APPLIED AND ENVIRONMENTALMICROBIOLOGYVOLUME 43* NUMBER 1 0 JANUARY 1982

EDITORIAL BOARDJames M. Tiedje, Editor-in-Chief (1985)Michigan State University, East Lansing

Robert T. Belly, Editor (1984)Eastman Kodak Company

Rochester, New York

A. L. Demain, Editor (1982)Massachusetts Institute of Technology, Cambridge

Martin S. Favero, Editor (1985)Centers for Disease Control,

Phoenix, Arizona

Robert B. Hespeil, Editor (1985)University of Illinois, UrbanaJohn J. Iandolo, Editor (1986)

Kansas State UniversityManhattan, Kansas

Ronald Atlas (1983)Richard H. Baltz (1984)Richard Bartha (1982)Barry L. Batzing (1983)Larry W. Belser (1983)Joan W. Bennett (1984)Merlin Bergdoll (1984)Jean-Marc BolUag (1983)Charles Boylen (1982)John A. Breznak (1983)Lee A. Bulia, Jr. (1983)Victor Cabelli (1982)Douglas E. Caldwell (1983)Paul E. Came (1982)Tom D. Y. Chin (1983)Richard T. J. Clarke (1984)Michael A. Cole (1982)Richard A. Consigli (1982)Ronald L. Crawford (1983)Frank Dazzo (1982)Burk A. Dehority (1983)Hugh W. Ducklow (1983)Richard Elander (1982)Douglas Eveleigh (1982)Samuel R. Farrah (1983)William R. Finnerty (1983)Heinz G. Floss (1983)Dennis Focht (1982)Edwin E. Geldreich (1982)Charles Gerba (1982)

Richard E. Goldstrand (1982)C. P. Leslie Grady, Jr. (1982)Charles Hagedorn m (1982)George Hegeman (1983)Bruce Hamilton (1984)John C. Hoff (1982)John Johnson (1982)David M. Karl (1982)Edward Katz (1982)Roger Knowles (1982)Linda L. Lasure (1983)Paul Lemke (1982)Eivind B. Lillehoj (1984)Carol Litchfield (1983)Robert J. Maier (1984)Allen J. Markovetz (1983)Scott E. Martin (1984)Prakash Masurekar (1982)MoUlie McBride (1984)Gordon A. McFeters (1984)Larry L. McKay (1983)Terry L. Miller (1982)Aaron L. Mills (1984)Thomas Montvllle (1983)Richard Morita (1982)Claude H. Nash H (1984)Kenneth W. Nickerson (1984)Betty H. Olson (1982)Ronald Oremland (1982)Hans W. Paerl (1984)

Michael W. Pariza (1984)Frederick C. Pearson (1982)Hap Pritchard (1982)Donald J. Reasoner (1982)C. A. Reddy (1982)Douglas Ribbons (1982)Donald C. Robertson (1984)Antonio H. Romano (1983)John P. Rosazza (1982)Abigail A. Salyers (1983)Dwayne Savage (1982)Robert D. Schwartz (1982)Oldrich K. Sebek (1983)Surendra N. Sehgal (1983)Jim Spain (1984)David C. Stemnberg (1983)Hiroshi Suglyama (1984)Anne 0. Summers (1982)Jon H. Tuttle (1983)Richard Unz (1984)Claude Vezina (1982)Edward Voss (1984)David M. Ward (1983)David White (1982)R. P. Williams (1984)Meyer J. Wolin (1982)Richard T. Wright (1983)William Yotis (1982)Stanley A. Zahler (1982)Alexander Zehnder (1982)

Helen R. Whiteley, Chairman, Publications BoardWalter G. Peter mI, Director, PublicationsLinda M. Illig, Managing Editor, Journals

Perry Turner, Production Editor

Applied and Environmental Microbiology (ISSN 0099-2240), a publication of the American Society for Microbiology, 1913 ISt., NW, Washington, DC 20006, is devoted to the advancement and dissemination of applied knowledge as well as ecologicalknowledge, both applied and fundamental, concerning microorganisms. Instructions to authors are published in the January issueeach year; reprints are available from the editors and the Publications Office. Applied and Environmental Microbiology ispublished monthly, two volumes per year. The nonmember subscription price is $120 per year; single copies are $12. Themember subscription price is $25 (foreign, $30 [surface rate]) per year; single copies are $6. Correspondence relating tosubscriptions, nonreceipt of journals, reprints, defective copies, availability of back issues, and lost or late proofs, should bedirected to the ASM Publications Office, 1913 I St., NW, Washington, DC 20006 (area 202-833-9680).

