Bacterial Growth and Form

439 Pages·1995·13.983 MB·English

by  Arthur L. Koch (auth.)

Thin and Thick and Thick and not
stretchable stretchable stretchable
E.coli E. hirae
I
i M. xanthus
Plasmids
--..... Viruses
........... Insertion
Sequences
Abiotic synthesis
Hyper-Cycles
Vesicle Formation Chemiosmosis
PRE-CELLULAR 'LIFE'
BACTERIAL
GROWTB
andFoRM
L.
ARTHUR KocH
INDIANA UNIVERSITY
BLOOMINGTON INDIANA
SPRINGER-SCIENCE+BUSINESS MEDIA, B.V.
Art direction: Andrea Meyer, emDASH inc.
Cover design: Saeed Sayrafiezadeh, emDASH inc.
Copyright © 1995
Springer Science+Business Media Dordrecht
Originally published by Chapman & Hali in 1995
Softcover reprint of the hardcover 1 st edition 1995
AII rights reserved. No part of this work covered by the copyright hereon may be reproduced or used in any
form or by any means-graphic, electronic, or mechanical, induding photocopying, recording, taping, or
information storage and retrieval systems-without the written permission of the publisher.
1 2 3 4 5 6 7 8 9 10 XXX 01 00 99 97 96 95
Library of Congress Cataloglng-in-PubUcadon Data
Koch, Arthur L.,
Baeterial Growth and Fonn / Arthur L. Koch.
p. em.
Includes bibliographieal referenees and index.
ISBN 978-1-4613-5717-9 ISBN 978-1-4615-1779-5 (eBook)
DOI 10.1007/978-1-4615-1779-5
1. Baeterial growth. 1. Title.
QR84.5.K63 1995
589.9'031-de20 94-38640
CIP
Brltish Library Cataloguing in Publicadon Data available
Contents
A Profile of Arthur L. Koch, Professor of Biology, Indiana University vii
Preface Xl
Prologue X111
1. From the First Cell to the Last Universal Ancestor 1
2. The Contrast of the Cellular Abilities of Eukaryotes and Prokaryotes 26
3. Bacterial Growth 37
4. The Synthesis of Functional Bacterial Wall 88
5. Thrgor Pressure of Bacterial Cells 118
6. The Surface Stress Theory: Nonvitalism in Action 143
7. The Mechanical Aspects of Cell Division 173
8. The Gram-Positive Coccus: Enterococcus hirae 197
9. Gram-Positive Rod-Shaped Organism: Bacillus subtilis 219
10. The Gram-Negative Rod: Escherichia coli 250
11. Apical Growth of Streptomyces and Fungi 306
12. Twisting and Rotation During the Growth of Gram-Positive Rods 326
13. The Structural and Physiological Roles of the Layers of the
Envelope of Gram-Negative Bacteria 339
14. Gliding Motility, Protonmotive Force, and Flagellar Rotation 361
15. Prokaryotic Perspective 378
Bibliography 393
Index 417
v
A Profile of Arthur L. Koch
Professor of Biology, Indiana University
Arthur Koch calls himself an ecologist. Those who know him recognize Arthur
as one of the true renaissance scientists of the several past decades. His accom
plishments span evolution, cell biology, prokaryotic genetics, bacterial growth,
bacterial transport mechanisms, mutagenesis, and more recently the development
of the concepts of surface stress on microbial morphology. His book is concerned
with all of the above. It seems that all of Arthur's past research has led to the
genesis of this book.
Arthur was born in Saint Paul, Minnesota, and served in the Pacific Theatre
in World War II as a member of the U.S. Navy. After military service he gained
a B.S. in chemistry from Caltech and a Ph.D. in biochemistry from the University
of Chicago. After four years in the Division of Biology and Medicine at the
Argonne National Laboratory, Arthur joined the faculty at the University of
Horida School of Medicine. In 1967 Arthur became a professor of Biology at
Indiana University, where he remains.
Arthur has won numerous awards, including a Guggenheim Fellowship. He
has been a generous donor of ideas to many of us. His enthusiasm for studying
the growth of bacteria is readily apparent in casual conversations. Even though
Arthur has a superior intellect, he has always encouraged and brought out the
best in others.
