JOURNAL OF VIROLOGY

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JOURNAL OF VIROLOGYVOLUME 280NUMBER 3*DECEMBER 1978EDITORIAL BOARDRobert R. Wagner, Editor-in-Chief (1982)University of Virginia School of Medicine,CharlottesvilleLennart Phiipson, Editor (1979)Biomedical CenterUppsala, SwedenDwight L. Anderson, Editor (1983)School of Dentistry,University of Minnesota,MinneapolisEdward M. Scolnick, Editor (1982)National Cancer InstituteBethesda, Md.David T. Denhardt, Editor (1982)McGill UniversityMontreal, Que., CanadaJohn P. Bader (1979)David Baltimore (1978)Amiya K. Banerjee (1979)David H. L. Bishop (1979)David Botstein (1979)Michael A. Bratt (1980)Dennis T. Brown (1978)Purnell Choppin (1980)John M. Coffin (1980)Richard W. Compans (1979)Nicholas R. Cozzarelli (1980)Walter Doerfler (1980)Elvera Ehrenfeld (1980)Suzanne U. Emerson (1980)Erwin J. Fleissner (1980)Robert M. Friedman (1979)Harold S. Ginsberg (1979)Angus Graham (1978)Hidesaburo Hanafusa (1980)John J. Holland (1978)Nancy Hopkins (1980)Calderon Howe (1979)Alice S. Huang (1978)Paul J. Kaesberg (1979)Joseph Kates (1979)D. C. Kelly (1979)George Khoury (1978)David W. Kingsbury (1979)Lloyd M. Kozloff (1979)Robert M. Krug (1980)Myron Levine (1979)David M. Livingston (1980)Karl Lonberg-Hoim (1978)Ronald B. Luftig (1978)Brian McAuslan (1979)Malcoim A. Martin (1978)Christopher K. Mathews (1979)Robert C. Miller (1980)Gisela Mosig (1979)Bernard Moss (1980)Joseph S. Pagano (1979)J. Thomas Parsons (1980)Ulf G. Petterson (1980)Peter G. W. Plagemann (1980)Robert E. Pollack (1978)Craig R. Pringle (1980)Fred Rapp (1978)Dan S. Ray (1980)Bernard E. Reilly (1980)William S. Robinson (1980)Bernard Roizman (1979)Roland R. Rueckert (1979)Norman P. Salzman (1978)Joseph Sambrook (1979)Sondra Schlesinger (1980)Phillip A. Sharp (1979)Aaron J. Shatkin (1979)Lee D. Simon (1978)Kai Simons (1978)Alan E. Smith (1980)Patricia G. Spear (1978)Donald F. Summers (1979)William C. Summers (1980)Natalie M. Teich (1979)Howard M. Temin (1979)David C. Ward (1980)Robin Weiss (1979)John S. Wiberg (1979)Ernest Winocour (1979)Julius Youngner (1979)Robert A. Day, Managing EditorGisella Pollock, Assistant Managing EditorSusan Birch, Production Editor1913 I St., N. W., Washington, D.C. 20006EX OFFICIOEdwin H. Lennette, President (1978-1979)J. Mehsen Joseph, SecretaryThe Journal of Virology, a publication of the American Societyfor Microbiology, 1913 I St., N.W., Washington, D.C. 20006, isdevoted to the dissemination of fundamental knowledge concerning viruses of bacteria, plants, and animals. Investigators areinvited to submit reports of original research in all areas of basicvirology, including biochemistry, biophysics, genetics, immunology, morphology, and physiology. Instructions to Authors arepublished in the January issue each year. The Journal is issuedmonthly, four volumes per year. The nonmember subscriptionprice is 90 per year; single copies are 8. The member subscription price is 21 per year. Correspondence relating to subscriptions, reprints, defective copies, availability of back issues, lostor late proofs, disposition