EDITORIAL BOARD L. LEON CAMPBELL, Editor-in-Chief University ofIllinois, Urbana L. S. BARON, Editor (1969) L. JOE BERRY, Editor (1968) Ifalter Reed Army Institute ofResearch, Washington, D.C. Bryn Mawr College, Bryn Mawr, Pa. R. D. DeMOSS, Editor (1970) University ofIllinois, Urbana

R. L. Anderson (1969) Herman Friedman (1969) Robert C. Parlett (1969) Robert Austian (1969) Lorraine Friedman (1968) Harry D. Peck (1970) W. Lane Barksdale (1969) Ruth E. Gordon (1969) Michael J. Pelczar, Jr. (1969) S. S. Barkulis (1969) Joseph S. Gots (1969) H. J. Phaff (1969) Alan W. Berheimer (1969) Leonard Hayflick (1969) Roy Repaske (1969) Robert W. Bernlohr (1970) James G. Hirsch (1969) H. V. Rickenberg (1970) S. Gaylen Bradley (1969) John G. Holt (1969) C. F. Robinow (1969) Arnold F. Brodie (1970) Paul Howard-Flanders (1969) W. R. Romig (1969) Allan Campbell (1969) Jerard Hurwitz (1969) NM R. J. Salton (1969) George B. Chapman (1969) Arthur G. Johnson (1969) Russell W. Schaedler (1969) V. P. Cirillo (1970) Elliot Juni (1967) W. R. Sistrom (1969) A. J. Clark (1969) R. E. Kallio (1969) Louis DS. Smith (1968) V. W. Cochrane (1969) Carl Lamanna (1970) Paul F. Smith (1970) S. F. Conti (1969) Maurice Landy (1970) Roger Y. Stanier (1970) C. D. Cox (1969) H. S. Levinson (1969) M. P. Starr (1970) William J. Cromartie (1969) H. C. Lichstein (1970) B. A. D. Stocker (1969) Roy Curtiss m (1969) W. R. Lockhart (1969) Bernard S. Strauss (1969) A. L. Demain (1969) Donald G. Lundgren (1968) Curtis B. Thorne (1969) Eliznbe McCoy (1969) John A. DeMoss (1969) H. E. Umbarger (1970) George B. Mackaness (1969) Michael Doudoroff (1969) Lewis W. Wannamaker (1969) Stanley Marcus (1970) Martin Dworkin (1970) Arthur Weissbach (1969) Ellis Englesberg (1969) Julius Marmur (1968) A. G. Marr (1969) R. W. Wheat (1969) Frank B. Engley, Jr. (1969) M. L. Morse (1969) Helen R. Whiteley (1969) Stanley Falkow (1969) Frederick C. Neidhardt (1969) J. B. Wilson (1969) William W. Ferguson (1969) Evelyn L. Ogiuky (1969) Willis A. Wood (1969) Leonard Frank (1970) Kenaeth Paigen (1969) Barbara E. Wright (1968) Martin Freundlich (1970) James T. Park (1969) Frank E. Young (1970)

ROBERT A. DAY, Managing Editor, 4715 Cordell Ave., Bethesda, Md.

EX OFFICIO S. E. LURIA, President (1967-1968) DENNIS W. WATSON, Vice-President (1967-1968) Massachusetts Institute ofTechnology, Cambridge University ofMinnesota, Minneapolis DONALD E. SHAY, Secretary H. B. WOODRUFF, Treasurer University ofMaryland, Baltimore Merck & Co., Inc., Rahway, N.J.