Second-class postage paid at Washington, DC 20006, and at additional mailing offices.POSTMASTER: Send address changes to Applied and Environmental Microbiology, 1913 I St., NW,Washington, DC 20006.Made in the United States of AmericaCopyright 0 1982, American Society for Microbiology. El * : IIt?u Ai aE Lf (; I it.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 articlemay be made for personal use or for personal use of specific clients. This consent is given on the condition, however, that thecopier pay the stated per-copy fee through the Copyright Clearance Center, Inc., P.O. Box 765, Schenectady, NY 12301, forcopying beyond that permitted by Sections 107 and 108 of the U.S. Copyright Law. This consent does not extend to other kindsof copying, such as copying for general distribution, for advertising or promotional purposes, for creating new collective works,or for resale.

Author IndexAlvarez, Maria E., 237Aranha, H., 257Arceneaux, J. E. L., 257

Baross, John A., 39Barraquio, W. L., 124Barrion, M., 124Bausum, Howard T., 28Behbehani, M. J., 255Bender, G. L., 233Bland, Patricia T., 218Bolen, Joseph B., 193Boone, David R., 57Bottazzi, Vittorio, 50Brinkley, Allen W., 86Brown, David A., 65Bryant, M. P., 70Byers, B. R., 257

Cabelli, Victor J., 90Cameron, Susan C., 97Carman, George M., 81Cavari, Benzion, ICen, Y., 233Consigli, Richard A., 193Conway de Macario, Everly, 227Cooney, Charles L., 75Cornell, John H., 144Cousminer, J. Jeffrey, 81Cuhel, Russell L., 151, 160

Dahle, Arne B., 169de Guzman, M. R., 124

England, Albert C., III, 240Eyssen, Hendrik J., 185

Federle, Thomas W., 129Fischer, J. R., 136, 247Fraser, David W., 240

Garfinkle, Jill A., 193Genthner, B. R. Sharak, 70Gorman, George W., 240Gottesman, Andrew R., 86Gresshoff, P. M., 233

Hayashi, Kazuya, 245Hoff, John C., 250Huijghebaert, Suzanne M., 185

Iannotti, E. L., 136, 247

Jannasch, Holger W., 151, 160Jordan, H. V., 255

Kaplan, Arthur M., 144Kaplan, David L., 144Keleti, Georg, 104Keller, Frederick A., Jr., 117Kenyon, Kathryn F., 28Kirchman, David, 200

Laake, Morten, 169LaCroix, Steven J., 90LeChevallier, Mark W., 97Lodge, J. S., 257

Macario, Alberto J. L., 227Mackel, Don C., 240Mallison, George F., 240Markwell, Deborah D., 110Mazza, G., 177McFeters, Gordon A., 97McKinley, Vicky L., 129Mertens, Jan A., 185Miller, Terry L., 227Mitchell, Ralph, 200Mizunuma, Takeji, 245Morelli, Lorenzo, 50Morita, Richard Y., 39

Morrison, N. A., 233Mott, Glen E., 86

Noda, Fumio, 245

O'Brien, R. T., 237

Paerl, Hans W., 210, 218Pedersen, Karsten, 6Peters, Susan, 39

Rice, Eugene W., 250Rolfe, B. G., 233

Santoro, D. L., 255Schaefer, Eugene J., 75Schaefer, Frank W., III, 250Schaub, Stephen A., 28Schiemann, D. A., 14Schwartz, Robert D., 117Scott, K. F., 233Shine, J., 233Shortridge, Kennedy F., 110Sievers, D. M., 136, 247Skaliy, Peter, 240Small, Mitchell J., 28Strachan, R. C., 257Sykora, Jan L., 104

Taylor, Craig D., 151, 160Toerien, Danie F., ITrinick, M. J., 233

Vescovo, Marisa, 50Vestal, J. Robie, 129

Watanabe, I., 124Wei, Cheng-I, 252Wolin, M. J., 227

Zaniewski, Richard L., 81

APPLIED AND ENVIRONMENTAL MICROBIOLOGY, Jan. 1982

APPLIED AND ENVIRONMENTAL MICROBIOLOGY

INSTRUCTIONS TO AUTHORS

HOW TO SUBMIT MANUSCRIPTSSubmit manuscripts directly to the ASM Pub-

lications Office, 1913 I St., NW, Washington,DC 20006. The manuscript should be accompa-nied by a covering letter stating the following:the journal to which the manuscript is beingsubmitted, the most appropriate section of thejournal, the address and telephone number ofthe corresponding author, and the former manu-script number (if it is being resubmitted).Submit two complete copies of each manu-

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usage should have their manuscripts checked bysomeone proficient in the English language.Manuscripts may be rejected on the basis ofpoor English or lack of conformity to acceptedstandards of style.