of submitted manuscripts, and generaleditorial matters should be directed to the ASM PublicationsOffice, 1913 I St., N.W., Washington, DC 20006 (area 202 8339680).Second-class postage paid at Washington, DC 20006, and atWillis A. Wood, President-Elect (1978-1979)Brinton M. Miller, Treasureradditional mailing offices.Made in the U.S.A.Copyright i 1978, American Society for Microbiology.All Rights Reserved.The code at the top of the first page of an article in this journalindicates the copyright owner's consent that copies of the articlemay be made for personal use, or for personal use of specificclients. 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, New York 12301,for copying beyond that permitted by Sections 107 or 108 of theU.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 IndexAbramczuk, Jan, 936Acheson, Nicholas H., 885Ackley, C. M., 997Arlinghaus, Ralph B., 929Bauman, Viestur, 717, 725Beemon, Karen, 957Bossart, Walter, 905Brennan, Victoria, 1002Brown, Neal C., 865Bruenn, Jeremy A., 1002Butel, Janet S., 743Calegari, L., 786Casella, Edith, 753Chabot, Francis, 972Chakravorty, Maharani, 736Champion, Margaret, 972Cheevers, W. P., 997Chen, Teh-Chung, 679Chinnadurai, G., 1011Chu, Herman, 877Coffin, John M., 972Crawford, T. B., 997Dasgupta, Biplab, 736Debbia, E., 772, 786Decl6ve, A., 819Dishler, Andris, 717, 725Dosmar, Michael, 753Drillien, R., 843Dudley, Jaquelin P., 743Dunker, Reinhard, 865Edens, Luppo, 835Fan, Hung, 802Fontana, R., 772Ford, John P., 795Fujinaga, K., 1011Gallo, Robert C., 686Green, M., 1011Gren, Elmar J., 717, 725Guntaka, Ramareddy V., 810,1015Hassell, John A., 671, 945Hastie, Nicholas D., 1002Hitzeman, Ronald A., 697Hsu, Ming-Ta, 795Hsu, Teresa W., 810, 1015Hunter, Tony, 957Ito, Junetsu, 895Jaenisch, Rudolf, 802Kalyanaraman, V. S., 686Kaplan, H. S., 819Kawakami, Thomas G., 767Kirn, A., 843Klessig, Daniel F., 945Konings, Ruud N. H., 835Lieberman, M., 819Lubeck, Michael D., 710McCrae, M. A., 828McGrath, M. S., 819McIntosh, Phyllis K., 865MacIsaac, Paul, 802Mark, George E., 810Markewych, Orest, 753Matoba, Sam I., 767Maul, Gerd G., 936Mellor, Andrew, 992Palese, Peter, 710Paoletti, Enzo, 905Pennington, T. H., 828Pichler, Mary E., 917Price, Alan R., 697Pruzzo, C., 772, 786Puimpen, Paul, 717, 725Ray, Dan S., 679Rho, M. H., 1011Rose, Jeffrey M., 743Rovera, Giovanni, 936Sabran, Jan L., 810Sarngadharan, M. G., 686Satta, G., 772, 786Schoenmakers, John G. G., 835Schulman, Jerome L., 710Sefton, Bartholomew M., 957Serwer, Philip, 917Smith, Alan E., 992Socher, Susan H., 743Spehner, D., 843Sun, Lily, 767Taylor, John M., 810, 1015Tershak, Daniel R., 1006Truitt, Christine L., 877van der Eb, A. J., 1011Vorbrodt, Andrzej, 936Walker, James R., 877Mi6ville, Frangoise, 885Mulder, Carel, 865Murphy, Edwin C., Jr., 929Weber, Joseph, 671Weissman, I. L., 819Witmer, Heman, 753Wu, Ray, 851Nuss, Donald L., 905Yang, Robert C. A., 851

