macnab (1991)Abdul Matin (1990)Philip Matsumura (1989)Larry McKay (1990)John Mekalanos (1991)S. Mizushima (1991)Edward A. Morgan (1990)Gisela Mosig (1991)Hiroshi Nikaido (1989)Staffan Normark (1991)Dennis Ohman (1991)William J. Paranchych (1991)John S. Parkinson (1990)Allen T. Phillips (1991)Anthony Pugsley (1991)Linda Randall (1990)Charles 0. Rock (1990)

Barry P. Rosen (1989)Lucia B. Rothman-Denes

(1989)Rudiger Schmitt (1989)June R. Scott (1990)Peter Setlow (1990)Howard A. Shuman (1991)Thomas J. Silhavy (1990)Gerald Smith (1991)Issar Smith (1990)Catherine Squires (1990)Gary Stacey (1991)Valley Stewart (1991)Anne 0. Summers (1990)Robert Switzer (1990)Godfried D. Vogels (1990)Judy D. Wall (1990)Barry Wanner (1990)Bernard Weisblum (1989)William B. Whitman

(1991)Peter A. Williams (1989)Malcolm Winkler (1991)David Womble (1989)David R. Woods (1989)Henry C. Wu (1990)Duane C. Yoch (1989)Ryland Young (1990)Howard Zalkin (1991)David Zusman (1991)

Helen R. Whiteley, Chairman, Publications Board Kirk Jensen, Director of PublicationsSara Joslyn, Production Editor

The Journal of Bacteriology (ISSN 0021-9193), a publication of the American Society for Microbiology, 1913 I St., N.W.,Washington, DC 20006-5107, is devoted to the advancement and dissemination of fundamental knowledge concerning bacteriaand other microorganisms. Instructions to authors are published in the January issue each year; reprints are available fromthe editors and the Publications Department. The Journal is published monthly, one volume per year. The nonmembersubscription price is $340 per year; single copies are $30. The member subscription price is $41 (foreign, $71 [air dropshipping]) per year; single copies are $8. Correspondence relating to subscriptions, reprints, defective copies, availability ofback issues, lost or late proofs, disposition of submitted manuscripts, and general editorial matters should be directed to theASM Publications Department, 1913 I St., N.W., Washington, DC 20006-5107 (phone: 202 833-9680).Claims for missing issues from residents of the United States, Canada, and Mexico must be submitted within 3 months afterpublication of the issues; residents of all other countries must submit claims within 6 months of publication of the issues.Claims for issues missing because of failure to report an address change or for issues "missing from files" will not be allowed.Second-class postage paid at Washington, DC 20006, and at additional mailing offices.POSTMASTER: Send address changes to Journal of Bacteriology, ASM, 1913 I St., N.W., Washington, DC 20006-5107.Made in the United States of America. Printed on acid-free paper.Copyright ©) 1989, American Society for Microbiology. la:mIL1 9W JEkAlfl A P 3t.All 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, thatthe copier pay the stated per-copy fee through the Copyright Clearance Center Inc., 21 Congress St., Salem, MA 01970, forcopying beyond that permitted by Sections 107 and 108 of the U.S. Copyright Law. This consent does not extend to otherkinds of copying, such as copying for general distribution, for advertising or promotional purposes, for creating new collectiveworks, or for resale.

Linda M. Illig, Managing Editor, Journals

Page 2: JOURNAL OF BACTERIOLOGY · JOURNALOFBACTERIOLOGY VOLUME 171 * JANUARY 1989 * NUMBER1 SamuelKaplan, Editor in Chief(1992) University ofIllinois, Urbana TerranceJ. Beveridge, Editor(1992)

1989APPLICATION FOR FULL MEMIBERSHIP IN THEAMERICAN SOCIETY FOR MICROBOLOGY

1913 I Strat, NW * Washington, D.C 20006 * (202) 833.9680COMPLETE ALL INFORMATION REQUESTED AND RETURN FORM WITH REMITTANCE IN U.S. FUNDS

Eligibility ASM omkxes to full membership aryone who is interested in its objectives and has a minimum of a bachelor's degree or equivalentin microbiology or a related field.

Initiation Memberships are iniatd and reneioed in January each year. Unless the an it to the y, m ership nomina-tions received prior to Septembt r 1 are credited to the current year, and back issues of the selectd publications for the currentyear are furnished, if available. Nominations receIved after September 1 will bem efecti e following January.

NAME MS. I(CIRCLE ONE) MRS. MR. FIRST INIlTIAL LAST

MAIL NAMEAS YOU WANT IT TO APPEAR ON YOUR MAIUNG LABEL

ADDRESSWHERE YOU WANT TO RECEIVE YOUR SUBSCRIPTIONS

STATEIPROINCE ZIWPOSTAL CODE COUNTRPHONE( ) ( ) YEAR OF BIRTH SEX_

O--FFICE (01) HOME (02)HIGHEST DEGREE SUBJECT AREA YEAR EARNED _

EMPLOYER PRESENT POSITION

SIGNATURE OF APPLICANT DATE*NOMINATED BY_______ ______MEMBER #1

SIGNATURE OF ASM MEMBER*If you are not associated with an ASM nominating member, you can still apply for membership and we will contact you.

Member How did you learn about the ASM? (Check one):Information LI A colleague O An advertisement in a journal O Presenting a paper at an ASM meeting

[1 A professor [ Direct mail inquiry O An ASM BranchO An ASM journal O A workshop, conference or meeting O Student membership in ASM

O None of the aboveDues Annual dues for 1989 are $65. Dues include ASM News (monthly) and a $43 credit which may be deducted from the total cost

of the journal(s) you purchase at the special membership rates indicated below.

Journals Please check:OL Enclosed is my dues payment (U.S. dollar only) ................................................... 865LO Please send me the following ASM journal(s) at Member Price(s):

U.S. Non-U.S. AmountAntimicrobial Agents and Chemotherapy ....................... $35 $61 .......

$ AA.

Applied and Environmental Microbiology .3.......................61..61 ....... AEMolecular and Cellular Biology .............................. 43.71 ....... CB

Clinical Microbiology Reviews ....................16...6 ..... 32 .._.....

