JOURNAL OF BACTERIOLOGYVOLUME 165 JANUARY 1986 NUMBER 1

Simon Silver, Editor in Chief (1987)Washington UniversitySt. Louis, Mo.

James D. Friesen, Editor (1987)University of TorontoToronto, Canada

Stanley C. Holt, Editor (1987)The University of Texas HealthScience Center, San Antonio

Samuel Kaplan, Editor (1988)University of Illinois, Urbana

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

Richard M. Losick, Editor (1988)Harvard UniversityCambridge, Mass.

L. Nicholas Ornston, Editor (1987)Yale UniversityNew Haven, Conn.

Robert H. Rownd, Editor (1990)Northwestern Medical SchoolChicago, Ill.

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

TechnologyCambridge, Mass.

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

Health and HumanDevelopment

Bethesda, Md.

EDITORIAL BOARDDavid Apirion (1988)Stuart J. Austin (1987)Frederick M. Ausubel (1986)Gad Avigad (1986)Barbara'Bachmann (1987)Manfred E. Bayer (1988)Margaret, H. Bayer (1986)Claire M. Berg (1986)Robert W. Bernlohr (1988)Terry J. Beveridge (1988)Edwin Boatman (1986)Joseph M. Calvo (1987)A. M. Chakrabarty (1986)Keith F. Chater (1988)Terence G. Cooper (1987)John E. Cronan, Jr. (1986)Walter B. Dempsey (1986)Patrick Dennis (1988)W. Ford Doolittle (1988)

David A. Dubnau (1986)Wolfgang Epstein (1987)David Figurski (1987)Timothy J. Foster (1986)Masamnitsu Futai (1988)Robert Gennis (1988)Jane Gibson (1988)E. Peter Greenberg (1988)Barry G. Hall (1988)R. E. W. Hancock (1987)Robert Haselkorn (1987)Gerald L. Hazelbauer (1987)Charles L. Hershberger (1987)Penny J. Hitchcock (1987)Martha M. Howe (1987)Karin Ihler (1987)A. W. B. Johnston (1986)Eva R. Kashket (1987)Wil N. Konings (1987)

Jordan Konisky (1987)Dennis J. Kopecko (1987)Terry Krulwich (1987)Lasse Lindahl (1987)Jack London (1987)Paul S. Lovett (1987)Ben J. J. Lugtenberg (1986)Robert Macnab (1988)Francis L. M,acrina (1986)Sarah A. McIntire (1986)Larry McKay (1987)S. Mizushima (1988)Edward A. Morgan (1987)R. G. E. Murray (1986)Robert A. Niederman (1988)Patrick J. Piggot (1987)Barry Polisky (1987)Linda Randall (1987)Marilyn Roberts (1986)

Palmer Rogers (1987)Barry P. Rosen (1986)Rudiger Schmitt (1986)June R. Scott (1987)Jane K. Setlow (1987)Peter Setlow (1987)Lucille Shapiro (1986)Issar Smith (1987)Catherine Squires (1987)Robert Switzer (1987)Alexander Tomasz (1986)Godfried D. Vogels (1987)Barry Wanner (1987)Bernard Weisblum (1986)David Womble (1986)Hienry C. Wu (1987)Duane C. Yoch (1986)Howard Zalkin (1988)

Helen R. Whiteley, Chairman, Publications BoardLinda M. Illig, Managing Editor, Journals Dennis G. Burke, Production Editor

The Journal of Bacteriology (ISSN 0021-9193), a publication of the American Society for Microbiology, 1913 I St., NWWashington, DC 20006, is devoted to the advancement and dissemination of fundamen,tal knowledge concerniing bacteria andother microorganisms. Instructions to authors are published in the January issue each year; reprints are available from theeditors and the Publications Department. The Journal is published monthly, and the twelve numbers are divided into fourvolumes per year. The nonmember subscription price is $280 per year; single copies are $21. The member subscription price is$41 (foreign, $54 [surface rate]) per year; single copies are $7. Correspondence relating to subscriptions, reprints, defectivecopies, availability of back issues, lost or late proofs, disposition of submitted manuscripts, and general editorial mattersshould be directed to the ASM Publications Department, 1913 I St., NW, Washington, DC 20006 (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.Clainis for issues missing because offailure 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., NW, Washington, DC 20006.Made in the United States of America.Copyright © 1986, American Society for Microbiology. at: /l?" 9 t Pl{8J; ( I'1kt.All Rights Reserved.