EDITORIAL POLICYManuscripts submitted to Applied and Envi-

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CopyrightTo maintain and protect the Society's owner-

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applied research as well as both applied andbasic ecological research on bacteria and othermicroorganisms, including fungi, protozoa, andother simple eucaryotic organisms. Topics thatare considered include microbiology in relation

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If you have questions about these guidelines,please contact the editor-in-chief of the journalyou are considering.Note that a manuscript rejected by one ASM

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i

INSTRUCTIONS TO AUTHORS

(4th ed. 1978; CBE Secretariat, 9650 RockvillePike, Bethesda, Md.), Robert A. Day's How toWrite and Publish a Scientific Paper (ISI Press,1979), and Scientific Writing for Graduate Stu-dents (CBE Secretariat), as interpreted andmodified by the editors and the ASM Publica-tions Office. The editors and the PublicationsOffice reserve the privilege of editing manu-scripts to conform with the stylistic conventionsset forth in the aforesaid publications and inthese instructions.

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Regular PapersRegular full-length papers should include the

elements described in this section.

Title. Each manuscript should present theresults of an independent, cohesive study; thus,numbered series titles are not permitted. Exer-cise care in composing a main title. Avoid themain title/subtitle arrangement. On the titlepage, include: title, running title (not to exceed46 characters and spaces), full name (includingfirst name and middle initial) of each author, ad-dress(es) of the institution(s) at which the workwas performed, and each author's affiliation or afootnote indicating the present address of anyauthor no longer at the institution where thework was performed. Place an asterisk after thename of the author to whom inquiries regardingthe paper should be directed, and give thatauthor's telephone number.

Abstract. Limit the abstract to 250 words orfewer. Summarize the basic content of the pa-per. Avoid abbreviations, diagrams, and refer-ences. When it is essential to include a refer-ence, use the full literature citation but omit thearticle title. Because the abstract will be pub-lished separately by abstracting services, it mustbe complete and understandable without refer-ence to the text.

Introduction. The introduction should supplysufficient background information to allow thereader to understand and evaluate the results ofthe present study without needing to refer toprevious publications on the topic. The intro-duction should also provide the rationale for thepresent study. Use only those references re-quired to provide the most salient backgroundrather than an exhaustive review of the topic.

Materials and Methods. The methods sectionshould include sufficient technical informationso that the experiments can be repeated. Forcommonly used materials and methods (e.g.,commonly used media, protein determinations),

..

INSTRUCTIONS TO AUTHORS

a simple reference is sufficient. If several alter-native methodologies are commonly employed,it is useful to identify the method briefly as wellas to cite the reference. For example, it ispreferable to state, "cells were broken by ultra-sonic treatment as previously described (9),"rather than to state, "cells were broken aspreviously described (9)." You should allow thereader to assess the methodology without con-stant reference to previous publications. De-scribe new methods completely, and givesources of unusual chemicals, equipment, ormicrobial strains. When large numbers of micro-bial strains or mutants are used in a study,include strain tables identifying the sources andproperties of the strains, mutants, bacterio-phages, plasmids, etc.A method, strain, etc., used in only one of

several experiments reported in the papershould be described in the Results section or, ifbrief enough, may be included in a table footnoteor figure legend.

Results. In the Results section, include onlythe results of the experiments; reserve extensiveinterpretation of the results for the Discussionsection. Present the results as concisely as pos-sible in one of the following: text, table(s), orfigure(s). However, avoid extensive use ofgraphs to present data that might be more con-cisely presented in the text or tables. For exam-ple, except in unusual cases, double-reciprocalplots used to determine apparent Km valuesshould not be presented as graphs; instead, thevalues should be stated in the text. Similarly,graphs illustrating other methods commonlyused to derive kinetic or physical constants(e.g., reduced viscosity plots, plots used todetermine sedimentation velocity) need not beshown except in unusual circumstances. Limitphotographs (particularly photomicrographs andelectron micrographs) to those that are absolute-ly necessary to demonstrate the experimentalfindings. Number figures and tables in the orderin which they are cited in the text, and be sure tocite all figures and tables.

Discussion. The Discussion should provide aninterpretation of the results in relation to previ-ously published work and to the experimentalsystem at hand and should not contain extensiverepetition of the Results section or reiteration ofthe introduction. In short papers, the Resultsand Discussion sections may be combined.

Acknowledgments. Acknowledgments for fi-nancial assistance and for personal assistanceare given in two separate paragraphs. The usualformat for acknowledgment of grant support isas follows: "This work was supported by Public

Health Service grant CA-01234 from the Nation-al Cancer Institute."

Appendixes. Appendixes, which contain sup-plementary material to aid the reader, are per-mitted. Titles, authors, and Literature Citedsections that are distinct from those of theprimary article are not allowed. If it is notfeasible to list the author(s) of the appendix inthe by-line or the Acknowledgment section ofthe primary article, rewrite the appendix so thatit can be considered for publication as an inde-pendent article, either full length or Note style.