Infection and Immunity ................................ 41.70 ......._IA

International Journal of Systematic Bacteriology.... 35....... 35 .... ___... IJ

Journal of Bacteriology ................................ 41.71 ......._JB

Journal of Clinical Microbiology ..............................35.661 .... _____..._ JCJournal of Virology ................................ 41.71 ......._

Microbiological Reviewse.vi ews........................ 16... 32 ... ____.... MRTotal Joumal Fee $ -

Subtract your $43 Member Journals Credit - $43Subtotal (if less than zero, enter zero) $Add your $65 Memberhip Dues + $65

Total (Dues plus Journals). If total is less than $65.00, enter $65 $

PAYMENT IN U.S. DOLLARS MUST ACCOMPANY APPLICATIONA membership card, voting registration form, ASM Placement Service information and the journal(s) of your choice wiH be sent within90 days upon completion of processing. ASM dues are tax deductible to the extent permitted by law. ASM designates $12 of yourdues for ASM News. Rates are for 1989 only.Applicants must remit in U.S. dollars by check or draft payable to ASM through a U.S. bank located within the Continental U.S. Appli-cants from Canada may use check made out in U.S. dollars and drawn on a Canadian bank or applicants may choose to pay withVISA, MasterCard, or American Express. If that is your preference, please fill in the box below.

O VISA # EXPIRATIONO MASTERCARD # I I I I I I I I I I DATE: MOI.YRO AMEX # MOYR

LTODAY'S DATEMONTH DAY IYEAR SIGNATURE MINIMUM CHARGE

$15.0 JB 1/89MONTHDAYYEAR~~~~~~~~~~~~~~~~~IJ 18

Page 3: JOURNAL OF BACTERIOLOGY · JOURNALOFBACTERIOLOGY VOLUME 171 * JANUARY 1989 * NUMBER1 SamuelKaplan, Editor in Chief(1992) University ofIllinois, Urbana TerranceJ. Beveridge, Editor(1992)

JOURNAL OF BACTERIOLOGY, Jan. 1989

JOURNAL OF BACTERIOLOGY

INSTRUCTIONS TO AUTHORS

HOW TO SUBMIT MANUSCRIPTSSubmit manuscripts directly to the ASM Publications

Department, 1913 I St., N.W., Washington, DC 20006-5107. Since all submissions must be processed throughthis office, alternate routings, such as to an editor, willdelay initiation of the review process. The manuscriptmust be accompanied by a covering letter stating thefollowing: the journal to which the manuscript is beingsubmitted, the most appropriate section of the journal,the address and telephone number of the correspond-ing author, and the former ASM manuscript numberand year if it is a resubmission. In addition, includewritten assurance that permission to cite personal com-munications and preprints has been granted.Authors may suggest an appropriate editor for new

submissions. If we are unable to comply with such arequest, the corresponding author will be notifiedbefore the manuscript is assigned to another editor.Submit two complete copies of each manuscript,

including figures and tables. Type every portion of themanuscript double spaced, including figure legends,table footnotes, and Literature Cited, and number allpages in sequence, including the abstract, figure leg-ends, and tables. Place the last two items after theLiterature Cited section. See p. v-vi for detailedinstructions about illustrations.

Copies of "in press" and "submitted" manuscriptsthat are important for judgment of the present manu-script should be enclosed to facilitate the review. Onecopy of each such manuscript should be provided witheach copy of the new manuscript.Authors who are unsure of proper English usage

should have their manuscripts checked by someoneproficient in the English language. Manuscripts thatare deficient in this respect may be returned to theauthor before review.

EDITORIAL POLICYManuscripts submitted to thejournal must represent

reports of original research. All authors of a manu-script must have agreed to its submission and areequally responsible for its content, including appropri-ate citations and acknowledgments. By submission ofa manuscript to the journal, the authors guarantee thatthe manuscript, or one substantially the same, was notpublished previously, is not being considered or pub-lished elsewhere, and was not rejected on scientificgrounds by another ASM journal.

Failure to comply with the above-mentioned policymay result in a 3- to 5-year suspension of publishingprivileges in ASM journals. (For further details, seeASM News, 50:260-263, 1984.)

Primary PublicationThe American Society for Microbiology accepts the

definition of primary publication as defined in How toWrite and Publish a Scientific Paper, third edition, byRobert A. Day, to wit: ". . . (i) the first publication oforiginal research results, (ii) in a form whereby peersof the author can repeat the experiments and test theconclusions, and (iii) in a journal or other sourcedocument [emphasis added] readily available withinthe scientific community."A scientific paper published in a conference report,

symposium proceeding, technical bulletin, or anyother retrievable source is unacceptable for submis-sion to an ASM journal on grounds of prior publica-tion. A preliminary disclosure of research findingspublished in abstract form as an adjunct to a meeting,e.g., part of a program, is not considered "priorpublication" because it does not meet the criteria for ascientific paper.

It is incumbent upon the author to acknowledge anyprior publication of the data contained in a manuscriptsubmitted to an ASM journal even though he or shemay not consider such publication in violation ofASMpolicy. A copy of the relevant work should accompanythe paper.

PermissionsIt is the author's responsibility to obtain permission

from the copyright owner to reproduce figures, tables,or quotations of more than 12 lines of text taken intactfrom previous publications, either his own or those ofanother author. Note that the journal or publisher (notthe author) is the copyright owner; however, as amatter of courtesy the author's permission should beobtained as well.

AuthorshipAn author is one who made a substantial contribu-

tion to the "overall design and execution of theexperiments"; therefore, ASM considers all coauthorsequally responsible for the entire paper. Individualswho provided assistance, e.g., supplied strains orreagents or critiqued the paper, should not be listed asauthors but may be recognized in the Acknowledg-ment section.

Page ChargesIt is anticipated that page charges, currently $20 per

printed page (price subject to change), will be paid byauthors whose research was supported by grants (de-partmental, governmental, institutional, etc.) or con-tracts or whose research was done as part of theirofficial duties. A bill for page charges is sent with thepage proofs and reprint order form.

If the research was not supported by any of the

i

Page 4: JOURNAL OF BACTERIOLOGY · JOURNALOFBACTERIOLOGY VOLUME 171 * JANUARY 1989 * NUMBER1 SamuelKaplan, Editor in Chief(1992) University ofIllinois, Urbana TerranceJ. Beveridge, Editor(1992)

INSTRUCTIONS TO AUTHORS

means described above, a request to waive the chargesmay be sent to Kirk Jensen, Director of Publications,American Society for Microbiology, 1913 I St., N.W.,Washington, DC 20006-5107, with the submittedmanuscript. This request, which must be separatefrom the covering letter, must state that the work wasnot supported and should be accompanied by a copy ofthe Acknowledgment section.

Minireviews (see p. v) are not subject to pagecharges.

CopyrightTo maintain and protect the Society's ownership

and rights and to protect the original authors frommisappropriations of their published work, ASM re-quires authors to sign a copyright transfer agreement.This agreement is sent to the submitting author whenthe manuscript is accepted for publication. Unless thisagreement is executed, ASM will not publish the manu-script. (U.S. government employees may file a state-ment attesting that a manuscript was prepared as partof their official duties. If they elect to do so, theyshould not sign the ASM copyright transfer agree-ment.)