ACKNOWLEDGMENTThe following have servedhelp is greatly appreciated.Stuart A. AaronsonAlice Adams-LindahlSankar AdhyaNina AgabianYosef AloniChristian AnfinsenG. AppleyardR. ArlinghausJ. Thomas AugustAndrew L. BallM. A. BaludaY. BarenholzRobert BassinNara BattulaThomas BenjaminRolf BenzingerPaul BergArnold BerkK. L. BernsMichael BishopPaul BlackRobert BlumenthalDani BolognesiD. BotsteinDavid BradleyGaylen S. BradleyJ. C. BrownAhmad Bukhari IClarence S. BullerB. W. BurgeJanet ButelR. CallahanAllan M. CampbellRobert CardiffEster ChangR. M. ChanockCharles ColeRoger ColeGeoff CooperL. CrawfordR. L. CrowellRichard P. CunninghamJames E. DahlbergSamuel DalesEllen DaniellNorman DavidsonV. DefendiE. DeHarvenRavi DharClive DicksonT. 0. DienerJohn E. DowdingE. J. DuboviPeter H. DuesbergJohn J. DunnWalter EckhartRobert Eisenmanasinvited special reviewers for the Journal during 1978, and theirLynn W. EnquistK. R. EriksenR. L. EriksonHung FanAnthony FarasBernard M. FieldsG. R. FinkPeter FischingerJane FlintDana FowlkesHeinz Fraenkel-ConratRichard M. FranklinM. FriedRobert C. GalloA. T. GanesanMartin GellertRaymond GildenMarc GirardNigel GodsonRobert GoldbergR. N. GourlayAllan GranoffMacDonald GreenMaurice GreenTerry GrodzickerWalter R. GuildGary N. GussinGordon HagarBerge HamparK. A. HarrapEchols HarrisonDonald HarterJanet HartleyWilliam HaseltineAshley HasseM. HatanakaMasaki HayashiWilliam HaywardRoger HendrixKathleen HerculesBernard HirtJohn A. HolowczakJames HopperJerand HorwitzMarshall S. HorwitzA. F. HowatsonMartha HowePeter HowleyJoel HubermanKlaus HummlerMargaret HuntRichard ImblumJunetsu ItoWarren JelinekLeevi K1ariiainenRobert KamenYong C. KangAlbert S. KaplanThomas Kelly, Jr.Ian KerrEli KeshetElliott D. KieffErwin D. KilbourneSaul KitNancy KlecknerEva KleinDaniel KolakofskyHilary KoprowskiStuart KornfeldMichael KuehlEdward KuffC. G. KurlandElizabeth KutterMichael LaiOtto E. LandmanRobert LazzarmniSeymour LederbergJonathan LeisPeter LengyetRichard E. LernerJay LevyAndrew LewisJim LewisF. LillyThomas LindahlJean LindenmannAlan LissRoyce Lockart, Jr.Harvey LodishDouglas LowyPhilip McAllisterWilliam McAllisterBrian McAuslanJames McCorquodaleJ. McDougallJames McSharryGoran MagnussonJacob MaizelManley MandelJack ManiloffDante MarcianiPaul A. MarksRobert G. MartinVivien MauntnerD. H. MetzG. MillerLois K. MillerStewart MillwardSankar MitraDan H. MooreN. F. MooreRichard MoyerHellos MurialdoFrederick Murphy

Kenneth MurrayDaniel NathansPaul E. NiemanEdward NilesErling NorrbyRobert NowinskiBo ObergJohn ObijeskiPeter PaleseTakis PapasWade P. ParksSarah A. ParsonsIra PastanEric PatzerMark L. PearsonHhkan PerssonPer PetersonRalph PettersonElmer PfeffercornLudvik PrevecAlan R. PriceA. M. PrinceCarol PrivesCharles RandallWilliam RawlsM. E. ReichmannAlan ReinDavid RekoshMichel RevelSolon RhodeMarc RhodesDaniel RifkinGordon RingoldJames A. RobbJeffrey W. RobertsRichard RobertsJ. A. RoseJohn K. RoseJeffrey RossEllen RothenbergSandra RuscettiPaul SadowskiKaoru SaigoMargarita SalasLois A. SalzmanPriscilla SchafferAndreas ScheidMilton SchlesingerRobert SchloemerJeffrey SchlomCarl SchnaitmanC. ScholtissekJune ScottDouglas ScrabaArup SenJane SetlowD. B. ShahAaron J. ShatkinJoel SheffieldPeter SheldrickTom ShenkFred ShermanThomas ShihDavid ShubSaul SilversteinRobert W. SimpsonAnna