Literature Cited. Arrange the Literature Citedsection in alphabetical order, by first author, andnumber consecutively. (Abbreviate journalnames according to the Bibliographic Guide forEditors & Authors, American Chemical Society,1974, or Serial Sources for the BIOSIS DataBase, BioSciences Information Service, 1981.)Cite each listed reference by number in the text.The following types of references are not valid

for listing: unpublished data, personal communi-cations, manuscripts in preparation, manu-scripts submitted, "in press" references,pamphlets, abstracts, patents, theses, disserta-tions, and any other material that has not beensubjected to peer review. References to suchsources should be made parenthetically in thetext. An "in press" reference to an ASM publi-cation should state the control number (e.g.,AEM 576) or the name of the publication, if it isa book.Follow the styles shown in the examples be-

low.1. Armstrong, J. E., and J. A. Calder. 1978. Inhibi-

tion of light-induced pH increase and 02 evolutionof marine microalgae by water-soluble componentsof crude and refined oils. Appl. Environ. Micro-biol. 35:858-862.

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

3. Finegold, S. M., W. E. Shepherd, and E. H.Spaulding. 1977. Cumitech 5, Practical anaerobicbacteriology. Coordinating ed., W. E. Shepherd.American Society for Microbiology, Washington,D.C.

4. Gill, T. J., III. 1976. Principles of radio-immunoassay, p. 169-171. In N. R. Rose and H.Friedman (ed.), Manual of clinical immunology.American Society for Microbiology, Washington,D.C.

5. Leadbetter, E. R. 1974. Order II. Cytophagalesnomen novum, p. 99. In R. E. Buchanan and N. E.Gibbons (ed.), Bergey's manual of determinativebacteriology, 8th ed. The Williams & Wilkins Co.,Baltimore.

. .

INSTRUCTIONS TO AUTHORS

6. Sacks, L. E. 1972. Influence of intra- and extra-cellular cations on the germination of bacterialspores, p. 437-442. In H. 0. Halvorson, R. Han-son, and L. L. Campbell (ed.), Spores V. AmericanSociety for Microbiology, Washington, D.C.

Parenthetical references in the text should becited as follows:... and protects the organisms against oxygentoxicity (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 andC. W. Slayman, Abstr. Annu. Meet. Am. Soc.Microbiol. 1976, K15, p. 139).

NotesSubmit Notes in the same way as full-length

papers. They receive the same review, and theyare not published more rapidly than full-lengthpapers nor are they considered preliminary com-munications. The Note format is intended forthe presentation of brief observations that do notwarrant full-length papers.Each Note must have an abstract of no more

than 50 words. Do not use section headings inthe body of the Note; report methods, results,and discussion in a single section. The text is notto exceed 1,000 words, and the number of fig-ures and tables should be kept to a minimum.Present acknowledgments as in full-length pa-pers, but do not use a heading. The LiteratureCited section is identical to that of full-lengthpapers.

ILLUSTRATIONS AND TABLES

PhotographsWhen submitting electron micrographs, pho-

tographs of polyacrylamide gels, etc., keep inmind the journal page size: 6.5 cm for a singlecolumn and 14 cm for a double column (maxi-mum). Include only the significant portion of theillustration. Each must be of sufficient contrastto withstand the inevitable loss of contrast anddetail inherent in the printing process. Electronand light micrographs must be first-generatior.copies of the original negatives. Indicate with ascale marker on the electron micrograph themagnification of each photomicrograph. Do notmount figures on cardboard. Composite figuresmay be mounted on bond paper. A complete setof photographs, not photocopies, must accom-pany each copy of the manuscript.Color photographs are discouraged. Howev-

er, if they are necessary, include three copies sothat a cost estimate for printing may be ob-tained. The cost of printing color photographsmust be borne by the author.

DrawingsSubmit graphs, charts, diagrams, and other

drawings as photographs made from finisheddrawings not requiring additional artwork ortypesetting. No part of the graph or drawingshould be typewritten. Use a lettering set orother professional-quality device for all labeling.Most graphs will be reduced to one-columnwidth (6.5 cm), and all elements in the drawingshould be prepared to withstand this reduction.Avoid very heavy letters, which tend to close upwhen reduced, and unusual symbols, which theprinter may not be able to reproduce in thelegend. Symbols and lettering should be of ap-propriate size; do not use large letters and smallsymbols or vice versa. Direct readouts fromcomputers, recorders, etc., are not usually ac-ceptable; such materials should be redrawn.

In figure ordinate and abscissa scales (as wellas table column headings), avoid ambiguous useof numbers with exponents. Usually, it prefera-ble to use the International System of Units (p,for 10-6, m for 10-3, k for 103, M for 106, etc.). Acomplete listing of SI symbols can be found inthe International Union of Pure and AppliedChemistry (IUPAC) "Manual of Symbols andTerminology for Physicochemical Quantitiesand Units" (Pure Appl. Chem. 21:3-44, 1970).Thus, a representation of 20,000 dpm on a figureordinate is to be made by the number 20 accom-panied by the label kdpm.When powers of 10 must be employed, the

editorial style of the journal follows the CBEStyle Manual recommendation, which differsfrom the conventions used by several otherjournals. The CBE Style Manual suggests thatthe exponent power be associated with the num-ber shown. In representing 2 x 107 cells per ml,the correct designation would be 2, labeled as107 cells per ml, not cells per ml x 10-7.Likewise, an enzyme activity of 0.06 U/mlwould be shown as 6, accompanied by the label10-2 U/ml. The preferred designation would be60 mU/ml (milliunits per milliliter).