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Author IndexAhmad, Suhail, 146Allen, Geoffrey, 21Anraku, Yasuhiro, 28Aronson, Arthur, 258

Balganesh, Meenakshi, 308Barbas, Julio A., 269Bialy, Harvey, 321Bitoun. Remy, 175Brown, Stuart W., 82Burnett, Thomas J., 139Butler, J. Scott, 198

Cameron, David R., 181Carmel, Gilles, 181Cejudo, Francisco J., 240Chung, Thomas D. Y., 283Cohen, Reuben A., 41Coucheron, Dag H., 336Coulton, James W., 181Cox, John, 328

Dailey, Harry A., 1D'Ari, Richard, 66Davidson, Barrie E., 233Debro, LaJoyce, 258DeFeyter, Robert C., 226,

233, 331Dfaz, Javier, 269Donald, Robert G. K., 72Dondon, Jacques, 198Downs, Diana, 193Drake, Harold L., 315DuPlain, Louise, 88

Eisner, Robin L., 204English, Jami E., 155

Fairweather, Neil F., 21Fillingame, Robert H., 244Finnerty, W. R., 301Fitz-James, Phillip C., 258Fitzmaurice, Wayne P., 297Fleming, Jennie E., 1Fry, Brian, 328Fukuchi, Akira, 334

Furuichi, Teiichi, 94

Galizzi, Alessandro, 123Gest, Howard, 328Gianni, Maurizio, 123Gilleece, Elaine S., 47Goldberg, Alfred L., 193Gottesman, Max, 167Grunberg-Manago, Marianne,

198

Hackstadt, Ted, 13Hageman, James H., 139Harbin, Bertille M., 1Haug, A., 116Hayes, J. M., 328Hazelbauer, Gerald L., 312Hermolin, Joe, 244Hernandez, Georgina, 133Hirato, Toru, 324Hopkins, John D., 47Horitsu, Hiroyuki, 334Hubbard, Michael J., 61

Ishiguro, Naotaka, 324Ivey, D. Mack, 252

Jaffe, Aline, 66Jarvis, Erich D., 204Jean, Richard, 181Jensen, Roy A., 146

Kamerud, John Q., 293Kawai, Keiichi, 334Kayanuma, Tsuneo, 94Kjosbakken, Johs, 336Kogoma, Tokio, 321Komano, Teruya, 94Koshland, Daniel E., Jr., 276Kossmann, Marina, 34Kubo, Akihiro, 94

LaFauci, Giuseppe, 204LaPointe, Jacques, 88Lers, Amnon, 175Levy, Stuart B., 82

Ljungdahl, Lars G., 252Loroch, Albert I., 72Ludwig, Robert A., 72, 304Lyness, Valerie A., 21

Mahajna, Jamal, 167Manilofff, Jack, 219Manson, Michael D., 34Markie, David, 61Mason, Patrizia, 181Matzanke, Berthold F., 283Mayer, Kenneth H., 47McGroarty, Estelle J., 116McKenney, Denise, 6Melton, Thoyd, 6Miyamoto, Shigemi, 28Mora, Jaime, 133Mora, Yolanda, 133Mutoh, Norihiro, 161

Nees, David W., 72Nisioka, Taizo, 94Nogi, Yasuhisa, 101Norris, Victor, 66Nowak, Jan A., 219

O'Brien, Thomas F., 47Ohsumi, Yoshinori, 28Ohya, Yoshikazu, 28Oppenheim, Amos B., 167Ornston, L. Nicholas, 288

Paneque, Antonio, 240Park, Chankyu, 312Parke, Donna, 288Peterson, Arnold A., 116Pickard, Derek J., 21Pittard, James, 226, 233, 331Pollack, J. Dennis, 53Porter, Bruce, 244Poulter, Russell T. M., 61Proulx, Mario, 88

Rashtchian, Ayoub, 82Rattray, Alison, 167Raymond, Christopher K., 72

Raymond, Kenneth N., 283Reichler, Judith, 82Rightmire, Berit, 146Rode, Harold N., 181Rodrfguez-Tebar, Alfredo, 269Roon, Robert J., 293Roth, John, 193Rothstein, David M., 319Rudner, Rivka, 204Russo, Andrew F., 276

Sato, Gihei, 324Savage, M. Dean, 315Scocca, John J., 297Setlow, Jane K., 308Shankweiler, Gary W., 139Shinagawa, Morikazu, 324Simon, Melvin I., 161Singer, John T., 301Sladek, Todd L., 219Springer, Mathias, 198Sun, Tai-ping, 107Syvanen, Michael, 47

