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The Bacteria, 1880

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ALBERT R. MANN LIBRARY


AT

CORNELL UNIVERSITY

Cornell University Library

QR

41.M192
The
Bacteria.

3 1924 003 200 288

The

original of this

book

is in

the Cornell University Library.

There are no known copyright

restrictions in
text.

the United States on the use of the

http://www.archive.org/details/cu31924003200288

THE BACTERIA.
BY

DR.

ANTOINE MAGNIN,

LICENCIATE OF NATURAL SCIENCES, CHIEF OF THE PKACTICAL LABORS IN NATURAL HISTORY TO THE FACULTY OF MEDICINE OF LTON8, LAUREATE OF THE FACULTY OF MEDICINE OF PARIS (SILVER MEDAL, 18T6), GENERAL SECRETARY OF THE BOTANICAL SOCIETY OF LYONS, MEMBER OF THE BOTANICAL SOCIETY OF FRANCE, ETC.

TRANSLATED BY

GEOEGE

M.

STERNBEEG,
U. a.

M.D.,

SURGEON

ARMY.

BOSTON:
LITTLE, BROWN,

AND COMPANY.

1880.

Copyright, 1880,

By George

SI.

Sternberg.

University Press:

John Wilson and Son, Cambridge.

PREFACE BY TRANSLATOR.

edge of the bacteria, by Dr. Magnin, of great assistance

which I have been engaged during the past year under the auspices
of the National

to me, in pursuing the investigations in

Board

of Health,

it

has seemed to

me

that a translation of the


cation in this

work into English and its publicountry would be productive of good in


one,
it

more ways than

and of the advancement of


cannot
fail to

science.

To

the naturalist,

be of value, as the

most approved
with a

classification, that of

Cohn,

is

given,

full description of species.

To

give additional

value to this portion of the work, figures of

many

of the

best-known forms, drawn from various foreign sources,

and reproductions

of

some of

my own

photo-mico-

graphs (by permission of the National Board of Health),

have been introduced.


If

we

are to judge from the scanty literature of the

subject in this country, the

amount

of interest

which
of

has been aroused by the revelation of a

new world
is

micro-organisms, and by the momentous questions which

have been raised in connection with them,


that
is

far

below

awakened

in

Germany, France, and England. This


;

not, however, really the case


in

for,

while

we have but

few active workers

the difficult fields of inquiry

PREFACE BY TRANSLATOR.
attractive, especially for the

which have proved so man and the French


1

Ger-

savants, there

is

nevertheless a

wide-spread interest in these investigations, and a desire


to

know

their results.

a difficulty

met with which has no doubt discouraged many, and


But, just here,
are
results

we

perhaps caused some to drop the whole subject in disgust.

The

have been so contradictory, and so


impossible

many would-be
to arrive at

savants have uttered opinions entirely


it

opposed the one to the other, that we find any


definite opinion,

not knowing
affairs, it

whom

to

believe.

This being the condition of


it is

seems to

me

that

necessary for us to
first

commence

investigating

for ourselves,

making

ourselves familiar with

what
the

has been done abroad, and then avoiding,

if possible,

quicksands into which unfortunate science has too often

been dragged by

her votaries.

One

great trouble
is

which we have experienced


ferent

in this country

in judg-

ing of the comparative value of the observations of dif-

men who
article
;

are equally

unknown

to us.

A very
careless

plausible

may be

written

by a very

observer

or a very cautious observer

may

fail to

give

confidence in his results, because of a certain degree of

confusion in his language.


devised, carefully executed,
sion, as is

When

experiments are well

and described with precidone by such men as Pasteur and Tyndall, we


to attach great

cannot

fail

weight to the conclusions

reached.

And when
Koch

so accomplished a microscopist as

thing, or has

such and such a made such and such measurements, we cannot doubt the reliability of the observation. But sometimes we are deceived by giving credence to a man
or

Cohn

asserts that he has seen

who

has achieved reputation in one line of study, but of

PREFACE BY TRANSLATOR.

whose skill and training in the use of the microscope we have no means of judging. Such a man may be a great
surgeon, or a great clinician, or a great chemist, and yet

be a mere tyro with the microscope.


see
it

When,

then,

we

announced that Dr. So-and-so


in blood, or

failed to discover

any micrococci in pus,


a certain source,

what

not, taken

from

we

are justified in asking,

first,

what

power did the learned doctor use ? second, is he capable of distinguishing micrococci in fluids which contain them beyond question ? Or, if he does discover them, we may ask if he is accustomed to making a differential diagnosis between micrococci and inorganic granular
material,
or

unorganized granules of organic origin.

This

is

a decision which the most accomplished microis

scopist

sometimes unable to make, except by the aid

of chemical tests and culture experiments.

To

avoid this want of confidence in results, which has

naturally

grown out

of carelessly
it

made
all

observations and

contradictory statements,

is

desirable that full

and

minute

details should be

given of

observations and

experiments made, and, whenever possible, that photo-

micographs should
to contain any

be

made

of

all

micro-organisms

described, or of a thin stratum of a liquid asserted not


;

as,

when

a sufficiently high

power

is

used,

this settles

the question of their presence

or

absence, beyond dispute, and enables other students to

make comparisons and measurements which cannot


to

fail

promote the interests of true science.

The National Board


has the credit of
first

of Health of the United States

adopting this method of recording

the results of scientific investigation, in this direction,


as a constant

and unimpeachable record of what has

PREFACE BY TRANSLATOR.

been seen by the investigator.

The commission sent

to

Havana
fever,

last

summer

for the investigation of yellow-

was instructed to pursue this method, and was accompanied by a photographer and supplied with all
the necessary appliances for carrying these instructions

into effect.

the

The superficial reader may find much to criticise in work of Dr. Magnin, but I am convinced that those who read it carefully cannot fail to be pleased
scientific

with the truly

spirit in

which

it is

written

the fairness with which conflicting opinions are stated

the caution manifested as to the drawing of definite


conclusions where questions are
and, above
all,

still

under discussion

the extent of his literary researches and

the systematic
sults.

way

in

which he has arranged the

re-

For the

naturalist, for the physician, or for the

non-

professional

man

of general culture,

who
up

desires to

have
re-

accessible in a condensed form the

most important

sults achieved in this line of inquiry

to the present
;

day, this volume cannot

fail to

be of value

while for

the student and the investigator in search of fuller


information, the

summary given

of the labors of nubibliog-

merous individuals, together with the copious


be of service.
pleasure for

raphy, which I have brought up to date, will doubtless


Believing this to be true,
to devote a portion of
it

has been a

me

my summer vacaG. M.
S.

tion to the translation of this little volume.

Salem, Mass., August

1,

1880.

TABLE OF CONTENTS.
PAGE

Introduction Historical

11

13

PART
CHAPTER
1.

FIRST.

MORPHOLOGY OF THE BACTERIA.


I.

Organization.
28 28 29
31

Of

the Bacteria in General Forms


Dimensions
Colors

Movements
Structure

32

35
35
36 39

Cell-Membrane Protoplasm
Cilia
2.

Different
Form Form Form
of

Modes of Association

43

of Little Chain (Torula)


of Zooglcea

43 44 45

Mycoderma, &c
II.

CHAPTER

1.

Classification

and Description.
48
53

Place of the Bacteria

Among
2.

Organized Beings

In the Vegetable Kingdom

55
59
60

Classification
Characters Generic and Specific
Classification of

Cohn

65

TABLE OF CONTENTS.
3.

Description of
Micrococcus

Genera and Species

....

PAGE 65
71

Spherobacteria

72 78
80 80
'

Monads
Microbacteria

Bacterium

Desmobacteria
Bacillus

86 87 90 91 92

Leptothrix

Spirobacteria
Vibrio
'

Spirillum

94

PART SECOND.
PHYSIOLOGY OF THE BACTERIA.

CHAPTER
1.

I.

Development

in General.

Origin of Bacteria
Heterogenesis

101 102 103 103


106

Dissemination
In Air

In Water In the Human. Organism


2.

107
Ill
Ill

Nutrition and Respiration Aliments Water


:

Nitrogen

112 113
115

Carbon

Oxygen
Temperature Other Agents
3.

118 121
123 123 126 130
133

Reproduction
Fission

Spores

Sporangia

Polymorphism


TABLE OF CONTENTS.
9

CHAPTER
1.

II.

Development

in

Different Media.

R6le

op Bacteria in Fermentations

....

paoe 137
139 142

Acetic Fermentation

Ammoniacal Fermentation Lactic and Butyric Fermentation


2.

144
.

R6le

in

Putrefaction and Nitrification

148 152 152

3.

Role in Virulent Affections


Septicemia

Charbon
Variola
Scarlatina

158
167
169 169

Measles
Diphtheria

170
171 171

Typhoid Fever Glanders, Farcy


Endocarditis
4.

173 175 176 178 185 188


191

Role

in Surgical Lesions Exposed to the Air


Closed

Therapeutic Deductions

Conclusions Bibliography

Index

223

LIST OF PLATES.
PLATE
I.

OPPOSITE PAGE

The

Cilia of B. termo

and S.

voluntatis.

(Drysdale

and Dallinger)
II.

40

Different

Modes

of

Grouping of the Bacteria

...
. .

47
58

III.

Saccharomycetes and Schizomyce'tes


Disease Ferments of
Different
Different

IV.

Wort and

Beer.

(Pasteur)

84
95

V.
VI.

Forms

of Bacteria. of

(Cohn)

Forms

Bacteria.

(From Photo-micro98

graphs by Dr. Sternberg)


VII.

Dissemination of the Bacteria.

(From

PhotOTinicro-

graphs by Dr. Sternberg)


VIII.

100
(Photo127

Reproduction of Bacillus Ulna, by Spores.

micrographs by Dr. Sternberg)

IX.

Formation of Spores in Bacillus, &c.


Spirillum Obermineri

(Koch)

153

X.

and

Bacillus Anthracis.

(Koch).

268

THE BACTERIA.
INTRODUCTION.
" Corruptio unius
est generatio alterius."

Luceetics,

De Serum Natura.

Of all the studies which have for their object the inferior organisms, those which relate to the bacteria offer, without contradiction, the greatest
interest, as

they touch the most divers problems,


true, are

which,
least

it

is

the most difficult and the

known

in biology.

The

history of these mi-

nute organisms is, in truth, related to that of spontaneous generation, to that of the fermentations, to the pathogeny and therapeutics of a great number of virulent and contagious affections, and, in a more general manner, to all the unknown which, notwithstanding the efforts of modern science, still surrounds the origin of life and its preservation.
If

the relation of these inferior organisms to


is

the origin of living beings

yet obscure, their


is

rdle in the preservation of life


It is

better known.

known

that organic matter, once produced


solid, so to

and become

speak, cannot again enter


it

into the general current until

has undergone

12

THE BACTERIA.

new

transformations, metamorphoses produced, according to some savants, favored, according to

without contradiction, accompanied by the development of bacteria; 1 and, without wishing to attribute to these organisms a finality which is repugnant to our monistic conception of the universe, it may be said that it is thanks to
others,

but,

them

that the continuation of

life is

possible

on

the surface of the globe.

But,
field is

if

these studies are full of interest, their

so vast

that

we cannot

flatter

ourselves
it

that

we have

passed over the whole of

with

equal care.

The

little

time that has been ac-

corded us for the composition of this thesis will be our excuse for the inevitable imperfections

which

will doubtless

be found in our work.

1 The bacteria : such is the subject which has been imposed upon us but it is certainly useless to give the reasons which have caused us to study not only the bacteria properly so called, taking the word in its most restricted sense, but all the organisms which are comprised under the names o bacteria, vibrios, schizomycetes, schizophytes, etc.

HISTORICAL.

The

bacteria are the lowest organisms, situated


limit of the

upon the

two kingdoms, animal and

vegetable, and are thus defined by the botanists

who have most


them
:

recently occupied themselves with

" Cells deprived of chlorophyll, of globular, oblong, or cylindrical form, sometimes sinuous and twisted, reproducing themselves exclusively by
transverse division,
families,
1

living isolated or in cellular

and having affinities which approach them to the algas and especially to the oscillatoriae."
But, before arriving at this degree of relative

precision, the history of the bacteria has passed

At one through the most diverse vicissitudes. time considered as animals, at another taken for vegetables, transported from the algae to the fungi, one author has even gone so far as to refuse to

them

This diversity minuteness of their dimensions and the difficulties with which their observation is surrounded.
of opinions
is

2 the nature of living beings.

due

to the

Reproduction by spores has been proved to occur in Bacillus subtilis, seems altogether probable that other species are reproduced in the same way. G. M. S. 2 Polotebnow.
1

and

it

14

THE BACTERIA.

Although an historical statement of the progress of our knowledge of these minute organisms has been given in several publications, we think it best to make here a new historical summary, which will be completed by an indication of the principal papers relating to them which have been published
recently.

The

first

observer

who

perceived bacteria was

Leeuwenhoeck.

As

early as 1675, while examin-

ing by chance with his magnifying glasses a drop of putrid water, the father of microscopy remarked with profound astonishment that it contained a multitude of little globules, which moved

with

agility.

The following year he recognized


;

the presence of bacteria in faeces and in tartar

from the teeth


it
is

and, if he has not named them, easy to assure one's self by the description

which he has giyen of their form and of their movements, and by the figures which accompany
these descriptions, 1 that the organisms observed by him are truly Bacteria, Vibrios, and perhaps

even Leptothrix. In 1773 0. F. Muller endeavored to classify these organisms. He made of them a group of infusoria, under the name of Infusoria crassiuscula, and established two genera, the g. Monas and Vibrio ; the first characterized as follows "vermis inconspicuus, simplicissimus, pellucidus,

punctiformis," comprising the following species: Monas termo, atomus, punctum, ocellus, lens, mica,
1

Leeuwenhoeck. Opera omnia, Lugd. Batav., 1722,

11, p. 40, fig.

A to G.

HISTORICAL.
tranquilla,
is

15
uva, which
it

lamellula, pulvisculus,

impossible to identify with the species at pres-

ent recognized.

The genus Vibrio


specific

" vermis incon-

spicuus, simplicissimus, teres, elongatus" enclosing

names, with the true bacteria, some organisms belonging to other classes of the animal and vegetable kingdoms. In the classification of the infusoria given byBory de Saint-Vincent in the " Encyclopedic MeHhodique" (1824) and afterwards in the "Dicthirty-five

under

tionaire Classique d'Histoire Naturelle " (1830) the

bacteria are distributed in two different families


of the microscopic

gymnodae, the monadaires and

the vibrionides.
called, of

Besides the monads, properly so

which the Monas termo has been pre-

served by the greater part of the bacterologists, the monadaires include some veritable infusoria,

which have no relation with the monads. It was the same with the vibrionides, of which the genera Vibrio and Mellanella included some beings very
different in
their

organization.
bacteria,

Indeedj beside

some

veritable

vibrios,

constituting

the genus Mellanella,


as

and spirilla, Bory placed


Anguillida

some nematoid worms, such


of vinegar.

the

With Ehrenberg (1838) and Dujardin (1841) the family of the vibrioniens was established upon more homogeneous, and their species upon distinctions truly scientific. But these two observers, followed in this by M. Davaine, deny
characters

completely the

affinities of

the elongated bacteria

16
(Bacterium,

THE BACTERIA.
Vibrio,
;

with the punctiform is necessary to come to the time of MM. Hallier, Hoffmann, Cohn, and the greater number of recent botanists, in order to see these two forms brought together anew.
etc.)

bacteria

(Monas)

and

it

In he

fact,

Ehrenberg defines his vibrioniens, which arranges between the volvocineae and the
filiform,

closteria " animals,

distinctly

or appar-

no mucous membrane, naked, without external organs, with the body (like monads) uniform and united in chains or filiform seently polygastric,
ries,

as

a result of

incomplete

division."

He

included in this

class* all

filiform

bodies gifted
of
articles,

with proper
dividing
1.

movement and formed them into four genera


:

Bacterium: filaments linear and inflexible; three


Vibrio
:

species.
2.

filaments linear, snakelike, flexible


filaments spiral, inflexible

nine

species.
3.

Spirillum

three spe

cies.

4.

Spirochete

filaments spiral, flexible; one species.

the Spirodiscus fulvus, with filaments in a helix, inflexible, disposed in contiguous layers, has not been seen since Ehrenberg. Let us add that
fifth

genus, including

but

one

species,

Ehrenberg often attributed to them a complex structure, stomachs more or less numerous, a proboscis,
all

cilia

characters
to

serving as organs of locomotion, that more recent observers have


Nevertheless,

failed

find.

we must make an

HISTORICAL.

17

cilia, of which the exbeen recently verified in the case of several of the bacteria by divers botanists, among others by MM. Cohn and Eug. Warming.

exception in favor of the


istence has

Dujardin (1841), in his " Histoire Naturelle des Zoophytes," preserved the family of the vibrioniens of Ehrenberg ing

among
:

them

as follows

the infusoria, characteriz" fiiliform animals, extremely

slender, without appreciable organization, without

He made but few which the principal consisted in uniting Spirochcela with Spirillum, Dujardin. Rejecting the character that Ehrenberg drew from
visible

locomotive organs."

modifications, of

the rigidity of the


tilis,

spirilla,

the Spiroehceta plica-

Ehrb. became the Spirillum plicatile, Duj. but, as will be seen later, this change has not been maintained. Dujardin, then, classed the bacteria in
1.

Bacterium
Vibrio
:

filaments rigid, with a vacillating

movement.
2.

filaments flexible, with an undulatory


filaments spiral,

movement.
3.

Spirillum

movement

rotary.

Until this time the bacteria had been considered


as animals placed at the foot of the series.

Subin

sequently the

tendency to place

them

the

vegetable kingdom became more and more pronounced.

Already, since 1853, M. Ch. Robin had pointed out the relationship of the bacteria and of the
2

18
vibrios

THE BACTERIA.

This opinion, which with Leptothrix. was not favorably received by the authors who adopted nearly all of the generic groups of Ehrenberg and Dujardin, is to-day accepted by many botanists, above all since the labors of Cohn. (See

below:

classification.)

At

all

events,

it

is

to

M.

Davaine (1859) that we are indebted for clearly


pointing out that the vibrioniens are vegetables, nearly allied to the algae, and especially to the
confervae.

This same author, having observed some motionless bacteria, thought it necessary to give
this character great
lish

consideration,

and to estab-

a fourth group, the genus Bacteridium, which he added to the three others admitted by Dujardin ; but in this creation he was less happy than
in
his placing the vibrioniens
;

among

the vege-

tables

for

acter of mobility or of immobility

and that it or upon certain conditions relating to the medium in which it is placed. The most recent complete exposition of the classification and of the ideas of M. Davaine is found in the " Dictionnaire Encyclop. des Sciences Me"dieales," art. BacteVies (1868).

on that this charis not absolute, depends upon the age of the bacterium
shall see further

we

be

summed up
\

as follows

It

may

Filaments straight
or bent, but not
in a spiral
.

Moving

sponta- ) Kigid.
. .

Bacterium.
Vibrio.

V
)

neously
Motionless

Flexible

Bacteridium.
Spirillum.

Filaments spiral

HISTORICAL.

19
six species,

The genus Bacterium comprises


B. termo,
din,

catenula,

punctum,

triloculare, or ariicula-

tum, already described by Ehrenberg and Dujar-

and B. putredinis and capitatum, new species of M. Davaine, established, the first for a bacterium producing rot in plants, the second for a species,

swollen at the extremity, observed in some


includes

macerations.

The genus Vibrio


V.
lineola,

twelve species,

tremulans, rugula, prolifer,

serpens,

bacillus,

synxanthus, and syncyanus of previous


the V.
lactic,

authors and
fermentations.

butyric,

and

tartaric

right, discovered

by M. Pasteur

in these different

In the genus Bacteridium, M. Davaine places


five

new

species,

the " Bacteridies charbonneuse,


the

intestinale,

du

levain, glair euse, et des infusions."

He
M.

includes also the ferment which, according to

Pasteur,

occasions

" sickness of

turned

wine."
Finally, the genus Spirillum includes the species S. undula, tenue, volutans of

rufum and leucomcenum


Duj.

of Perty,

Ehrenberg, 8. and S. plieatile,

From
enters

this

moment

the history of the bacteria

upon a new phase. The labors of M. Pasteur upon the inferior organisms and their role in fermentation, the researches of MM. Davaine and Hallier upon the bacterium of charbon, and the
micrococci of contagious maladies,
tion of chemists
call

the atten-

and

of pathologists to these or-

20

THE BACTERIA.
Their

ganisms and especially to the bacteria.


origin, their
arities

evolution, the

physiological peculi-

of

their

the object of

and reproduction, are numerous labors, and give rise to


nutrition
to

passionate discussions relating

the subject of

spontaneous generation, polymorphism of fungi, theories of fermentation, and the pathology of For this reason virulent and infectious maladies. an exposition of these researches, often contradictory,
is

extremely

difficult.