ScopeThe Journal of Bacteriology publishes descriptions

of basic research on bacteria and other microorgan-isms, including fungi and other unicellular eucaryoticorganisms. Topics that are considered include struc-ture and function, biochemistry, enzymology, metab-olism and its regulation, molecular biology, genetics,plasmids and transposons, general microbiology, plantmicrobiology, chemical or physical characterization ofmicrobial structures or products, and basic biologicalproperties of organisms.ASM publishes a number of different journals cov-

ering various aspects of microbiology. Each journalhas a prescribed scope that must be considered indetermining the most appropriate journal for eachmanuscript. The following guidelines should be ofassistance.

(i) The Journal ofBacteriology will consider papersthat describe the use of antibiotics and antimicrobialagents as tools for elucidating the basic biologicalprocesses of microorganisms. However, papers deal-ing with antimicrobial agents, including manuscriptsdealing with the susceptibility, resistance, biosyn-thesis and metabolism of such agents, are more appro-priate for Antimicrobial Agents and Chemotherapy.

(ii) The Journal ofBacteriology will consider manu-scripts that emphasize the interrelationship betweenthe bacteriophage and the host cell, manuscripts aboutwork in which viruses were used as tools for elucidat-ing the structure or biological processes of microor-ganisms, and manuscripts that concern phages that arerelated to transposable elements or plasmids. Papersdescribing phage structure and those that describephage models that are pertinent for virus infection of

eucaryotic cells are more appropriate for the Journalof Virology.

(iii) Manuscripts describing new or novel methodsor improvements in media and culture conditions willnot be considered by the Journal of Bacteriologyunless they are applied to the study of basic problemsin microbiology. Such manuscripts are more appropri-ate for Applied and Environmental Microbiology or forthe Journal of Clinical Microbiology.

(iv) Manuscripts dealing with ecology or environ-mental studies, or with the application of microorgan-isms to agricultural or industrial processes, are moreappropriate for Applied and Environmental Microbiol-ogy.

(v) Manuscripts dealing with the immune system orwith topics of medical interest are more appropriatefor Infection and Immunity.

(vi) In most cases, reports that emphasize methodsand nucleotide sequence data alone (without experi-mental documentation of the functional and evolution-ary significance of the sequence) will not be consid-ered by the Journal of Bacteriology.

(vii) Papers that include extensive taxonomic mate-rial (e.g., descriptions of new taxa) should be sub-mitted to the International Journal of SystematicBacteriology, which is published by ASM for theInternational Union of Microbiological Societies.

(viii) Molecular and Cellular Biology includes stud-ies on the molecular biology of eucaryotic microbes.The scope statements for Molecular and CellularBiology and the Journal of Bacteriology are comple-mentary. They provide authors with appropriate jour-nals for the publication of research covering all aspectsof eucaryotic microbiology. When a research reportwould be equally appropriate for either journal, theauthor's preference will be followed.

Questions about these guidelines may be directed tothe editor in chief of the journal being considered.A manuscript rejected by one ASM journal on

scientific grounds or on the basis of its general suit-ability for publication is considered rejected by allother ASM journals.

Culture DepositionThe Journal of Bacteriology encourages authors to

deposit important strains in publicly accessible culturecollections and to refer to the collections and strainnumbers in the text. Since the authenticity of subcul-tures of culture collection specimens that are distrib-uted by individuals cannot be assured, authors shouldindicate laboratory strain designations and donorsource as well as original culture collection identifica-tion numbers. When authors describe mutants forwhich genetic stock repositories have not been estab-lished or strains that have not been deposited inpublicly accessible collections, the- Journal expectsthat the authors will make such strains available toother microbiologists.

. .

Page 5: JOURNAL OF BACTERIOLOGY · JOURNALOFBACTERIOLOGY VOLUME 171 * JANUARY 1989 * NUMBER1 SamuelKaplan, Editor in Chief(1992) University ofIllinois, Urbana TerranceJ. Beveridge, Editor(1992)

INSTRUCTIONS TO AUTHORS

Nucleotide SequencesASM requires that the primary nucleotide and/or

amino acid sequence data contained in a paper bedeposited in a data bank such as GenBank or EMBLData Library. A GenBank form will be sent with theacceptance letter to the corresponding author. Gen-Bank may also be contacted at: GenBank Submis-sions, Mail Stop K710, Los Alamos National Labora-tory, Los Alamos, NM 87545, U.S.A.; telephone:(505) 665-2177; electronic mail: gb-sub%[email protected] EMBL Data Library may be contacted at: EMBLData Library Submissions, Postfach 10.2209, D-6900Heidelberg, F.R.G.; telephone: (06221) 387 257; tele-fax: (496221) 387 306; computer network: (BITNET/EARN): [email protected] numbers must be included in the manu-

script or be added to the proofs.

Editorial StyleThe editorial style of ASM journals conforms to the

CBE Style Manual (5th ed., 1983; Council of BiologyEditors, Inc., 9650 Rockville Pike, Bethesda, Md.),ASM Style Manualfor Journals and Books (AmericanSociety for Microbiology, 1985), Robert A. Day's Howto Write and Publish a Scientific Paper (3rd ed., 1988,Oryx Press), and Scientific Writing for Graduate Stu-dents (Council of Biology Editors, Inc., 1968), asinterpreted and modified by the editors and the ASMPublications Department.

Review ProcessAll manuscripts are subjected to peer review by the

editors, members of the editorial board, or qualified adhoc reviewers. When a manuscript is submitted to theJournal, it is given a manuscript control number and isassigned to one of the editors. The authors are notifiedof this number and the editor to whom the manuscripthas been assigned. (It is the responsibility of thecorresponding author to inform the coauthors of themanuscript's status throughout the review and publi-cation processes.) The reviewers operate under strictguidelines set forth in "Guidelines for Reviewers" andare expected to complete their reviews expeditiously.Authors are notified, generally within 4 to 6 weeksafter submission, of acceptance, rejection, or the needfor modification. When a manuscript is returned to theauthor for modification, it should be returned to theeditor within 2 months; otherwise it may be consideredwithdrawn.

Notification of AcceptanceWhen an editor has decided that a manuscript is

acceptable for publication on the basis of scientificmerit, it is sent to the Publications Department, whereit is checked by the production editor. If the manu-

script has been prepared according to the criteria setforth in these instructions, it is scheduled for the nextavailable issue and an acceptance letter that indicatesthe month of publication, approximate page proofdates, and section is mailed to the corresponding

author. The editorial staff of the ASM PublicationsDepartment completes the editing of the manuscript tobring it into conformity with prescribed standards.