Thomson, Robert O., 21

Valla, Svein, 336van Tuijl, Johanna J., 301Vary, Patricia S., 155Vazquez, David, 269

Wachi, Sayuri, 334Waldman, Alan S., 297Waxman, Lloyd, 193Webster, Robert E., 107White, Laura K., 244White, Robert H., 215Widom, Russell L., 204Williams, Marshall V., 53Winkelmann, Gunther, 283

Yamamoto, Kazumi, 334Yelton, David B., 41

Zamir, Ada, 175

JOURNAL OF BACTERIOLOGY, Jan. 1986

JOURNAL OF BACTERIOLOGY

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INSTRUCTIONS TO AUTHORS

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experiments reported in the paper may be described in

. .

INSTRUCTIONS TO AUTHORS

the Results section or, if very brief (one or twosentences), 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 toshow 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.

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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, 1984).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, and material thathas not been subjected to peer review. References'tosuch sources should be made parenthetically in thetext. An "in press" reference to an ASM journalincluded in Literature Cited should state the controlnumber (e.g., JB 976-86) or the month of publication,so that the copy editor can verify the reference andinclude it in the listed references.

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-bition by endotoxin i-n 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).

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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

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IV

INSTRUCTIONS TO AUTHORS

use a heading. The Literature Cited section is identicalto that of full-length papers.

MinireviewsMinireviews are brief summaries (limit of six printed

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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, diagrams, and other drawings

as glossy photographs made from finished drawings notrequiring additional artwork or typesetting. No part ofthe graph or drawing should be typewritten. Use alettering set or other professional-quality device for alllabeling. Both axes of graphs must be labeled'. Mostgraphs 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. Direct readouts from comput-ers, recorders, etc., are not usually acceptable; suchmaterials should be redrawn. One of the two sets ofdrawings may consist of photocopies: the other, how-ever, must consist of photographs.

In figure ordinate and abscissa scales (as well astable 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 for103, 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 employed, the editorial

style of the Journal follows the CBE Style Manualrecommendation, which differs from the conventionsused by several other journals. The CBE Style Manualsuggests 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 x10-4'"). 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 alegend only if the discussion is very brief (one or two

v

INSTRUCTIONS TO AUTHORS

sentences). Define all symbols and abbreviations usedin the figure that have not been defined elsewhere.

Camera-Ready CopyDrawings, tables, chemical formulas, etc., that can

be photographically reproduced for publication with-out further typesetting or artwork are referred to as"camera ready." Such copy may also be prepared forcomplicated mathematical or physical formulas, por-tions of genetic maps, diagrams, and flow schemes. Itshould not be hand lettered. Camera-ready copy mustbe carefully prepared to conform to the style of theJournal. The advantage in submitting camera-readycopy is that the material will appear exactly as envi-sioned by the author, and no second proofreading isnecessary. This is particularly advantageous whenthere are long, complicated tables and when the divi-sion of material and spacing are important.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. A well-constructed table is shown below.

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

(NIH 12) (NIH 191)

N sourcea Sp act Sp actTotal (U/mg Total (U/mg

enzymeb of enzyme ofprotein) 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.

NOMENCLATUREChemical and Biochemical NomenclatureThe recognized authority for the names of chemical

compounds is Chemical Abstracts (Chemical Ab-stracts Service, Ohio State University, Columbus) andits indexes. For guidelines to the use of biochemicalterminology, consult the following: Biochemical No-menclature and Related Documents, 1978, reprintedfor The Biochemical Society, London; the instructionsto authors of the Journal ofBiological Chemistry and

the Archives of Biochemistry and Biophysics (firstissues of each year); and the Handbook of Biochem-istry and Molecular Biology (G. D. Fasman, ed., 3rded., 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., 1979).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 usedfor 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; strain designations and numbersare not.The spelling of bacterial names should follow the

Approved Lists ofBacterial Names (American Societyfor Microbiology, 1980), the subsequent validationlists and relevant articles published in the Interna-tional Journal of Systematic Bacteriology since 1980,or Bergey's Manual ofSystematic Bacteriology (N. R.Krieg and J. G. Holt, ed., The Williams & WilkinsCo., 1984). If there is reason to use a name that doesnot have standing in nomenclature, the name shouldbe enclosed in quotation marks and an appropriatestatement concerning the nomenclatural status of thename should be made in the text (for an example, seeInt. J. Syst. Bacteriol. 30:547-556, 1980).