We will make
upon the

it

suc-

cinctly, insisting especially

labors relating

to the classification of the bacteria,

and reserving
their

to ourselves the privilege of returning to the his-

tory of several points,

when we approach

study in the special chapters of this thesis. The first important memoir published
that of

after

M. Davaine upon the bacteria is that of M. Hoffmann, in 1869. He demonstrates First,


:

that the bacteria are plants, having a very distinct


cellular organization
;

second, that they can only


linear bacteria,

form and and the latter into microbacteria, mesobacteria, and megabacteria (M. Hoffmann includes with the linear bacteria, Vibrio, Bacterium, and Leptothrix, which are bacteria united in a chapletj) third,
be
classified

in accordance with their

size,

at

first

into

monads and

that mobility or immobility


acter,

not a specific charthe same species under the influence of changes of temperature, of
is

but

may

present

itself in

density of medium, etc.


also

M. Hoffmann

shidies

the origin of the bacteria, and rejects the hypothesis of a spontaneous generation. As to

HISTORICAL.
their role in the
of organic bodies

21

phenomena

of the decomposition

and in fermentations, M. Hoffmann confesses " that, with the exception of yeast, and of the acetic and butyric ferments, all the rest is still enveloped in obscurity." M. Cohn is the naturalist who, in our days, has occupied himself the most with the bacteria. In 1853, he published his first researches upon this subject. The genera Zoogloea, which he established at this time for the bacteria arranged in ge-

more or less crowded was not a happy creation. It was adopted at first by M. Eabenhorst who, in his work on the fresh-water algse of Europe, places them after the palmellaceas, while he classes the other bacteria, Vibrio and Spirillum, in the family of the oscillatorige. The Zooglcea are later abandoned by their author as a generic group, and are preserved
latinous masses, diffused or

together,

only as the
stages

name of one of the diverse transitory through which the bacteria pass in the
of

course

their

evolution (Zoogloea, Leptothrix,

Torula).

Twenty years
the

later the same savant commenced publication of a series of " Memoirs " upon

these organisms (in his " Beitrage zur Biologie der

Pflanzen").

In the

first

paper the author gives an

exposition of his researches upon the organization,


classification of the bacteria, and upon their action as ferments. M. Cohn considers them as a well-defined group,

development, and

the

schizospores, belonging to the algae, at the


of the series of the

commencement

phycochroma-


22
cese,

THE BACTERIA,

with several families with which the different genera of bacteria have many affinities. He recognized, however, that the absence of chlorophyll approaches them, at least from a functional point
of view, to the fungi.

Upon

this point

we may

say that for other botanists this character is decisive, and the bacteria are classed as fungi. M. N'ageli, who takes this view, describes them

under the name


the
bacteria
:

of Schizomycetes.

Cohn

divides
six

into

four

tribes,

comprising

genera

1.

2.
3.

4.

The Sphcerobacteria or globular B. The Microhaeteria or rod B. The Desmobacteria or filamentous B. The Spirobacteria or Spiral B.

We

will return to this classification.

In 1874, M. Th. Billroth, in his researches upon the Coceobacteria septica, expressed opinions entirely different

from those
bacteria

of

Cohn.

According

to

Billroth,

the

differ

considerably in

form according to the medium in which they are placed and divers circumstances. He claims that they constitute but a single species, the CoceobacThis vegetable organism can preunder the form of globular articles (coccos) or under that of rods (bacterie). These two forms may reproduce themselves by becoming elongated and dividing transversely, or may pass Billroth claims to have the one into the other. found both forms united in a single filament, a
teria septica.

sent

itself

HISTORICAL.
fact

23

which in

his

opinion demonstrates conclu-

their relationship. Each of these two forms can also present variations of size, in accordance with which he establishes the following

sively

divisions

Micrococcos Mesococcos Megacoccos

Microbacteria.

Mesobacteria.

Megabacteria.

And

varieties of association
:

following names

which give

rise to the

Monococcos
Diplocoecos Strep tococcos Gliacoecos
Petalococcos

Monobacteria.
Diplobacteria.

Strep tobacteria.
Gliabacteria.

Petalobacteria.

Ascoecos.

The following year (1875), Cohn, in the second part of his " Researches " upon the bacteria, criticised the opinions expressed

by Billroth

in the pre-

ceding memoir. Cohn believes that we should regard as distinct genera and species all the bacteria having a particular form and acting differently as ferments, so long as the proof of their
identity has not been demonstrated in an evident

manner.

Coming back

also to the affinities

and

classification of these

organisms, he insists
to

anew

upon
teria

their near relationship

the Phycochro-

maceae;
as

and, no longer distinguishing the bac-

special

family,

different genera in a group,

he distributes his which he calls Schi-

zophytes, which includes the greater part of the

24

THE BACTERIA.

Chrococcece and of the Osdllarice.

We

will re-

turn to this subject

when we speak

of the clas-

sification of the bacteria.

In 1876, appeared in the same number of Cohn's " Beitr'age " two important papers. The first, by Cohn, treats of. the influence of temperature upon the bacteria, of their origin, of the formation of spores in the Bacillus of hay infusion,

and

of

charbon.

gives the result of his

by Koch, researches upon the bacsecond,

The

teria of charbon, the Bacillus anthracis.

Koch has been

able

by

skilful

cultivation

to

follow the complete development of this Bacillus, and to witness the formation of spores, of which the vitality is very great, and which are the principal agents of

the transmission of this terrible

malady.
I

must

still

indicate, in addition to these special

works, a quantity of notes and of memoirs scattered through the reviews and periodical publications.

be found in the bibliography appended to this work. I must also cite the recent work of M. Nageli upon "The Inferior Fungi
list will

The

and

their

Bole in Infectious Maladies."

The

learned professor of Munich has studied the di-

produce decompositions. He the Mucorini, Saccharomycetes, and the the Schyzomicetes, which correspond to the bacteria. According to Nageli,
verse fungi which
divides

them

into three groups,

HISTORICAL.

25

the bacteria are fungi which produce putrefaction.

In presence of these opinions, so diverse, as to the nature of the bacteria and their classification, we will finish by saying with Conn " So long as the makers of microscopes do not place at our disposal much higher powers, and, as far as possible, without immersion, we will find
:

ourselves, in the
situation of

domain

of the bacteria, in the

a traveller

who wanders

in an un-

known country at the hour of twilight, at the moment when the light of day no longer suffices to enable him clearly to distinguish objects, and when he is conscious that, notwithstanding all his
precautions, he
is

liable to lose his

way."

PART

FIRST.

MORPHOLOGY OF THE BACTERIA.

CHAPTER

I.

ORGANIZATION OF THE BACTERIA.

When
eye.

bacteria

develop in a liquid in a

suffi-

cient quantity, they

become
either

visible to the

naked

They appear
little
its

as

a slight cloud, or
liquid, or

gathered in

masses in the

forming

a pellicle upon
in the liquid.

surface, or as a deposit

upon the

walls of the vessel and

upon the objects contained However, we must hasten to say

with M. Cohn, that the fact of the absence of all turbidity in a liquid does not exclude the possibility of the presence of bacteria. In liquids more dense than water (serum, lymph, etc.), when the
refractive

power

of these corpuscles

is

the same as

that of the liquid, their presence

may
will

revealed by the naked eye.

We
to

not be add that


their
is

sometimes

their

color

serves

indicate

presence in a liquid, although this color

often

very feeble, and can only be perceived when a considerable thickness of the liquid is examined. If we examine these clouds, these accumulations, these deposits, with the microscope, we see that

28

MORPHOLOGY OF THE BACTERIA.


little

they are formed of a myriad of


with motion, sometimes colored.

bodies iso-

lated or grouped, globular or linear, gifted or not

These variations constitute so many characters which require to be studied with some detail.

1.

Bacteria in General.
bacteria, as understood to-day

Form.
most
ies,

The

by

botanists,

when

considered in their separate

state, are of

two principal forms, or monads, and bodies more or

globular bodless filiform, or

bacteria properly so called.

The globular

bacteria comprise organisms round-

ed, ovoid, sometimes elongating themselves into a The Monas crepusculum of tube (Warming). Ehrenberg may be taken as a type. This form includes also the Micrococcus of Hallier, the Microsporon of Klebs, the round forms of the Amylobacter of M. Trecul, and perhaps the Microzyma of M. Be"champ. We will see farther on that

these are very probably phases of development of

the spores of bacteria, properly so called.

The

bacteria, not

globular, present a greater

diversity of

form

they

may

be straight, unduin

lating, or twisted in a spiral.

The

rectilinear

bacteria are

cylindrical throughout their


this case

some exactly whole extent and in


;

they form very short cylinders, as in the Bacterium, Cohn, or cylinders of which the length
is

several times as great as the thickness, as in


;

the Bacteridies {Bacillus ulna Cohn)

others are


ORGANIZATION OF THE BACTERIA.
29

swollen in the middle, with their extremities rounded) such as certain forms of Vibrio serpens ("Warming) ; others again are fusiform, swollen in the middle and attenuated at the extremities,

Bacterium fusiforme (Warming) rectilinear bacteria swollen at the two extremities are met during the life of certain species, B. lineola and B. termo, for example, above all when they are
;

transported to a more
ly,

favorable

medium
;

this
final-

modification usually precedes segmentation

one meets sometimes bacteria swollen at one extremity only; the swollen part presents often a clear point and sometimes an evident spore we
:

shall see later the signification of this peculiarity.

"With these claviform bacteria

we may

include the

Bacterium capitatum Dav., the Eelobacteria of Billroth, and certain Amylobacter, with heads of the Ficus carica, etc. (Ch. Eobin).

The undulating

bacteria constitute the Vibrios


etc.).

properly so called (V. rugula, serpens,

The
or less

spiral bacteria of

which the turns are more

elongated

are

named

Spirillum, Spiro-

chceta, etc.

Dimensions.
oscillate

The
it

dimensions of the bacteria


limits, but in' be said that they are the

between the most variable

a general
smallest of

way
all

may

microscopic beings.

Some

of

them

are situated at the extreme limit of our highest

magnifying powers j and their proportions, as to

30

MORPHOLOGY OF THE BACTERIA.


comprised within the

length and thickness, are

limits of errors of observation.

The globular bacteria are the smallest, and the dimensions of some species are so minute that
they cannot be measured directly.

The largest are the Spirillum, which attain a length of T2j of a millimetre. Between these two extremes, there are all intermediary sizes possible. The dimensions of some of the bacteria are given
below:

Monas
to

vinosa, 0.5 to 1

/i,

in diameter; length 3

4 p. Bacterium termo, breadth 0.6 to 0.8


Vibrio lineola, breadth 0.5 to 1
/a
;

\l\

length 2

to 3 p.

Bacillus ulna,

B.

anthracis,

0.7 to 1

p
fi

length 3 to 8 p. 5 to 8" fi.

to 2

/j, ;

,,10

Spirillumvolutans,

to 50 10 to 40

fi.

fi.

Several

authors,

considering
to

exclusively
their size.

this

character of dimensions, have divided the monera

and the bacteria according

Thus

Hoffmann recognizes
macrobacteria.
fies

in addition to the monera,

only the microbacteria, the mesobacteria, and the

In the same way Billroth classimonads according to their dimensions into micro, meso, mega coccos, and the bacteria into
the

micro, meso,

mega
differ

bacteria.

Finally, Klebs sep-

arates the Micrococcos

which do not

from the Microsporines, from them except by their smaller dimensions, both forms being able to pass
to the state, of bacteria (rods).

ORGANIZATION OF THE BACTERIA.


Cdlor.

31

The phenomena

relating to the color

of bacteria have only recently been pointed out.

" But little attention has been given tq the color of the bacteria, regarded generally as colorless," said M. de Seynes in 1874 ; and recently M. de Lanessan, " The bacteria are ordinarily quite color-

However, M. Cohn had already insisted upon the globular bacteria chromogenes, or of pigmentary fermentation, and upon the colors produced by different monads, which have long since been studied by microscopists.
less."

Upon
teria

this subject, let us

observe that the bac-

to two very diforganisms always known as such, but which were not formerly included with the bacteria, as the different monads, which have become the Micrococcus prodigiosus,

which are colored belong


First,

ferent groups.

colored

cyaneus, aurantiacus, Cohn, etc.; the second group includes

the bacteria properly so called, which

absorb the coloring matter of vegetables upon

which they are fixed as parasites, or of the media in which they live. This is the case with the bacteria observed by M. de Seynes upon the Penicillum glaucum, and perhaps with the Vibrio synxanthus and syncyanus^hreub., which give to milk

We

a yellow or blue color according to the species. will return to this subject when we speak of

the nutrition of the bacteria.

As

to

the

purple-colored monads, they have

been especially studied as early as 1838 by Dunal, then by Morren and Ehrenberg, and in our own day by Eay-Lankester, Cohn, Klein, and finally

32

MORPHOLOGY OF THE BACTERIA.'

by Warming and
rious

in sea-water, in hot sulphur springs, water containing animal or vegetable matThey appear someter in a state of putrefaction. times upon bread, meats, and in general upon

media

Giard.

They

are found in va-

in fresh

cooked food placed in a humid atmosphere. The different colors which they present are red, yelIt is probably to anallow, orange, and blue. ogous organisms that we must attribute the blue color presented by pus under certain circumstances, the green and blue color studied by M. Chalvet, and the orange-yellow, bright red, and blue colors observed by C. Eberth in perspiration.

In Norway, red bacteria appear in summer in such masses that the borders of the sea are sometimes colored of an intense red (Warming).

Movement.
ferent states.
it is

The bacteria
They

are

met

in

two
;

dif-

are active or motionless

but
that

now

well settled for the greater

number

the same species

may

present

itself

sometimes in

a state of repose, sometimes in a state of move-

ment.

a movement

two kinds, and a movement of translation. The first is sometimes nothing more than a molecular or brownien movement, which occurs in the smallest forms. But at other times it is more extended, and consists in a movement of rotation round the axis, or a bending of the body. This flexibility is, above all,
bacteria are of
of the corpuscle

The movements of the

upon

itself

ORGANIZATION OF THE BACTERIA.

33

observed in the long forms, the Bacillus, the Vibrions, etc.


it is

As

to
;

the
at

movement

of translation,

very variable
it
is

rapid,

one time slow, at another in relation with the length and form

of the bacterium.

the modifications of
lines
:

M. Cohn has well described all movement in the following

" Almost

all

the bacteria possess two different


characterized

modes of life, movement.


" In
certain

by repose and by

they are excessively mobile ; and when they swarm in a drop of water, they present an attractive spectacle, similar to that of a swarm of gnats, or an ant-hill. The bacteria advance, swimming, then retreat without turning about, or even describe circular lines. At one time they advance with the rapidity of an arrow, at another, they turn upon themselves like a top; sometimes they remain motionless for a long time, and then dart off The long rod-bacteria twist their like a flash. swimming, sometimes slowly, sometimes bodies in with address and agility, as if they tried to force It for themselves a passage through obstacles. is thus that the fish seeks its way through aquatic They remain sometimes quiet, as if to replants.
conditions,

pose an instant: suddenly the


to oscillate,

little

rod commences

and then to swim briskly backwards, to again throw itself forward some instants after. All of these movements are accompanied by a second movement analogous to that of a screw which moves in a nut. When the vibrios in the
3

34

MORPHOLOGY OF THE BACTERIA.

shape of a gimlet turn rapidly round their axis, they produce a singular illusion one would believe that they twisted like an eel, although they are extremely rigid." The causes of these movements have been sought,
:

supposed animal nature of the bacand the movements assimilated, consequently, to voluntary movements but this opinion can no longer be sustained, as similar movements are to be seen in a great number of vegetable organisms, such as the diatoms, the oscillatoriaa, the spores of They have also been algae and some fungi, etc. attributed to the existence of locomotor appendices (Ehrenberg); but, although the cilia, denied at first by most microscopists, have been seen since in nearly all the bacteria, the botanists who have best studied them, M. Warming, for example, recat first, in the
teria,
;

it is scarcely probable that these organs are the cause of their movements, for " one meets some examples in which the body remains motionless while the cilia are in violent agitation, and others in which the body moves while the cilia remain inert, or dragging behind." The movements appear to depend rather upon

ognize that

the nutrition, or respiration, and especially upon the presence of oxygen (Cohn) ; indeed when this

wanting the bacteria become motionless. Immobility may also be produced by want of
gas
is

nutriment, poisoning by different toxic substances,


(chloroform, iodine,
ters derived
etc.),

dessication, etc.

The attempt has been made

to use the charac-

from the existence or absence of

ORGANIZATION OF THE BACTERIA.

35

motion, and the form of the bacteria, in order to


classify

them

clearly that these transitory

but what has just been said shows phenomena cannot be


specific characters.

taken for generic or


Structure.

It was for a long time believed that the bacteria were constituted of amorphous masses

of protoplasm, or of solid rods.

The researches

of

Hoffmann have shown


lar

that they have a true celludescribe, then, succescilia,

structure.

We

shall

sively, their

membrane, the contents, and the

which

may

be considered as belonging to the pro-

toplasm.

Cell-membrane.

The

extreme minuteness of

the bacteria usually prevents a direct demonstration of the cell-membrane, and the existence of

envelope has not, heretofore, be^en clearly demonstrated except by indirect proofs chemical reactions, for example.
this
;

Thus Hoffmann
lar envelope

verifies the existence of a cellu-

when

" the contents, which

is

a trans-

parent plasma, are partly coagulated, as sometimes

happens, or disappear, and are then replaced by air which shows precisely the form of the normal Warming, also, has not been able bacterian cell." to see the membrane, " which only appears distinctly

when

a vacuole has formed just against the

periphery."

On

the other hand, the action of chemical agents

upon bacteria proves that they have an envelope of cellulose, which is colored by tincture of iodine

36
is

MORPHOLOGY OF THE BACTERIA.

not destroyed by caustic potash, ammonia, or even acids and resists putrefaction for an ex;

ceedingly long time.


bles the

In this respect,

it

resem-

membrane

of cellulose of vegetable cells

(Cohn).

We should add that Cohn claims to have succeeded with high powers in seeing directly the cell-membrane. On the other hand, Warming has never succeeded in so doing. The last observer remarks also that the resistence of bacteria to acids, to alkalis, etc., does not seem to prove the existence of a membrane, " inasmuch as this may
be the result of a particular condition of the plasma, which in all the bacteria is of a more consistent nature than in other plants."
Finally, the
teria,

membrane may
flexible

be, in certain bac-

tender,
of

ments

torsion.

and susceptible of moveIn others, it is rigid and

incapable of bending.

Cohn

thinks also that

it

may

swell and dissolve into mucilage, a fact

which

would explain the origin


Zooglcea.

of this substance in the

Protoplasm.

The

contents of the

cell is

nitrogenous substance, generally colorless,


highly refractive than water.

more

In the smallest species, this protoplasm appears homogeneous but in the bacteria of medium size, and above all in the large species, the contents of the cell encloses portions more highly refractive, vacuoles, special granules, and sometimes diverse
;

coloring matters.

ORGANIZATION OF THE BACTERIA.

37

Cohn has first pointed out the movements of the protoplasm, in which currents occur, above all in
the central portion, the peripheral portion remain-

ing homogeneous and motionless. The vacuoles are also found in the central portion Warming,
;

however, who has observed them in Monas Okenii, Vibrio rugula, V. serpens and Spirillum undula var. littoreum, has sometimes seen them in the middle of the plasma, at another near the exterior wall. The granules which are seen in the protoplasm were considered by Ehrenberg as stomachal vesicles or ovules. They are of two sorts the one, highly refractive and not bordered by a dark circle, are considered by Warming as nothing more than mere compact- masses of protoplasm; the second, also highly refractive, but surrounded by a dark circle, resemble drops of oil, and have been taken for fat granules but the recent researches of Cramer, Cohn, and Warming have proved that some of them, at least, are formed of crystalline sulphur. They are not soluble either in hydrochloric acid or in water, but they are dissolved in absolute alcohol, in hot caustic potash and sulphite of soda, in nitric acid and chlorate of potash at ordinary temperatures, and in bisulphide of carbon, when the membrane, which is permeable with difficulty, has been previously destroyed by sulphuric acid. Although their small dimensions and great refractive power prevent them from being distinguished with certainty as crystals of sulphur, as they are doubly refractive to polarized light their crystalline nature cannot be doubted.
;

38

MORPHOLOGY OF THE BACTERIA.

These globules of sulphur have been observed in Monas Okenii, Bacterium sulphuratum, Ophidomonas, and the different species of Beggiatoa, both in fresh water, in putrid sea-water, and in thermal sulphur waters. It will be seen when we speak of the physiology of these organisms what their role is in the elimination of sulphur and the formation of sulphuretted hydrogen. We have said, in speaking of the colored bacteria, that some borrow their color from the surrounding medium, and that others, on the contrary, have a color of their own. The protoplasm of the latter contains a granular coloring matter, which
is

ordinarily yellow, blue, or red.