SkalkaJ. J. SkehelPeter SnudstadJohn SpencerFrank StahlJoan SteitzJohn R. StephensonVictor StollarNat L. SternbergWilliam Stewart IIJames H. StraussWilliam StudierJ. H. Subak-SharpeM. D. SummersJohn SussenbachJ. F. SzilagyiKenneth TakemotoIgor TammValerie TateJohn TaylorPeter TegtmeyerRay TennantEthel TessmanIrwin TessmanRobert ThackClark TibbettsT. W. TinsleyGeorge TodaroDavid TroxlerPeter van der VlietGeorge VandewoudeH. E. VarmusJan VilcekPeter VogtHuber WarnerKlaus WeberPhil WeckRoger WeilRobert WeinbergHarold WeintraubArthur WeissbachSherman WeissmanGail WertzHeiner WestphalHelen R. WhiteleyReed WicknerJohn S. WilbergN. M. WilkeJ. WilliamsGary A. WilsonHerbert H. WinklerErnst WinnackerOwen WitteNelson WivelWilliam WoldKeith YamamotoClifford YehleHoward A. YoungJames F. ZisslerHarald zur HausenHans Zweerink

1979 Full Membership American Society for Microbiology1913 I St., NW, Washington, DC 20006The Society welcomes to full membership any person who is interested in its objects, and who holds a bachelor'sdegree in microbiology or a related field (or who has had training and experience equivalent to that represented by abachelor's degree).The minimum annual membership assessment is 28 which includes a 3 membership fee, 4 for subscription toASM News, and 21 toward subscriptions to the Society's scientific journals. The Society publishes eight scientificjournals (listed below). Members may subscribe to one or more of these journals at special member rates; 21 of theminimum dues payment of 28 may be credited towards journal subscriptions. In addition, the Society publishes themonthly ASM News, which is sent to all members and contains news and announcements of interest to all microbiologists.Memberships are initiated and renewed as of 1 January each year. Unless there are directions to the contrary, membership nominations received prior to 1 November will be credited to the current year, and back issues of the selectedpublications for the current year will be furnished if available. Nominations received after 1 November will becomeeffective on the following 1 January.First NameInitialLast NameAddressArea CodeCityHighest DegreeZip CodeStateYearMajorPhone No.Yr. BirthSexGranting InstitutionPositionExperienceSignature of NomineeNominated by(Signature of Society member)I. 1. Membership Fee ( 3.00) and ASM News ( 4.00) . 7.002. Additional journals may be subscribed to at the following rates. Please enter thehigher rate (column B) if you reside outside the U.S. and its possessions.ADfBEl 30 19Journal of Bacteriology (JB) . 1. 22Antimicrobial Agents and Chemotherapy (AAC) . 2. 16Applied and Environmental Microbiology (AEM) . 3. 17 22International Journal of Systematic Bacteriology (IJSB) . 4. 15 15Journal of Virology (JVI) . 5. 21 26Journal of Clinical Microbiology (JCM) . 6. 16 19Microbiological Reviews (MR) . 7. 9 10Infection and Immunity (IAI) . 9. 21 263. Total dues and journals from lines 1 and 2 (MINIMUM PAYMENT 28). .-.II D Check this block if you prefer to receive only ASM News and remit 28.RATES ARE FOR 1979 ONLYPlease enclose payment with application.

