Table Of ContentA TEXT-BOOK
OF BACTERIOLOGY
BY
R. W. FAIRBROTHER
M.D., D.Sc, F.R.G.P.
Director of the Clinical Laboratory, Manchester
Royal Infirmary; Special Lecturer in Bacteriology,
University of Manchester; Lt.-Col., R.A.M.C.; Late
Research Fellow in Bacteriology, Lister Institute, London.
FOURTH EDITION
{Third Impression)
LONDON
WILLIAM HEINEMANN · MEDICAL BOOKS · LTD
1945
First Edition, February 1937
Reprinted, July 1937·
Second Edition, May 1938
Reprinted, November 1938
Reprinted, April 1939
Third Edition, July 1940
Reprinted, April 1941
Fourth Edition, September 1942
Reprinted, June 1943
Reprinted, January 1945
Printed in Great Britain by
The Whitefriars Press Ltd., London and Tonbridgt.
PREFACE TO FIRST EDITION
THIS book is an outline of the medical aspects of bacteriology.
The field of bacteriology has been so greatly extended during the
past 10 or 15 years that no account of the whole of it, even in
outline, is possible in the short time available in the medical
curriculum. The medical student must therefore restrict his
attention to those branches of the subject that are of medical
importance, viz., bacteria as agents of disease in man, and the
application of bacteriological methods in the prevention, diagnosis
and treatment of disease. Few of the available text-books stress
these points adequately, and the presentation of principles is
often interrupted by masses of technical detail needed only in the
laboratory.
In the present book an attempt is made to meet these criticisms.
In the absence of a universally accepted bacterial nomenclature,
that recommended by the Society of American Bacteriologists is
followed, and the biological relationship of allied organisms is
emphasized.
The bacteriological works consulted during the writing of this
book are too numerous to list in full, but my great indebtedness
to the " Principles of Bacteriology," by W. W. C. Topley and
G. S. Wilson, and to the " System of Bacteriology " published by
the Medical Research Council, demands specific acknowledgment.
I wish to thank Professor H. B. Maitland, Dr. D. T. Robinson
and Dr. R. Whitehead for advice and criticism, Dr. W. Susman
and Mr. V. H. Moore for the various photographs, and my Wife
for the drawings, the coloured plates and much general assistance.
DEPT. ΟΓ BACTERIOLOGY, R· W. Γ .
UNIVERSITY OF MANCHESTER. January, 1937.
PREFACE TO FOURTH EDITION
GREAT interest has been stimulated in chemotherapy following
the striking success of the sulphonamide compounds in the
treatment of many bacterial diseases. Voluminous literature has
accumulated, much of which is, however, inaccessible to the
student. A new chapter has consequently been introduced into
this edition in order to present in a simple form the salient features
of the recent developments in this subject.
Several chapters have also been revised, particularly those
dealing with cerebrospinal meningitis, tuberculosis, viruses,
rickettsise and water.
MANCHESTER, 1942. R. W. F.
PREFACE TO FIRST EDITION
THIS book is an outline of the medical aspects of bacteriology.
The field of bacteriology has been so greatly extended during the
past 10 or 15 years that no account of the whole of it, even in
outline, is possible in the short time available in the medical
curriculum. The medical student must therefore restrict his
attention to those branches of the subject that are of medical
importance, viz., bacteria as agents of disease in man, and the
application of bacteriological methods in the prevention, diagnosis
and treatment of disease. Few of the available text-books stress
these points adequately, and the presentation of principles is
often interrupted by masses of technical detail needed only in the
laboratory.
In the present book an attempt is made to meet these criticisms.
In the absence of a universally accepted bacterial nomenclature,
that recommended by the Society of American Bacteriologists is
followed, and the biological relationship of allied organisms is
emphasized.
The bacteriological works consulted during the writing of this
book are too numerous to list in full, but my great indebtedness
to the " Principles of Bacteriology," by W. W. C. Topley and
G. S. Wilson, and to the " System of Bacteriology " published by
the Medical Research Council, demands specific acknowledgment.
I wish to thank Professor H. B. Maitland, Dr. D. T. Robinson
and Dr. R. Whitehead for advice and criticism, Dr. W. Susman
and Mr. V. H. Moore for the various photographs, and my Wife
for the drawings, the coloured plates and much general assistance.