Figure LegendsFigure legends may be placed beneath the

photocopy of a drawing for the convenience ofreviewers. (In addition, however, a complete setof photographs or drawings, with legends onseparate pages, must accompany each copy ofthe manuscript.) Legends should provideenough information so that the figure is under-standable without frequent reference to the text.

IV

INSTRUCTIONS TO AUTHORS

However, do not repeat experimental methodsin the legend. Define all symbols and abbrevia-tions used in the figure. Common abbreviationsand others used frequently in preceding textneed not be redefined in the legend.

Tables

Type each table on a separate page. Arrangethe data so that columns of like material readdown, not across. The headings should be suffi-ciently clear so that the meaning of the data willbe understandable without reference to the text.See "Abbreviations" in these instructions forthose that should be used in tables. Explanatoryfootnotes are acceptable, but more extensivetable "legends" are not. Footnotes should notinclude detailed descriptions of the experiment.A well-constructed table is shown below.

TABLE 1. Distribution of protein and ATPase infractions of dialyzed membranesa

ATPaseMembranes Fraction

from U/mg of Total Uprotein

Control Depleted 0.036 2.3membrane

Concentrated 0.134 4.82supernatant

El treated Depleted 0.034 1.98membrane

Concentrated 0.11 4.6supernatant

a Specific activities of ATPase of nondepleted mem-branes from control and treated bacteria were 0.21 and0.20, respectively.

Camera-Ready Copy

Drawings, tables, chemical formulas, etc.,that can be photographically reproduced forpublication without further typesetting orartwork are referred to as "camera ready."Such copy may also be prepared for complicatedmathematical or physical formulas, portions ofgenetic maps, diagrams, and flow schemes. Itshould not be hand lettered. Camera-ready copymust be carefully prepared to conform with thestyle of AEM. The advantage to submitting cam-era copy is that the material will appear exactlyas envisioned by the author, and no secondproofreading is necessary. This is particularlyadvantageois when there are long, complicatedtables and when the division of material andspace are important.

NOMENCLATURE

Chemical and Biochemical NomenclatureThe recognized authority for the names of

chemical compounds is Chemical Abstracts(Chemical Abstracts Service, Ohio State Uni-versity, Columbus) and its indexes. For bio-chemical terminology, including abbreviationsand symbols, consult the following: Internation-al Union of Biochemistry Biochemical Nomen-clature and Related Documents (1978; reprintedfor The Biochemical Society, London, En-gland), instructions to authors of the Journal ofBiological Chemistry and Archives ofBiochem-istry and Biophysics (first issues of each year),and the Handbook ofBiochemistry and Molecu-lar Biology (G. D. Fasman, ed., 3rd ed., 1976,CRC Press, Inc.).Do not express molecular weights in daltons;

molecular weight is a unitless ratio. Molecularmass is expressed in daltons.For enzymes, use the recommended (trivial)

name assigned by the Nomenclature Committeeof the International Union of Biochemistry asdescribed in Enzyme Nomenclature 1978 (Aca-demic Press, Inc., 1979). If a nonrecommendedname is used, place the proper (trivial) name inparentheses at first use in the abstract and text.Use the EC number when it has been assigned,and express enzyme activity either in katals(preferred) or in the older system of ",umol/min."

Nomenclature of MicroorganismsBinary names, consisting of a generic name

and a specific epithet (e.g., Escherichia coli),must be used for all microorganisms. Names ofgenera and higher categories may be used alone,but a specific epithet must be preceded by ageneric name the first time is is used in a paper.Thereafter, the generic name should be abbrevi-ated to the initial capital letter (e.g., E. coli),provided there can be no confusion with othergenera used in the paper. Names of all taxa(phyla [for fungi, divisions], classes, orders,families, genera, species, subspecies) are print-ed in italics; strain designations and numbers arenot.

In general, the nomenclature of bacteriashould follow that presented in Bergey's Manualof Determinative Bacteriology (8th ed., TheWilliams & Wilkins Co., 1974). If you wish tochallenge this nomenclature, you may express ajudgment, but the name given in Bergey's Man-ual should follow in parentheses the first timethe name is used in both the text and theabstract. Only those names which were includedin the Approved Lists of Bacterial Names (Int.J. Syst. Bacteriol. 30:225-420, 1980) and those

v

INSTRUCTIONS TO AUTHORS

which have been validly published in the Inter-national Journal of Systematic Bacteriol-ogy since 1 January 1980 have standing in no-menclature. It is acceptable to use a namewithout standing in nomenclature to facilitatecommunication, but it is recommended that thefirst time such a name is used it should befollowed by a statement indicating its status(e.g., "not on Approved List of BacterialNames").