The red

color-

is most common, and this has been best and appears to be the best known. One of these colors which gives a pink tint (peach color) to Bacterium rubescens, Ray-Lank. Clathrocystis roseopersicina, Cohn) 3Ionas vinosa, (

ing matter
studied,

Cohn; M. gracilis, Warming; Rhabdomonas rosea, Cohn; M. Warmingii, Cohn; Ophidomonas sanguinea, Ehrb. ; Merismopedia littoralis, Rabenh. etc., has been studied by RayEhrb.,

M.

Okenii,

Lankaster,
in

who has given


ether,

to

it

the

name

of bac-

terio-purpurine.
alcohol,
oils,

It is insoluble in water, soluble

fatty

carbolic

acid,

glycerine,
it

and

characteristics

which make

resemble

spectrum. Other red coloring matters which appear to be different have been found in Monas prodigiosa, Ehrb.; Bacillus ruber, Cohn; and Micrococcus fulThese should not be confounded with vus, Cohn.
chlorophyll.

It has also a characteristic


ORGANIZATION OF THE BACTERIA.

39

the purple coloring matter of other algae, as that of the Porphyridium cruentum, which comes from a mixture of chlorophyll and of phycoerythrine.

The
In

bacteria never contain chlorophyll.


this connection, it is interesting to recall the

protoplasmic constitution of the Amylobacter of Tre'cul. These organisms are, according to Van

Tieghem, bacteria, to which he has given the name of Bacillus Amylobacter, and which does not differ from B. subtilis, except by a specific character, extremely transitory, the presence of amorphous starch, formed and stored in reserve during the period of growth, to be again used later, and con-

sumed during the process


Cilia.

of reproduction.

These appendages which were described


in the Bacterium
trilocular

by Ehrenberg
permit
vis

have

not been seen since.

To-day, recent researches

doubt in Bacillus, Bacterium, Spiall the true bacteria, rillum. They have been perceived in a great
to say that cilia exist without

number
Vibrio

of forms,

Spirillum volutans, Sp. undula,


Cohnii,

rugula, Spiromonas

pens, and several species of Bacillus.


in the smallest of the bacteria that
it

Vibrio serIt is only has hitherto

been impossible to demonstrate their presence. They have, however, been recently seen by Dallinger and Drysdale in Bacterium termo.

Warm-

ing has perceived as

many

as

two or three on one

extremity in Ophidomonas sanguinea Spirillum


volutans var. robusium, and Vibrio rugula.

PLATE

I.

Taken from " Monthly Microscopical Journal" of

Sept. 1st, 1875.

B. termo magnified with the same power as which diameters). B. termo, seen with flagellum at one end, the light comFig. ing in the direction of the arrow. The same object when moved at right angles to Fig.
Fig
1.

a.

6,

is

a specimen of Spirillum volutans, showing flagella at each end. Fig. 2. B. termo, as seen with a power of about 600 diameters. Fig. 3. The same as seen with $ and second eye-piece (3,700
4.

5.

it

its

former position, the light coming from the same direction, causing the sight of the flagellum to be lost. Fig. 6 represents one B. termo which was in a still condition, but one flagellum moving. The light came in the direction of the arrow. When the end marked 2 b was in focus, a flagellum was seen, but none at the end c. When the end marked 1 a was focused carefully, the flagellum at that end was seen, and lost at the

end

d.

Fig. 7. The true form of B. termo. Fig. 8. The form as shown by the " supplementary stage " illumination before flagella were found, showing the pointed termination of the body at a, b.

The Monthly Microscopical Journal Sep 1 1.1875

Plate

:v_

-7*"""

<xxz>

"V

a<

FtAGELLA ON
By W.
H.Dallinger,

FRM.S

BACTERIUM TERMO and J.J. Drysdale, M D.ER.M.S

ORGANIZATION OF THE BACTEBIA.

41

EXTRACT FROM PAPER "ON THE EXISTENCE OF FLAGELLA IN BACTERIUM TERMO," BY W. H. DALLINGER, F.R.M.S., AND J. J. DRYSDALE, M.D.,
F.R.M.S.
" In the summer of 1872, some very fine specimens of S. volutans came under our notice, and were carefully examined. We were enabled fully to confirm Cohn's discovery, and demonstrated repeatedly the presence of a pair of swiftly lashing flagella. The drawing at b, Fig. 1, was made from a specimen magnified 1,300 diameters (diminished by |). " Having closed for the present our Monad researches, we have been stimulated by the hope that the experience gained by these might enable us to prosecute similar investigations into the true life history of bacWe have commenced the work this summer, and, guided by the teria. analogy of 5. volutans, we have been led to make several continuous efforts to find whether or not there existed a flagellum or flagella in B. termo. The task, of course, under the best circumstances, must be a a difficult one, from the extreme minuteness of the object. We tried each of Powell and Lealand's powers successively, from the i\ to the -^, but with no definite result. Repeatedly we both saw vortical action at both the distal and proximal end of the termo, but could not absolutely see the organ causing it. But in the process of our investigations we made very close and careful observations on the fission of this form we do not purpose now to describe the process, but merely to point out a. phenomenon that further confirmed our suspicion of the presence of an invisible filament. In separating into two, the jointed rod of sarcode which is in process of division shakes to and fro at the constriction, as if the constricted part were a hinge and at length a clear separation takes place to quite the length of the original termo (sometimes longer), and there is no visible connection between them ; nevertheless they act as one crea:

ture, so that if one moves in any direction, the other goes with it, just as the two parts did before separation; showing that, although we cannot see the con-

nection, there

filament, such as

must be one; and the presumption was that it was a fine we detected in the fission of some monads. 1 We could
;

make no
was
'

called to the

further progress in the question apparently but our attention new Jth objective prepared by Messrs. Powell and

We used it at first with Lealand, with which we were soon supplied. the supplementary stage ' for very oblique illumination, supplied by the same makers, and this has the advantage of throwing the light in only from one direction. We were soon convinced of the exquisite performance of the glass when used as an immersion. Amphipleura pellucida was not merely seen to be striated clearly and sharply, but the stria}
" M. M.
J., vol. *., p. 55;

and

vol. xi., p. 8."

42

MORPHOLOGY OF THE BACTERIA.

were resolved into beads with the third and fourth eye-pieces. In like manner the fine striae in Surirella gemma were instantly shown to be beaded, with perfect and brilliant definition, with the second eye-piece. Navicula rhomboides and an extremely delicate specimen of Pleurosigma attenuatum which had resisted everything below a ^th immersion, showed beaded striae perfectly. We were therefore encouraged to try again to
discover flagella in the termo. Some of our specimens, nourished in Cohn's nutritive fluid, were placed in a drop of distilled water, and put upon the supplementary stage on an ordinary slide covered with the thinnest cover. The utmost delicacy and tact in manipulation of the
light
is

the great desideratum

but, with this,

enough may be secured

to

be made to enter the objective at almost every angle, but it is always projected in a direction at right angles to the stage ; and the first thing we observed when the objects were sufficiently slow in their movements, and at right angles to the light, was that the ends of the termo, which we (and all other observers, as far as we know) had taken for round, proved themselves to be conical, terminating in a sharp point. The usual appearance of B. termo, as seen with a magnification of about 600 diameters, is seen in Fig. 2 whilst the same seen with a magnifying power of 3,700 diameters [Jtrth and second eye-piece) is seen in Fig. 3, where a globular granule is seen in the end of each half. But with the method above referred to, the best conditions being secured, the two ends of the bacterium were distinctly pointed, as seen at a b, Fig. 8, and after nearly five hours of incessant endeavor a flagellum was distinctly seen at one end of each of two termos which were moving slowly across the field. The discovery was not sudden and transient, but lasted for at least twenty minutes. The exquisitely delicate flagellum was lashing rapidly the whole time and one of its frequent conditions is shown in Fig. 4, the arrow indicating the direction of the light but if the termo turned round at right angles, as in Fig. 5, all trace of the flagellum was gone, showing that its discovery depended entirely, all things being equal, upon its position in regard to the light. " But this observation was made only by one of us, the other not being present and in pursuance of our plan we determined to see it again, convincing ourselves separately, and then together. After many hours of labor, this was accomplished and Fig. 6 shows one of two instances which we both saw together at the same time and in the same instrument. It was lying still, obliquely across the field, the light coming in the direction of the arrow. Both ends were not perfectly in focus at the same time, but in focusing the end marked 2 b (Fig. 6) the flagellum was distinctly seen, and was seen also to coil and lash but no flagellum was then seen at the end c of the same object ; but by bringing it into delicate focus it presented the aspect seen at 1 a (Fig. 6), which really represents the same object at the same time, only with the other end in the focus, while the end marked d corresponding to 2 b of Fig. 6 was in its turn slightly out of focus, and the flagellum lost to view. This observation, made together, was as satisfactory as could be desired and it

work with the fourth

eye-piece.

The

light

may

ORGANIZATION OF THE BACTERIA.


was thus demonstrated that there was a flagellum
dinary B. termo.
" It will of course be understood that
;

43

at both ends of the or-

it is by no means an easy matter and whenever we repeat it, it must always be with nearly the same amount of trouble and patience, although we

to secure the demonstration

front and (occasionally) behind the termo, as

can now with the ordinary condenser detect the vortical action, both in we never did before. But the demonstration of the ultimate structure of a fixed object as for must be looked upon as a matter of instance Amphipleura pellucida great ease in comparison and that for many reasons, the foremost being the motion and the minuteness of the object, the swift play of the flagella, their similarity in optical properties to the fluid in which bacteria live, the difficulty of retaining them in focus, and of getting them in such a position in relation to the light as to make demonstration possible. Of course, all this would be removed if dead bacteria would answer, but they very rarely (indeed only once) have done so with us. The flagellum needs to be in slow motion to properly show itself for even with the more delicate and minute monads it is a difficult thing to show the flagella in dead forms, since in the majority of cases they appear to be attracted round the body of the creature."

2.

Of
states,

the Different Modes of Grouping of the Bacteria.

The
two

bacteria are found in different media

in

free, isolated (unicellular bacteria),

or united several together, either in chains, in


filaments, or in masses of greater or less extent,

and sometimes by the aid of a mucous substance which they are imbedded. The free unicellular bacteria are found in the When they are Spirillum, Bacillus, Monas, etc. united, they are grouped in the following modes
in
:

The usual method


bacteria
is

].

Form

of a

little

chain:

Torula, Leptothrix.

of multiplication

among
;

the

by fission

(" scissiparite")

each cor-

puscle divides transversely, and gives birth to two

44

MORPHOLOGY OF THE BACTERIA.


individuals,

which sometimes become separated completely the one from the other, to form unicellular bacteria, sometimes remain united; and

new

segmentation again occurring in each portion, a chain is formed of articles more or less numerous.

When
teria,

these chains are formed of spherical bac-

they have been called torulce; if they are formed of filiform bacteria, they correspond to The morphological difference leptothrix (Robin).

between the torula and the


the fact that in the
first

leptothrix consists in

the articles are separated


is

by

constrictions, while this


It is also to

not the case in the

be remarked, according to Cohn, that the microbacteria never take either Warming states, however, that of these forms. he has met the torula form in Bacterium lineola, B. catenula, and B. termo (?). Billroth has called these two forms of bacteria He has even sireplococcos and streptobacteria. considered it necessary to create the words diplococcos and diplobacteria for organisms constituted only of two articles.
second.
2.

Form

of Zooglaza.

Generally, when bacte-

ria are

rapidly multiplying, they remain grouped

in masses, swarms, or Zooglcea.


dition,

In the latter conthey are closely pressed against each other in the midst of a viscous substance, hyaline, homogeneous, colorless, and constituting masses

more

or

less

diffused

or

definite,

in

irregular

globes, bunches, or tubes,

swimming

in the water

or near

its

surface.
cells

When

the bacteria multiply

abundantly, the

become removed from each

ORGANIZATION OF THE BACTERIA.


other, so as to leave
vals.

45
inter-

between them greater

The masses sometimes

attain a diameter of

several centimetres.

The

gelatinous substance in which the bacteria

are included seems to be produced

by

a thicken-

ing and jellification of this cell-membrane, or by a secretion from their protoplasm, but the latter

view seems more plausible than the former (De


Lanessan).
It is

commonly the

spherical bacteria (Micro-

coccus) and the microbacteria (Bacterium) which are found in this state.
spirilla are never found in gelatinous masses (Cohn). Ray-Lankester, however, claims to have met the Spirillum tenue, in the form of zoogloea, and Klein the Spirillum undula and rosaceum (Warming). The form of Zoogloea, properly so called, gelatinous and thick, has never been found by Warm-

The

filiform bacteria

and the

ing in infusions of sea-water.

According to the terminology of Billroth the are called gliacoccos and glidbacteria (from y\la, mucus substance). 3. Form of Mycoderma. In certain cases, the bacteria unite on the surface of the water, or of the liquid in which they are developed, to form a
zoogloea

membrane. This production mycoderma by Pasteur is a sort of zooglcea, but differs from it by the absence of the intermediary mucus substance. The bacteria are, however, motionless, although living, since they come
thick layer, a sort of
called

to the surface to be in contact with oxygen,


is

which

necessary to them.

,46

MORPHOLOGY OF THE BACTERIA.

The petalococcos and

petalobacteria of Billroth

correspond with the mycoderma of Pasteur. 4. Swarms. We have seen that the filiform and spiral bacteria do not, usually, form zooglcea.

These microphytes are either disseminated and free, or united in swarms. This formation may be seen, for that matter, in all the bacteria, when, thanks to abundant nourishment, they multiply rapidly and gather together in considerable masses. They are very active in these swarms, whilst in the zooglcea the corpuscles are motionless, because of
the intermediary glairy substance.

Pulverulent precipitate. When the nutritive elements are exhausted in a liquid, the bacteria
cease to multiply, fall to the bottom of the receptacle,

deposit formed in this

ness
teria

becomes clear. The manner may acquire a thickvery appreciable to the naked eye. The bac-

and the

liquid gradually

in a state of

which form this precipitate are not dead, but temporary repose and if a new supply of nutritive material is added to the liquid, they are seen to multiply anew, until this has been exhausted (Cohn).
;

Plate

II.

'.-.iwj y
3Hf
*

% ^

v"

Fig.

Fig

^p

-,>

mm
3.

Fig.

Fig. 4.

PLATE

II.

DIFFERENT MODES OF GROUPING.

From photo-micographs made

in

Havana and New Orleans ;

copied by

permission of the National Board of Health.

Fig. 1. Torula form of spherical bacteria (Micoderma aceti Pasteur) from rotten banana, New Orleans, April, 1880. 1500 diameters by Zeiss's -^ in. objective.

cell

Fig. 2. Zoogloza form of spherical bacteria developed in culturecontaining blood of leper. 600 diameters.

Mycoderma, from surface of foul gutter-water. New Fig. 3. Orleans, April, 1880. 400 diameters by Becks, in. objective. Fig. 4. Leptothrix form of desmobacteria developed in yellow-

fever urine, exposed in laboratory, Havana, July, 1879.

400

diameters by Becks, \ in. objective.

CHAPTER

IT.

CLASSIFICATION OF THE BACTERIA.

1.

Position

of the Bacteria.

The
for

place of the bacteria in the scale of beings,

demands to be not only for the naturalists, who only view the question from a systematic point of view, but above all for the biologists who study the role of these organisms in the chema long time undetermined,
;

established with precision

phenomena with which they According to Ch. Robin, not to define the animal or vegetable nature of these organisms, " is for them as grave as it would be for a chemist to leave undecided the question as to whether it was nitrogen or hydrogen, urea or stearine, which he had obtained from a tissue, or of which he is following the combinations in certain
ical

or pathological

are associated.

operations."

This determination
there are
still

is,

to-day, possible

and,

if

some

differences of opinion

naturalists as to the place of the bacteria

among among

the cryptogams, there


their vegetable nature.

is

but one opinion as to

It is surprising to see a savant like M. Pasteur " not to pronounce positively upon the vegetable

CLASSIFICATION OF THE BACTERIA.

49

or animal nature of several of the ferments which

he has studied," and of which some belong to the


bacteria.

We

shall

first

indicate rapidly the characters

which permit
if

us, at first, to recognize certain spe-

cies of bacteria as organized beings, to

determine
finally to

they are animal or vegetable, and

classify

them

either

among

the algae or

among

the

fungi.

Distinction
stances.

The question

of Bacteria

from Inorganic Subwhether bacteria are

as to

organized beings can only be raised in relation to the smallest species, those Micrococci which are scarcely perceptible with the highest powers the
;

organized nature of the other organisms of the

same group has never been questioned, even by


the earliest observers,

who

all,

since

Leeunhoeck,

have, without exception, taken them for animals But the smallest forms of bacteria or vegetables.

may

be confounded with various matters, with organic particles, molecular granules, fat globules, " These productions, which are found in conetc.
siderable quantity in the liquids or in the tissues
of animal or vegetable origin, often resemble so
closely, in form, size,

and grouping, the spherical

bacteria, that

it

is

absolutely impossible to guard

one's self against confusion, unless the most mi-

nute precautions are taken in making the observations " (Cohn).

The

detritus, the

amorphous powder

of precipi-

tated molecules of inorganic substances, even

when

50

MORPHOLOGY OF THE BACTERIA.

they exhibit the brownien movement, are t easilyenough distinguished from Micrococci by optical
their angular form, their less refractive power, and finally by their reaction with certain chemical agents above all if they are mineral
signs,
;

substances, crystalline bodies, etc.


It will

not be the same with molecular granules

of organic nature.
ters, their

They have

as

common

charac-

power, movements.
less regular,

rounded form, their notable refractive Nevertheless, their form is

more angular, their color variable, their power always less. In doubtful cases, Tiegel has given a method which enables us to distinguish them from Micrococci. It consists in warming the glass slide which supports the correfractive

they are " Coccos," they are seen to move in a manifest manner. This does not occur in the case of molecular granpuscles under examination,
if

ules.

It is
difficult

these

productions which render

it

very

phenomena which occur during the coagulation of milk. The caseine septo observe the

arates in the form of extremely minute globules having a very rapid molecular movement. But we may distinguish these from bacteria by the use of liquor potassae, which dissolves the former without attacking the latter. As another example of pseudobacteria, I will mention, after Cohn, the form which fibrine assumes when it separates from the plasma of the blood. It disposes itself in very slender filaments, closely resembling filamentous bacteria.

CLASSIFICATION OF THE BACTERIA.

51

Fat globules, which are found of all sizes, are often of the same dimensions as Micrococcus, and are very difficult to distinguish from the latter. The difference in refractive power is slight, and the action of re-agents, such as ether, is not certain in

mucilaginous solutions.

Hiller,

who

has

paid especial attention to the means of recognizing


bacteria, divides

them

into
:

two groups
1.

A. The optical signs


teristic

comprising

The

charac-

vegetable form, rods, leptothrix, this he


little

recognizes as of

use, as in this case there

is

no doubt; 2. The characteristic movements of the monads; 3. The mode of growth and of multiplication 4. The mode of junction of the granules. B. The chemical signs 1. False zoogloea become softened and diffluent under the action of liq. potassae, and are coagulated by the direct applica; :

tion of alcohol

2.

In sections of tissues, after an


liq.

hour of maceration in
the

potassae, diluted y^th,

monads

are colored

brown by

iodine, while fat

granules are not.

But, in truth, the method of cultivation, extolled by Cohn and Wollf, is the best means of " The distinction of distinguishing the bacteria.
pseudobacteria," says the first of these authors, " from veritable globular bacteria is a problem

which our microscopists cannot


a study of their
distinction can be

resolve, in every
It is only

case, with the desirable certainty.

by

mode
made.

of

development that this The globules which di-

vide

and develop in form of chains are organized beings ; when this does not occur, we are dealing

with pseudobacteria."

52
This
Nageli,
is

MORPHOLOGY OF THE BACTERIA.


not, however, exactly the to consider

opinion of
as the

who seems

movement

surest distinctive characteristic. " There are," he says, " but three

distinctive

signs which enable us to recognize with some certainty that granules under observation are organisms, spontaneous movement, multiplication, and equality of dimensions, united with regularity

of form.

"The most certain character is movement in a straight or curved line, a movement which inorganic granules never present. One should take care not to be deceived by movements which are caused by currents in the liquid under observation. Nor should one allow himself to be deceived by the tremulous motion, called molecular movement, in which the granules do not really change their position. These movements are seen in most cells, and even in those of the Schizomycetes, and inorganic bodies themselves present it.

" Multiplication

is

character less important

than movement.
are

found united in

When among granules some pairs, it may be supposed

with probability that division and multiplication are taking place. When rods are bent at an angle,

one may predict

their

division in

two

parts.
dif-

" Finally, as to size and form.


ferent size and of a

Granules of

more or less irregular form ought not to be considered as belonging to the group of segmented fungi; if, on the contrary,
is

the granules offer dimensions perfectly equal, and

a spherical or oval form, the distinction

more

CLASSIFICATION OF THE BACTERIA.