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 ASMPublications Department.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, and serious grammatical errors. "Inpress" references for which page numbers have be-come available should be placed in the LiteratureCited section as "a" numbers (e.g., 12a). Do notrenumber references.

Questions about late proofs and problems in theproofs should be directed to the ASM PublicationsDepartment, telephone (202) 833-9680.

ReprintsReprints (in multiples of 100) may be purchased by

contributors. 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 literature citation but omit the article title. Becausethe abstract will be published separately by abstracting

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INSTRUCTIONS TO AUTHORS

services, it must be complete and understandablewithout 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 the ratio-nale for the present study. Use only those referencesrequired to provide the most salient background ratherthan an exhaustive 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.Enzyme purifications should be described in this

section, but the results of such procedures should bedescribed in the Results section.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. The Results section should include theresults of the experiments. Reserve extensive interpre-tation of the results for the Discussion section. Presentthe results as concisely as possible in one of thefollowing: text, table(s), or figure(s). Avoid extensiveuse of graphs to present data that might be moreconcisely presented in the text or tables. For example,except in unusual cases, double-reciprocal plots usedto determine apparent Km values should not be pre-sented as graphs; instead, the values should be statedin the text. Similarly, graphs illustrating other methodscommonly used (e.g., calibration plots for molecularweight by gel filtration or electrophoresis) need not beshown except in unusual circumstances. Limit photo-graphs (particularly photomicrographs and electronmicrographs) to those that are absolutely necessary to

show the experimental findings. Number figures andtables in the order in which they are cited in 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 are given inseparate paragraphs. The usual format for acknowl-edgment of grant support is as follows: "This workwas supported by Public Health Service grant CA-01234 from the National Cancer Institute."

Appendixes. Appendixes, which contain supplemen-tary material to aid the reader, are permitted. Titles,authors, and Literature Cited sections that are distinctfrom those of the primary article are not allowed. If itis not feasible to list the author(s) of the appendix inthe by-line or the Acknowledgment section of theprimary article, rewrite the appendix so that it can beconsidered for publication as an independent article,either full-length or Note style. Equations, tables, andfigures should be labeled with the letter "A" precedingthe numeral to distinguish them from those cited in themain body of the text.

Literature Cited. The Literature Cited section mustinclude all relevant published work, and all listedreferences must be cited in the text. Arrange theLiterature Cited section in alphabetical order, by firstauthor, and number consecutively. Abbreviate journalnames according to Serial Sources for the BIOSISData Base (BioSciences Information Service, 1988).Cite each listed reference in the text by number.The following types of references are not valid for

listing: unpublished data, personal communications,manuscripts in preparation, manuscripts submitted,"in press" references, pamphlets, abstracts, patents,theses, dissertations, newsletters, letters to the editor,editorials, and material that has not been subjected topeer review. References to such sources should bemade parenthetically in the text. An "in press" refer-ence to an ASM journal included in Literature Citedshould state the control number (e.g., JB 976-89) or themonth of publication, so that the copy editor canverify the reference and include it in the listed refer-ences.

Follow the styles shown in the examples below.

1. Anagnostopoulos, C., and J. Spizizen. 1961. Requirementsfor 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 for enzyme inhi-

IV

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INSTRUCTIONS TO AUTHORS

bition by endotoxin in mice, p. 321-325. In D. Schles-singer (ed.), Microbiology-1977. American Society forMicrobiology, Washington, D.C.

3. Finegold, S. M., W. E. Shepherd, and E. H. Spaulding.1977. Cumitech 5, Practical anaerobic bacteriology. Co-ordinating ed., W. E. Shepherd. American Society forMicrobiology, Washington, D.C.

4. 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, 8thed. The Williams & Wilkins Co., Baltimore.

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

Parenthetical references in the text should be citedas follows:... and protects the organisms against oxygen toxic-ity (H. P. Misra and I. Fridovich, Fed. Proc. 35:1686,1976).... system was used (W. E. Scowcroft, A. H. Gibson,and J. D. Pagan, Biochem. Biophys. Res. Commun.,in press).... in linkage group XIV (R. D. Smyth, Ph.D. thesis,University of California, Los Angeles, 1972).... in poly mitochondria (S. E. Mainzer and C. W.Slayman, Abstr. Annu. Meet. Am. Soc. Microbiol.1976, K15, p. 139).

NotesThe Note format is intended for the presentation of

brief observations that do not warrant full-length pa-pers. Submit Notes in the same way as full-lengthpapers. They receive the same review, and they arenot considered preliminary communications.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 is not to exceed 1,000 words, and the number offigures and tables should be kept to a minimum.Materials and methods should be described in the text,not in the figure legends or table footnotes. Presentacknowledgments as in full-length papers, but do notuse a heading. The Literature Cited section is identicalto that of full-length papers.

MinireviewsMinireviews are brief summaries (limit of six printed

pages) of developments in fast-moving areas. Theymust be based on published articles: they may addressany subject within the scope of JB. Minireviews maybe either solicited or proffered by authors respondingto a recognized need. Irrespective of origin, minire-views are subject to editorial review.

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 PublicationsDepartment.

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, chairmanof the Publications Board, and director of publicationswill not be involved. Letters from both authors mustaccompany the author's correction sent to the Publi-cations Department.

DisclaimersStatements disclaiming governmental or any other

type of endorsement or approval will be deleted by thePublications Department.

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 size: 3516 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 necessary. 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"printer'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 so that acost estimate for printing may be obtained. The cost ofprinting color photographs must be borne by theauthor.

DrawingsSubmit graphs, charts, sequences, complicated

chemical or mathematical formulas, diagrams, and

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INSTRUCTIONS TO AUTHORS

other 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 belarge enough to withstand this reduction. Avoid heavyletters, 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 theappropriate SI symbols (,u for 10-6, m for 10-3, k for10, M for 106, etc.). A complete listing of SI symbolscan be found in the IUPAC "Manual of Symbols andTerminology for Physicochemical Quantities andUnits" (Pure Appl. Chem. 21:3-44, 1970). Thus, rep-

resentation of 20,000 cpm on a figure ordinate shouldbe made by the number 20, accompanied by the labelkcpm.When powers of 10 must be used, the journal

requires that the exponent power be associated with thenumber shown. In representing 20,000 cells per ml,the numeral on the ordinate would be "2" and thelabel would be "104 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. The preferred designation would be 60 mU/ml(milliunits per milliliter).

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 a

legend only if the discussion is very brief (one or twosentences). Define all symbols and abbreviations usedin the figure that have not been defined elsewhere.

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 Abbreviations inthese instructions for those that should be used intables. Explanatory footnotes are acceptable, butmore-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.