It is recommended that a strain be depositedin a recognized culture collection when thatstrain is necessary for the description of a new

taxon (see Bacteriological Code, 1975 Revision,American Society for Microbiology, 1975).

Since the classification of fungi is not com-plete, it is the responsibility of the author todetermine the currently accepted binomial for agiven yeast or mold. Some sources for thespelling of these names include The Yeasts (J.Lodder, ed., North-Holland Publishing Co.,1970) and Ainsworth and Bisby's Dictionary ofthe Fungi, Including the Lichens, 6th ed. (Com-monwealth Mycological Institute, Kew, Surrey,England, 1971).Names used for viruses should be those ap-

proved by the International Committee on Tax-onomy of Viruses (ICTV) and published in the3rd Report of the ICTV, "Classification andNomenclature of Viruses" (Intervirology, vol.12, no. 3-5, 1979). If desired, synonyms may beadded parenthetically when the name is firstmentioned. Approved generic (or group) andfamily names may also be used.

Microorganisms, viruses, and plasmids shouldbe given strain designations consisting of lettersand serial numbers. It is generally advisable toinclude a worker's initials or a descriptive sym-bol of locale, laboratory, etc., in the designation.Each new strain, mutant, isolate, or derivativeshould be given a new (serial) designation. Sucha designation should be distinct from those ofthe genotype and phenotype, and genotypic andphenotypic symbols should not be included.A registry of plasmid designations is main-

tained by the Plasmid Reference Center, Depart-ment of Medical Microbiology, Stanford Univer-sity, Stanford, CA 94305.

Genetic NomenclatureBacteria. The genetic properties of bacteria

are described in terms of phenotypes and geno-

types. The phenotypic designation describes theobservable properties of an organism. The geno-

type refers to the genetic constitution of an

organism, usually in reference to some standardwild type. Use the recommendations of De-merec et al. (Genetics 54:61-74, 1966) as guide inemploying these terms.

(i) Phenotypic designations must be em-ployed when mutant loci have not been identi-fied or mapped. Phenotypic designations gener-ally consist of three-letter symbols; these are notitalicized, and the first letter of the symbol iscapitalized. It is preferable to use roman orarabic numerals (instead of letters) to identify aseries of related phenotypes. Thus, a series ofbacteriocin-tolerant mutants might be designat-ed TolI, TolIl, TolIl, etc., or a series of nucleicacid polymerase mutants might be designatedPoll, Pol2, Pol3, etc. Wild-type characteristicscan be designated as Tol+ or Pol+ and, whennecessary for clarity, negative superscripts(Tol- Pol-) can be used to designate mutantcharacteristics. Superscript letters may be usedto further delineate phenotypes (e.g., Strs forstreptomycin sensitivity). Phenotypic designa-tions should be defined.

(ii) Genotypic designations are similarly indi-cated by three-letter locus symbols. In contrastto phenotypic designations, these are lowercaseitalic (e.g., ara his rps). If several loci governrelated functions, these are distinguished byitalicized capital letters following the locus sym-bol (e.g., araA araB araC). Promoter, termina-tor, and operator sites should be indicated asdescribed by Bachmann and Low (Microbiol.Rev. 44:1-56, 1980): e.g., lacZp, lacAT, andlacZo.

(iii) Wild-type alleles are indicated with asuperscript plus (ara+ his'). Where the geno-type of an organism is being specified (e.g., in astrain table), a superscript minus is not used toindicate a mutant locus. Elsewhere, a super-script minus may be used to distinguish betweenthe symbol of a mutant allele and that of agenetic locus. However, this distinction is bestmade in the context, and thus one refers to anara mutant rather than an ara- strain.

(iv) Mutation sites are designated by placingserial isolation numbers (allele numbers) afterthe locus symbol (e.g., araAl araA2). If it is notknown in which of several related loci the muta-tion has occurred, a hyphen is used instead ofthe capital letter (e.g., ara-23). It is essential inpapers reporting the isolation of new mutantsthat allele numbers be given to the mutations.For Escherichia coli, there is a registry of suchnumbers: E. coli Genetic Stock Center, Depart-ment of Human Genetics, Yale UniversitySchool of Medicine, P.O. Box 3333, New Ha-ven, CT 06510. For Salmonella, the registry is:Salmonella Genetic Stock Center, Departmentof Biology, University of Calgary, Calgary, Al-berta, Canada T2N 1N4.

(v) The use of superscripts with genotypes(other than + to indicate wild-type alleles)should be avoided. Designations indicating am-ber mutations, temperature-sensitive mutations,

Vi

INSTRUCTIONS TO AUTHORS

and indications of phenotype should follow theallele number [e.g., araA230(Am) hisD2J(Ts)].