Arthur Koch's appetite for prokaryotic cell biology emerges in all chapters of
this book. The theme of surface stress, like a theme in a great symphonic
work, reoccurs time after time. The surface stress theory constitutes a significant
achievement in bacterial physiology. In this book, the role of surface stress is
defined with respect to cell morphologies, to the regulation of autolysins, to the
method that some bacteria use to precisely bisect their septa, to the method some
bacteria use to maintain constant wall thicknesses, to prospective sites for new
antibiotics, to the method that a prokaryote uses to segregate its growing chromo-
vii
viii / Profile of the Author
some(s), to the dynamics of cell wall turnover, and to flagellar movement and
gliding motility.
The surface stress theory states that microbial morphologies are predictable
when just a few facts are known about their division patterns. The principles of
engineering applied to the construction of dams or boilers are also applicable to
the construction of a bacillus or a coccus. The concept "make before break" in
reference to assembly and turnover of cell walls of bacteria is analogous to the
concept of constructing cofferdams before permanent dams are built. The equa
tions developed for surface tension are the same for a bacterium as for a soap
bubble or a chemical reactor unit. In his work, Arthur has relied on equations
developed by D' Arcy Wentworth Thompson. His thesis is that bacterial growth
(and of course that of all walled microorganisms) depends on how the bacterial
surface responds to turgor pressure. When a bacterium takes in nutrients, it must
either expand its surface area or develop a higher turgor pressure. The way in
which the bacterium responds to turgor ultimately dictates its shape. This is, of
course, an oversimplification, but it permits the many variations of shapes found
in the microbial world. The fundamental equation
PaV=TAA,
where P is pressure (or force per unit of surface), aV is volume change, pay
is the work of expansion, T is surface tension (a unit of force per distance), and
AA is the change in surface area, can be modified to accommodate spheres, rods,
or capped rods under pressure. The text reveals how a simple equation can
undergo permutations, become complicated, and ultimately be applied to all
kinds of microbial shapes. It is challenging to follow the author's arguments about
turgor and shapes. The reward is a refreshing understanding of the relationships
between turgor pressures, surface expansion, and the forces that determine micro
bial morphologies.
As surface stress theory has emerged these past few years, several corollaries
have developed. Surface stress states that a new wall must be added before an
old wall is turned over by autolysins (make before break). Surface stress requires
that new wall units be inserted into the pre-existing wall, to become functional
as wall only when they are crosslinked and able to bear turgor. Surface stress
demands that surface expansion (elongation) of rod-shaped bacteria occur by the
diffuse intercalation of new wall into old wall in cell cylinders. When this new
wall is crosslinked and able to bear pressure it is thought to stretch. The addition
and subsequent stretching of wall at many sites ensures elongation of the cell.
Surface stress rules out the elongation of cell cylinders of rod-shaped bacteria
by equatorial growth zones. Surface stress can now account for the growth
patterns of Escherichia coli, Bacillus subtilis, the blunt-ended B. anthracis, and
streptococci. Koch has provided the only serious work to explain the morpholo
gies of bacteria first described by van Leeuwenhoek over three hundred years
ago.
Profile of the Author I ix
Another theme intercalated into this book is one that has been popularized by
Koch; namely, that bacteria undergo periods of feasting, only to be confronted
with periods of fasting. The feast-famine lifestyle has resulted in evolutionary
adaptations and to a large extent makes certain patterns of bacterial metabolism
and behavior predictable. Starvation results in small cells, causing the ratio of
(he surface area to volume to increase, optimizing the uptake of nutrients. The
surface stress theory allows for an understanding of modulation of surface-to
volume ratios at various growth rates and defines the need to couple surface
expansion with the environment of the bacterium.
We now invite you to partake of the richness of this book. We both have been
blessed to know Arthur personally. Those of you who will never be able to meet
Arthur will get to know him well by studying this book. At the same time you
will gain a new appreciation of microbial physiology.