Tables that can be photographically reproduced forpublication without further typesetting or artwork are

TABLE 1. Induction of creatinine deiminase in C. neoformansand C. bacillisporus

C. neoformans C. bacillisporus(NIH 12) (NIH 191)

N sourceaTotal Sp act Total Sp actenyeb (U/mg of

enye (U/mg ofNsourcenzymeb protein) enzyme protein)

Ammonia 0.58 0.32 0.50 0.28Glutamic acid 5.36 1.48 2.18 0.61Aspartic acid 2.72 0.15 1.47 0.06Arginine 3.58 2.18 3.38 2.19Creatinine 97.30 58.40 104.00 58.30

a The inoculum was grown in glucose broth with ammonium sulfate,washed twice, and then transferred into the media with the N sources listedabove.

b Enzyme units in cell extract obtained from ca. 1010 cells.

referred 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.

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 (10th ed., 1983; Merck &Co., Inc., Rahway, N.J.) is also an excellent source.For guidelines to the use of biochemical terminology,consult the following: Biochemical Nomenclature andRelated Documents, 1978, reprinted for The Biochem-ical Society, London; the instructions to authors of theJournal of Biological Chemistry and the Archives ofBiochemistry and Biophysics (first issues of eachyear); and the Handbook ofBiochemistry and Molec-ular Biology (G. D. Fasman, ed., 3rd ed., CRC Press,Inc., 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 MicroorganismsBinary names, consisting of a generic name and a

specific epithet (e.g., Escherichia coli), must be used

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INSTRUCTIONS TO AUTHORS

for all microorganisms. Names of higher categoriesmay be used alone, but specific and subspecific epi-thets may not. A specific epithet must be preceded bya generic name the first time it is used in a paper.Thereafter, the generic name should be abbreviated tothe initial capital letter (e.g., E. coli), provided therecan be no confusion with other genera used in thepaper. Names of all taxa (phyla [for fungi, divisions],classes, orders, families, genera, species, subspecies)are printed in italics and should be underlined in themanuscript; strain designations and numbers are not.The spelling of bacterial names should follow the

Approved Lists ofBacterial Names (American Societyfor Microbiology, 1980) and the validation lists andrelevant articles published in the International Journalof Systematic Bacteriology since 1980. If there isreason to use a name that does not have standing innomenclature, the name should be enclosed in quota-tion marks and an appropriate statement concerningthe nomenclatural status of the name should be madein the text (for an example, see Int. J. Syst. Bacteriol.30:547-556, 1980).

Since the classification of fungi is not complete, it isthe responsibility of the author to determine the ac-cepted binomial for a given yeast or mold. Somesources for the spelling of these names include TheYeasts: a Taxonomic Study, 3rd ed. (N. J. W. Kreger-van Rij, ed., Elsevier Science Publishers B.V., 1984)and Ainsworth and Bisby's Dictionary of the Fungi,Including the Lichens, 7th ed. (Commonwealth Myco-logical Institute, Kew, Surrey, England, 1983).

Microorganisms, viruses, and plasmids should begiven designations consisting of letters and serial num-bers. It is generally advisable to include a worker'sinitials or a descriptive symbol of locale, laboratory,etc., in the designation. Each new strain, mutant,isolate, or derivative should be given a new (serial)designation. This designation should be distinct fromthose of the genotype and phenotype, and genotypicand phenotypic symbols should not be included.

Genetic NomenclatureBacteria. The genetic properties of bacteria are

described in terms of phenotypes and genotypes. Thephenotypic designation describes the observable prop-erties of an organism. The genotype refers to thegenetic constitution of an organism, usually in refer-ence to some standard wild type. Use the recommen-dations of Demerec et al. (Genetics 54:61-76, 1966) asa guide to the use of these terms.

(i) Phenotypic designations must be used whenmutant loci have not been identified or mapped. Phe-notypic designations generally consist of three-let-tersymbols; 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, nucleic acidpolymerase mutants might be designated Poll, Pol2,Pol3, etc. Wild-type characteristics can be designatedwith a superscript plus (Pol+) and, when necessary for

clarity, negative superscripts (Pol-) can be used todesignate mutant characteristics. Lowercase super-script letters may be used to further delineate pheno-types (e.g., Strs for streptomycin sensitivity). Pheno-typic designations should be defined.

(ii) Genotypic designations are sirnilarly indicatedby three-letter locus symbols. In contrast to pheno-typic designations, these are lowercase italic (e.g., arahis rps). If several loci govern related functions, theseare distinguished by italicized capital letters followingthe locus symbol (e.g., araA araB araC). Promoter,terminator, and operator sites should be indicated asdescribed by Bachmann and Low (Microbiol. Rev.44:1-56, 1980), e.g., lacZp, lacAt, and lacZo.

(iii) Wild-type alleles are indicated with a super-script plus (ara+ his'). When the genotype of anorganismn is being specified in a table, a superscriptminus is not used to indicate a mutant locus. In thetext, this distinction is best made in context, and thusone refers to an ara mutant rather than an ara- strain.

(iv) Mutation sites are designated by placing serialisolation numbers (allele numbers) after the locussymbol (e.g., araAl araA2). If only a single such locusexists or if it is not known in which of several relatedloci the mutation has occurred, a hyphen is usedinstead of the capital letter (e.g., ara-23). It is essentialin papers reporting the isolation of new mutants thatallele numbers be given to the mutations. ForEscherichia coli, there is a registry of such numbers:E. coli Genetic Stock Center, Department of Biology,Yale University, P. 0. Box 6666, New Haven, CT06511-7444. For Salmonella, the registry is: Salmo-nella Genetic Stock Center, Department of Biology,University of Calgary, Calgary, Alberta, T2N 1N4Canada. For Bacillus, the registry is: the BacillusGenetic Stock Center, Ohio State University, Colum-bus. A registry of allele numbers and insertion ele-ments (omega [Q] numbers) for chromosomal muta-tions and chromosomal insertions of transposons andother insertion elements has been established in con-junction with the ISP collection of Staphylococcusaureus at Iowa State University. Blocks of allelenumbers and fl numbers are assigned to laboratorieson request. Requests for blocks of numbers and addi-tional information can be obtained from Peter A.Pattee, Department of Microbiology, Iowa State Uni-versity, Ames, IA 50011. A registry of plasmid desig-nations is maintained by the Plasmid Reference Cen-ter, Department of Medical Microbiology, StanfordUniversity School of Medicine, Stanford, CA 94305.

(v) The use of superscripts with genotypes (otherthan + to indicate wild-type alleles) should beavoided. Designations indicating amber mutations(Am), temperature-sensitive mutations (Ts), constitu-tive mutations (Con), cold-sensitive mutations (Cs),and production of a hybrid protein (Hyb) should followthe allele number [e.g., araA230(Am) hisD21(Ts)]. Allother such designations of phenotype must be definedat the first occurrence. If superscripts must be used,

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INSTRUCTIONS TO AUTHORS

they must be approved by the editor and they must bedefined at the first occurrence.