(vi) Deletions are indicated by the symbol Aplaced before the deleted gene or region, e.g.,AtrpA432, A(aroP-aceE)419, or Ahis(dhuA hisJhisQ)1256. Similarly, other symbols can be used(with appropriate definition). Thus, a fusion ofthe ara and lac operons can be shown as 4)(ara-lac)95. Similarly, '?(araB'-lacZ+)96 indicatesthat the fusion results in a truncated araBgene fused to an intact lacZ, and F(malE-lacZ)97(Hyb) shows that a hybrid protein issynthesized. An insertion of an E. coli his geneinto plasmid pSC101 at zero kilobases (0 kb) isshown as pSC101 fQ(Okb::K-12hisB)4. An alter-native designation of an insertion can be used insimple cases, galT236::TnS. The number 236refers to the locus of the insertion and, if thestrain carries an additional gal mutation, it islisted separately. Additional examples, whichutilize a slightly different format, are to be foundin the papers by Campbell et al. and Novick etal., cited below. It is important in reportingconstruction of strains in which a mobile ele-ment was inserted and subsequently deleted thatthis last fact be noted in the strain table. This canbe done by listing the genotype of the strain usedas 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 set-ting of parenthetical remarks within the geno-type or dividing the genotype into constituentelements, parentheses and square brackets areused without special meaning; square bracketsare used outside parentheses. To indicate thepresence of an episome, parentheses (or brack-ets) are used (X, F+). Reference to an integratedepisome is indicated as described above forinserted elements, and an exogenote is shownas, for example, W3110/F'8(gal+).

(vii) Keep in mind the distinction between amutation (an alteration of the primary sequenceof the genetic material) and a mutant (a straincarrying one or more mutations). One mayspeak about the mapping of a mutation, but onecannot map a mutant. Likewise, a mutant has nogenetic locus-only a phenotype.

(viii) Avoid the use of a genotype as a name(e.g., "subsequent use of leuC6 for transduc-tion"). If a strain designation has not beenchosen, select an appropriate word combination(e.g., "either strain PA3092 or another straincontaining the leuC6 mutation").Any deviations from standard genetic nomen-

clature should be defined in Materials and Meth-ods or in a table of strains. For more detailedinformation about genetic maps of locus sym-bols in current use, consult reviews by Bach-mann and Low (Microbiol. Rev. 44:1-56, 1980)for E. coli K-12, Sanderson and Hartman (Mi-

crobiol. Rev. 42:471-519, 1978) for Salmonellatyphimurium, Holloway et al. (Microbiol. Rev.43:73-102, 1979) for Pseudomonas, and Hennerand Hoch (Microbiol. Rev. 44:57-82, 1980) forBacillus subtilis. For yeasts, Chlamydomonas,and several fungal species, symbols such asthose given in Handbook ofMicrobiology (A. I.Laskin and H. A. Lechevalier, ed., CRC Press,Inc., 1974) should be employed.

Viruses. The rules for genetic nomenclature ofviruses (bacteriophages) differ from those formicroorganisms. In most instances, viruses haveno phenotype, since they have no metabolismoutside host cells. Therefore, distinctions be-tween phenotype and genotype cannot be made.Superscripts are employed to indicate hybridgenomes. Genetic symbols may be one, two, orthree letters. For example, a mutant strain of Xmight be designated as X Aamll int2 redl14c1857; this strain carries mutations in 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 Xwhich carries the immunity region (imm) ofphage 21 and the attachment (att) region ofphage 434. Host DNA insertions into virusesshould be delineated by square brackets, and thegenetic symbols and designations for such in-serted DNA should conform to those employedfor the host genome. Genetic symbols for phageA can be found in Szybalski and Szybalski (Gene7:217-270, 1979) and in Echols and Murialdo(Microbiol. Rev. 42:577-591, 1978).

Transposable elements, plasmids, and restric-tion enzymes. Nomenclature of transposable ele-ments (insertion sequences, transposons, phageMu, etc.) should follow the recommendations ofCampbell et al. (Gene 5:197-206, 1979), with themodifications given in section vi. The system ofdesignating transposon insertions at sites wherethere are no known loci, e.g., zef-123::TnS, hasbeen described by Chumley et al. (Genetics91:639-655, 1979). The nomenclature recom-mendations of Novick et al. (Bacteriol. Rev.40:168-189, 1976) for plasmids and plasmid-specified activities, of Low (Bacteriol. Rev.36:587-607, 1972) for F-prime factors, and ofRoberts (Nucleic Acids Res. 9:r75-r96, 1981) forrestriction enzymes and DNA fragments derivedfrom treatment with these enzymes should beused. Recombinant DNA molecules constructedin vitro follow the nomenclature for insertions ingeneral. DNA inserted into recombinant DNAmolecules should be described by using the genesymbols and conventions of the organism fromwhich the DNA was obtained.

.ii

INSTRUCTIONS TO AUTHORS

ABBREVIATIONS AND CONVENTIONS

Verb TenseUse the past tense in referring to results

recorded in the present paper. Use the presenttense in discussing previously established find-ings and generally accepted phenomena.