DEPT. ΟΓ BACTERIOLOGY, R· W. Γ .
UNIVERSITY OF MANCHESTER. January, 1937.
PREFACE TO FOURTH EDITION
GREAT interest has been stimulated in chemotherapy following
the striking success of the sulphonamide compounds in the
treatment of many bacterial diseases. Voluminous literature has
accumulated, much of which is, however, inaccessible to the
student. A new chapter has consequently been introduced into
this edition in order to present in a simple form the salient features
of the recent developments in this subject.
Several chapters have also been revised, particularly those
dealing with cerebrospinal meningitis, tuberculosis, viruses,
rickettsise and water.
MANCHESTER, 1942. R. W. F.
A TEXT-BOOK
OF BACTERIOLOGY
PART I
GENERAL BACTERIOLOGY
CHAPTER I
HISTORICAL SURVEY
ALTHOUGH there is no doubt that bacteria were observed almost
three centuries ago, their significance in the production of disease
was not established until the brilliant work of Louis Pasteur, a
French chemist, some eighty years ago. Previous to this period
much had been written about the nature of disease and the
spontaneous generation of living things, but these views were
largely the speculations of philosophers and in only a few cases
were they subsequently proved to be correct. These speculations,
however, were mainly responsible for the development of bacterio-
logy, and a brief review of their salient features is both interesting
and instructive.
It has been long recognized that disease might be communicated
from one individual to another ; there are numerous references
in the works of the earlier writers to indicate this, particularly in
connection with plague and small-pox, which frequently appeared
in epidemic form during the Middle Ages. The nature of the
infecting material was not, however, determined ; Fracastor of
Verona (1546) put forward the view that disease was spread by
particles too small to be seen and termed the responsible agent
a contagium vivum. He divided the contagious diseases into
two groups : those transmitted per contactum, i.e., from person
to person, and those transmitted per fomitem, i.e., through the
agency of some outside object. The first person to suggest, on
experimental grounds, that disease might be caused by visible
T.B. B
2 A TEXT-BOOK OF BACTERIOLOGY
living bodies was a monk, Kircher. Kircher, in 1671, using simple
lenses of low magnification, stated that he had observed peculiar
" worms " in the blood of persons suffering from plague.
Although he considered that these were the agents responsible
for the disease, there seems to be little doubt that the objects he
saw were simply arrangements of the red blood corpuscles.
A short time afterwards the existence of minute living bodies
was definitely established. Leeuwenhoek, a Dutch draper whose
hobby was the preparation of lenses, introduced the simple
microscope during the last quarter oî the seventeenth century.
Using these instruments, with which magnifications of from 40
to 300 diameter were obtained, he observed minute bodies which
he termed "animalcules". These were present in considerable
numbers in rain water and scrapings collected from the spaces
between the teeth. Some of the bodies were shaped like bent
sticks and others like spirals, while many were actively motile.
Leeuwenhoek described his results in numerous letters to the
Royal Society of London, but he did not appreciate that the
animalcules might have some connection with the production of
disease. There appears to be no doubt that Leeuwenhoek actually
saw the larger bacteria, which are normally present in water
and the mouth. An Austrian, Plenciz, in 1762 postulated the
germ theory and the specificity of disease. Plenciz, as a result
of experimental work of doubtful value, concluded that each
disease had its own specific agent which multiplied in the body
and so produced disease.
At this time, the middle of the eighteenth century, a consider-
able amount of controversy was taking place on the question of
spontaneous generation. After Leeuwenhoek had described the
peculiar minute bodies, or animalcules, their presence in various
situations was confirmed without difficulty. The adherents of
the theory of spontaneous generation considered that by an
investigation of the animalcules proof of this theory could be
definitely established. Needham, about 1750, communicated
to the Royal Society the results of his experiments on the genera-
tion of these minute bodies. He heated meat-broth at tempera-
tures, as he thought, sufficient to destroy living things and left
it tightly sealed for some days. On subsequent examination
Needham found that the broth was swarming with the animalcules.
By that and similar experiments he considered that the spon-
taneous origin of the animalcules was proved.