Subscripts may be used in two situations. Subscriptsmay be used to distinguish between genes (having thesame name) from different organisms or strains, e.g.,

hisE. coli or hisK12 for the his gene of E. coli or strainK-12 in another species or strain, respectively. Anabbreviation could also be used if it were explained.Similarly, a subscript can also be used to distinguishbetween genetic elements that have the same name.

For example, the promoters of the gln operon can bedesignated glnAp1 and glnAp2.

(vi) Deletions are indicated by the symbol A placedbefore the deleted gene or region, e.g., AtrpA432,A(aroP-aceE)419, or Ahis(dhuA hisJ hisQ)1256. Simi-larly, other symbols can be used (with appropriatedefinition). Thus, a fusion of the ara and lac operons

can be shown as '1(ara-lac)95. Similarly, 4F(araB'-lacZ+)96 indicates that the fusion results in a truncatedaraB gene fused to an intact lacZ, and 4§(malE-lacZ)97(Hyb) shows that a hybrid protein is synthe-sized. An inversion is shown as IN(rrnD-rrnE)J. Aninsertion of an E. coli his gene into plasmid pSC101 atzero kilobases (O kb) is shown as pSC101 Ql(Okb::K-12hisB)4. An alternative designation of an insertioncan be used in simple cases, e.g., galT236::Tn5. Thenumber 236 refers to the locus of the insertion, and ifthe strain carries an additional gal mutation, it is listedseparately. Additional examples, which utilize a

slightly different format, can be found in the papers byCampbell et al. and Novick et al. cited below. It isimportant in reporting the construction of strains inwhich a mobile element was inserted and subsequentlydeleted that this latter fact be noted in the strain table.This can be done by listing the genotype of the strainused as an intermediate, in a table footnote, or by a

direct or parenthetical remark in the genotype, e.g.,(F-), AMu cts, mal::AMu cts::lac. In setting paren-

thetical remarks within the genotype or dividing thegenotype into constituent elements, parentheses andsquare brackets are used without special meaning;square brackets are used outside parentheses. Toindicate the presence of an episome, parentheses (orbrackets) are used (k, F+). Reference to an integratedepisome is indicated as described for inserted ele-ments, and an exogenote is shown as, for example,W3110/F'8(gal+).Any deviations from standard genetic nomenclature

should be explained in Materials and Methods or in a

table of strains. For more detailed information aboutthe symbols in current use, consult Bachmann(MicrQbiol. Rev. 47:180-230, 1983) for E. coli K-12,Sanderson and Roth (Microbiol. Rev. 52:485-532,1988) for Salmonella typhimurium, Holloway et al.(Microbiol. Rev. 43:73-102, 1979) for Pseudomonas,Piggot and Hoch (Microbiol. Rev. 49:158-179, 1985)for Bacillus subtilis, Perkins et al. (Microbiol. Rev.46:426-570, 1982) for Neurospora crassa, andMortimer and Schild (Microbiol. Rev. 49:181-213,1985) for Saccharomyces cerevisiae.

"Mutant" vs. "mutation." Keep in mind the distinc-tion between a mutation (an alteration of the primarysequence of the genetic material) and a mutant (astrain carrying one or more mutations). One mayspeak about the mapping of a mutation, but one cannotmap a mutant. Likewise, a mutant has no geneticlocus, only a phenotype.

Strain designations. Do not use the genotype as aname (e.g., "subsequent use of leuC6 for transduc-tion"). If a strain designation has not been chosen,select an appropriate word combination (e.g., "an-other strain containing the leuC6 mutation").

Bacteriophages. The genetic nomenclature forphages differs from that for bacteria. In most in-stances, phages have no phenotype, since they haveno metabolism outside host cells. Therefore, distinc-tions between phenotype and genotype are not made.Superscripts are used to indicate hybrid genomes.Genetic symbols may be one, two, or three letters. Forexample, a mutant strain of X might be designated asAamll int2 redll4 c1857; this strain carries mutationsin genes cI, int, and red and an amber-suppressible(am) mutation in gene A. A strain designated X att434imm21 would represent a hybrid of phage A whic.Acarries the immunity region (imm) of phage 21 and theattachment (att) region of phage 434. Host DNAinsertions into phages should be delineated by squarebrackets, and the genetic symbols and designations forsuch inserted DNA should conform to those used forthe host genome. Genetic symbols for phage A can befound in Szybalski and Szybalski (Gene 7:217-270,1979) and in Echols and Murialdo (Microbiol. Rev.42:577-591, 1978).

Transposable elements, plasmids, and restriction en-zymes. Nomenclature of transposable elements (inser-tion sequences, transposons, phage Mu, etc.) shouldfollow the recommendations of Campbell et al. (Gene5:197-206, 1979), with the modifications given in sec-tion vi. The system of designating transposon inser-tions at sites where there are no known loci, e.g.,zef-123::TnS, has been described by Chumley et al.(Genetics 91:639-655, 1979). The nomenclature rec-ommendations of Novick et al. (Bacteriol. Rev.40:168-189, 1976) for plasmids and plasmid-specifiedactivities, of Low (Bacteriol. Rev. 36:587-607, 1972)for F-prime factors, and of Roberts (Nucleic AcidsRes. 9:r75-r96, 1981) for restriction enzymes andDNA fragments derived from treatment with theseenzymes should be used. Recombinant DNA mole-cules constructed in vitro follow the nomenclature forinsertions in general. DNA inserted into recombinantDNA molecules should be described by using the genesymbols and conventions for the organism from whichthe DNA was obtained. The Plasmid Reference Cen-ter, Stanford University School of Medicine, Stanford,CA 94304, assigns Tn and IS numbers to avoid con-

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INSTRUCTIONS TO AUTHORS

flicting and repetitive use and also clears noncon-flicting plasmid prefix designations.