1979 Student Membership American Society for Microbiology1913 I St., NW, Washington, DC 20006Any regularly matriculated student major in microbiology or a related field is eligible for election as a Student Member.Student Members have all privileges of membership except the right to vote and hold office in the Society.Student Members receive the monthly ASM News and are entitled to subscribe to the Society's journals at memberrates.Memberships are initiated and nenewed as of 1 January each year. Unless there are directions to the contrary, membershipnominations received prior to 1 November will be credited to the current year, and back issues of the selected publicationsfor the current year will be furnished if available. Nominations received after 1 November will become effective on thefollowing 1 January.NAkMEADDFRESSIIL L-1 L.l I %JLIIJ.jZip CodeStateCityPhone No.Yr. of BirthHighest DegreeSexSchoolMajor field of studySignature of NomineeSignature of chairman of major dept.**Nominated by:(1)(Society members)(2)If the department chairman is a member of the Society, only one additional nominating signature is required.1. 1979 Dues (includes ASM News) . 2.502. Additional journals may be subscribed to at the following rates. Please enter thehigher rate (column B) if you reside outside the U.S. and its possessions.Journal of Bacteriology (JB).Antimicrobial Agents and Chemotherapy (AAC).Applied and Environmental Microbiology (AEM).International Journal of Systematic Bacteriology (IJSB).Journal of Virology (JVI).Journal of Clinical Microbiology (JCM).Microbiological Reviews (MR).Infection and Immunity (IAI).3. TOTAL and Remit.RATES ARE FOR 1979 ONLYPlease enclose payment with application.1.2.3.4.5.6.7.9.A 22 16 17 15 21 16 9 21 B 30 19 22 15 26 19 10 26

STATEMENT OF OWNERSHIP, MANAGEMENT AND CIRCULATION,-i.o,.1 OIWc-oeo-C-1LJournal of Virolor.o.0Ie* ,E ''9'Voooontill1y121913NS. .'shinyton.1 St.SN. W-hinato.1913 T St.-DCDC20006S CO.-L.Tt -DD-E t or.O.j*S9(20006 .lLHE. G.IX ED--OAaAoric Societs for icrobioloxy. 1913 I St. NW, R9hington, DC 20006R. Waer, Uis.itS of Virginii Chrlottesoille, VA 22901R 6bert!.oo DA.A pa, 1913 I St., Ni. s.s.ihi.to, DC 20006Ari.D Society foricr-biolob y1913 I St., IH,Wsigston.DC20006i-.L .O-T OF GO.M. .ORTG"gg 0. O"NAt 61--- --PNone1.0. .0-LEll- 01 '. .O- -'I .'Z-OM .I.O.M. ". -1 .11-T.,--o cow-6-sc., .'as A- 22. 122. m.,swwlesoof R 6301usTgA." lLU5556ttot w lo wc9LTlol --o-&lo255566286555i58063013030tO tC * T§S DI -0,11I1101)iCGK li[t fl55860'5610* g, i sz1ET-.AOO- gtW-zC iOIOfF@CZ,tOt" VY COU-tCO O@ t-X* ET9t* A T715676'ZE.4NONlE 62866 6301 XZAwIendythmtT u * m- e. .b5.bn, -tGt taiom-,-- - d 'p trAMI.- 1.1121. I." S-o'.'.Fi3526 tPslI(SJe-ttU t or, r

AUTHOR INDEXVOLUME 28Abramczuk, Jan, 936Acheson, Nicholas H., 885Ackley, C. M., 997Acton, Ronald T., 84Allen, George P., 361Arlinghaus, Ralph B., 929Ball, L. Andrew, 324Bassin, Robert H., 656Bates, Robert C., 20Bauman, Viestur, 717, 725Beemon, Karen, 551, 957Belfort, Marlene, 270Berg, Paul, 475Bishop, David H. L., 417Bishop, J. Michael, 600Black, Lindsay W., 643Blair, Phyllis B., 611Blanton, R. A., 450Bond, Sheila B., 357Bone, Donald R., 14Boseley, Paul G., 171Bossart, Walter, 905Brahic, Michel, 543Brennan, Victoria, 1002Brown, Moira, 14Brown, Neal C., 865Bruenn, Jeremy A., 1002Bryant, Martin L., 300Butel, Janet S., 743Calegari, L., 786Carter, T. H., 450Casella, Edith, 753Cash, P., 199Chabot, Francis, 972Chakravorty, Maharani, 736Champion, Margaret, 