These views, however, did not satisfy an Italian priest,
HISTORICAL SURVEY 3
Spallanzani, who prepared similar flasks of meat-broth and
boiled 'them for varying periods of time. One set was sealed
and the other corked ; they were then stored for several days.
On examination it was found that the animalcules were present
in all the flasks except those which had been sealed after being
boiled for some time ; they were present in those flasks that had
only been boiled for a short time. Spallanzani considered that
the minute bodies could resist boiling for a short time and that
spontaneous generation had not taken place ; the living bodies
had simply developed from the living bodies already present in
the medium. Needham in reply to these criticisms stated that
the experiments of Spallanzani were valueless, as by the pro-
longed heating of the meat-broth some hypothetical substance,
which he termed " vegetative force ", had been destroyed. This
vegetative force he considered was necessary for the production
and development of the animalcules. This objection was over-
come by Spallanzani ; he found that the animalcules would grow
readily in meat-broth that had been subjected to high tempera-
tures for prolonged periods of time. As a result of further
experiments Spallanzani was able satisfactorily to counter the
hypotheses advanced by the adherents of the doctrine of
spontaneous generation. This theory subsequently became more
and more discredited, although some naturalists continued to
uphold it for many years.
The development of the microscope was also advancing rapidly,
and higher magnifications were soon available for the study of
minute bodies. It must be realized that, at this stage, the only
means of examination was by the observation through these
early microscopes of unstained material. In spite of these
difficult technical conditions definite progress was made ;
Ehrenberg, 1838, described various forms and classified four
groups : vibrio, bacterium, spirillum and spirochseta! A short
time previously, 1837, Latour and Schwann had found yeasts in
beer and wine and considered that they were responsible for the
fermentation process.
Although some of these early observations were undoubtedly
accurate, many conflicting statements were also made on most
unsatisfactory experimental findings. This haphazard manner
of investigation was criticized appropriately by Henle, a famous
anatomist, in 1840. Henle emphasized that, before aetiological
relationship of a micro-organism to a disease process could be
established it must be found constantly in cases of the disease
1 2
4 A TEXT-BOOK OF BACTERIOLOGY
and it must be isolated and tested by inoculation into healthy
animals. These arguments were amplified later by Koch in his
now-famous postulates.
At this time the question of spontaneous generation again
came to the fore. The observations of Schwann and Latour,
that yeasts were responsible for the fermentation process in beer
and wine, were subjected to much criticism. Liebig, a celebrated
German chemist, stated that fermentation was purely a chemical
process and that the ferment was obtained from the dead protein
already present in the reacting system. In view of his reputation,
Liebig's views were almost generally accepted ; this did much to
retard progress.
It was at this stage, 1856, that Pasteur entered the realms of
bacteriology. Originally a chemist, Pasteur undertook an
investigation into the cause of some faulty fermentation which
was seriously affecting the wine industry of Lille. As a result of
his work, Liebig's theory of fermentation was finally and com-
pletely disproved ; Pasteur found that yeasts were responsible
for the fermentation process and that sugar could be converted
into alcohol in the absence of any protein in the medium. He also
showed that faulty fermentation was the result of the presence
of some contaminating rod-shaped micro-organisms. The theory
of spontaneous generation was also finally crushed by further
researches of Pasteur, in which innumerable flasks with wide,
constricted, twisted and elongated necks were made in order to
show that the contaminating micro-organisms had their origin
in the air. The effect of heat on these bodies was also examined,
and he found that certain organisms were able to withstand
boiling for some time ; these were doubtless the spore-bearing
bacteria. Some organisms were found to grow in the absence
of oxygen while others required its presence. The use of cotton-
wool plugs in order to prevent organisms from entering flasks
already sterilized was also demonstrated about this time by
Schroeder and Dusch. The relationship of micro-organisms to
the processes of fermentation and putrefaction thus became
definitely established, but it is interesting to note that up to this
time conclusive evidence that these minute bodies were agents
of disease had not been obtained.
After his brilliant work on the question of fermentation, Pasteur
(1865) was next persuaded to investigate a disease of silkworms,
pébrine, which was crippling the silk industry of France.
