JOURNAL OF BACTERIOLOGY VOLUME 164 o DECEMBER 1985 * NUMBER 12 Simon Silver, Editor in Chief (1987) Robert H. Rownd, Editor (1990) Washington University June J. Lascelles, Editor (1989) Northwestern Medical School St. Louis, Mo. University of California, Los Angeles Chicago, Ill. James D. Friesen, Editor (1987) Richard M. Losick, Editor (1988) Graham C. Walker, Editor (1990) University of Toronto Harvard University Massachusetts Institute of Toronto, Canada Cambridge, Mass. Technology Stanley C. Holt, Editor (1987) L. Nicholas Ornston, Editor (1987) Cambridge, Mass. The University of Texas Health Yale University Robert A. Weisberg, Editor (1990) Science Center, San Antonio New Haven, Conn. National Institute of Child Samuel Kaplan, Editor (1988) Health and Human University of Illinois, Urbana Development Bethesda, Md. EDITORIAL BOARD David Apirion (1985) A. Eisenstark (1985) K. W. Knox (1985) Palmer Rogers (1987) Arthur I. Aronson (1985) Wolfgang Epstein (1987) Wil N. Konings (1987) Barry P. Rosen (1986) Stuart J. Austin (1987) David Figurski (1987) Jordan Konisky (1987) Harry Rosenberg (1,985) Frederick M. Ausubel (1986) Timothy J. Foster (1986) Dennis J. Kopecko (1987) Rudiger Schmitt (1986) Gad Avigad (1986) John D. Foulds (1985) Terry Krulwich (1987) June R. Scott (1987) Barbara Bachmann (1987) Masamitsu Futai (1985) Thomas G. Lessie (1985) Jane K. Setlow (1987) Manfred E. Bayer (1985) Ann K. Ganesan (1985) Lasse Lindahl (1987) Peter Setlow (1987) Margaret H. Bayer (1986) Robert Gennis (1985) Jack London. (1987) Lucille Shapiro (1986) Claire M. Berg (1986) Jane Gibson (1985) Paul S. Lovett (1987) Issar Smith (1987) Robert-W. Bernlohr (1985) E. Peter Greenberg (1985) Ben J. J. Lugtenberg (1986) Paul F. Smith (1985) Terry J. Beveridge (1985) Barry G. Hall (1985) Robert Macnab (1985) Brian G. Spratt (1985) Edwin Boatman (1986) R. E. W. Hancock (1987) Francis L. Macrina (1986) Catherine Squires (1987) I. D. J. Burdett (1985) Robert Haselkorn (1987) Ann Matthysse (1985) I. W. Sutherland (1985) Joseph- M. Calvo. (1987) Gerald L. Hazelbauer (1987) Sarah A. Mclntire (1986) Robert Switzer (1987) A. M. Chakrabarty (1986) Charles E. Helmstetter (1985) Larry McKay (1987) Alexander Tomasz (1986) Keith F. Chater (1985) Ulf Henning (1985) S. Mizushima (1985) James L. Van Etten (1985) Terence G. Cooper (1987) Charles L. Hershberger (1987) Edward A. Morgan (1987) Godfried D. Vogels (1987) John E. Cronan, Jr. (1986) Penny J. Hitchcock (1987) David Mount (1985) Barry Wanner (1987) Walter B. Dempsey (1986) Bruce Holloway (1985) R. G. E. Murray (1986) Bernard Weisblum (1986) Patrick Dennis (1985) Martha M. Howe (1987) Robert A. Niederman (1985) David Womble (1986) W. D. Donachie (1985) Karin Ihlet (1987) Patrick J. Piggot (1987) Henry C. Wu (1987) W. Ford Doolittle (1985) A. W. B. Johnston (1986) Barry Polisky (1987). Duane C. Yoch (1986) J. Allan Downie (1985) Robert J. Kadner (1985) Linda Randall (1987) Howard Zalkin (1985) Gerhart Drews (1985) Eva R. Kashket (1987) William S. Reznikoff (1985) David A. Dubnau (1986) Edward Kellenberger (1985) Marilyn Roberts (1986) Helen R. Whiteley, Chairman, Publications Board Linda 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., NW Washington, DC 20006, is devoted to the advancement and dissemination offundamental 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. Jouirnal is published monthly, and the twelve, numbers are divided into four volumes per year. The nonmember subscription price is $249 per year; single copies are $21. The member subscription price is $37 (foreign, $49 [surface rate]) per year; single copies are $7. 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., NW, Washington, DC 20006 (phone: 202 833-9680). Claims fo'r 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 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 © 1985, American Society for Microbiology. El fi'etM1;A A,J(L I Jrtf 1,; 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. Author Index Ainsworth, Teresa, 1233 Garciarrubio, Alejandro, 1032 Maillet, Fabienne, 1200 Sato, Gihei, 977 Atlan, Daniele, 1376 Garnerone, Anne-Marie, 1200 Malamy, Michael H., 1248 Sawers, R. Gary, 1324 Gibson, Janet L., 1188 Mandecki, Wlodek, 1353 Schell, Maria G., 1301 Ballantine, Stuart P., 1324 Gilchrist, Carol A., 1004 Manson, Michael D., 1057 Shah, Vinod K., 1081 Barros, M. E. C., 1386 Gokul, Nanda K., 1049 Marrero, Robert, 1283 Shapiro, James A., 1171 Barroso, G., 1094 Goldman, Robert A., 1353 Maurer, Kick, 1064 Shen, Gwo-Jenn, 1153 Bassett, Carole L., 1362 Gots, Joseph S., 1288 Maurizi, Michael R., 1124 Silver, Simon, 983 Bastarrachea, Fernando, 1032 Gottesman, Susan, 1124 McGroarty, Estelle J., 1256 Singer, M. E., 1011, 1017 Batut, Jacques, 1200 Grisolia, Vincenza, 1317 Mendelson, Neil H., 1136, Sirotkin, Karl M., 1301 Berg, Claire M., 1350 Gunge, Norio, 1373 1141 Sjoberg, Britt-Marie, 1194 Bhandari, Basant, 1383 Gunsalus, Robert P., 1100 Mieschendahl, Martin, 1366 Sjogren, Agneta, 1278 Bialkowska-Hobrzanska, Misra, Tapan K., 983 Smith, Linda Tombras, 1088 Hanna, 1004 Haapasalo, Markus, 1278 Molin, S0ren, 1262 Snyder, Irvin S., 1071 Bjork, Glenn R., 1117 Hahne, Solveig, 1194 MouchEs, C., 1094 S0gaard-Andersen, Liselotte, Bohach, Gregory A., 1071 Haldenwang, William G., 1356 Muller-Hill, Benno, 1366 1262 Booth, Ian R., 1218, 1224 Halverson, Larry J., 1301 Song, Seung-dal, 1271 Borgia, Peter T., 1039, 1049 Hancock, Robert E. W., 1256 Nappo, Anna Guilia, 1317 Spitzer, Eric D., 994 Bove, J. M., 1094 Harms, Nellie, 1064 Natvig, Donald O., 1309 Stacey, Gary, 1301 Boxer, David H., 1324 Hartmann, Anton, 1271 Nelson, Kristin K., 1301 Stouthamer, Adrian H., 1064 Brill, Winston J., 1081 Higgins, Christopher F., 1218, Nicholas, D. J. D., 1383 Stuber, Dietrich, 1117 