53

uncertain

they

may

belong to the schizomycetes

or be of inorganic nature."

Place of the bacteria among organized beings. Distinction between animals and vegetables. The

characters which serve to distinguish the inferior

animal organisms from the inferior vegetable organisms are of two orders, optical and chemical. A. The optical characters are drawn from the general form, the movements, and the mode of
reproduction.

The morphological

characters have

no value

except among the larger species of bacteria. If we bring together a Spirillum and a Spirulina, Kiitz., their affinities will be apparent to every
one.
It is

not the same for the large species of

Bacillus, of which the relations with the Oscilla-

The rod form seems very spenecessarily imply the vegenot does cial, but table nature of the organisms which possess it.
toria are evident.
it

Finally, the spherical bacteria,

crococcus,

resemble

Monas
their

entirely

by

and Miform some


It

infusorial animals.

is

Movement is not a more special now well proved that it does not
and that
it is

character.

sively to animals,

belong exclumet with in a ceral-

tain

number
fact,

of the inferior vegetables.

In

the anatomical characters are not


reliable
first,
;

ways absolutely alone that Cohn

but it is from these then Davaine, have recog-

nized the bacteria as vegetables.


B. Chemical characters.

Robin depends upon

54

MOEPHOLOGY OF THE BACTERIA.

these characters to demonstrate the vegetable nature of the bacteria. He takes for point of de-

parture the notions of general physiology as given by De Blainville in the following points
:

1.

We

find in animals various elementary sub-

stances of
ciprocally.
2.

the same kind

as

in plants,

and

re-

The ternary compounds predominate, how;

and the quarternary, nitrogenized, are more abundant, on the contrary, in animals. 3. In both, the fundamental cellular structure at least originally for the greater is the same number, and always in the most simple of organever, in plants
;

ized beings, etc.

" It results from this, then," continues M. Robin, " that so long as there is no digestive tube one can only distinguish plants from animals by the study of their elementary principles, and of the chemical reactions which these exhibit in general

by the

study, in particular, of the reactions which

the predominance of ternary cellulose principles

over all others gives to plants, and that of nitrogenized principles in animals, at all periods of
their existence."

Starting from this basis, Robin

made numerous

attempts to find in liquor ammonia, concentrated,


as prepared for use in laboratories, a reagent for

corpuscles of a vegetable nature.

In

effect,

am-

monia
mals,

dissolves the eggs, the embryos, of all ani-

the

bodies

of

all

the
etc.,

inferior

infusoria,

attacks the spermatozoa,

whilst

it

leaves ab-

solutely intact all the varieties of

cellulose

and

CLASSIFICATION OF THE BACTERIA.


the anatomical reproductive

55

elements of plants,

whether As to

it is

used cold or boiling.


other

the

chemical characters praised

during recent years,

we will content ourselves with mentioning concentrated acetic acid, which causes all animal tissues to become pale, whilst it is without action on bacteria (Luckonvsky) iodine, and sulphuric acid (Letzerich), etc. Hematoxyline (Luckonvsky) and fuschine (Hoff;

mann)

color the bacteria deeply.


to give

One ought,
microscopic
etc.,

then,

no longer
malcules,
error.

to

the

bacteria, as

do some
ani-

recent authors, the

names

of

infusoria,

microzoa,

and

other

expressions without precision, or consecrating an

Let us add that some naturalists of high

re-

pute, Hackel for example, have created for these

minute beings, monera, protoplasts, flagellata, diatoms, etc., an intermediary kingdom between the animal and vegetable, the Protista. Place of the Bacteria in the Vegetable Series. The vegetable nature of the bacteria once established, it remains now to determine to what class of vegetables they belong. Are they, algae or are they fungi ? This is the question which divides the naturalists. It is true that it is to-day very difficult to find a characteristic of these two classes of vegetables, both having, in a general manner, identical forms,

similar reproductive apparatus, etc.

and,

if

it

is

impossible to confound a Basidiomycete with a


Floridese, for example,
it

is

not the same

when

56

MORPHOLOGY OF THE BACTERIA.


Species.
is

one studies the inferior


acter

The 'only

char-

which appears general chlorophyll in the algae and


fungi.

its

the presence of absence in the


are obliged to

But,
it

if

we adopt
all
its

this distinctive character^

and apply
separate

in

rigor,

we

in

the inferior algae some forms very

nearly related, and which do not differ from their relations except in this particular. And this is ex-

what happens in the case of the bacteria. In truth, the bacteria, although entirely without chlorophyll, have numerous affinities as to form, movement, etc., with the oscillatoriacece^ and, according as one or the other of these characters have appeared to predominate, the bacteria
actly

have been classed


It
is

as algae or as fungi.

thus that Davaine, Rabenhorst, then Cohn,


all

Struck above
of grouping,

by the resemblance
of

of form,

mode

and

multiplication,

the bacteria
all,

among

the algae.

have placed Cohn insists, above


;

upon the
and

affinities of

the filiform bacteria with


of the micrococ-

the beggiatoa and the leptothrix


cus,

of the bacterium^ with the chroocoocacece.

He

at first placed
;

them

at the

commencement

of

this last series

but

we

shall see further

on that

in his last publications he has disseminated

them
insist

among

the oscillatoriaceae and the chroococcaceae.

Eobin and Nageli, on the other hand,

rather upon the affinities of the bacteria with the

yeast plants, which are incontestably fungi, and

they include them in this class. Robin says expressly " All the corpuscles deScribed under the name of Bacterium termo> B,
:

CLASSIFICATION OF THE BACTERIA.

57

punctum,

etc.,

Zoogloea, Micrococcus, and

many

others, are vegetable cells, spores of fungi, of several distinct species certainly
;

spores, or repro-

ductive bodies of the


another, either

first

order, derived one


fission,

from

by germination,

or from a

mycelium
the

reproductive bodies, in a word, of the

order of those which Tulasne has arranged under

name

of conidia, etc."

Nageli establishes in the inferior fungi which produce decompositions three very natural groups.
1.
2.

Saccharomycetes, or budding fungi, which produce the fermentation of wine, beer, etc.
3.

The The
The

Mucorini, or mould fungi;

Schizomycetes, or fission fungi,

which pro-

duce putrefactive^processes. This last group is formed of our bacteria (Micrococcus, Bacterium), etc.

Sachs solves the question by uniting the algae and fungi in a single group, the thallophytes, in

which he establishes two series exactly parallel, one comprising the forms with chlorophyll; the other, the forms which are deprived of it, and preserving in a transverse direction the morphological affinities of these organisms.

As
think

this classification is
it

yet but

little

known, we
:

best to give

it

in the following table

THALLOPHYTES.
Forms
with chlorophyll.

Forms

without chlorophyll.

Cl.

1.

Peotophytes.
A'. Schizomycetes (Bacteria).
B'.

A. Cyanophyceae
B. Palmellacese.

(Oscil-

latoriacese, etc.).

Saccharomycetes

(Ferments).

PLATE

III.

Saccharomycetes and Schizomycetes (Nageli), developed in urine (of yellow-fever patient) exposed in laboratory of the Yellow-fever

Commission, Havana,
the

My,

1879.

Reproduced by permission of

National Board of Health.


1.

Fig.

and

Tolles's amplifier.

Fig. 2.
objective.

Photo-micrograph made with Beck's objective 400 diameters. Photo-micrograph made with Zeiss's ^g-in. horn. im.
-in.

1,450 diameters.

Plate

III.

Q
i

'

f--

m
'
<

^L.
Fi<;

HcliotyiM' Printing

<-'u.,

BhfIii

classification of the bacteria.


Cl.

59

2.

Zygospores.
A'. Myxomycetes.
B'.

A. Volvocineee.
B. Conjugueee and Diatoms.

Zygomycetes,

Cl.

3.

Oospores.
Saprolegnise.

A. Sphseroplese.
B. Coeloplastese.
C. GEdogonise.

Peronosporese.
4.

Cl.
B. Floridese.
C. Characese.

Carpospores.
A'. Ascomycetes.
B'. CEcidiomycetes.

A. Coleochaetese.

C.

Basidiomycetes.

Our preferences are


fication,

for this last

mode

of classi-

to follow the classification of Cohn, the

but obliged, in the description of species, most comhas been given hitherto,
for the present.

plete which

we must

abandon

it

2.

Classification

Generic and Specific Characters.


;

classifications of the bacteria of given an abstract in the historical which we have part of this work, show how variable have been the ideas of the microscopists as to the nature of

The numerous

these organisms.

Before giving the most recent, those among will have to choose, it is best to study the characters upon which authors have depended for grouping the bacteria in genera and species,

which we

and

to estimate the value of these characters.

60

MORPHOLOGY OF THE BACTERIA.

These have 1. Generic and specific characters. been drawn from the dimensions, form, movement and evolution of the bacteria. The size, which, according to Cohn, is the dominating distinctive character,
nable,
is

often indetermi-

even in employing the highest powers. Besides, for a great number of neighboring forms, the differences of measurement given as distinctive are so slight that they cannot serve in practice. Thus, according to Dujardin, the Bacterium termo has a length of 1, 7 /*, and the B.punctum of 1, 7 to 0.6 Another difficulty presents itself when we examine bacteria formed of several articles. Shall
/jl.

we

consider the length of a single article or of the

chain, which consists of a

number

of articles, a

number ordinarily variable ? The form of the bacteria and


colonies, also
offer differences,

their union in

which have been utilized; but do they depend upon differences truly specific, or do they come from foreign influences, from phases of development of the same organism? Even when one uses these as distinctive specific characters, the form is sometimes of little assistance since if one refers to the descriptions of Dujardin, the Bacterium termo will be found to have a cylindrical body swollen in the middle, and the B. punctum an elongated ovoid body. As to movement, we have seen that the phenomena of mobility or of immobility sometimes present themselves in the same species, according to age or changes in the medium. We have left, the mode of development, the
;

CLASSIFICATION OF THE BACTEEIA.

61
or

phenomena

of

reproduction

by

fission

by
to

spores, as the only character

which can serve

establish our natural genera; but, unfortunately,


this has

only been ascertained for a small number


of bacteria cannot

of bacteria, the Bacillus anthracis, for example.

The genera
significance as
tables
;

have the same

among animals and superior vegethey can only be established in accordance with the most prominent characters, reserving the feeble modifications of these generic forms as
specific characters.

Are
The

there distinct, well-defined, species

among

the Bacteria ?

microscopists have given the most diverse


Miiller,

opinions upon this subject.


tinction of the

Ehrenberg,
dis-

Dujardin, Davaine, have admitted the specific

numerous vibrioniens which they have described. Davaine, however, raises some
doubts as to the absolute value of the species " Those which are deestablished in his time. classifiers," the he says, " ought scribed to-day by to be considered as the expression of types under

which are hidden a certain number


species."

of

distinct

Cohn
in

dwells

still

more upon the

impossibility,

which we are to-day, of distinguishing with certainty genera and species among the bacteria. However, he is convinced that the bacteria are divided into species as distinctly as the other plants and inferior organisms. It is only the imperfection of our means of observation which makes it impossible to recognize these differences.

This

is

above

62
all true,

MORPHOLOGY OF THE BACTERIA.


he says, of the
spirilla,

which are not only

distinguished from the rod bacteria, properly so


called; but

differences as constant as
of alga or of infusoria.

which present in their species some any well-defined species


Billroth, Robin, Nageli, etc.,

Hallier,

Hoffmann,

consider the different forms of bacteria in a very

According to them they are not autonomous species, but phases of development of one or of several species. According to Hallier, we may see, a propos of the polymorphism of the bacteria, the singular transformations which he has obtained by their
different fashion.
cultivation.

According to Billroth, the bacteria belong to a


single species of plants, the Coccobacteria septica,

with the exception of the Spirillum and Spirochceta, in regard to which Billroth is not willing to give

an opinion.
certain

This view has been adopted


of microscopists,

number

and above

all

by a by

the pathologists, such as Frisch, Tiegel, etc.

Micrococcus, Vibrio, Bacterium and Leptoihrix, but considers them the distinct and successive phases in
also admits the genetic relation of

Robin

the evolution of several species


described under the

1st.

Corpuscles

name

of
;

Bacterium termo,
2d.

punctum,

etc.,

Micrococcus
;

Mycelial

fila-

ments, Vibrio,
bacteria, etc.
;

etc.

3.

Bacteria, Bacteridies, Micro-

4th. Leptoihrix

and forms more

advanced. The opinion of Nageli corresponds very nearly with the preceding. " As much as I am con-

CLASSIFICATION OF THE BACTERIA.

63

vineed," he says, " that the schizomycetes cannot

be grouped in accordance with their action as ferments and their exterior forms, and that altogether too many species have been distinguished so, on the other hand, it seems to me very improbable that all the schizomycetes constitute a single natu;

ral species.

"I

am rather inclined to

suppose that there exists

a small number of species, which have little in common with the genera and species admitted to-day, and of which each runs through a cycle of determined forms sufficiently numerous.
of the veritable species of schizomycetes is not limited to presenting itself under the different forms of Micrococcus, Bacterium, Vibrio, and Spirillum, but can also show itself as the agent of
acidification of milk, of putrefaction, and as the agent producing several maladies.'However, Nageli recognizes that it is necessary to distinguish these forms, notably those of Micrococcus, Vibrio, Bacterium, and Spirillum, without, however, losing from view the fact that the organisms

among them

Each

thus classified have a very inconstant constitution, and pass continually from one form to another.
Finally, other savants such as
less

M. Pasteur, take

account of the structural characters than of the physiological functions, regarding as a particular species every form of bacterium which is born

determined medium, or which causes a special kind of fermentation. Nageli opposes to this view the following
constantly in a
objections.
First,

he has verified the presence,

in

64

MOBPHOLGGY OF THE BACTERIA.

the same decomposition, of several different forms


of schizomycetes.

On

the other hand, in decom-

positions quite different,

we may

observe schizo-

mycetes entirely similar as to their exterior form. Finally, we may change the mode of action of a
schizomycete in subjecting it to a certain treatment. One sees that it is truly difficult to form

an opinion

as to the value of these species purely

physiological.
in order to establish genera

may be used and species in the group of the bacteria are of small number and of very unequal value. Some, characters of form, of dimension, of movement, etc., are often difficult to determine, or have only a relative value others, characters drawn from development and reproduction, are only known in so few species that they
To sum
up, the characters which
;

cannot be made to serve as a basis of


tion.

classifica-

One
there
one.

will
is

not be surprised, then, to find that


classification

no natural
it is

of the bacteria,

and that

impossible for the naturalists to give

All those that can be established are pro-

visory, being only based

upon the morphology of

these organisms.

Following the example of all the botanists, we will use an analogous classification, but without wishing to prejudge in any particular
the genealogical relationship of the different organisms, which
era and species.

we

shall consider as distinct

gen-

CLASSIFICATION OF THE BACTEEIA.

65

3.

Classification

and Description of the Genera and Species of the Bacteria.

have seen in the historical portion of this work, a propos of the classifications which have been given of the bacteria, that, in 1872, M. Cohn, recognizing the numerous relations, absence of chlorophyll, mode of nutrition, etc., which make these organisms a natural family, divided them
into four tribes
1.
:

We

2.
3.

4.

The The The The

Spherobacteria, or spherical bacteria.


Microbacteria, or B. in short rods. Desmobaeteria, or B. in straight filaments.
Spirobacteria, or B. in spiral filaments.

In the spherobacteria, Cohn has only adopted one genus, the g. Micrococcus, of which the species are divided into three series, the pigmentary M., or chromogenes, the M. of fermentations, or zymogenes, and the M. of contagious affections,

or pathogenes.

The microbacteria
terium, with

include only the genus Bac-

B.

two species, B. termo, Dujardin, and Cohn. The desmobaeteria comprehend the g. Bacillus
lineola,
;

and Vibrio

the

first

established

by Cohn
of the

for the

B. subiilis, Cohn (with B. anthracis as a variety) and the B. the second, characterized by unduulna, Cohn lating filaments, is reduced to V. rugula and serrectilinear filaments is
;

composed

pens, Auct.

66

MORPHOLOGY OF THE BACTERIA.


Finally, the spiral filaments of the spirobacte-

Spirillum and Spirochasla, which might be united in a single genus comprising Sp. plicatile, tenue, undula, and volutans.
ria characterize the gr.

Since then, Cohn, struck with the affinities which each of the preceding genera presents with several genera of oscillatoriaceae and of chroococcese, from which the bacteria only differ by the

absence of chlorophyll, has established a class of Schizophytes, which includes all the inferior vegetable organisms, provided or not with chlorophyll,

multiplying by fission. We give below the complete table

2. Classification

of
1.

the Schizophytes, Cohn.

TRIBE

GLtEOGENES.
an
intercellular substance.

Cells free or united in glairy families hy

A.

Cells free or united

by 2
.

or
.

by 4

Cells spherical

Cells cylindrical

Chroococcxjs, Nag. Synechococcus, Nag.

B. Cells united in glairy families,

amorphous in
repose
a.
:

state

of

Cellular

membrane, con-

founded with the intercellular substance


1.

Cells without phycochrome, very small

Cells spherical

Micrococcus,

Hallier.

::

CLASSIFICATION OF THE BACTERIA.


Cells cylindrical
2.
.

67

Bacterium, Duj.

Cells with phyco-

chrome, larger
Cells spherical
. .

Cells cylindrical
t.

Aphanocapsa, Nag. Aphanothece, Nag.

Intercellular

substance

formed of several membranes enclosed one within the other


Cells spherical
.

Cells cylindrical

Glceocapsa, Kg. Glceothbce, Nag.

C. Cells united in glairy families

of definite form

a.

Families of a single layer


of cells disposed in plates
1.

2.

Cells in fours formMerismopedia, Meyen. ing a plane surface Cells without regular arrangement, forming
.

a curved surface
Cells spherical, families

with reticu.
.

lated rupture

Clathrocystis, Henfr.

Cells cylindrical, cu-

neiform, families

divided
striction
b.

by con. . .

CcELOSPHJSRnJM, Nag.

Families with several layers of cells, united in spherical corpuscles


1.

Number
termined

of cells de:

Cells spherical, colorless,

arranged
.
. .

in fours

Sarcina, Goods.

68

MORPHOLOGY OF THE BACTERIA.


Cells cylindrical, cu-

neiform, with phy-

cochrome, without regular ar-

rangement
2.

GOMPHOSPILERIA, Kg.

Number
nate
:

of cells very

great and indetermiCells colorless, very

small
Cells

....
by
. .

Ascococcus,

Billr.

colored

phycochrome and larger

>
)

Polycystis, Kg. Coccochloris, Spr. Polycocctjs, Kg.

TRIBE

2.

NEMATOGENES.

Cells disposed in filaments.

A. Filaments not branched


a.

Filaments free
laced.
1.

or

inter-

Filaments cylindrical,
colorless, articulations

not very distinct Filaments very slender, short


. . .

Bacillus, Cohn.

2.

Filaments very fine, long Leptothrix, Kg. Filaments larger, long Beggiatoa, Trev. Filaments cylindrical, \ with phycochrome, I TT

....

articles well defined, (

Hwheothrix, Kg.

without cellular reproduction


.

^laria,

Bosc.

: :

CLASSIFICATION OF THE BACTERIA.


3.

69

Filaments cylindrical,
articulated, with

co-

nidia

Filaments colorless Crenothrix, Cohn. Filaments with phy-

cochrome
4.

Cham.esiphoN'.

Filaments spiral without phycochrome


Filaments, short,
light,

sinuous

Vibrio, Ehr.

Filaments, short, spiral, rigid


. .
.

Spirillum, Ehr.

Filaments, long, spiral, flexible


.

Spirochete, Ehr.

with phycochrome Filaments long, spiral, flexible


. .

Spirulina, Link.

5.

Filaments in chaplet Filaments, without

phycochrome
. .

Streptococcus,

Billr.

6.

Filaments with phy- Anabena, Bory. cochrome f Spermosika, Kg. Filament flagelliform,
)
.

slender

Mastigothrix,

etc.

Filaments united into glairy families by an intercellular substance 1. Filaments cylindrical,


colorless
2.

Myconostoc, Cohn.
) I
,

Filaments cylindriJ , ... cal, with phyco-

Chthonoblastus.
Limnoclide, Kg.
etc.

>
\ ;
.

Tr

chrome
3.

4.

Filaments in chaplet Nostoc, Filaments flagelliform,


slender

Rxvularia,

etc.


70

MORPHOLOGY OF THE BACTERIA.

B. Filaments with false ramification


1.
:

Filaments
cal, colorless

cylindri. .

Cladothrix, Cohn. Streptothrix, Cohn.


Calothrix, As. _ SCYTONEMA, Ag.
.

2.

cal,
,

Filaments cylindri, vi pnyco-u with


. . . .

) f

}
\ ;
.

chrome

3. Filament in chaplets 4 Filaments flagellitorm, slender towards

Merizomyria, Kg.
Schizosiphon, ScHIZ0SIPH0N5

i
l

K g.