AbbreviationsIt is strongly recommended that all abbrevia-

tions except those listed below be introduced inthe first paragraph in Materials and Methods.Alternatively, define each abbreviation and in-troduce it in parentheses the first time it is used;e.g., "cultures were grown in Eagle minimalessential medium (MEM)." Generally, eliminateabbreviations that are not used at least five timesin the text (including tables and figure legends).Abbreviations should be used primarily as an aidto the reader rather than as a convenience to theauthor, and therefore their use should be limit-ed. Abbreviations other than those recommend-ed by the IUPAC-IUB (Biochemical Nomencla-ture and Related Documents, 1978) should beused only when a case can be made for necessi-ty, such as in tables and figures.

It is often possible to use pronouns or toparaphrase a long word after its first use (e.g.,"the drug," "the substrate"). Standard chemicalsymbols, numerical multiples (e.g., Me2SO fordimethyl sulfoxide), and trivial names or theirsymbols (folate, Ala, Leu, etc.) may be used forterms that appear in full in the neighboring text.

In addition to abbreviations for standard unitsof measurement and chemical symbols of theelements, the following should be used withoutdefinition in the title, abstract, text, figure leg-ends, and tables: DNA (deoxyribonucleic acid);cDNA (complementary DNA); RNA (ribonucle-ic acid); RNase (ribonuclease); DNase (deoxyri-bonuclease); rRNA (ribosomal RNA); mRNA(messenger RNA); tRNA (transfer RNA); AMP,ADP, ATP, dAMP, GTP, etc. (for the respective5' phosphates of adenosine or other nucleo-sides); 2'-AMP, 3'-AMP, and 5'-AMP (the 2'-,3'-, and 5'-, when needed for contrast, phos-phates of the nucleosides); NAD+ (nicotinamideadenine dinucleotide, oxidized); NADH (nico-tinamide adenine dinucleotide, reduced); NADP(nicotinamide adenine dinucleotide phosphate);NADPH (nicotinamide adenine dinucleotidephosphate, reduced); Pi (orthophosphate); PP,(pyrophosphate); UV (ultraviolet); PFU(plaque-forming units); Tris [tris(hydroxymeth-yl)aminomethane]; DEAE (diethylaminoethyl);and EDTA (ethylenediaminetetraacetate). Ab-breviations for cell lines (e.g., HeLa cells) alsoneed not be defined.The following abbreviations should be used

without definition in tables:

amt (amount)approx (approximately)avg (average)concn (concentration)diam (diameter)expt (experimentht (height)mo (month)mol wt (molecular weight)no. (number)prepn (preparation)SD (standard deviation)

Reporting Numerical Dat

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

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

Standard metric units are used for reportinglength, weight, and volume. For these units andfor molarity, use the prefixes m, ,u, n, and p (for0o-, 10-6, 10-9, and 10-12, respectively). Like-

wise, use the prefix k (for 103). Avoid compoundprefixes, such as m,u or RR,. Parts per million(ppm) may be used when that is the commonmeasure for the science in that field; however, itmust be defined parenthetically at first use (e.g.,,ug/ml). Units of temperature are presented asfollows: 37°C or 324 K.When fractions are used to express such units

as enzymatic activities, it is preferable to usewhole units, such as g or min, in the denomina-tor instead offractional or multiple units, such as,ug or 10 min. For example, "pmol/min" wouldbe preferable to "pmol/10 min," and ",umol/g"would be preferable to "nmol/,g."

It is also preferable that an unambiguousform, such as exponential notation, be used inplace of multiple slashes; for example, ",umolg-1 mmin-" is preferable to ",umol/g per min."

See the CBE Style Manual for more detailedinformation regarding the reporting of numbers.Also contained in this source is information onthe appropriate SI units to be used for reportingillumination, energy, frequency, pressure, andother physical terms. Always report numericaldata in the applicable SI units.

Isotopically Labeled CompoundsFor simple molecules, isotopic labeling is indi-

cated in the chemical formula (e.g., 14CO2,3H235. J235S04). Brackets are not employedwhen the isotopic symbol is attached to a wordthat is not a specific chemical name (e.g., 13li-labeled protein, 14C-amino acids, 3H-ligands,etc.).For specific chemicals, the symbol for the

isotope introduced is placed in brackets directlypreceding the part of the name that describes thelabeled entity. Note that configuration symbolsand modifiers precede the isotopic symbol. Thefollowing examples illustrate correct usage.

. .i.

INSTRUCTIONS TO AUTHORS

[t4C]ureaL-[methyl-'4C]methionine[2,3-3H]serine[a-14C]lysine[X-32P]ATP

UDP-[U-14C]glucoseE. coli [32P]DNAfructose 1,6[1-32P]diphos-

phate

AEM follows the same conventions for isotopiclabeling as the Journal ofBiological Chemistry,and more detailed information can be found inthe instructions to authors of that journal (firstissue of each year).

ix

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