Although mishaps and setbacks were frequent, Pasteur, after
HISTORICAL SURVEY 5
five years' work, had settled the problem and re-established the
silk industry on a prosperous basis. He found that the disease
of the worms was due to some living microscopical organism and
that the disease was transmitted from one animal to another.
By obtaining healthy silkworms and subsequently removing any
showing signs of infection Pasteur was able to maintain a healthy
stock, although he was surrounded by infected farms. The
relationship of living microscopical objects to the disease was
thus proved ; it is a point of interest that this disease was of
protozoal and not bacterial origin.
These important fundamental observations of Pasteur were
not allowed to pass unnoticed by the outside world. Although
there were many who decried these new and revolutionary ideas,
there were others who realized the true significance of this work.
Lister, then Professor of Surgery at Glasgow University, con-
sidered that the micro-organisms, so prevalent in the air, might
be responsible for the frequent occurrence of the inflammation
and suppuration after surgical operations. In spite of much
criticism and opposition, he, in 1867, revolutionized surgery by
introducing an antiseptic technique. In this method sprays or
washings of carbolic acid were directed during the operation on
the wound, which was subsequently protected from the air by
suitable dressings. The results which followed this procedure
were remarkably successful and the practice became increasingly
popular ; there followed a striking reduction in the incidence of
post-operative sepsis. This was an important demonstration
of the rôle played by micro-organisms in the causation of disease.
A later advance was the replacement of antiseptic surgery by
aseptic surgery.
The study of the micro-organisms, although a matter of some
difficulty, was now becoming increasingly popular. Many varied
forms were seen and described. A further and more systematic
method of classification was elaborated by Cohn (1872) ; in this
were included the following generic names : micrococcus,
bacterium, bacillus, vibrio and spirochseta. The name
bacteria was also used for the first time to designate the
micro-organisms multiplying by fission. This classification was
based on morphological observations, and it is interesting to note
that many of the terms are used to to-day in the sense originally
employed by Cohn, who also described the appearance of spores
in a non-pathogenic bacillus and noted the increased resistance
to heat and disinfectants possessed by the spore-bearers.
6 A TEXT-BOOK OF BACTERIOLOGY
About this time the relationship of a micro-organism to an
important animal disease, splenic fever or anthrax, was being
investigated. Pollender, 1849, and Davaine, 1850, had described
the presence of a rod-shaped organism in the blood of animals
suffering from splenic fever. Some time later Davaine repro-
duced the disease in healthy animals by injecting them with blood
collected from infected animals. The connection between this
organism and the disease, however, was not definitely established
until the thorough and brilliant work of a medical practitioner
in Germany, Robert Koch. Koch examined the organisms
present in the blood of infected animals, reproduced the disease
in healthy mice by inoculating them with infected blood and also
prepared cultures by inoculating the aqueous humour of a
bullock's eye with the infected blood on a hollow ground slide.
The growth of the organisms in culture was observed, and after
several subcultures the organism was still found to be virulent
for the mouse. The production of spores by the rods and the
vegetation of the spores into rods were also observed and described.
In this manner, in 1876, the aetiology of anthrax was established
beyond any shadow of doubt. This observation was confirmed
and elaborated shortly afterwards by Pasteur, who introduced a
method of active immunization whereby the resistance of animals
to the disease was increased. Pasteur had previously found by
accident that the inoculation of chickens with relatively avirulent
cultures of the organism of chicken cholera did not produce the
disease and that a later inoculation of these birds with a virulent
strain was also unsuccessful. This result indicated to Pa&teur
that the original avirulent culture had protected the chickens
against subsequent infection with the virulent organisms. This
was a step of great fundamental importance ; although prophy-
lactic immunization against small-pox by means of cow-pox
material had been developed and practised by Jenner in 1800,
this procedure was, however, entirely empirical.
Pasteur next endeavoured to apply his discovery to splenic
fever. Here, owing to the presence of the resistant virulent
spores, the difficulty was to obtain a relatively avirulent culture.
This, however, was overcome by growing the organisms at a
temperature higher than that optimal for growth and at which
spores did not develop. This attenuated culture was given,
followed after a short interval by a second inoculation of a less
attenuated culture. The value of these injections in protecting
sheep was demonstrated publicly by Pasteur at Pouilly-le-Fort