Geocyclus, Kg.

the extremity

An

inspection of this table shows that each of

the genera of the ancient group of the bacteria

has been placed beside some genus of oscillatoriacese, which it resembles by its organization, Micrococcus and Bacterium, beside Aphanothece

and Aphanocapsa ; Bacillus, beside Leptothrix and Beggiatoa ; Vibrio and Spirillum, beside Spirulina.

These
a

affinities are

undeniable, and the advan;

tages of such a classification are manifest

but, in
it.

work

like this,

we cannot

think of employing

We

preserve, then, in a distinct group the schizo-

phytes deprived of chlorophyll, which may be arranged in the four primary divisions of Cohn with the exception of Sarcina, Ascococcus, Crenoihrix, etc., and the other genera created recently

by

this botanist.

Thus we
1.

will describe successively

The

Spherobacteria of

Cohn

and beside them

the different

Monas recently

studied,

the Microcoe-

CLASSIFICATION OF THE BACTERIA.


cms described
adies.
2. 3.

71

by

Hallier in several infectious mal-

The The
The

Microlaceria.

Desmobacteria, including Bacillus, Lepto-

ihrix, Beggiatoa,
4.

and Crenothrix.

Spirobacteria, including the three genera,

Vibrio, Spirillum,
5.

and

Spirochceta.

Finally,

we

will give

some account of the Mer-

ismopedia, Sarcina, Ascococcus, Streptococcus, Myconostoc, Cladothrix,

and

Streptothrix.

1.

Sphekobacteria, Cohn.

The

spherical bacteria are characterized

by their

rounded or oval form, their small size, often less than 1 /i. They are ordinarily isolated, often in pairs (diplococcus), sometimes in a chain of several torula of Cohn), the myarticles (streptococcus cothrix of Hallier and Itzigsohn, or in the form of zooglma when they are young and actively multiplying, and that of mycoderma, when they are gathered upon the surface of liquids. They have no spontaneous movement, but a simple molecular

trepidation.

" The spherical bacteria are ferFunctions ments, not producing putrefaction, but substitutions of another kind " (Cohn). Obs. According to the facts observed by Koch, Cohn, Pasteur, Toussaint, upon the development of certain bacteria, it is very probable that some
:

at least of the spherobacteria are spores of Bacillus or of other bacteria


;

at least, the micrococci

and these spores are

identical in

form and

aspect.


72

MORPHOLOGY OF THE BACTERIA.

The spherobacteria include only the genus Micrococcus.

g.

Micrococcus, Cohn
sphceria,

(Hallier emend.

Microdivision

Cohn, ante).

Cells colorless, or scarcely colored,

very small,

globular or oval, forming


filaments of

by transverse

two or several articles, in form of chaplet, or united in numerous cellular families, or in gelatinous masses, all deprived of movement.

The
logical

species are divided

into

three physio-

groups

a.
b.
o.

M. Chromogenes. M. Zymogenes. M. Pathogenes.


Micrococcus Chromogenes.

Section

(a)

The pigmentary bacteria grow in the state upon the surface of the substances which furnish them with nutriment. They are always alkaline all are avid of oxygen their
of Zooglcea
; ;

morphological characters are identical, and one

can only distinguish them by their different


coloring properties.

cies

According to Cohn, they are veritable spefor 1. Their pigments offer the greatest diversity as to chemical action and by spectro;

scopic analysis, etc. in the

2.

Each

species cultivated

most diverse media produces always the

same coloring matter.

CLASSIFICATION OF THE BACTERIA.

73

They

are divided into two categories, accordis

ing as the pigment


1.

soluble or not in water.

Coloring matter insoluble.

M. Prodigiosus, Cohn (Monas prodigiosa, Ehrb.

Palmella
A
in

prodigiosa, Mont.

Bacteridium

prodigiosum, Schroeter).
red gelatinous mass, pink carmine, developing upon cooked alimentary substances placed
air, never before cooking. has also been observed in red milk, attributed incorrectly to lesions of the teats, etc. (Cohn).

damp

It

M.

luteus,

Cohn {Bacteridium

luteum, Schroeter).

A yellow gelatinous mass


and Cohn upon potatoes.
2.

studied

by Schroeter

Coloring matter, soluble.

M. aurantiacus, Cohn {Bacteridium auriantiacum,


Schroeter).

more or less extended, by Schroeter, upon golden yellow, slices of cooked potato; by Cohn, upon hard
Little drops, or stains,

cultivated

white of egg.

M. chlorinus, Cohn.

A glairy yellowish-green pigment found upon hard white of egg, not reddened by acids, but
loses its color.

M. cyaneus, Cohn (Bacteridium cyaneum, Schroeter).

Pigment deep

blue, observed

by

Schroeter

74

MORPHOLOGY OF THE BACTERIA.


and cultivated by Cohn in solutions. This coloring matter is reddened by acids, and restored to blue by alkalies, just as that which forms when lichens are macerated in presence of ammonia.

upon cooked

potato,

nutritious

M. violaceus, Cohn (Bacteridium violaceum, Schrceter).

Violet-blue masses or glairy stains


elliptical corpuscles larger

formed of

digiosus, observed first

than those of M. proby Dr. Schneider, then

by Schroeter on cooked
Later,

potato.

Cohn has described the two following


which should be included in

new
this

species (1876),

group

M. Candidus, Cohn. Stains and points white as snow, observed upon slices of cooked potato. M.
fulvus,

Cohn.

Little rust-colored drops, consisting of cells,

globular or united in pairs, in a tenacious intercellular substance, diameter 1.5 /*, observed by Eidam, then by Kirchner, upon horse dung. It is also to the genus Micrococcus that we

rmist refer the little

globular bacteria, gifted

with movement, found by Eberth in white, yellow, and red perspiration, and by Chalvet in the pus on the edges of certain wounds, but which should not be confounded with the blue color produced by a Bacterium.

CLASSIFICATION OF THE BACTERIA.

75

Section

(b):

Micrococcus Ztmogenes.

Globular bacteria producing fermentations of


diverse nature.

M. crepusculum, Cohn (Monas crepusculum,


Globular
cells, colorless,

Ehrb.).
all

developing in

in-

fusions of animal and vegetable matter under-

going decomposition.

M.

urese,

Cohn.
cells, isolated,
/a

Oval
1.2 to 2

diameter 1.5 (Cohn) or united by 2,


In urine of which

/*.

(Pasteur),
{to-

4, to 8

rula), in a line, straight, curved, zigzag, or

even

in cross form.

it

transforms

the urea into carbonate of

ammonia

(Pasteur).

Torula which appears identical with the preceding Micrococcus, produces the decomposition of hippuric acid into benzoic acid and gly-

collamine (Van Tieghem).

M. of stringy wine,
Globular
cells

etc.

of 2

/*

diameter, in chaplets,

found in stringy wine, perhaps identical with


the preceding (Pasteur).

Torulaceas quite similar

is

found in certain

fermentations of tartrate of ammonia and of

beer yeast, with or without the addition of car-

bonate of potash (Pasteur).


Section (c)
:

Micrococcus Pathogenes.

Spherical bacteria found in affections of a contagious nature.

76

MORPHOLOGY OF THE BACTERIA.

M. vaccinae, Cohn (Microsphcera Vaccince, Cohn).

Very small
and

micrococci,

0.5

/4

scarcely, iso-

lated or united in pairs in recent vaccine virus


in the pus of variola pustules.

By

cultiva-

from two to eight cells may be obtained, then masses containing sixteen to thirty-two cells of 10 /4 and more diameter. The M. of vaccine virus and of variola are identical, and Cohn regards them as different races of the same species.
tion, chaplets of

M. diphtheriticus, Cohn. Granular cells, ovoid, measuring from 0.35 to to 1.1/x., isolated or more often united in twos or in a chaplet of four to six cells sometimes multiplying in colonies and extending themselves in all the diseased tissues, decomposing and destroying them (GErtel).
;

M. septicus, Cohn (Microsporon septicus, Klebs). Little rounded cells, of 0.5 /t, motionless and crowded in masses or united in chaplets, in the
secretion
of

wounds

in

cases

of

septicemia

(Klebs), in zooglcea in callous ulcers, in isolated


cells,

united in pairs, or in chaplets in the seof epidemic puerperal fever (Waldyer), in


etc.,

rum
all

the tissues, vessels,

in cases of

pyemia

and septicemia.
M. bombycis, Cohn (Mycrozyma hombycis, champ).
Cells with a diameter of 1
/4,

Be"-

ordinarily united

in chaplets of two, three, four, five, or more, in


CLASSIFICATION OF THE BACTERIA.

77

the intestine of silkworms sick with " la flacherie" (Pebrine).

p.

In a more recent work, Cohn (Beitrage, 1875, 201) gives them an oval form and a diameter
fi,

of 0.5

at the outside.

Hallier has described

many

other Micrococci in

diverse contagious or virulent affections.

We
: '

will

only refer to them in a summary manner

M. of

of animals, Hallier. Small M. endowed with active movement, furnished with a very delicate caudal appendage, sometimes united in the form of little elongated rods, found in pustules, spontaneous or inoculated, in the lymphatic canals and the ganglia of animals attacked with variola.
the variola

M. of

rugeola, Hallier.
colorless M.,

Very small

having often a caudal

prolongation, in the sputa and blood of the sick.

M. of

scarlatina, Hallier.

M., free or in colonies, on the surface or in the interior of the blood corpuscles, or in
chains.

M. of

epidemic diarrhoea, Hallier.


vibrios,
cells,

M. in intestinal matters with and monads. (?)


1

"It

is

quite probable that Hallier comprises, in part, under the

same organisms that I call spherical bacteria but the doctrine of Hallier concerning Micrococcus, as has already been pointed out by Hoffmann and de Barry, is so covered by inexact assertions and improbable hypotheses, that it is impossible to draw any conCohn, Beit. II, p. 148. clusions from the facts he has observed."

name

of Micrococcus the

78

MORPHOLOGY OF THE BACTERIA.

M. of exantkematous typhus, Hallier. M. relatively large, brown having a rapid


1

movement, sometimes in chains (Mycothrix), in


the blood.

M. of

intestinal typhus, Hallier.


;

M. very small, in repose in the blood larger,, endowed with active motions, and furnished
with contractile appendices in the dejections.

M. of

glanders, Ziirn.

Cells free or attached to the blood globules, or

even penetrating into their interior, sometimes M. in in chains (Mycothrix) in the blood. chains, very numerous, and endowed with rapid movements, in the lymphatic ganglia, the mucus of the frontal sinuses, and in the chancroid
ulcers.

M. of

syphilis, Hallier.
colorless, free or in globules, in

M. numerous,

gonorrhoea, the primitive ulcer, and the blood of

persons suffering from constitutional syphilis.

Monads.

Beside the Spherobacteria are placed the Monads, not the organisms described

under

this

name
micro-

by the
phytes,

older
spores,

microscopists,

comprising
animals,

and

infusorial

but the

Monas
day.
sides

as understood

by

botanists of the present

Thus limited, the Monads include also, besome microphytes related to the Spherobacteria, and differing from them by their greater dimensions, some organisms of doubtful affinities.

CLASSIFICATION OF THE BACTERIA.

79

As in the case of the Micrococci it is very probable that the Monads are only the spores, or
lower forms of bacteria higher in the
places the
scale.

Cohn
ru-

Monas vinosa

of

Ehrenberg with the

Clathrocystis roseopersicina,

Cohn (Bacterium
it

bescens, Ray-Lank.), considering


latter.

a spore of the

Monas

vinosa, Ehrb.
/x,

Cells spherical, oval, regular, of 2.5

often united

in pairs, formed of a pink substance with granules of

a deeper color, having spontaneous movements. Hab., waters containing decomposing vegetable matters (Ehrb. 1838, Ch. Morren 1841, Perty 1852, Cohn

1875).

M.

Okenii, Ehrb.

Cells cylindrical; average length 7 to 15 fi (Cohn), (Warming), 10 fM (Ehrb.), sometimes from 60 to 80 red color, having beautiful a rapid of a diameter 5 /x gyratory movement, with a cilium at the posterior extremity or two cilia at the two extremities. Hab., stagnant water (Ehrb. 1836, Eichwald, Weiss, Cohn,
//.
;

etc.).

M. Warmingii, Cohn.
extremities some
Cell cylindrical, pink, containing at deep-red granules
fi,

its
;

two rounded

length 15 to

20
of

width 8
cilium.

fi

movement

uncertain, having a vi-

bratile

Hab., brackish water on the coast

Norway (Warming).

M.

gracilis,

Warming.
cylindrical, pink,
;

Cells straight,

two extremities length 60 /z, thickness 2 ment slow. Hab., fresh water.

rounded at the movefi


;

80

MORPHOLOGY OF THE BACTERIA.


rosea, Cohn.
;

Rhabdomonas

Cells pale pink, isolated, fusiform

eight times as

long as broad, having a length of 20 to 30 /a, and a width of 3.8 to 5 fi; having a slow oscillatory movement, the pink substance containing numerous granSab., stagnant ules of darker color and vacuoles.
water.

Ophidomonas sanguinea, Ehrb.


Cells pale

pink, spiral, rigid,

movement

active

thickness 3

/a,

length of one turn of the spiral, 9 to

12

fj,.

Sab., brackenish waters of

Denmark (Warm-

ing).

Spiromonas Cohnii, Wanning.


Cells spiral, flattened
to 3.5
fi,
;

1 turn of

spiral,

diam. 1.2

width 1.2 to 4

jj,.

Sab'., coast of

Denmark.

2.

MlCROBACTERIA, Cohn.

Rod-bacteria, cells cylindrical, short, having spon-

taneous movement.

A single
g.

genus,

Bacterium.
or elliptical, free or united

Bacterium,

Duj. emend.

Cells cylindrical

in pairs during their division, rarely in fours,

never in chains (leptothrix or torula), sometimes in zooglaia (differing from the Z. of spherical B. by a more abundant and firmer intercellular substance), having spontaneous movements, oscillatory and very active, especially in media rich in alimentary material and in presence of oxygen.

;;

CLASSIFICATION OF THE BACTERIA.

81

We

might, as in the Spherobacteria, divide


:

the rod -bacteria into three groups


teria of putrefaction,

1.

the bac-

B. termo, B.
;

lineola,

and

their varieties

2.

the Bacteria of the lactic and


3.

acetic fermentations, etc. of colored milk

Chromogenes, B.

and pus.
(

B. termo, Ehrb. 1830, Duj.


ex. p. 1838;

Vibrio lineola, Ehrb.

Monas

termo, Miiller).

Cells cylindrical, slightly swollen in the middle,


isolated,

sometimes united in
;

pairs,
/j.,

two

to five times

as long as wide
1.8
fi
;

length 2 to 3
oscillatory.

thickness 0.6 to

movements

Appears at the end of a very short time in infusions of animal and vegetable substances multiplies with rapidity in numerous zoogloza ; then disappears as other species, to which it According serves as nutriment, are developed.
all

to recent observations, this bacterium has cilia

(Dallinger, Drysdale, Warming).


also

Warming

has

found it in the state of torula. B. termo is the veritable agent, the first cause, of putrefaction, it is the true ferment saprogene
(Cohn).

M. "Warming

has recently described two allied forms


cells larger,

/*,

B. griseum,
B. littoreum,
length 2 to 6

more rounded

length 2.5 to 4

thickness 1.8 to 2.5 p.

In infusions of fresh and

salt water.

cells elliptical or elongated, slightly

rounded

^,

thickness 1.2 to 2.4 p.

Coasts of Denmark.

B. lineola,

Cohn

Duj., Miiller,

ex p., V. tremulans, Ehrb., Bacterium


(

Vibrio

lineola,

Ehrb.

triloculare, Ehrb).

82

MORPHOLOGY OF THE BACTERIA.


Cells cylindrical, straight, rarely a little twisted, larger than the cells of B. termo, isolated or united

in pairs, sometimes in
3.8 to 5.25
/a,

fours,

never more
/j, ;

length

thickness attains 1.25

movements

like those of B. termo,

but a

little

more

active.

Is

found in various vegetable and animal

in-

fusions of fresh or salt water, often takes the

form
their
in the

of zooglcea containing motionless rods in

mucus
form

substance.

Warming has met


is

it

of chains
Its

composed

of eight to ten

cells (torula).

protoplasm

dotted with re-

fractive granules.
It is not known whether B. lineola constitutes a specific ferment (Cohn).

The B. fusiform, Warming,


the form of
ities
;

differs

from the preceding by


plasma not punc-

its

body, which
ft,

is

attenuated at the two extrem;

length 2 to 5

width 0.5 to 0.8 ^

tated.

Beside these species, which have been well studied, may be placed the following, which

demand new

investigations

B. punctum, Ehrb.
Elongated rods, oval,
colorless,

having slow

movements,
length 5.2
/a,

oscillating, often

united in pairs;

thickness 1.7

/a.

Diverse infusions

of animal substances.

B. catenula, Duj.

Body

filiform,

cylindrical,
;

often

united
/x,,

in

three, four, or five


0.4 to 0.5
/jl.

length 3 to 4

thickness
fe-

In fetid infusions, in typhoid

ver (Coze and Feltz).

CLASSIFICATION OF THE BACTERIA.


Vibrio
lactic,

83

Pasteur.

almost globular, very short, a little swollen at the extremities; length of an article,
1.6
/u,,

"Articles

of a series, 50

//,."

This vibrio seems to resemble B. catenula or B. termo. It is developed, according to Pasteur, in sweetened liquids, where it causes the formation of lactic acid and the coagulation of the casein of milk. According to other researches, coagulation of casein results from the influence of a soluble ferment (zymase), and not from an organized ferment.

Acetic ferment {Mycoderma


aceti, Ktg.).

aceti, Pasteur,

Ulvina

" Articles short, constricted, two to three times as long as broad length 1.5 /u,, often united in long chains, forming pellicles on the surface of a liquid."
;

very similar to the preIt must not be confounded with the ceding. Mycoderma vini, which may develop in the same media, but which is a fungus of the group
This species
is

also

of Saccharomycetes.

The acid fermentation of


due
to a

beer seems to be form of Bacterium resembling B. termo


little

(Cohn), but a

larger than the type.

Cohn

has found
teria,

it

in beer undergoing acid fermenta-

tion, beside oval

saccharomyces, elliptical bachaving movement, often united in pairs,

rarely in fours, etc.

Vibrio tartaric right (Pasteur).


Bacteria similar to those of the lactic fermentation,

PLATE
From

IV.
Macmillan
Sf Co.,

" Pasteur's Studies on Fermentation."

London, 1879.

"The engraving represents the different diseased ferments, together with some cells of alcoholic yeast, to show the relative size of these organisms."
Fig. 1 represents the ferments of turned beer, as it is called. These are filaments, simple or articulated into chains of different size, and having a diameter of about the thousandth part of a millimetre (about ^ i o oo inch). Under a very high power they are seen to be composed of many series of shorter filaments, immovable in their articulations, which are scarcely visible. In No. 2 are given the lactic ferments of wort and beer. These are and contracted in their middle. They are generally detached, but sometimes occur in chains of two or three. Their diameter is a little greater than that of No. 1.
small, fine,

In No. 3 are given the ferments of putrid wort or beer. These are mobile filaments, whose movements are more or less rapid, according to the temperature. Their diameter varies, but is for the most part greater than that of the filaments of Nos. 1 and 2. They generally appear at the commencement of fermentation, when it is slow, and are almost invariably the results of very defective working.
In No. 4 are given the ferments of viscous wort, and those of ropy beer, which the French csdl Jilante. They form chaplets of nearly spherical grains. These ferments rarely occur in wort, still less frequently in beer.
No. 5 represents the ferments of pungent, sour beer, which possesses an acetic odor. These ferments occur in the shape of chaplets, and consist of the mi/coderma aceti, which bears a close resemblance to lactic ferments (No. 2), especially in the early stages of development. Their physiological functions are widely different, in spite of this similarity.

The ferments given in No. 7 characterize beer of a peculiar acidity, which reminds one more or less of unripe, acid fruit, with an odor sui generis. These ferments occur in the form of grains which resemble little spherical points, placed two together or forming squares. They are generally found with the filaments of No. 1, and are more to be feared than the latter, which cause no very great deterioration in the quality of beer, when alone. When No. 7 is present, by itself or with No. 1, the beer acquires a sour taste and smell that render it detestable. We have met with this ferment existing in beer unaccompanied by other ferments, and have been convinced of its fatal effects.
No. 6 represents one of the deposits belonging to wort. not be confounded with the deposits of diseased ferments.
This must

The

latter

are always visibly organized, whilst the former is shapeless, although it would not always be easy to decide between the two characters, if several samples of both descriptions were not present. This shapeless deposit

from
it.

Among the shapeless granules of No. 6 may be discerned little spheres of different sizes and perfect regularity. These are balls of resinous and coloring matter that are frequently found in old beer at the bottom of bottles and casks. They resemble organized products, but are nothing of the kind.

interferes with wort during its cooling. It is generally absent beer, because it remains in the backs or on the coolers, or it may get entangled in the yeast during fermentation, and disappear with

Plate

IV.