ABBREVIATIONS AND CONVENTIONS

Verb TenseUse the past tense to narrate particular events in the

past, including the procedures, observations, and dataof the study that you are reporting. Use the presenttense for your own general conclusions, the conclu-sions of previous researchers, and generally acceptedfacts. Thus, most of the abstract, Materials and Meth-ods, and Results sections will be in the past tense, andmost of the introduction and some of the Discussionwill be in 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," "Fig. 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 leastfive times in *the text (including tables and figurelegends).Not requiring introduction. In addition to abbrevia-

tions for standard units of measurement and chemicalsymbols of the elements, the following should be usedwithout definition in the title, abstract, text, figurelegends, and tables: DNA (deoxyribonucleic acid);cDNA (complementary DNA); RNA (ribonucleicacid); cRNA (complementary RNA); RNase (ribonucle-ase); DNase (deoxyribonuclease); rRNA (ribosomal

RNA); mRNA (messenger RNA); tRNA (transferRNA); AMP, ADP, ATP, dAMP, ddATP, GTP, etc.(for the respective 5' phosphates of adenosine or othernucleosides) (add 2'-, 3'-, or 5'- when needed forcontrast); ATPase, dGTPase, etc. (adenosine triphos-phatase, deoxyguanosine triphosphatase, etc.); NAD(nicotinamide adenine dinucleotide); NAD+ (nicotin-amide adenine dinucleotide, oxidized); NADH (nico-tinamide adenine dinucleotide, reduced); NADP(nicotinamide adenine dinucleotide phosphate);NADPH (nicotinamide adenine dinucleotide phos-phate, reduced); poly(A), poly(dT), etc. (polyadenylicacid, polydeoxythymidylic acid, etc.); oligo(dT), etc.(oligodeoxythymidylic acid, etc.); Pi (orthophos-phate); PPi (pyrophosphate); UV (ultraviolet); PFU(plaque-forming units); CFU (colony-forming units);MIC (minimal inhibitory concentration); MBC (min-imal bactericidal concentration); Tris [tris(hydroxy-methyl)aminomethane]; DEAE (diethylaminoethyl);A260 (absorbance at 260 nm); and EDTA (ethylene-diaminetetraacetic acid). Abbreviations for cell lines(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)mo (month)mol wt (molecular weight)no. (number)prepn (preparation)SD (standard deviation)

Reporting Numerical Data

SE (standard error)SEM (standard error of themean)

sp act (specific activity)sp gr (specific gravity)temp (temperature)tr (trace)vol (volume)vs (versus)wk (week)wt (weight)yr (year)

Standard metric units are used for reporting length,weight, and volume. For these units and for molarity,use the prefixes m, ,i, n, and p for 10-3, 10-6, 10-9,and 10- 2, respectively. Likewise, use the prefix k for103. Avoid compound prefixes such as m,u or p,u. Usejig/ml or pug/g in place of the ambiguous ppm. Units oftemperature 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 denominatorinstead of fractional or multiple units, such as jig or 10min. For example, "pmol/min" would be preferable to"nmol/10 min," and "[tmol/g" would be preferable to"nmol/,ug." It is also preferable that an unambiguousform such as the exponential notation be used insteadof multiple slashes; for example, "jimol g-3 min-" ispreferable to ",umol/g per 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 SI units for thereporting of illumination, energy, frequency, pressure,

IX

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INSTRUCTIONS TO AUTHORS

and other physical terms. Always report numericaldata in the appropriate SI unit.

entity. Note that configuration symbols and modifiersprecede the isotopic symbol. The following examplesillustrate correct usage:

Isotopically Labeled CompoundsFor simple molecules, isotopic labeling is indicated

in the chemical formula (e.g., C02, 3H20, H235S04).Brackets are not used when the isotopic symbol isattached to a word that is not a specific chemicalname (e.g., 11-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 labeled

['4C]ureaL-[methyl-4C]fmethionine[2,3-3H]serine[a-14C]lysine

[ey32P]ATPUDP-[U-14C]glucoseE. coli [32P]DNAfructose 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).

x

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Author IndexAbeles, Ann L., 43Adams, Camellia W., 473Adams, Lee F., 521Aiba, Atsu, 198Alam, Jawed, 162Alam, Kiswar Y., 342Allen, Judith E., 285Allen, Kenneth E., 53Angerer, Annemarie, 238Arp, Daniel J., 430Artz, Stanley, 538Asoh, Sadamitsu, 558Austin, Stuart J., 43

Balakrishnan, Ramaswami,488

Bassford, Philip J., Jr., 402,503

Battisti, Laurie, 104Baumann, Maritta, 308Baumeister, W., 190Beatty, J. Thomas, 473Belas, Robert, 602Bercovier, H., 70, 581Bergemann, Andrew D., 593Bernard, Theophile, 531Bieger, Charles D., 141Blum, Paul, 538Bolten, Barbara M., 488Borotz, Susan E., 573Brandner, Janine P., 360Braun, Volkmar, 238Brefort, Georges, 547Briggs, Kathleen, 547Burger-Wiersma, Tineke, 498Byers, David M., 59

Cameron, Beatrice, 547Cava, Joseph R., 8Cavard, Daniele, 410Chang, Ming, 172Cho, Dan-Sung C., 24Clark, David P., 342Clark-Curtiss, J. E., 70Click, Eva Marie, 616Cohen, Stanley N., 473Cornelis, Guy, 254Court, Donald L., 585Crawford, Irving P., 172Crouzet, Joel, 547Curtis, Stephanie, 162

Daimon, Hirohiko, 620Danneel, Hans-Jurgen, 308Daum, Henry A., III, 205Dean, David A., 503Decaris, Bernard, 419Demuyter, Philippe, 419de Rouvroit, Catherine

Lambert, 254Dicker, David T., 349Donohue, Timothy J., 360,436

Dooley, J. S. G., 190Doyle, Charles M., 430Driessen, Arnold J. M., 280,

292Dykstra, Christine C., 565

Eberhard, Anatol E., 1Echols, Harrison, 588Egilmez, Nejat K., 37Elias, Pappi M., 8

Engel, Joanne N., 335Engelhardt, H., 190Escamilla, Edgardo, 465

Fikes, John D., 402, 503Finch, Lloyd R., 593Forrest, Mary E., 473Fortson, Mark, 565Fosnaugh, Kathy, 606Foster, Rosemary, 272Free, Stephen J., 53Fuchs, James A., 162Fukui, Tetsuya, 184

Ganem, Don, 335Garcia-Bustos, Jose, 114Gehring, Kalle, 503Gerami-Nejad, Maryam, 162Gibson, Jane, 1Giffhorn, Friedrich, 308Glaser, David, 349Gleason, Florence K., 162Goldberg, Edward B., 488Golden, Susan S., 24Goosen, Nora, 447Gots, Joseph S., 205Green, Brian D., 104Greenberg, E. P., 606Grossman, Nili, 74, 80Guerinot, Mary Lou, 573Gutierrez, Claude, 511

Hadero, Ayele, 172Hardigree, Alice A., 230Harker, Alan R., 314Harwood, Caroline S., 1Heinzel, Peter, 321Helmann, John D., 222Higgins, Michael L., 349Hirayama, Noriaki, 596Holzschu, Donald, 538Horsman, Harold P. A., 447Howard, S. Peter, 410Huinen, Rend G. M., 447

