Table Of ContentBacterial Conjugation
Bacterial Conjugation
Edited by
Don B. Clewell
The University of Michigan
Ann Arbor, Michigan
Springer Science+Business Media, LLC
Library of Congress Cataloglng-in-Publication Data
Bacterial conjugation / edited by Don B. Clewell.
p. cm.
Includes bibliographical references and index.
ISBN 978-0-306-44376-3 ISBN 978-1-4757-9357-4 (eBook)
DOI 10.1007/978-1-4757-9357-4
1. Conjugation (Biology) 2. Plasmids. I. Clewell, Don B.
QR86,5.B33 1993
5B9.9'01662--dc20 93-7458
CIP
This limited facsimile edition has been issued
for the purpose of keeping this title available
to the scientific community.
1098765432
ISBN 978-0-306-44376-3
© 1993 Springer Science+Business Media New York
Originally published by Plenum Press, New York in 1993
All rights reserved
No part of this book may be reproduced, stored in a retrieval system, or transmitted
in any form or by any means, electronic, mechanical, photocopying, microfilming,
recording, or otherwise, without written permission from the Publisher
Contributors
Gordon L. Archer • Departments of Microbiology and Immunology and Medicine,
Medical College of Virginia, Virginia Commonwealth University, Richmond, Virginia
23298
Don B. Clewell • Department of Biologic and Materials Sciences, School of Dentistry and
Department of Microbiology and Immunology, School of Medicine, The University of
Michigan, Ann Arbor, Michigan 48109-0402
Walter B. Dempsey • University of Texas Health Science Center and Department of
Veterans Affairs, Veterans Affairs Medical Center, Dallas, Texas 75216
Stephen K. Farrand • Department of Plant Pathology and Microbiology, University of
illinois at Urbana-Champaign, Urbana, illinois 61801
Susan E. Flannagan • Biologic and Materials Sciences, School of Dentistry, The Univer
sityof Michigan, Ann Arbor, Michigan 48109-0402
Laura S. Frost· Department of Microbiology, M330 Biological Sciences Building,
University of Alberta, Edmonton, Alberta T6G 2E9, Canada
Donald G. Guiney • Department of Medicine and Center for Molecular Genetics, Univer
sity of California, San Diego, San Diego, California 92103
Dieter Haas • Mikrobiologisches Institut, Eidgenossische Technische Hochschule,
CH-8092 Zurich, Switzerland
v
vi Contributors
Donald R. Helinski' Department of Biology and Center for Molecular Genetics, Univer
sity of California, San Diego, La Jolla, California 92093-0634
David A. Hopwood • John Innes Institute, John Innes Centre, Norwich NR4 7UH,
England
Karin Ippen-Ihler • Medical Microbiology and Immunology, Health Science Center,
Texas A & M University, College Station, Texas 77843-1114
Clorence I. Kado • Davis Crown Gall Group, Department of Plant Pathology, University
of California, Davis, Davis, California 95616
Tobias Kieser' John Innes Institute, John Innes Centre, Norwich NR4 7UH, England
Barbara Lewis KitteU • Department of Biology and Center for Molecular Genetics,
University of California, San Diego, La Jolla, California 92093-0634
Erich Lanko • Max-Planck-Institut fUr Molekulare Genetik, Abteilung Schuster, D-l000
Berlin 33, Federal Republic of Germany
Francis L. Macrina • Department of Microbiology and Immunology, Medical College of
Virginia, Virginia Commonwealth University, Richmond, Virginia 23298-0678
Cornelia Reimmann • Mikrobiologisches Institut, Eidgenossische Technische Hoch
schule, CH-8092 Zurich, Switzerland
Ronald A. Sku"ay • School of Biological Sciences, University of Sydney, Sydney, New
South Wales 2006, Australia
Brian Wilkins' Department of Genetics, University of Leicester, Leicester LEI 4RH,
United Kingdom
Neil Willetts • Biotech Australia Proprietary, Limited, Roseville, New South Wales 2069,
Australia
Preface
Bacterial plasmids originating in a wide range of genera are being studied from a variety of
perspectives in hundreds of laboratories around the globe. These elements are well known
for carrying "special" genes that confer important survival properties, frequently neces
sary under atypical conditions. Classic examples of plasmid-borne genes are those provid
ing bacterial resistance to toxic substances such as antibiotics, metal ions, and bacte
riophage. Often included are those determining bacteriocins, which may give the bacterium
an advantage in a highly competitive environment. Genes offering metabolic alternatives
to the cell under nutritionally stressed conditions are also commonly found on plasmids,
as are determinants important to colonization and pathogenesis.