Beliotypc Printing Co.. Bostor

CLASSIFICATION OF THE BACTERIA.


with globular articles, short chains of 50 fi.
;

85
united in

diameter 1

/x,

Decomposes racemic
tartaric acid.

acid, causing

the right

tartaric acid to disappear,

and

setting free left

MlCROBACTEEIA ChROMOGENES.

B. xanthinum, Schrceter ( Vibrio synxanthus, Ehrb.).


" Bodies cylindrical, slightly flexible, formed of corpuscles rarely exceeding five in

an
it

article, 0.7 to 1 p.

number length of In tainted cow's milk, to which


;

gives a yellow color."

B. syncyanum, Schrceter ( Vibrio syncyanus, Ehrb.). This Bacterium, which has the same characters as the preceding, has been observed in sour milk, to which it gives a blue color. B. oeruginosum, Schrceter. In greenish blue pus. These B. chromogenes resemble entirely the According lactic vibrios, B. termo or catenula. to Robin, colored milk contains colorless vibrios, and the coloration is due to an alga similar to
Leptomitus.

B. brunneum, Schrceter.

Rods

in a

brown coloring matter

in infusions

of rotten corn.

Following the colored Microbacteria, I place

two species of Bacterium recently described by Ray-Lankester and Warming.


B. rubescens, Ray-Lank., 1873. Under this name Ray-Lankester has

described


86

MORPHOLOGY OF THE BACTERIA.

some phases of development of Clathrocystis roseopersicina of Cohn. Now Cohn is inclined to regard the Monas vinosa, Ehrb. as the wandering cells of On the other hand Warming has deClathrocystis.
scribed his
:

B. sulfuratum, Warming, 1876, giving

it

for synonymes,

Monas vinosa, Ehrb.; M. erubescens, Ehrb.; M. WarmIt follows, ingii, Cohn; RTiabdomonas rosea, Cohn. then, that the Monas which we have described with
the Spherobaeteria should be referred to a Bacterium
called sulphuratum
identical with

by Warming, but which B. rubescens of Ray-Lankester.


3.

is

also

Desmobacteria.

Filiform bacteria, composed of elongated cylindrical articles, isolated, or in

chains

more

or less

extended, resulting from transverse division.

Un-

der this form they correspond to leptothrix, Auct. (differing from torula in that the filaments are not
constricted at the point of junction of the articulations) filaments sometimes united in swarms, Movements and state of renever in zooglma. pose alternating and depending upon the presence or absence of oxygen, the reaction of the medium, and other conditions unknown. Some forms never exhibit movement. Bacteridie of Davaine (Cohn). We will only preserve in the Desmobacteria the genus Bacillus, Cohn. The vibrios are rather allied to Spirillum because of their undulating filaments.
;

However,

after the

exposition of the different


will

species of Bacillus,

we

say something of three

genera of colorless

oscillatoriaceae,

which are nearly

CLASSIFICATION OF THE BACTERIA.


related to them,

87

the

Leptothrix, Beggiatoa, and

Crenothrix.
1.

Fil.

with indistinct articulations

Fil.

very slender, short


very slender, long

Fil.

Fil. thick,
2.

broad

Fil. articulated distinctly

.... .... ....

Bacillus. Leptothrix.

Beggiatoa. Crenothrix.

g. Bacillus, Cohn.

The

bacilli are characterized

by slender

fila-

ments, straight, short or of moderate length, rigid or flexible, endowed or not with motion.

chromogene, the B. ruber of to this genus that we should refer the Amylobacter of Trecul.
species
is

One

Cohn.

Finally it

is

B. subtilis,

Cohn {Vibrio

subtilis,

Ehrb.; Ferment

butyrique, Pasteur).

Filaments very slender and elongated, formed of a


single cell having usually a length of 6
articles of
/x,

or of
fi,

two

which the
/a),

total length is then' 12

or of

three (length 16

or of a greater

number (some-

with a total length of 40 to 60 and 130 /x) thickness not measurable, well-defined movements of flexion, active or passive, and of transreproduction by fission lation forward or backward and by globular or oval spores developing themselves in the interior of the articles (Cohn). Is found in
times as
as twenty,
; ;

many

stagnant water.

Plays a great role in the butyric fermentation


(Pasteur).

This B. exists in rennet, can support

a temperature of 105, and live in a medium deprived of free oxygen, in which case it takes
a form cephalee, containing persistent spores,

88
which,

MORPHOLOGY OF THE BACTERIA.

when

set free, give birth to other rods

of Bacillus (Cohn).

B. anthracis,
vaine).

Cohn (Bacteridie charbonneuse, Da-

Species very similiar to the preceding, generally

longer and always motionless length 4 to 12 and even 50 p, thickness, scarcely appreciable, 0.8 to
;

14

jx

(Bollinger).

The

B.

anthracis

is

developed in charbon

(malignant pustule of man, sang de rate of sheep, maladie de sang of cattle, fievre charbonneuse of horses), and in the rabbit, the rat,
etc.
;

never in the dog, the cat, the birds, and coldblooded animals. It is found above all in the Cultivated in suitable media, capillary vessels. such as the aqueous humor of the eye of the ox, the Bacillus of anthrax develops spores in the interior of its filaments, which may germinate and reproduce rods (Koch). According to recent observations not yet published, by cultivating the B. Anthracis in the blood of the dog, a development of veritable sporangia may be obtained, containing from
three to six spores (Toussaint).

B. amylobacter,

Van Tieghem (Amylobacter,


Tre"cul).

TJro-

cephalum and Clostridium


forms,

B. occurring, like the preceding, under various in pointed cylindrical filaments of 6.6 to

26 /a in length and 1.1 //, in thickness, or in form of tadpole, with a spore in the terminal swelling, or of a spindle, with a spore in the middle. In fact, it
does not differ from B.
subtilis,

except by the appear-

CLASSIFICATION OF THE BACTERIA.


ance of starch in
its

89

period of multiplication.

protoplasm at the end of the These B. are sometimes endowed with movement (Nylander).
It

develops in vegetable tissues, which


putrefaction,

fall

into

spontaneously according to

Tre'cul, or

anism
of

introduced from without by a mechunknown. This is the essential agent vegetable putrefaction (Van Tieghem).
still
(

B. ulna, Cohn
one,

Vibrio bacillus, Ehrb.).


rigid,

Filaments articulated, thick, and

two

to four articles, straight or

formed of broken in zig-

length of an article 10 fi, length of a filament of four articles 42 fi ; slow movements of rotation and

zag

of progression.

Develops in various infusions of fresh or salt In certain cultivations, Cohn has seen large globules (spores ?) form in the protoplasm. Warming believes that he has seen cilia.
water.

B. ruber, Cohn.

Long rods, isolated or movement very active

united in two or four,


in a red

stance, vermillion, developed

mucous subupon grains of rice.

Observed by Franck and Cohn.


Bacteridies,

Davaine has described five additional species of which appear to be bacilli. They are
:

La

Bacteridie intestinal.

Filaments straight, thick from 10 to 40 fi in length. In the intestines of birds.

La

B.

du

levain.
fi,

Filaments slender and short, of 10 to 20

divided

90

MORPHOLOGY OF THE BACTERIA.


into two, sometimes three to four articles, identical

with the B. of charbon (Davaine).

La

B. glaireuse.
Filaments extremely slender, straight or elbowed
in length.

attaining 10

/*

La

B. du vin towmi.
Filaments very slender, of variable length,
flexible,

indistinctly articulated.

La B.

des infusions.
/*.

Filaments of 10 to 20
g.

In various infusions.

Leptothrix, Ktz.

The Leptothrip
der,

differ

from Bacilli by their

filaments being very long, adherent, very slen-

and

indistinctly articulated.

Their forms are numerous.


are the principal
:

The

following

L. rigidula, Ktz.

Length 100 to 150 /*, diameter 1.3 to 1.9/*. stagnant water, adherent to other vegetables.
L. crcespitosa, Ktz.

In

Length 100 humid rocks.

to 200

/*,

diameter 2.4 to 2.8

/*.

Upon

L. brevissima, Ktz.

Length 75 to 100 nant water.


L. pusilla, Rabh.

/*,

diameter 2.7 to 3.5

/*.

In stag-

Length 60

to 70

/*,

diameter 0.5 to 0.6

/*.

L. parasitica, Ktz.

Length 90

to

150

ji,

diameter 1

/*.

L. radians Ktz., and L. spissa, Rabh.


Parasites

upon marine

algse.

CLASSIFICATION OF THE BACTERIA.


g.

91

Beggiatoa, Trev.
ter, rigid,

Filaments very slender, surrounded by mucous mathaving oscillatory movements. Protoplasm

white, enclosing numerous granules, which recent


observations have demonstrated to be crystalline sul-

phur (Cramer, Cohn).

The Beggiatoa are found most abundantly in thermal sulphur waters,. where they constitute flocculi, which have been named Glairine, Baregine. They often live in water not containing free oxygen. They play a great role in the elimination of sulphur and the disengagement of sulphuretted hydrogen in thermal waters. Their principal species are
:

B.

alba,

Trev.

A whitish mucous mass enclosing colorless filaments,


having a diameter of 3.5 to 4 stagnant waters.
B. arachnoidea, Rabh.
Flocculi very minute,
as broad,
fi.

In most thermal and

snow

5.4 to 7

//,.

white, filaments as long In the thermal waters of

Europe.

B.

nivea,

species, living in the

Rabh. B. leptomitiformis, Trev., nearly related same conditions.


;

Cohn and Warming have


B. mirabilis, Cohn,
to

also described

measuring 20

articles scarcely flexible,

40

p.

B. minima, Warming, a very small


length 40
/*,

species,

very flexible

thickness 1.8 to 2 p.
4.

Spirobacteria.

This tribe includes the bacteria with undulating


filaments, or filaments in spirals,

more or

less de-

92

MORPHOLOGY OF THE BACTERIA.

veloped, from the Vibrio rugula, which only presents a single curve in
of Spirillum
its

centre, to certain species

which have numerous turns of the spiral. In several species, cilia, or a flagellum, have recently been observed. We divide them into three genera
:

Fil. short, slightly sinuous


Fil. short, spiral, rigid

Fil. long, spiral, flexible

.... ....

Vibrio. Spirillum.

Spirochete.

g. Vibrio, Auct. emend.

Body
lated,

filiform,

more

or less distinctly articu-

having serpentine movements. This genus forms the transition between the Desmobacteria and Spirillum "from which it cannot be separated " (Warming).
always
undulating,
Fil. thick, with a single
Fil. slender,

curve

...
.

with several undulations

V. rugula. V. serpens.

V. rugula, Miiller (V.


ture, feeble

lineola, Duj. ex parte).

Filament presenting in its centre a single curvabut distinct length 8 to 16 /u.. The shortest are slightly curved (= 6 Warming), the larger, which may attain 17.6 fi (Cohn), 35 /a (Warm.), are about to divide. Movements of rotation more or of progression less rapid around their longer axis forward, giving then the idea of a serpentine movement: having a cilium (Warming).
; fx,
;

V. rugula

is

infusions, in deposits
tinal

commonly found in swarms, in upon the teeth, in intesdis-

matters (Leewenhoeck), in choleraic

charges (Pouchet).

CLASSIFICATION OF THE BACTERIA.

93

V. serpens,

Miiller.
less in

Filament one half

ceding, rigid, annulate, having

diameter than the pretwo or three regular


;

undulations, at least two in the shortest height of one turn of the undulations 8 to 12 /j,, diameter 1 to
3
fj,,

total length

11 to 25

/m,

thickness 0.7

fi

move-

ments analogous ium (Warm.).

to those of B. subtilis ; having a cil-

In numerous swarms in infusions, river water,


etc.

g. Spirochaete, Ehrb.
S. plicatilis,

Ehrb.

Filament not extensible, twisted in a long helix,


flexible,

the turns of the spiral near together

suscep-

tible of twisting

upon

its

axis

and

of
p.

an undulatory

movement

total length
;

130 to 200

Rare species
sea-water, etc.
S. Obermeieri,

in infusions, stagnant water,

Cohn.

Does not differ from the preceding, either in size, conformation, or in its movements, but by its habitat and physiological peculiarities.

In the blood of persons attacked by recurrent


fever (Obermeier, 1872, Weigert, Bisch-Hirschfeld, etc.)

during the period of access, never

during the remission.


S. gigantea,

Warming.

Found upon

the coasts of

Den-

mark

thickness of body, 3 p, height of spiral 25 p, diam-

eter 7 to 9 p.

; ;

94
g.

MORPHOLOGY OF THE BACTERIA.


Spirillum, Ehrb.

Filament spiral, rigid; turns of and regular.


S. tenue,

spiral

short

Ehrb.

Filament slightly tortuous, three to four turns of the spiral length and diameter of a single turn, 2 to 3 fi. When the filament has a turn and a half, it resembles an fl the filaments of two to five turns have a length of 4 to 15 /a spiral movement very rapid.
;
;

In infusions,
S. undula,

etc.

Ehrb. (Vibrio prolifer, Ehrb.)

Filament larger, turns of the spiral wider apart (from 3 to 5 /a) having usually one half a turn to one full turn, rarely one and a half, two, or three length 8 to 10 fi, breadth 5 /x, thickness of filament 1.3 fi having a very rapid spiral movement.
; ;

Fetid animal and vegetable infusions and running water.

The
color.

S. rufum, Pertz only differs from this

by

its

reddish

S. volutans,

Ehrb.

Filament large and thick, turns of spiral regular,


well separated, and 13

in height

number

of turns

two, three, and three and a half, rarely six and seven total length 25 to 30 ft, thickness 1.5 fi, breadth 6.6 fi;

movement sometimes
(Cohn, Warming).

rapid, sometimes motionless a well-defined ciliurn, already seen by Ehrenberg

found in vegetable and animal infusions, in sea-water, and in


This giant of the bacteria
is

fresh water.

FromMicr.Jourrr.Vol.XlH.lfS

P1V

A-MeiseUitli..

PLATE
From
Fig.
1.

V.

" Microscopical Journal."

Micrococcus

prodigiosus

(Monas

prodigiosa, Ehr.).
;

Spherical
the other
this

bacteria of the red pigment, aggregated in pairs and in fours


one.

pigment bacteria are not distinguishable with the microscope from

Micrococcus vaccinal. Fig. 2. Spherical bacteria, from pock-lymph in a state of growth, aggregated in short four to eight-jointed straight or bent chains, and forming also irregular cell-masses. Fig. 3. Zoogloea-form of micrococcus, pellicles or mucous strata characterized by granule-like closely set spherules. Fig. 4. Rosary chain (Torula-form) of Micrococcus urece, from the

urine. Rosary-chain and yeast-like cell-masses from the white deFig. posit of a solution of sugar of milk which had become sour. Saccharomyces (Cryptococcus Fersen.), a pulluFig.
5. 6.

glutinis

glutinis,

lating yeast potatoes.

which forms beautiful rose-colored patches on cooked

Bacterium-pellicle, formed by rod-shaped bacteria arranged Fig. one against the other in a linear fashion, from the surface of sour beer. Bacterium free motile form. Fig. Zoogloea-form of B. Fig. Motile filamentous Bacteria, with a spherical, or Fig. highly refringent " head," perhaps developed from gonidia. Bacillus short cylinders and longer, very Fig. motile filaments, some of which are in process of Bacillus single segments and longer threads, some Fig.
10.

Sarcina spec * from the blood of a healthy man,** from the Fig. 7. surface of a hen's egg grown over with Micrococcus luteus, forming yellow patches. Bacterium termo, free motile form. Fig. 8. Zoogloea-form of Bacterium termo. Fig. 9.

11. 12.

lineola,

lineola.

13.

elliptical

14.

subtilis,

flexible

division.

15.

ulna,

breaking up into segments.


Fig. 16. Fig.
bits,

Vibrio rugula, single or in process of division.

17.

Vibrio serpens, longer or shorter threads,

some dividing

into

at* two threads entwined.


tenue,

Fig. 18. Fig. 19. Fig. 20. Fig. 21. Fig. 22.

" Swarm " of the threads Spirillum single and felted into " swarms.'' Spirillum undula. Spirillum volutans* two twisted around one another. Spirochmte
V. serpens,

felted.

spirals

plicatilis.

lens No. IX. of

All the figures were drawn by Dr. Ferdinand Cohn with the immersion Hartnack Ocular III., representing a magnifying power

of 650 diameters.

96

MORPHOLOGY OF THE BACTERIA.


M. Warming
has recently described three
:

found upon the coast of Denmark


Sp. violaceum, height 8 to 10
;

new

species

p,

diameter 1 to 1.5

p, thick-

ness 3 to 4 p a cilium at each extremity. Sp. Rosenlergii, height of helix 6 to 7.5 p, thickness 1.5 to
2.6 p.
<S^>.

attenuation,

body very attenuated

at the

two extrem-

ities,

without a cilium.

We
g.

give below some details concerning the


:

other colorless Schizophytes


Sarcina, Goods.

The Sarcina, which


here, can be

it

is

useless to describe

considered as bacteria in which

the division occurs by two perpendicular partitions

in

such a manner that multiplication


is

takes place in two directions.

Sarcina

very nearly
it

allied to

Merismopedia,
of

from which
chlorophyll
;

only differs by the absence


siliceous skeleton allies it

its

with

the diatoms.
g. Ascococcus, Billr.

Cells

hyaline, small,

globular, closely united

in

globular or oval families, irregularly lobed and lobulated,

surrounded by a thick gelatinous envelope,

cartilaginous, forming a soft

membrane,

flaky, easily

separating.

A.

Billrothi,

Cohn.

Families in masses of 20 to 160 p, surrounded by a thick membrane of 15 p.

In a solution of tartaric acid exposed to the


g. Myconostoc,

air.

Cohn.

Filaments very slender, colorless, folded, rolled up in a mucous substance, united in very small globules.

CLASSIFICATION OF THE BACTERIA.

97

M.

gregarium, Cohn.

Unique
infusion.

species

found on the surface of

putrefying

g. Cladothrix,

Cohn.

Filaments in form of leptothrix, very slender, colorless, not articulated, rigid or a little undulating, falsely diehotomous.
Gl. dichotoma,

Cohn.

In foul water.

g. Streptothrix,

Cohn.

Filaments in form of leptothrix, very slender, colorless, not articulated, straight or slightly spiral, a little branched.
Sir. Farsteri,

Cohn.

In concretions in the lachrymal canal of man.

PLATE
From photo-micrographs made
produced by permission of
Fig.
1.

VI.
Re-

in

the

Havana and New Orleans. National Board of Health.

ana, Sept. 1879.


2.

Spirochmte From foul bilge-water, Havobjective. X 1,450 by Zeiss's Vibrios from water of harbor, Havana, near discharge Fig. of sewer, Aug., 1879. X 1,450 diameters by Zeiss's ^ objective. Sarcina From standing water in flower-vase, Fig.
(plicatile?).
-fa

in.

in.

3.

(sp. ?).

Lafayette cemetery,

1880. 400 diameters by Beck's \ in. objective. From foul gutter-water, New Fig. 4. Spirillum (volutans ?). Orleans, May, 1880. 600 diameters.

New Orleans, April,

Plate

VI.

Fig.

4.

PART SECOND.
PHYSIOLOGY OF THE BACTERIA.

CHAPTER

I.

DEVELOPMENT OF THE BACTERIA.


The
bacteria are

now known
:

to us

from a mor-

proceed to study the life of these microscopic beings ; first, from a general point of view, that is to say, by studying their functions of nutrition and reproduction,
let us

phological point of view

independently of the special characters impressed upon these functions by certain media; then by
considering the relations which are established between the bacteria and the particular media in which they may be developed. The bacteria are of all beings the most widely diffused. We meet them everywhere, in the air, in water, upon the surface of solid bodies, in the interior of plants and animals. If we expose

a transparent liquid containing traces of organic substances, we find after a short time that it has

become clouded, and the microscope shows us that


it

contains myriads of these beings.

What

is

the source of these organisms so widely

disseminated, and which develop so rapidly?

This

PLATE

VII.

Dissemination of the Bactekia.

From photo-micrographs made

in New Orleans. Copied by permission of the National Board of Health. salt

Spirillum (Sp.?) from water of marsh., near Salem, 400 diameters by Beck's \ objective. Bacteria water (see note on page 107). X 1,000 diameters by Zeiss's fa objective. Fig. Leptothrix buccalis, from human mouth. X 1,000 diameters by Zeiss's fa objective. Bacteria from human Fig. X 1,500 diameters by
Fig. Mass. Fig.
1.

in.

2.

in distilled

in.

3.

epithelial- cell, etc.,


in.

4.

fceces.

Zeiss's fa in. objective.

Plate

VII.

7 f-4-*?&? &.
'

DEVELOPMENT OF THE BACTEKIA.


is

101

the

first

question which presents

itself,

a ques-

tion which has given rise to long discussions, in

the examination of which

we

shall only enter in

order to give a short historical statement.