Published monthly by the American Society for Microbiology. Manufactured and distributed by Waverly Press, Inc., and The Williams & Wilkins Co., at 428 E. Preston St., Baltimore, Md. 21202, U.S.A. Subscription price $30.00 per year, single issue $4.00 Second class postage paid at Baltimore, Md. Made in the of America AUTHOR INDEX Baker, Bruce, 388 Ide, Misao, 317 0ro, J., 333, 344, 349 Barker, L. F., 403 Bennett, E. 0., 344 Jackson, M., 323 Paigen, Kenneth, 493 Bonventre, Peter F., 306 Johnson, Charles E., 306 Patt, J. K., 403 Bowers, W. D., Jr. 482 Johnson, Emmett J., 409 Brenner, Don J., 486 Razin, Shmuel, 359 Klainer, Albert S., 378, 383 Remsen, Charles C., 422 Chai Ning-Chun, 415 Klein, Harold P., 475 Rogoff, M. H., 485 Clowes R. C. 365 Rottem, Shlomo, 359 Cohen-Bazire,Germaine, 458 Lark, Karl G., 415 Curtiss, Roy, mIf, 490 Levinson, Hillel S., 434, 441 Schlesinger, Sondra, 327 Lindorfer, Robert K., 300 Smith, K. ., 300 Deibel, R. H., 291 297 London, Jack, 458 Stallions, D. R., 490 Demain A. L., 323 Domer, Judith E., 466 McClary, D. 0., 482 Thayer, Donald W. 488 MacFarren, Adelaide C., 365 Tornabene, T. G. 333, 344, 349 Freer, John H., 441 McGinnis, James, 493 Trenner, N. R., 323 Freifelder, David, 396 Maheswaran, S. K., 300 Mallette, M. 283 Gelpi, E., 333 Markovitz, Alvin,F.1 388 Valois Frederica W., 422 Goodman N. S., 485 Martin, Malcolm A., 486 Volkmann, Carol, 475 Gorbach Sherwood, 378, 383 Mayeux, Jerry V., 409 Gottfriec, R. J., 485 Watson, Stanley W., 422 Nowlan, Sandra Simpson, 291, 297 Weibel, Joseph, 475 Hamilton, James G. 466 Weinstein Louis, 378 383 Harkin, James C., 4&6 Ogg, James E., 488 Williams, Beverly, 49i Holmes P. K., 434 Ogur M., 482 Hoyer, bill H., 486 Okabayashi, Tadashi, 317 Yoshimoto, Akihiro, 317 JOURNAL OF BACTERIOLOGY 5 SUSTAINING MEMBERS

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The Journal of Bacteriology a publication of the American Society for Microbiology, is devoted to the advancement and dissemination of fundamental knowledge concerning bacteria and other microorganisms. The Journal is published monthly and the twelve numbers are divided into two volumes (January-June and July-December). The subscription price is $30.00 per year, in- cluding Bacteriological Reviews. Single copies are $4.00. Members of the American Society for Mi- crobiology may receive the Journal as part oftheir dues.

INSTRUCTIONS TO AUTHORS Correspondence. Submit manuscripts in duplicate Figures. A complete set offigures, preferably glossy (original and one carbon) to Robert A. Day, Manag- photographs, should accompany each of the two ing Editor, American Society for Microbiology, 4715 copies of the manuscript. Graphs (submit as photo- Cordell Ave., Bethesda, Md. 20014. graphs) should be finished drawings, not needing General policy. Any manuscript submitted must be further artwork or type-setting. Absolutely no part of a report of unpublished original research, which is not a graph should be typewritten (except the legend, being considered for publication elsewhere. A manu- which should be typed on a separate page). AU letter- script accepted and published by the Journal must not ing should be done with a lettering set. 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When first low the recommendations of Demerec et aL (Genetics used in a manuscript or abstract, the name ofa species 54:61, 1966). The Journal reserves the privilege of should be written in full (with the generic name editing manuscripts to make them conform with the capitalized). After first use, the generic name may be adopted style. (Somewhat more complete "Instructions to Authors" are printed in the January and July issues abbreviated by use of its initial capital letter. In of the Journal.) general, the nomenclature for bacteria presented in Form oj manuscript. Manuscripts should be typed Bergey's Manual of Determinative Bacteriology (7th on bond paper (8% by 11 inches). All parts ofthe man- ed., 1957) is used. 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