R. J. Doyle, Professor University of Louisville
Lolita Daneo-Moore, Professor Temple University
Preface
In this book, the pronoun "we" is used frequently. It is neither the royal
nor the editorial "we"; rather, it means you and I. We are going to consider
bacteriological problems together. As much as possible, I will try not to lecture;
we will be thinking our way through the important biological problems. I assume
that you already know some microbiology. My most important goal in writing
the book is to make accessible the relevant thinking from fields of science other
than microbiology that are important to microbiology. The book is written for
people who already have a fascination with bacteria.
This book consists of topics that are largely omitted from microbiology text
books. It contains a good deal of my own work, both experimental and theoretical,
together with a lot of speculation. It is, however, only one-third of a text on
microbial physiology. The book by Neidhardt, Ingraham, and Schaechter (1990)
covers another third. The final third is covered in the recent textbook by White
(1995).
A list of all those who contributed to the ideas developed in this book would
number more than one hundred. The short list follows: Ronald Archibald, Dick
D' Ari, Terry Beveridge, Ian Burdett, Angelika Busch, Marta Carparr6s, Steve
Cooper, Lolita Daneo-Moore, Paul Demchick, Ron Doyle, Don Gilbert, Frank
Harold, Jean van Heijenoort, Michael Higgins, Joachim-Volker Holtje, Kathryn
Koch, Herb Kubitschek, Harold Labischinski, Stine Levy, Neil Mendelson,
Nanne Nanninga, David Nickens, Tom Olijhoek, Greg Payne, Miguel de Pedro,
Suzanne Pinette, Harold Pooley, Tina Romeis, Elio Schaechter, Heinz Schwarz,
Uli Schwarz, Gerry Schockman, Elaine Sonnenfeld, Marcus Templin, John
Thwaites, Frank Trueba, David White, Steve Woeste, and Conrad Woldringh.
This list does not include three orthopedic surgeons who kept me quiet enough
to start to write this book.
I would never have written this book if it had not been for Greg Payne. He
encouraged me, worked over my English, and did not worry about deadlines.
xi
xii / Preface
Many people, both known and unknown, helped me by correcting my grammar,
correcting my ideas, correcting my organization, and in the production of the
book. These include David White, Greg Payne, Lisa LaMagna, Torrey Adams,
George Hegeman, William Baldwin, and Frank Harold. A young undergraduate
wrote to tell me that I wrote very clearly and explained things very well. That
is a lie, but I answered by saying that she was correct, but only because there
had been twenty-seven drafts that had been corrected, criticized, complained
about, and rejected by friends and enemies. This book has been no different, it
needed all the help it could get.
PROLOGUE
Thinking about Bacteria
OUTliNE
WHAT PROKARYOTES CAN AND CANNOT DO
BACTERIA ARE THE SIMPLEST ORGANISMS THAT GROW AND
REPRODUCE IN A SELF-CONTAINED WAY
THE IMPORTANCE OF BEING SMALL
THE VARIOUS KINDS OF CELL REGULATION
NOW IS A GOOD TIME TO OVERVIEW GROWTH
PHILOSOPHY OF PRESENTATION TO BE USED IN THIS BOOK
THIS BOOK'S GOAL: TO BE THE INTERFACE BETWEEN MORPHOLOGY
AND CHEMISTRY
KEY IDEAS
Prokaryotes are simple, but sophisticated.
Prokaryotes occupy important niches.
"Minimalist" organisms do nearly everything needed to grow.
Many problems are solved by being small.
Prokaryotes have exquisite regulatory systems, different from those of eukaryotes.
Crucial to the biology of all life forms are osmotic pressure and turgor pressure.
The purpose of this book is to outline and defend an approach to thinking
about what bacteria were, what bacteria are, and how they do what they do.
Particular emphasis is directed to their ability to establish their shapes as they
grow and divide. In this prologue I will try to convince you that the study of
bacteria and their life strategies is important even during what is being called
the Golden Age of the Developmental and Cell Biology of Eukaryotic Organisms.
The major point is that prokaryotes [both eubacteria and archaebacteria (arch
aea)] do many of the same things that eukaryotes do, but with simpler equipment
that is utilized in extremely sophisticated ways. This idea is interwoven with
xiii