It is likely that in many, if not most, cases plasmids and their passenger determinants
represent DNA acquired recently by their bacterial hosts, and it is the characteristic
mobility of these elements that enables their efficient establishment in new bacterial cells
by the process known as conjugation. Whereas many plasmids are fully capable of
promoting their own conjugal transfer, others move only with help from coresident
elements. The ability of a plasmid to establish itself in a variety of different species is com
mon, and recent studies have shown that transfer can in some cases occur from bacterial
cells to eukaryotes such as yeast. Interkingdom transfer in nature is indeed evident from the
findings that certain diseases in plants are directly related to the introduction of bacterial
DNA by a conjugationlike process. It is noteworthy that the genes commonly found on
plasmids often exhibit their own form of mobility by being located on transposons able
to move from one DNA molecule to another intracellularly. Some transposons even have
self-conjugative potential and can transfer directly to another bacterial cell without having
to hitchhike on a plasmid; these are the so-called conjugative transposons.
Considering the general significance of conjugation, from a basic as well as clinical,
environmental, and evolutionary perspective, a volume that brings together a comprehen
sive collection of reviews focusing primarily on this subject would seem timely. It is hoped
vii
viii Preface
that this book will serve as a useful resource for individuals interested in the mysteries
of DNA transfer in the bacterial world.
Don B. Clewell
Ann Arbor, Michigan
Contents
I. Bacterial Conjugation: A Historical Perspective
Neil Willetts
1. Introduction ............................................... I
2. Recognition of F as a Physical and Genetic Entity ............... 2
3. Mutational Studies of Conjugation ............................ 3
4. Complementation Analysis and Mapping Studies. . .. .. . . . . . . . . . . . 3
5. tra Gene Proteins .......................................... 5
6. The Pilus ................................................. 5
7. Conjugational DNA Metabolism .............................. 7
8. Regulation of Conjugation ................................... 10
9. Conjugation: A Common Phenomenon ......................... 12
10. New Techniques for Old... . . . . . . ... ..... .. . . .. .. .. . . . . . . . . . . 14
II. A Personal Odyssey ........................................ 15
12. Conclusion ................................................ 16
References ................................................ 17
2. Genetic Organization of lhmsfer-Related Determinants on the Sex
Factor F and Related Plasmids
Karin Ippen-Ihler and Ronald A. Skurray
I. Introduction ............................................... 23
2. Genetic Characterization of the Transfer Region ................. 26
3. Transfer Gene Functions ..................................... 28
4. Transfer Loci Involved in Gene Regulation and Expression ........ 29
4.1. FinOP Regulation of traJ Expression ..................... 30
4.2. Other Fin Systems .................................... 30
ix
x Contents
4.3. Expression of tra Operon Genes. . . . . . . . . . . .. . . . . .. . . . . . . 31
5. Genes Involved in the Expression of Pilus Filament .............. 31
5.1. The F-Pilin Subunit ................................... 32
5.2. Pilus Assembly Proteins ............................... 34
6. Products Required for Aggregate Stabilization ................... 35
7. Surface Exclusion Genes .................................... 36
7.1. traT ................................................ 37
7.2. traS ................................................ 37
8. Genes Required for DNA Nicking, Displacement, and Transport. . . . 38
8.1. oriT ................................................ 38
8.2. tra! ........................................ . . . . . . . . 39
8.3. traY ................................................ 39
8.4: traM ............................................... 40
8.5. traD . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40
9. Genes of Unknown Function in the Transfer Region .............. 41
10. Genes in the Leading Region of Transfer ....................... 42
11. Conclusions ............................................... 43
References ................................................ 45
3. Key Regulatory Aspects of 1hlnsfer of F-Related Plasmids
Walter B. Dempsey
1. Introduction ................................................ 53
2. Properties of Rl00 .......................................... 54
2.1. Early Studies ......................................... 54
2.2. Statement of Problem .................................. 54
2.3. Proposed Biological Latching Relay ....................... 56
3. Components of the Biological Relay ............................ 57
3.1. RNA Secondary Structures .............................. 57
3.2. Protein Binding in the orzT Region ....................... 58
3.3. traM .............. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 58
3.4. traJ ............................................... . . 59
3.5. traY ................................................. 64
3.6. finP ... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 64
3.7. finO ................................................. 65
3.8. IHF and tra! .......................................... 66
3.9. Evidence that Upstream Transcripts Cross traJ .............. 66
4. Recapitulation or Sources of Speculations ....................... 68
5. Summary .................................................. 68
References ................................................. 70