Since the classification of fungi is not complete, it isthe responsibility of the author to determine the cur-rently accepted binomial for a given yeast or mold.Some sources for the spelling of these names includeThe Yeasts: 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 theFungi, Including the Lichens, 6th ed. (CommonwealthMycological Institute, Kew, Surrey, England, 1971).

Microorganisms, viruses, and plasmids should begiven strain designations consisting of letters andserial numbers. It is generally advisable to include aworker's initials or a descriptive symbol of locale,laboratory, etc., in the designation. Each new strain,mutant, isolate, or derivative should be given a new

Vi

INSTRUCTIONS TO AUTHORS

(serial) designation. This designation should be dis-tinct from those of the genotype and phenotype, andgenotypic and phenotypic symbols should not be in-cluded.

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 in employing these terms.

(i) Phenotypic designations must be employed whenmutant loci have not been identified or mapped. Phe-notypic 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 bacteri-ocin-tolerant mutants might be designated Toll, TolIl,TolIlI, etc., or a series of nucleic acid polymerasemutants might be designated Poll, Pol2, Pol3, etc.Wild-type characteristics can be designated with asuperscript plus (Tol+ or Pol+) and, when necessaryfor clarity, negative superscripts (Tol- Pol-) can beused to designate mutant characteristics. Lowercasesuperscript letters may be used to further delineatephenotypes (e.g., Strs for streptomycin sensitivity).Phenotypic designations should be defined.

(ii) Genotypic designations are similarly 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 anorganism is being specified (e.g., in a strain table), asuperscript minus is not used to indicate a mutantlocus. Elsewhere, a superscript minus may be used todistinguish between the symbol of a mutant allele andthat of a genetic locus. However, this distinction isbest made in the context, and thus one refers to an aramutant 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 HumanGenetics, Yale University School of Medicine, P. 0.

Box 3333, New Haven, CT 06510. For Salmonella, theregistry is: Salmonella Genetic Stock Center, Depart-ment of Biology, University of Calgary, Calgary,Alberta, T2N 1N4 Canada. For Bacillus, the registryis: the Bacillus Genetic Stock Center, Ohio StateUniversity, Columbus. A registry of allele numbersand insertion elements (omega [fQ] numbers) for chro-mosomal mutations and chromosomal insertions oftransposons and other insertion elements has beenestablished in conjunction with the ISP collection ofStaphylococcus aureus at Iowa State University.Blocks of allele numbers and fl numbers are assignedto laboratories on request. Requests for blocks ofnumbers and additional information can be obtainedfrom Peter A. Pattee, Department of Microbiology,Iowa State University, Ames, IA 50011. A registry ofplasmid designations is maintained by the PlasmidReference Center, Department of Medical Microbiol-ogy, Stanford University 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,temperature-sensitive mutations, indications of pheno-type, etc., should follow the allele number [e.g.,araA230(Am) hisD2J(Ts)] and should be defined at thefirst occurrence. If superscripts must be used, theymust be approved by the editor and they must bedefined at the first occurrence.

(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 operonscan be shown as 4'(ara-lac)95. Similarly, FD(araB'-lacZ+)96 indicates that the fusion results in a truncatedaraB gene fused to an intact lacZ, and 'F(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 fQ(Okb::K-12hisB)4. An alternative designation of an insertioncan be used in simple cases, e.g., galT236::TnS. Thenumber 236 refers to the locus of the insertion, and ifthe strain carries an additional gal mutation, it is listedseparately. Additional examples, which utilize aslightly 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 adirect 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 (or

.ii

INSTRUCTIONS TO AUTHORS

brackets) are used (X, 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 defined in Materials and Methods or in atable of strains. For more detailed information aboutthe symbols in current use, consult Bachmann(Microbiol. Rev. 47:180-230, 1983) for E. coli K-12,Sanderson and Roth (Microbiol. Rev. 47:410-453,1983) 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, and theHandbook ofGenetics, vol. 1 (R. C. King, ed., PlenumPublishing Corp., 1974) 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 microorganisms. In mostinstances, phages have no phenotype, since they haveno metabolism outside host cells. Therefore, distinc-tions between phenotype and genotype are not made.Superscripts are employed to indicate hybridgenomes. Genetic symbols may be one, two, or threeletters. For example, a mutant strain of X might bedesignated as Aamll int2 redll4 c1857; this straincarries mutations in genes cI, int, and red and anamber-suppressible (am) mutation in gene A. A straindesignated K att434 imm21 would represent a hybrid ofphage k which carries the immunity region (imm) ofphage 21 and the attachment (att) region of phage 434.Host DNA insertions into phages should be delineatedby square brackets, and the genetic symbols anddesignations for such inserted DNA should conform tothose employed for the host genome. Genetic symbolsfor phage K can be found 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::Tn5, 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, assignsTn and IS numbers to avoid conflicting and repetitiveuse.