1.

Origin
is

of the Bacteria.

The

origin of the bacteria, as of all the other

inferior organisms,
:

manners 1. For some, these organisms are produced by heterogenesis ; that is to say, by creation outright from mineral or organic substances (spontaneous
generation).
2.

conceived in three different

individuals like themselves,

According to others, they come directly from by one of the known


of generation,
it
is

modes
3.

fission, spores, etc.

Finally

believed that they are derived


existing,

from organisms already

and are nothing


of developlife

more than different states or phases ment of known species, of which the
not yet discovered.

cycle

is

examine the latter hypothesis, which is called polymorphism, when we what constitutes speak of the phenomena of reproduction. As to the two first, we will content ourselves with indicating the late works which have appeared for and against each; insisting above all upon the facts which relate to the proof of the presence of bacteria or their germs in the air, water, and
will

We

liquids or tissues of the

human

organism,

blood,

urine, etc.

102

PHYSIOLOGY OF THE BACTERIA.

Pougiven chet and of his pupils, and the arguments by MM. Tre'cul and Fremy, the last facts invoked in favor of heterogenesis are due to MM. Onimus,
Heterogenesis.
of

Since the experiments

Servel, Bastian, etc.

M. Onimus contends that the " proto-organisms

may be

born in media, protected against the air, which contain albuminoid substances." M. Martin sustains an analogous idea. According to him, the bacteria are derived from protein granules. According to Neusch, bacteria are produced in the interior of animal or vegetable cells without any lesion and without coming from the air. To demonstrate this he plunges divers fruits under water, in saline or acid liquids (phosphates, sulphates, carbonate of potassa, etc.), and he finds
there bacteria;
but, according to him, these are

not living organisms, properly so called, but ab-

normal cellular vegetations. M. Servel, decapitating some guinea-pigs, caused the heads, the livers, and the kidneys to fall into At the end a solution of chromic acid, 1 to 100. of several days, the superficial parts were hard : ened but the centre was softened, and filled with
;

bacteria.

The presence

of

bacteria in eggs has several

times been verified, and the favor of their theory.

heterogenists have

hastened to draw an argument from this fact in

M. Gayon explains the appearance of these organisms in the eggs of birds by their presence in the normal state in the
oviducts.

DEVELOPMENT OF THE BACTERIA.

103

Finally, Bastian, having succeeded in obtaining

bacteria in liquids which he believes deprived of

every germ, believes in their spontaneous generation. The following is a resume of his experiment Normal acid urine is brought to the boilingpoint, then a solution of potash (in sufficient quantity to neutralize the volume of urine employed) is also brought to the boiling-point after cooling, the two liquids are mixed, and the whole placed in an oven at 50. At the end of two or three
;

days, bacteria are developed.

Pasteur points out three causes of error in the

experiment of Bastian 1. The germs may come from the urine 2. The germs may come from the
:

solution

of

potash

3.

The germs may be

fur-

nished by the vessels employed in the experiment.

In support of this criticism, Pasteur has made some similar experiments, guarding against these causes
of error,

and has not obtained

bacteria.

Dissemination op Bacteria in Air and Water.


Air. The experiment of Pasteur for gathering He fixes a atmospheric germs is well known. window-blind. glass tube in an aperture made in a

The extremity
with the open

of

the tube, which communicates


closed with a plug of cotton,
is

air, is

to the other extremity

attached an aspirator.

When
filled

the air has filtered through the cotton for


this is

some hours,

examined, and

is

found to be

with germs.

104

PHYSIOLOGY OF THE BACTERIA.

Before Pasteur, Ehrenberg and G. de Claubry had already announced the presence in the air of the eggs of infusoria. Robin had also recognized that the atmosphere contains, in addition to all
sorts of
insects,

debris, .spores, pollen-grains, portions of

and rarely the eggs of, infusoria. More recently Maddox and Cunningham, by the aid of an aeroscope invented by the former, gathered

numerous microbes, as well as bacteroid particles. Tyndall, by causing a ray of light to enter a darkened chamber, has rendered visible all these minute corpuscles. His researches show that the optical examination of air enables us to determine in an exact manner the presence or absence of germs. Let us also mention the experiments recently made by Miquel in the park of Montsouris. This observer has found in the atmosphere a considerable number of germs. For the forms of which the diameter exceeds 2 fi, he has ascertained that " the average number of microbes in the air is feeble in winter and augments rapidly in spring,
etc.
;

2.

of these microbes;

That rain always diminishes the number 3. That rain-water introduced

with the greatest precautions, into flasks with slender curved necks, first heated to destroy germs, rarely contains rotifers, etc., but always contains
bacteria."

En
tion.

resume, the existence of germs can be dem1,

onstrated,

by

direct research

and

2,

by

cultiva-

Direct research

may

be made by the optical


of Tyndall), the

examination of the

air

(method

DEVELOPMENT OF THE BACTERIA.

105

microscopic examination of dust (method followed

by Marie-Davy,
particles obtained

Tissandier), the examination

of

by filtration, by gathering germs with an aeroscope, by condensation of atmospheric moisture upon refrigerating vases., etc. The cultivations consist in exposing to the air which
is

to

be examined some liquids in which all pre-existing germs have been destroyed (Pasteur, Tyndall, etc.). This method has shown that liquids exposed in an atmosphere deprived of all germs does not undergo putrefaction, but this occurs as soon as the access of air not deprived of germs is permitted (Tyndall). All of these methods give concordant results; deposits containing germs of various kinds are always obtained. But this objection presents itself
to the

mind

Do

the bacteria obtained


?

by

cultiva-

tion exist in the atmosphere

or do they

come

from germs which have developed rapidly upon


finding a favorable

medium ?

Prom

the experi-

Cohn, Miquel, etc., it is known that the atmosphere contains very few adult bacteria. Miquel in a recent communication says, in effect, that bacteria are rarely found in the air in a complete state, but rather under the form of shining points, difficult to distinguish directly one species from

ments

of

another.

Are not these

brilliant points

Micrococci?
spores,

speaking of the reproduction of the bacteria, develop at a certain


shall see in

In other terms, the organisms which, as

air contains

permanent

we

period of the existence of the adult forms, in their


interior,

which escape from the sporogenous

fila-

106

PHYSIOLOGY OF THE BACTERIA.

ment, are drawn into the air by the evaporation of the liquid containing them, or, after dessication, by the winds. These spores are the point of departure of epidemic foci, and their extreme lightness explains how readily they are" disseminated by the
winds.

Water.

Water contains considerable quantities


and especially of germs.
is

of bacteria

Their pres-

ence in atmospheric water

established

experiments of Lemaire and Gratiolet^

them by more recent


densers
filled

observers,

by means

and

by the
after

of con-

with

ice,

and placed in the

fields

and

for comparison in closed apartments.

Rindfleisch

has since expressed the opinion that the vapor of water does not contain spores or bacteria, and that telluric waters alone contain them but Billroth, Cohn, and others have proved that Rindfleisch was
;

too positive in his statement.


It is

not surprising that telluric waters contain

such a quantity of bacteria that their existence is admitted by all. The dust gathered upon the surface of stones, of leaves, of fruits, etc., shows

upon

microscopic examination an abundance of germs (Marie"-Davy, Tissander) ; the washing of these


objects and of the soil

by the

rain transports
rivers to

them

into the rivers

and from the

the sea,

which contains considerable quantities of them. Thus, a drop of water from the Seine, according to Pasteur and Joubert, is always fecund, and may
give birth to several species of bacteria.
tilled

The

dis-

water of laboratories also contains germs, and

DEVELOPMENT OE THE BACTERIA.

107

these of so small a diameter that they pass through


all filters.

arrested

Cohn has proved by a super position of


1

that

some are not


filters.

sixteen

The

only waters which do not contain them are those drawn from the very source of a spring.

Dissemination op Bacteria in the Organism.

Human

If bacteria are so generally disseminated in the

great external media,

it is

not surprising that they

are found on the surface of -the

human body and

in the interior of the organs in communication

with the exterior. But to account for their presence in the interior of organs we find ourselves in presence of two hypotheses one admitting the spontaneous production of these organisms in the interior of the tissues, the second explaining it by the introduction through the membranes of the
:

germs

of bacteria

from without.

1 Having been directed by the National Board of Health' to make some experiments with a view to confirming or disproving the results of Klebs and Crudelli, who claim to have found the germ of malarial fevers

in the atmosphere of the Pontine marshes near


larias), I

aspirated ten gallons of air on the edge of a

Rome (their Bacillus maswamp in the vicin-

New Orleans, through 4 c.c. of distilled water. Upon examining water with the microscope on the following morning, I was surprised to find a large number of actively moving bacteria and monads [Monan
ity of
this
lens).

To make

sure that these really

came from

the air, I

examined

my

distilled water,

which had been standing

in the laboratory for several

but occasionally opened as distilled water was in considerable numbers, not so numerous, however, as in the water through which swamp air had been drawn. As the germs were present in the distilled water, I presume that the passing of air through it for several hours, and the organic

weeks

(in a bottle, corked,

required) and found the

same forms present

these micro-organisms.

favored the development and multiplication of Subsequent experiments with freshly distilled water gave very different results as to the number of organisms found.

matter contained

in

it,

See

fig. 2,

plate

vii.

G. M.

S.

108

PHYSIOLOGY OF THE BACTERIA.

In truth, the cutaneous surfaces are penetrated with difficulty by germs, although the hairs upon the surface of the body serve to collect them.

The

short hairs in the nares prevent, to

some ex-

tent, the atmospheric

germs from penetrating into


is

the bronchi, but this protection

not sufficient
in the al-

and, notwithstanding the


veoli of the lungs,

mucus

of the nasal fossae

and of the pharynx, they may be found


if

we may

believe Rindfleisch

and Eberth. Do the bacteria pass into the blood ? They may be transported in food and drink into the alimentary canal, where an elevated temperature, the

presence of saliva,

etc.,

favor their dejuice

velopment.
moderate,
if

On

the other hand, the acid secretions

of the stomach, the bile,

and the pancreatic

they do not prevent, the multiplicabacteria in normal blood and

tion of these organisms.

The presence of

urine, or their occasional entrance into these fluids,

are important questions, which have induced


1

many

contradictory researches, but which are not yet


definitely settled.

is any organism in the blood of yellow-fever demonby the highest powers of the microscope as at present perfected, the photo-micrographs taken in Havana should show it. No such organism is shown in any preparation photographed immediately after collection. But in certain specimens kept under observation in culture cells, hyphomycetous fungi and spherical bacteria made their appearance after an interval of from one to seven days. The appearance of these organisms was, however, exceptional and in several specimens taken from the same individual at the same time, it occurred that in one or two a certain fungus made its appearance, and in others it did not. This fact shows that the method employed cannot be depended upon for the exclusion of atmos1

"If there

strable

pheric germs, but does not affect the value of the result in the considerable

number

of instances in which no development of organisms occurred

DEVELOPMENT OF THE BACTERIA.

109

Two
blood.
ination,

kinds of researches have been undertaken

for the purpose of discovering

germs

in

The

direct method, or microscopic

normal exam-

has given results very

much
is

disputed.

The blood
and which
coccus.

contains, indeed, a considerable

number

of little granules, of

which the nature

doubtful,

to distinguish from MicroThus, while Liiders asserts that normal blood contains germs, or spores, which only await a favorable alteration in the fluid in order to deit is difficult

velop themselves, Rindfleisch formally denies their


existence.

The

indirect method,

which

consists in cultivat-

which blood, in a moist state was kept under daily observation for a week or more. " The method employed seemed the only one practicable for obtaining blood from a large number of individuals without inflicting unwarrantin culture cells in

able pain and disturbance upon the sick.


patient's fingers

with

back from
glass

It was as follows One of the was carefully washed with a wet towel (wet sometimes alcohol and at others with water), and a puncture was made just of the matrix of the nail with a small triangular-pointed trocar hypodermic syringe case. As quickly as possible a number of thin covers were applied to the drop of blood which flowed. And these
:

were then inverted over shallow cells in clean glass slips, being attached usually by a circle of white zinc cement. In dry preparations, which are most suitable for photography, the small drop of blood was spread upon the thin glass cover by means of the end of a glass slip. " The thin glass covers were taken from a bottle of alcohol, and cleaned immediately before using; and usually the glass slips were heated shortly before applying the covers, for the purpose of destroying any atmospheric germs which might have lodged upon them. These precautions were not, however, sufficient to prevent the inoculation of certain specimens by germs floating in the atmosphere (Penicillium and micrococci) and in nearly every specimen the presence of epithelial cells, and occasionally a fibre of cotton or linen, gave evidence that under the circumstances such contamination was unavoidable. It is therefore believed that any organism developing in the blood of yellow-fever, or of other diseases collected by the method described, or by any similar method, can have no great significance, unless it is found to develop as a rule (not occasionally) in the blood of patients suffering from the dis;

110

PHYSIOLOGY OF THE BACTERIA.

ing normal blood in flasks perfectly closed, has also given some favorable results, such as those of

Nedvedsky, and and " conPasteur. According to Nedvedsky, the blood tains germs capable of undergoing in it, under certain circumstances, an ulterior development: these are the Hemococcos." If these germs do not
Hensen,
Tiegel,
Billroth,

and

some unfavorable

results, as those of Liiders

give birth, normally, to bacteria,

it is

because the

blood

is

as injurious to

them

as the

most advanced
If this

stages of putrefaction (Billroth).


esis is true, it

hypoth-

explains several embarrassing facts,

such as the existence of micrococci in the pus of


ease in question, and
is

proved by comparative

tests

not to develop in

the blood of healthy individuals, obtained at the

same time and by the

same method. " Tried by this test, it must be admitted that certain fungi and groups of micrococci, shown in photographs taken from specimens of yellowfever blood collected at the military hospital and preserved in culture cells, cannot reasonably be supposed to be peculiar to or to have any Preliminary Report of Havana Commiscausal relation to this disease." sion to National Board of Health. In subsequent observations upon the blood of malarial fever, of syphilis, and of leprosy, I have sometimes obtained a development of micrococci in culture cells where all possible precautions as to the exclusion of atmospheric germs had been taken, and in one case have seen

the development of Penicillium in another of Sarcina.


tion
is,

The

last observa-

so far as I

know

unique, and I have

still

in

my possession

the

containing numerous masses of Sarcina, presenting the This slide was put up characteristic arrangement of the cells in fours.
culture-slide

from intermittent fever in the CharEvery precaution was taken to exclude atmospheric germs. The patient's finger was washed with absolute alcohol just before making the puncture from which the little drop of blood was obtained. The question as to whether in this and similar cases the germs of the organism which develops come from the atmosphere or pre-existed in the blood is one to which I propose to give special attenat the bedside of a patient suffering
ity Hospital,

New

Orleans.

and, after further experiment, I shall discuss G. M. S. National Board of Health.


tion
;

it

in

my report to

the

DEVELOPMENT OF THE BACTERIA.


closed abcesses, in cysts, in urine bladder, etc.

Ill

drawn from the

2.

Nutrition
,

and Eespiration of the Bacteria.

cell

The bacteria, being organisms composed of a membrane of cellulose, and of protoplasmic


of

contents, deprived

chlorophyll,

nutriment and respire in the colorless vegetables and all the inferior animals deprived of special apparatus, that is to say, by endosmotic absorption. Although the media in which the bacteria develop are various, yet, from the point of view of the nutritive function, they act everywhere according to the same laws. No matter in what medium they live, they must have water, nitrogen, carbon, and oxygen, as well as certain mineral salts which enter, but in quantities exceedingly minute, into the chemical constitution of all organ-

must receive the same manner as all

ized bodies.

element is indispensable to the active life and development of the bacteria. Dessication arrests completely the movements of those which are mobile, and the functions of all the bacteria in general; but it does not kill them, at least if it be not prolonged beyond a certain time. The Micrococci of different kinds of virus are examples of the continued vitality of these organisms after dessication for a considerable time.
Water.

This

112

PHYSIOLOGY OF THE BACTERIA.


bacteria present in this respect

The

numerous va-

and the period of development which they have attained. In the state of permanent spores, they are extremely tenacious of vitality. They resist for a long time not only dessication, but a considerable elevation
riations according to the species
of temperature.

Among
uids,

the

the bacteria, some are developed in

greater number,

others upon
A

liq-

damp

surfaces.

The former can

live in fresh water, sea-

water, thermal waters, and the liquids of animal


or
is,

vegetable organisms, etc.

surprising fact

that the composition, so different, of fresh and

sea water appears to have no influence


bacteria.

We

find in both all the species,

upon the from

Bacterium termo to Spirillum volutans.


Nitrogen. Pasteur has demonstrated that it is not necessary that the nitrogen which is to serve as

nutriment to the bacteria should be in the form of albumen, but that these organisms can take possession of
it

in the

form

of

ammonia.
fol-

In
lows
:

fact, in

Pasteur's solution, composed as

Distilled water

100.
10.

Sugar candy
Tartrate of

Ashes of

ammonia one gramme

....
of yeast
.

1.

0.075.

the bacteria increase sometimes with such rapidity


that they interfere with the development of the
alcoholic ferment.

DEVELOPMENT OF THE BACTERIA.

113

Cohn, in order to better observe the phenomena and to get rid of the moulds, which the cane-sugar caused to develop too rapidly, employed the following culture-fluid
:

Distilled water

....
.

100.
1.
,

Tartrate of

ammonia
. .

Ashes of yeast

1.

Bacteria develop in this fluid wonderfully, which proves that sugar is not indispensable to them. One other solution often employed is that of Mayer. It has the advantage of not requiring the

employment

of ashes of yeast

Phosphate of potash Sulphate of magnesia Tribasic phosphate of lime Distilled water

... ...

0.1 0.1

gramme.

c.c.

0.1

20

Cohn adds

to this 0.2 gr. tartrate of

ammonia.

En

resume, the bacteria can take nitrogen, which

they need in order to form their protoplasm, either from albuminous compounds, which they decompose, as in putrefaction, or in the form of ammonia, or, perhaps, by borrowing it from nitric acid, but this last source is not well established
(Cohn).

Carbon. In addition to the sources common to other organisms, the bacteria can take this important element of their composition, under certain circumstances,

from the organic


8

acids.

Thus,

when we

cultivate bacteria in a solution containing

114

PHYSIOLOGY OF THE BACTERIA.

only tartrate of ammonia with a small quantity of


mineral
salts

(phosphoric acid, potash, sulphuric

and magnesia), they develop rapidly, taking their carbon from the tartaric acid. Cohn has endeavored to ascertain if other organic acids could be assimilated by the bacteria. By making use of succinate of ammonia, or neutral acetate of ammonia, he has been able to cultivate these microphytes. Besides, as Pasteur had already experimented with solutions containing lactates, and in which bacteria had developed until the salt had completely disappeared, we may admit that the
acid, lime,

bacteria can assimilate the organic acids,


succinic, acetic,

tartaric,

but tartaric acid seems to furnish the best alimentary solution. Other substances containing carbon are also aslactic
;

and

similated

by the

bacteria,

cane-sugar, milk-sugar,

glycerjne, and even cellulose (according to Mitscherlich).

Cohn
find

concludes, " that the bacteria multiply quite

normally, and in great quantity, whenever they

elements in solution which constitute ashes, and that they can take the carbon which
the

they need from any organic substance containing and their nitrogen from ammonia, urea, and probably from nitric acid. The bacteria, then, resemble green plants, in that they assimilate nitrogen contained in their cells by taking it from
it,

ammonia compounds, which animals cannot do. They differ from green plants in that they cannot draw their carbon from carbonic acid, and only
assimilate organic

substances containing carbon,

DEVELOPMENT OF THE BACTERIA.


above
atives
all
;

115

the hydrates of carbon and their derivin this respect they resemble animals."

and

Absorption.

How are these various substances


observations of

absorbed

The
etc.,

Grimm, Hoffmann,

de Seynes,
stances

these organisms absorb

permit us to assure ourselves that by endosmosis the sub-

upon which they are nourished. Grimm, upon examining with the microscope some particles of lemon containing bacteria and spores of algse, saw a certain number of the former
gather aro\md a spore, and
fix

themselves to

it

by one
trate
it

of their extremities.
;

They

did not pene-

but when they abandoned it, the spore had diminished in volume, and lost a portion of its contents, while the bacteria had taken a greenish
color.

Hoffmann has seen

that these
of

little

organisms,

carmine or of fuwhen schine, after a time are colored an intense red, while the mucus surrounding them remains colorplaced in a solution
less.

De

Seynes,

also,

from

his

observations

upon

the vibrios which accompanied some colored filaments of Penicillium glaucum, believes that bacteria are susceptible of absorbing coloring matters

from vegetables and from animals with which they


are in contact.