972Cheevers, W. P., 997Chen, Teh-Chung, 679Chenciner, Nicole, 95Chien, Yueh-hsiu, 352Chinnadurai, G., 1011Chiswell, D. J., 199Chiu, Nancy H., 590Chu, Herman, 877Coffin, John M., 972Collins, Peter L., 324Compans, Richard W., 84Cooper, Geoffrey M., 45Cox, Donald C., 444Crawford, T. B., 997Crombie, Ian, 14Cuzin, Frangois, 1, 421Dasgupta, Biplab, 736Davidson, Norman, 352Davis, Alan R., 375Davison, A. J., 499De, Barun K., 375Dean, Donald H., 403Debbia, E., 772, 786Decleve, A., 819Delagneau, Jean F., 518DePamphilis, Melvin L., 53Devare, Sushilkumar G., 457Diacumakos, Elaine G., 415Dishler, Andris, 717, 725Dittmar, Kurt J., 106Dosmar, Michael, 753Drillien, R., 843Dubois-Dalcq, Monique, 567Dudley, Jaquelin P., 743Duncan, Michael R., 444Dunker, Reinhard, 865Edens, Luppo, 835Edlund, Louise E., 84Ehrenfeld, Ellie, 387Fan, Hung, 802Fanshier, Lois, 600Fenger, Terry W., 292Filippi, Pierre, 543Flamand, Anne, 518Fontana, R., 772Ford, John P., 795Francke, Bertold, 14Friis, Robert R., 434Fujinaga, K., 1011Gallo, Robert C., 686Garro, Anthony J., 395Gentry, Glenn A., 361Gentsch, Jon R., 417Gershey, Edward L., 415Gibson, Wade, 240Given, Douglass, 524Grandgenett, Duane P., 279Green, M., 1011Greene, Maurice, 314Gren, Elmar J., 717, 725Groman, Neal, 28Guntaka, Ramareddy V., 810, 1015Guttman, Burton S., 262Harrison, S. C., 578Harvey, Robert, 154Hassell, John A., 671, 945Hastie, Nicholas D., 1002Hastings, Kenneth E. M., 490Hightower, Lawrence E., 324Hitzeman, Ronald A., 697Hooghe-Peters, Elisabeth, 567Howe, Calderon, 292Howley, Peter M., 337

iiAUTHOR INDEXHsu, Ming-Ta, 795Hsu, Teresa W., 810, 1015Huang, Eng-Shang, 661Hunter, Tony, 240, 551, 957Igarashi, K., 212, 219Ito, Junetsu, 895Jaenisch, Rudolf, 802Janda, Michael J., 375Jeng, Yun-hua, 314Jense, Holly, 387Jensen, Kaj Frank, 427Jones, Elaine V., 61Jones, K., 411Kalyanaraman, V. S., 686Kang, C. Yong, 34Kaplan, H. S., 819Kawakami, Thomas G., 767Kemp, Maurice C., 84Kieff, Elliott, 524Kierszenbaum, A. L., 661Kingsbury, D. W., 411Kirn, A., 843Klessig, Daniel F., 945Knauert, Fred, 387Kolakofsky, Daniel, 584Konings, Ruud N. H., 835Krug, Robert M., 75Kuff, Edward L., 66Kurokawa, T., 212, 219Kutter, Elizabeth M., 262J. VIROL.Murphy, Edwin C., Jr., 929Nayak, Debi P., 375Niiyama, Y., 219Nuss, Donald L., 905Oberg, Bo, 119Okenquist, Sharon, 45Palese, Peter, 710Panet, Amos, 434Paoletti, Enzo, 905Patton, John T., 20Paucha, Eva, 140, 154Pearson, Gary R., 344Pennington, T. H., 828Perbal, Bernard, 1Perkins, John B., 403Persson, Hakan, 119Philipson, Lennart, 119Pichler, Mary E., 917Plotch, Stephen J., 75Pollock, Thomas J., 408Preble, Olivia T., 6Preston, V. G., 499, 624Price, Alan R., 697Pridgen, C., 411Pringle, C. R., 199Pritchard, Cynthia, 20Pruzzo, C., 772, 786Pumpen, Paul, 717, 725Putzrath, Resha M., 254Qualtiere, Louis F., 344Lane, Mary-Ann, 611Law, Ming Fan, 395Lieberman, M., 819Lubeck, Michael D., 710Lueders, Kira K., 66Lynch, Sue, 584McCrae, M. A., 828McGowan, John J., 361McGrath, M. S., 819McIntosh, Phyllis K., 865MacIsaac, Paul, 802McQuain, C. O., 368Mak, S., 227Malloy, P., 199Maniloff, Jack, 2

JOURNAL OF VIROLOGY VOLUME 28 0 NUMBER 3 * DECEMBER 1978 EDITORIAL BOARD Robert R. Wagner,Editor-in-Chief (1982) University ofVirginia SchoolofMedicine, Charlottesville Dwight L. Anderson, Editor (1983) SchoolofDentistry, University ofMinnesota, Minneapolis DavidT. Denhardt,Editor (1982) McGill University Montreal, Que., Canada Lennart Phiipson .

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