Igarashi, Yasuo, 65Imanaka, Tadayuki, 369Inada, Toshifumi, 585Ippen-Ihler, Karin, 213Ishiguro, Naotaka, 620Ishii, Masaharu, 65Ishikawa, Kiichi, 184Ishizaki, Tomoyuki, 596Ito, Eiji, 424Ito, Koreaki, 585Iwasaki, Hiroyoshi, 424

Jacobs, Nicholas J., 573Jacobson, Gary R., 263Jazwinski, S. Michal, 37Jensen, Peter, 565Jimenez, Antonio, 321, 329Jung, Jae U., 511Jurgens, Uwe J., 498

Kadner, Robert J., 154Kamholz, John, 205Kaplan, Samuel, 360, 436Kay, W. W., 190Keyhani, Jacqueline, 205Kijne, Jan W., 569Kodama, Tohru, 65Komatsu, Keiko, 353

Konings, Wil N., 280, 292Konisky, Jordan, 93Kovacs, Kornel L., 430Kranz, Robert G., 456Kredich, Nicholas M., 130Kuhstoss, S., 16Kunai, Kenji, 558Kuramitsu, Howard K., 263Kuriki, Takashi, 369Kushner, Sidney R., 565Kuster, Uta, 308Kwan, Hoi-Shan, 538

Laakel, Mohamed, 419Larimer, Frank W., 230Lazdunski, Claude, 410Leblond, Pierre, 419Le Rudulier, Daniel, 531Lett, Marie-Claire, 353Levinson, Hillel S., 83Lipinska, Barbara, 488L6pez-Cabrera, Manuel, 321,

329Losick, Richard, 561Lowendorf, Henry S., 53Lugtenberg, Ben J. J., 569Lundrigan, Michael D., 154

Madduri, K., 299Mahan, Michael J., 612Mahler, Inga, 83, 222Mainzer, Stanley E., 244Makino, Souich, 353Martin, G. Steven, 272Mat-Jan, Fairoz, 342Matsuyama, Asahi, 577Matsuzawa, Hiroshi, 558McEwan, Alastair G., 360McNally, Mark T., 53Merkel, Susan M., 1Mett, Helmut, 483Michiels, Thomas, 254Misono, Haruo, 30Mitchenall, Lesley A., 124Miwa, Keiko, 184Mizobuchi, Kiyoshi, 198Moore, Melissa, 83Mortenson, Leonard E., 430Mount, David W., 303Moutier, Laurence, 419Mullin, David A., 383Muraiso, Kanae, 558Murooka, Yoshikatsu, 596

Nagasaki, Susumu, 30Nagata, Shinji, 30Nakamura, Yoshikazu, 585Nakanishi, Shigetada, 184Nakano, Eiichi, 577Nalty, Mark S., 24Nettleton, David O., 120Newton, Austin, 383, 392Nierlich, Donald P., 141Nikaido, Hiroshi, 503Nimi, Osamu, 596Ninfa, Alexander J., 383Noel, K. Dale, 8

Odani, Shoji, 184Ohta, Takahisa, 558Okada, Nobuhiko, 353Olsen, Ronald H., 314

Ordal, George W., 120Ossanna, Nina, 303Ostrowski, Jacek, 130

Panzer, Scott, 561Pardo, Jose Manuel, 329Pau, Richard N., 124Pdrez-Gonzalez, Josd

Antonio, 321, 329Perry, Jack R., 230Piepersberg, Wolfgang, 321Plunkett, Guy, III, 588Pocard, Jean-Alain, 531Poole, Robert K., 465Poolman, Bert, 244Poy, Florence, 263Pridmore, Sylvie, 547

Ramakrishnan, Girija, 383Rao, A. S. M. Krishna, 488Rao, R. Nagaraja, 16Razin, S., 581Reaves, Lucretia D., 43Richardson, M. A., 16Riggs, Daniel, 538Robson, Robert L., 124Ron, Eliora Z., 74, 80Rosner, Esther, 74Rosta, Sabine, 483Roth, John R., 612Royer, Theresa J., 244Russell, James B., 280

Saito, Terumi, 184Sakai, Takashi, 353Sakurai, Kyoko, 375Salyers, Abigail A., 148Sampei, Gen-ichi, 198Sanbongi, Yoshihiro, 65Sardinia, Lisa M., 335Sasakawa, Chihiro, 353Sasaki, Takuji, 375Sato, Gihei, 620Sato, Yutaka, 263Schmidt, Brian F., 244Schnaitman, Carl A., 616Schneider, Karl-Heinz, 308Seefeldt, Lance C., 430Seidler, Ramon J., 314Sela, S., 70, 581Setlow, Peter, 561Shimada, Akira, 424Shinagawa, Morikazu, 620Shinkawa, Hidenori, 596Silver, Simon, 83Simon, Horst, 308Simonet, Jean-Marc, 419Slayman, Carolyn W., 53Sluiters, Christine, 254Smid, Eddy J., 292Smit, Gerrit, 569Smith, John M., 205Smith, Linda Tombras, 531Sober6n, Mario, 465Sockett, R. Elizabeth, 436Sommer, Jurg M., 392Speer, Brenda S., 148Stephens, Richard S., 285Strobel, Herbert J., 280Stuttard, C., 299Sun, Dongxu, 561Suzuki, Jun-ichi, 184Suzuki, Kenji, 184

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ii

Takasuga, Akiko, 558Takekawa, Shiro, 375Thorne, C. B., 104Thorner, Jeremy, 272Tiedeman, Amelia A., 205Tigyi, Gabor, 430Tomasz, Alexander, 114Tomita, Kenkichi, 184Trust, Trevor J., 190Tsukagoshi, Norihiro, 375Turowski, Debra A., 8

Udaka, Shigezo, 375Uozumi, Nobuyuki, 375van de Putte, Pieter, 447Varga, Amy R., 436Villarejo, Mema R., 511Vining, L. C., 299Visick, Jonathan E., 521Volkert, Michael R., 99

Walsh, Christopher T., 83,222

Wang, Ying, 83, 222Watanabe, Wakako, 198Whiteley, H. R., 521Whitley, Jane C., 593Whitman, William B., 93Williams, Huw D., 465Williams, Michael G., 565Wood, Alvin G., 93Wu, June H., 213

Yamada, Masatoshi, 353

Yamagata, Hideo, 375Yamamoto, Hideko, 577Yamamoto, Jun, 184Yogev, D., 581Yokoyama, Kohei, 424Yonezawa, Junichi, 30Yoshikawa, Masanosuke, 353

Zimmermann, Luitgard, 238Zito, Edward T., 349