Bruni, Carmelo Bruno, 1317 1224 Sugisaki, Yuji, 1373 Bulthuis, Roel, 1294 Homma, Michio, 1370 Opgenorth, A., 1238 Burris, Robert H., 1271 Hovmoller, Sven, 1278 Tabita, F. Robert, 1188 Hyun, H. H., 1146, 1153, 1162 Paranchych, W., 1238 Takemura, Tomoaki, 1182 Cairney, John, 1218, 1224 Perombelon, Michel C. M., Tally, Francis P., 1248 Candresse, T., 1094 Idota, Yuji, 1182 1110 Tamura, Gakuzo, 1373 Carlomagno, Maria Stella, Ihara, Hayato, 1182 Peterson, Arnold A., 1256 Terzaghi, Betty, 1200 1317 lino, Tetsuo, 1370 Peterson, Bryan C., 1359 Thwaites, J. J., 1136 Caruthers, Marvin H., 1353 Imperial, Juan, 1081 Phillips, Gregory J., 1039, Trempy, Janine E., 1356 Chatterjee, Anadi N., 1337 Iritani, Sumiko, 1182 1049 Trisler, Patsy, 1124 Chu, Lien, 983 Ishiguro, Edward E., 1233, Phipps, B. M., 1332 Truchet, Georges, 1200 Covarrubias, Alejandra A., 1332 Platz, Anton, 1194 Trust, Trevor J., 1233, 1332 1032 Ishiguro, Naotaka, 977 Portalier, Raymond C., 1376 Tsukagoshi, Norihiro, 1182 Postma, Erik, 1294 Dahl, Michael K., 1057 Jones, Helen M., 1100 Potts, Malcolm, 1025 Udaka, Shigezo, 1182 Debelle, Frdddric, 1200 Powell, Bradford S., 1353 Ugalde, Rodolfo A., 1081 Denarid, Jean, 1200 Karamata, D., 1141 Denhardt, David T., 1004 Karlsson, Margareta, 1194 Ranta, Helena, 1278 Vasse, Jacques, 1200 de Vries, Gert E., 1064 Kay, William W., 1233, 1332 Ranta, Kari, 1278 Veltkamp, Eduard, 1064 de Vrij, Wim, 1294 Kessler, Andrew I., 1288 RayChaudhuri, Debabrata, Verma, Meera, 1381 Doker, Renate, 1211 Koch, H. Uwe, 1211 1337 Kogoma, Tokio, 1309 Rawlings, D. E., 1386 Wang, Ming-der, 1350 Egan, J. Barry, 1381 Robillard, Norman J., 1248 Weiss, Bernard, 994 Eriksson, Staffan, 1194 Konings, Wil N., 1294 Rokeach, Luis A., 1262 Kopec, Leslie K., 1283 Romero, David, 1032 Whalen, William A., 1350 Farr, Spencer Bryant, 1309 Kushner, Sidney R., 1362 Rosenberg, Charles, 1200 Woods, D. R., 1386 Farrants, George, 1278 Rownd, Robert H., 1359 Wroblewski, H., 1094 Favre, Didier, 1136, 1141 Laddaga, Richard, 983 Russell, Paul, 1301 Finlay, B. B., 1238 Lazzaroni, Jean Claude, 1376 Yamagata, Hideo, 1182 Finnerty, W. R., 1011, 1017 Lee, J. S., 1238 Saillard, C., 1094 Yamasaki, Makari, 1373 Fischer, Werner, 1211 Le6n, Patricia, 1032 Sakaguchi, Kenji, 1373 Yasbin, Ronald E., 1283 Forbes, Kenneth J., 1110 Lindstrom, Par H. R., 1117 Sasaki, Takuji, 1182 Frost, L. S., 1238 Lounatmaa, Kari, 1278 Sasatsu, Masansori, 983 Zeikus, J. G., 1146, 1153, 1162 ACKNOWLEDGMENT The following have served as invited special reviewers for the Journal during 1985, and their help is greatly appreciated. Mark Achtman H. D. Caldwell Henry L. Ehrlich M. Carolyn Hardegree Lloyd M. Kozloff Sankar Adhya David H. Calhoun S. Dusko Ehrlich Simon S. J. Hardy V. Krishnapillai Nina Agabian Ercole Canale-Parola Eric Eisenstadt A. R. Harker A. Kroger Mary M. Allen Frank Cannon Barry I. Eisenstein Franklin M. Harold Herbert E.