ABBREVIATIONS AND CONVENTIONS

Verb TenseUse the past tense to narrate particular events in the

past, including the procedures, observations, and dataof the study you are reporting. Use the present tensefor general statements, including your own generalconclusions, the conclusions of previous researchers,and generally accepted facts. In addition, the presenttense should be used for discourse having an immedi-ate effect on the reader ("the data indicate"; "Fig. 1shows").Abbreviations

General. It is strongly recommended that all abbre-viations except those listed below be introduced in thefirst paragraph in Materials and Methods. Alterna-tively, define each abbreviation and introduce it inparentheses the first time it is used; e.g., "cultureswere grown in Eagle minimal essential medium(MEM)." Generally, eliminate abbreviations that arenot used at least five times in the text (including tablesand figure legends). Abbreviations should be usedprimarily as an aid to the reader, rather than as aconvenience for the author, and therefore their useshould be limited. Abbreviations other than thoserecommended by the IUPAC-IUB (Biochemical No-menclature and Related Documents, 1978) should beused only when a case can be made for necessity, suchas 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.

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);

. .i.

INSTRUCTIONS TO AUTHORS

cDNA (complementary DNA); RNA (ribonucleicacid);cRNA (complementary RNA); RNase (ribonucle-ase); DNase (deoxyribonuclease); rRNA (ribosomalRNA); mRNA (messenger RNA); tRNA (transferRNA); AMP, ADP, ATP, dAMP, GTP, etc. (for therespective 5' phosphates of adenosine or other nucleo-sides) (add 2'-, 3'-, or 5'- when needed for contrast);ATPase, dGTPase, etc., (adenosine triphosphatase,deoxyguanosine triphosphatase, etc.); NAD (nicotin-amide adenine dinucleotide); NADI+ (nicotinamideadenine dinucleotide, oxidized); NADH (nicotinamideadenine dinucleotide, reduced); NADP (nicotinamideadenine dinucleotide phosphate); NADPH(nicotinamide adenine dinucleotide phosphate, re-duced)'; poly(A), poly(dT), etc. (polyadenylic acid,polydeoxythymidylic acid, etc.); oligo(dT), etc.(oligodeoxythymidylic acid, etc.); Pi (orthophosph-ate); PPi (pyrophosphate); UV (ultraviolet); PFU(plaque-forming units); CFU (colony-forming units);MIC (rninimal inhibitory -concentration); MBC (mini-mal 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 cells) 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)ht (height)mo (month)mol wt (molecular weight)no. (number)prepn (preparation)SD (standard deviation)

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)

103. Avoid compound prefixes such as m,u or jiR. Usejig/ml or ,ug/g in place of the ambiguous ppm. Units oftemperature are presented as follows: 3'7C 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 .g or 10min. For example, "pmol/min" would be preferable to"nmol/10 min," and "j.mol/g" would be preferable to"nmol/,g."

It is also preferable that an unambiguous form suchas the exponential notation be used instead of multipleslashes; for example, ",umol g min'-" is preferableto ",umol/g per min."

See the CBE Style Manual, 5th edition for moredetailed information about reporting numbers. Alsocontained in this source is information on SI units tobe used 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, isotopic labeling is indicated

in the chemical formula (e.g., 14C02, 3H20, H235S04).Brackets are not employed when the isotopic symbolis attached to a word that is not a specific chemicalname (e.g., 1311-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]ureaL-[methyl-14C]methionine[2,3-3H]serine[a-'4C]lysine

[y-32P]ATPUDP-[U-14C]glucoseE. coli [32P]DNAfructose 1,6-[1-32P]bisphosphate

Reporting Numerical DataStandard metric units are used for reporting length,

weight, and volume. For these units and for molarity,use the prefixes m, ,u, n, and p for i0-, 10-6, 1o-9,and 1012, respectively. Likewise, use the prefix k for

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

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