Oxygen.
First, it

The

role of

bacteria has given rise to

oxygen in the life of the numerous controversies.

seems a priori that the bacteria ought

116

PHYSIOLOGY OF THE BACTERIA.

to act like all other living beings,


like the other inferior

and to respire

organisms deprived of chlorophyll that is to say by absorbing oxygen and This is, indeed, the eliminating carbonic acid. But, opinion of a great number of botanists. bacteria. according to Pasteur, it is not so with the

When we
we

examine what occurs in putrefaction,


developed

find that at first certain species are

(Monas crepusculum, Bacterium termo, etc.), which absorb all the oxygen dissolved in the liquid, and come to the surface where they form a thick veil
after
this,

other species of vibrioniens

appear,

which are developed in a medium entirely deprived of free oxygen, by borrowing this gas from the fermentable matters contained in the liquid. These chemical decompositions constitute
putrefaction.

The

first of

these organisms, regarding the na-

ture of which Pasteur has long been uncertain,


: they live in contact with the air, and have need of oxygen. The second, anaerobies, not only have no need of oxygen, but are killed by it. These differences in the respiration of organisms belonging to the same group are not admitted

are aerobics

by a great number of recent observers. Hoffmann, among others, says expressly " These little
:

beings cannot live without

air,

I should say with-

out oxygen

if

this gas is

move and do not

multiply at
is

liquid full of bacteria

wanting, they cease to all. If a drop of placed upon a glass slip,

then covered by a piece of thin glass, the active

DEVELOPMENT OF THE BACTERIA.


bacteria will
of the cover
all

117

approach gradually to the margins it is there that at the end of several days, after the successive death of the greater number, some are still found endowed with
;

and

life

and movement. If a similar preparation is at the same time protected by an impermeable cement against dessication and against the introduction of atmospheric air,
all

movement among

the

bacteria will cease at the end of two minutes, provided, however, that no air bubble has been im-

prisoned with the liquid."

and development of bacteria is also very manifest in an experiment recently made, and not yet published, by Toussaint, who has been kind enough to communicate it to me. In studying the development of the spores of Bacillus anthracis in the moist chamber of
influence of
life

The

oxygen upon the

Ranvier,
those

Toussaint

has

observed

the

following

curious facts, which offer a striking analogy to

from Hoffwhich occupy the central portion of the moist chamber and which by reason of their situation receive very little oxygen from the groove, are soon arrested in their development while those which occupy the borders are long and heaped up in immense numbers, those in the centre remain small, formed of two, four, or five articles, which are easily separated from each other; they soon cease to grow and are not transformed into spores." Cohn is also as explicit. " There is no doubt,"
above
"

mentioned, borrowed
bacteria,

mann.

The

118

PHYSIOLOGY OF THE BACTERIA.

he says, " that the complete development of Bacillus, and above all reproduction by means of spores, is only made under the influence of free access
of air."

We might explain the contradictory facts of Pasteur by admitting, with Cohn, that the appearance of different roles played by the aerobies
(Bacterium) and the anaerobies (Bacillus) is simply due to a veritable struggle for existence which
takes place between the microbacteria and the

desmobacteria.

Action op Various Agents upon the Bacteria.


In
this

paragraph I

shall pass in

review the
of va-

action of temperature, of
rious antiseptics.

movement, and

It is very important to study which bacteria comport themselves under extreme variations of temperature. It is, indeed, upon the results furnished by these researches that a great part of the arguments opposed to the panspermatists by the heterogenists

Temperature.

the

manner

in

are based.

We
of

shall consider the

influence

upon bacteria

moderate temperatures and of extremes above and below zero. Moderate temperatures that is to say those which are comprised between 25 and 40 (77 to 104 Fah.) are generally favorable. The most favorable has been found to be 35 (95 Fah.)

(Onimus).

DEVELOPMENT OE THE BACTERIA.

119

of resistance to extreme temperavery variable, according to the species. Thus, according to Frisch, a temperature of 45 to 50 (113 to 122 Fall.) is sufficient to kill B. termo, u Barteriwhilst 80 (176 Fah.) does not kill the

The degree
is

tures

dies " (Bacillus).

The permanent spores are especially remarkable by the tolerance which they possess for high temperatures. They have been subjected to 100
(212 Fah.) (Schwann), 110 (Pasteur) and even 130 (Schrader) without losing their power of

germinating.

We

must, however, recognize that the results

of the experimenters offer the greatest diversity,

the result, according to Cohn, of the difficulty of


obtaining an equable distribution of the heat in
the media, which are generally bad conductors.

Cohn has

arrived at the following conclusions as

the result of numerous experiments


the Bacillus of
1.

hay infusions At a temperature of 45


the

made upon

to 50 (113 to 122

Fah.) other

Bacillus

still

multiplies

rapidly,

and
of

forms as usual membranes and spores, while .the


schizophytes
existing
in

the

infusion

hay are
plication.
2.

at this temperature incapable of

multi-

At a temperature
all

Fah.)

ceases.

50 to 55 (122 to 131 reproduction and development of Bacillus It neither forms pellicles or spores; the
of

filaments are killed, the spores,

on the contrary,

preserve, for a longer time (for at least seventeen

hours) the property of germinating.

120
3.

PHYSIOLOGY OF THE BACTERIA.

While infusions

of

hay are generally

sterilized

by a temperature

of 60 (140 Fah.) or more, pro-

longed during twenty-four hours, certain spores of Bacillus seem able to endure a temperature of 70 to 80 (158 to 176 Fah.) during three or four days without losing the power of germinating. By some experiments made with refrigerating
mixtures,
are not killed

Cohn has ascertained that the bacteria by very low temperatures, acting

18 for example during several hours, (0 Fah.). But they are benumbed at a temperature of 0 (32 Fah.) and probably at a temperature

even

little

higher, losing the

of reproduction,

ferments.
to

power of movement and and consequently their action as They preserve, however, their capacity
activity at a

resume their

more elevated tem-

perature.

Frisch has pushed the experiment still further than Cohn. By the evaporation of carbonic acid, 87 he has produced as low a temperature as a 123 Fah.) in liquids containing bacteria, with( out destroying the vitality of these organisms,

cos

development having subsequently occurred of cocand of bacteria. Congelation, then, cannot

serve to destroy the organized ferments.

Let us add, however, that if the passage to extreme temperatures is too sudden, there is then an
alteration (destruction ?) of these organisms (Schu-

macher).

Movement. We would not have consecrated a paragraph to the action of movement upon

DEVELOPMENT OF THE BACTERIA.


bacteria,
if

121

Crova had not recently asserted that

liquid containing completely arrests their development. This is an assertion in complete opposition to all that we know of the physiology of these organ-

movements impressed upon a


bacteria

isms,

and which

it

is difficult

fact that bacteria

may

to reconcile with the develop even in the torrent

of the circulation.

fluence of
bacteria.

Compressed Air. We have just seen the inair, and especially of oxygen, upon the

When

this

agent

is

in a certain state of

it acts in a different manner. M. Paul Bert has proved that under a tension of twentythree to twenty-four atmospheres all the putrefactive processes depending upon the development of

tension,

vibrios cease to occur. Since, the same savant has found that the anatomical elements and even the red blood globules are killed by oxygen.

These researches agree well enough with those Grossmann and Mayerhauser upon the life
bacteria
in
gas.

of of

From

their

numerous experi-

ments

it

appears that, under the influence of oxyis

gen, there

the bacteria; but

an exaggeration of the activity of if the oxygen is under a presdie,

sure of five to seven atmospheres, the bacteria live

from

six to

twenty hours, then

and cannot be

resuscitated

by atmospheric
definite

air.

Ozone causes a
arrest of

and almost instantaneous

movement.

122

PHYSIOLOGY OF THE BACTERIA.

Other gases studied by the same savants have given the following results Hydrogen at first causes an acceleration of movement, which is maintained for several days
:

then movement becomes


ceases altogether.

less active,

and

finally it

by

Contrary to the facts stated Carbonic Acid. Pasteur, this agent was found to paralyze the bacteria, and reduced them to complete immobility. If the carbonic acid is displaced by oxygen, the
bacteria resume their activity.

Chloroform.

This substance, according

to the

researches of Miintz, arrests the vital


of organized ferments.

phenomena
upon

Miintz uses this charac-

ter in order to recognize the soluble ferments,

which

it

has no action.

Since the labors of Dumas, Boracic Acid. which have demonstrated that boracic acid kills the inferior organisms by depriving them of their oxygen, this substance has been employed in various circumstances as an antiseptic.

Sulphate of Quinine.
is

The

action of quinine,

either in the state of chlorhydrate or of sulphate,

not yet well established.

The experiments

of

Binz, Manassein, Kroevitsch, Bochefontaine, etc.,

have, in truth, given contradictory results.

Carbolic Acid.
sei'n

The

have demonstrated

.that

experiments of Manas^th. per cent of car-

DEVELOPMENT OF THE BACTERIA.


bolic acid is sufficient to prevent all

123

development It is employed with success in anthrax, in the treatment of wounds, etc.


of living beings.

3.

Keproduction
fission,

of the Bacteria.

It
tiply

is

well established that the bacteria can mulalso

and reproduce themselves the formation of endogenous spores. Fission. The multiplication by fission

by

by

consists
a bac-

in a transverse division of the

cell.

When
its

terium has attained nearly double


length,

ordinary

we

see, in the larger species, that the proto-

plasm becomes clearer in the central portion, and a partition forms in the median line separating the contained protoplasm into two portions. The partition, at first very delicate, becomes thicker, divides into two, and the two articles separate. This phenomenon is produced more or less
quickly according to the nature of the medium,
its

richness in nutritive material, the temperature,

etc.

When

the growth

is

rapid, the

new

cells

form

more quickly than they separate, and are arranged Very often we only find them in in chaplets. this form, in strings of two to four cells coupled In some forms the transverse division together. is preceded by constriction near the middle of the Before the two new cells are separated, the cell.
bacterium in this case presents the appearance of
a figure 8, and seems to be a simple cell swollen
at the

two extremities.

124

PHYSIOLOGY OF THE BACTERIA.

Under other circumstances, and probably in consequence of a mucus transformation of the walls
mother oped by a mass
of the
ogloea.
cells,

the

new

bacteria are envel-

of glutinous substance.

We
of

described these masses under the

name

have Zo-

The
fission

conditions which favor multiplication


are, a certain

by

degree of temperature and

a sufficient quantity of nutritive material. The more rapid higher the temperature, the is the segmentation of the bacteria, the more rapid their
multiplication,
limit, variable

that

is

to

say,

up

to a certain

with the species and beyond which

the bacteria are destroyed.

The
ture
is

multiplication decreases

when

the tempera-

lower, and ceases entirely in the vicinity


influence of richness of nutriment
artificial cultivation.

of 0 (32 Fah.).

The
seen in
to

is

well

So long as the bacte-

ria find the necessary aliment, in sufficient quantity,

form new protoplasm, they multiply with acbut as soon as the organic matter is devoured, they cease to divide, fall to the bottom of the vessel, where they accumulate, motionless, and form a deposit more or less abundant.
tivity
;

The

multiplication of the bacteria


if

sion has for result,

by binary fisnothing occurs to interfere

with the most favorable conditions, the invasion of the medium by an incredible number of these little beings, of which we can only form an idea

by

calculation.

" Let us suppose," says Cohn, " that a bacterium

DEVELOPMENT OF THE BACTERIA.


divides into

125

two

in the space of an hour, then in

four at the end of a second hour, then in eight


at the

end

of three hours, in twenty-four hours


will already

the number

amount

to

more than

six-

teen millions and a half (16,777,220); at the end of two days this bacterium will have multiplied
to the incredible

number
it

the end of three days

281,500,000,000; at will have furnished fortyof

seven

trillions

at the

end

of

about a week, a

number which can only be represented by fifty-one


figures.

" In order to render these numbers more comprehensible, let us seek the volume and the weight

from the multiplication of a The individuals of the most single bacterium. common species of rod-bacteria present the form of a short cylinder having a diameter of a thousandth of a millimeter, and in the vicinity of one five hundredth of a millimetre in length. Let us represent to ourselves a cubic measure of a millimetre. This measure would contain, according to what we have just said, 633,000,000 of rod-bacteria without leaving any empty space. Now, at the end of twenty-four hours the bacteria coming from a single rod would occupy the fortieth part of a cubic millimeter; but at the end of the following day they would fill a space equal to 442,570 Let us of these cubes, or about a half a litre. admit that the space occupied by the sea is equal to two-thirds of the terrestrial surface, and that its mean depth is a mile, the capacity of the ocean

which

may

result

will be 928,000,000 of cubic miles.

The

multipli-

126

PHYSIOLOGY OF THE BACTERIA.

cation being continued with the

the bacteria issuing from a single


the ocean in five days."

same conditions, germ would fill

Reproduction by Spores.

The
mode

multiplication
of propagation

by

fission,

known

to the earliest microscopists, has

been

until recently the only

admitted by the authors.


in the excellent article

Thus M. de Lanessan, which he has devoted to

the bacteria, says that the marvellous resources of

modern

science have not yet permitted us to rec-

ognize any other organisms.

mode

of propagation for these

However, M. Ch. Robin had already, in 1853,


indicated the presence in Leptothrix buccalis of
little

round bodies, " which are perhaps spores."

Pasteur has since, in 1865, recognized that " the vibrios of putrefaction and of butyric fermentation present a sort of ovule, or ovoid corpuscle, which
refracts

light

strongly, either

in

the extremity
Later, the same

or in the body of the articles."


savant,

more

explicitly, says clearly that these or-

ganisms have two modes of reproduction,


fission

by

and by interior spores (" noyaux "). Towards the same epoch, Hoffmann also pointed out a reproduction by free cellular formation in some bacteria. But we must come to the labors of Cohn, Billroth, and Koch, in order to find precise observations in this regard.

The formation
in Bacillus

of

spores

has

been

observed

subtilis

by Koch, and
Tieghem.

in

by Cohn, Bacillus anthracis Bacillus Amylobdcter by Van

Plate

VIII.

Fig.

Pig.

^m

PLATE

VIII.

Formation op Spores in Bacillus.


From photo-micrographs made
in

Havana and

in

New

Orleans.

Re-

produced by permission of the National Board of Health.


Figs. 1 and
eters
2.

Bacillus (ulna
in.

?)

patient (post-mortem) five days after collection.

by
2.

Zeiss's

Fig. Fig.

1.

Fig. 3. Orleans, April, 1880.


large cells are

^ objective. Rods joined in leptothrix chain. A single rod showing spore at one extremity. Spores of Bacillus developed rotten potato,
in

found in blood of yellow- fever X 3,000 diam-

New
The

1,500

by

Zeiss's

in. objective.

some species of Saccharomycete, which was also present in the same specimen. Development of bacilli from spores, from culture exFig. 4. periment with fish gelatine solution. X 1,500 diameters by Zeiss's

in. objective.

128
Cohn,

PHYSIOLOGY OF THE BACTERIA.

who had

in his first publications refused

to the bacteria the property of reproduction


spores, thinking that the facts observed

by

by Hoff-

mann

related to different beings, has verified the

experiments of Koch upon the development of B. anthracis, and has himself demonstrated similar phenomena in B. subtilis. In culture experiments made with infusion of
hay,

we

see, at a

certain
of

moment,

in the

homo-

geneous filaments

the Bacilli very refractive

Each of corpuscles making their appearance. them becomes a spore, oblong or in the form
of

a short filament, highly refractive, and with


outlines.

well-defined

We

find

the

spores ar-

ranged in a simple series in the filaments. As soon as the formation of spores has terminated, the filaments can generally no longer be distinguished, and one would say that the spores were completely free in the mucus but their linear arrangement shows always that they are produced Little by little these in the interior of filaments. and the dissolve, being reduced to a fine powder spores fall to the bottom of the liquid, where they The germination of the are found in abundance. spore does not seem to occur in the same medium but if we take a spore from the deposit formed in an infusion of boiled hay, and transport it into a new infusion, we see the spore swell up, and a short tube form itself at one of its extremities at this moment it resembles a bacterium with a head. Soon the very refractive body disappears, the tube
;

stretches out into a short rod of Bacillus,

com-

DEVELOPMENT OF THE BACTERIA.


mences to move, and becomes jointed by
verse division.

129
trans-

Koch, in cultivating the bacteria of charbon in aqueous humor from the eye of the ox, has observed some facts exactly similar, both as to production of spores in linear series in the filaments
of Bacillus anthracis

and

as to the germination of

the spore and the birth of a


of

new
:

rod.

According to Amylobacter
first,

Van Tieghem,
is

as follows

the development " The development

of a Bacillus includes four successive periods.

In

the

the body, cylindrical

and slender, recently

developed from a spore, stretches out rapidly, the articles soon separate and is partitioned united in long filaments or remain subtilis), (B. This is the stage of growth and (B. anthracis).
;

multiplication,

two things which at bottom are


articles previously

but one. "Secondly, the

formed, having

ceased to elongate and divide, increase sensibly in magnitude, becoming the seat of interior chemical and this increase in size operates transformations
;

according to circumstances, in three different manSometimes ners, with some intermediate forms.
it

occurs uniformly throughout the length of the


;

which remains cylindrical sometimes it is localized, either at one extremity, which is swollen
article,

a tadpole, or in the middle of the article, which swells to a spindle shape. This is the stage
like

of enlargement, or of

nutrition, solitary

and

si-

multaneous, which prepares the following state. "In the third period or phase of reproduction

130
there
is

PHYSIOLOGY OF THE BACTERIA.

nourished a spherical or ovoid spore, homogeneous, highly refractive, having a dark outline. At the same time, the protoplasm which occupies the rest of

formed in each

article

so

the cavity disappears


spore from the

little

by

little,

and

is

re-

placed by a hyaline liquid, which separates the

membrane

this dissolves

in its
If the

turn, and finally the spore

is

set at liberty.

article is swollen in tadpole shape, it is in the ter-

minal swelling that the spore has birth ; if it is spindle-shaped, it is near the middle if it is cylin;

drical,

it

may

be at any point whatever, but

is

usually near one extremity.


free

The spore when

set

under favorable circumstances. At a point where its circumference becomes pale,


germinates
it

gives out a little tube slightly more slender than itself, which elongates rapidly and divides. This fourth period of development or germinative

phase brings us back to our point of departure." only do the bacteria develop spores in the interior of their filaments,
Sporangia.

Finally, not

we may also observe formation veritable the of a sporangium containslightly modified in form, but

ing
of

many spores. The unpublished observations M. Touissant, Professor of Physiology in the


result,

Veterinary School of Toulouse, give this which he has kindly communicated to me.
in

In cultivating spores of the bacteria of charbon the serum of the blood of the dog, under the

microscope, in

the warm chamber of Ranvier, Toussaint has seen the filaments take a transverse

DEVELOPMENT OF THE BACTERIA.

131

diameter almost double the ordinary diameter, then the protoplasm of the filament to gather together at certain points, a fact clearly made out, as in the parts where the protoplasm was wanting the bacteria had lost all refractive power.

Finally, at a later period the points occupied

by augment considerably in volume, and form some ovoid organs- more or less
the condensed protoplasm
elongated, or swollen into a ball, or in the form

gourd at one extremity. In the interior of these sporangia, from three to six spores afterward form, clearly defined and highly refractive then, finally, by breaking up of the membranous enveof a
;

lope the spores

become

free.

Toussaint has also followed in the same apparatus

moist and warm chamber


spores are at
first

of

Ranvier

the

mode

of germination of the spores. ing are the most important facts


:

The

follow-

and animated by brownien movements; at the end of half an hour to an hour, at a temperature of 37 to 40, in urine, aqueous humor, or serum, the spores lose their refractive power, and their brownthen the ien movements cease almost entirely spore assumes an aspect slightly granular, it becomes elongated in the direction of its greatest diameter (they are oval). After two hours of cultivation, the bacterium has two or three times the dimensions of the primitive spore the elongation makes rapid progress, and four to six hours from the commencement of the cultivation, some may
highly refractive
;

The

132

PHYSIOLOGY OF THE BACTERIA.

be found to occupy the entire field of the microscope. From this moment the phenomena which occur differ according to the conditions in which the bacteria are placed. Upon the edge of the
air-groove in the moist chamber, the bacteria de-

velop very rapidly, forming an interlaced mass and in sixteen to eighteen hours, spores may be

seen to appear in their interior,

above

all, if

the

preparation has been exposed to light.

Often, in

this case, the transverse partitions of the filament

cannot be seen. If, on the contrary, the bacterium has not been exposed to light, the spores are a longer time in showing themselves (ten or twelve hours more), and almost always division of the
filament precedes their formation.

In that case,

a spore usually appears at each end of the seg-

ment
to

in such a

manner

that the spores belonging


to each

two successive segments are nearer

other than those in the same segment.


also,

Often,

a spore aborts in a segment (Toussaint).

have seen above, in speaking of the ressame observer has noted in the course of his experiments some phenomena proving the evident influence of oxygen upon the development of Bacillus. It is the same
piration of bacteria, that the
for the formation of spores.

We

And upon

this point

Toussaint makes the very just remark that the phenomena occur in a different manner in culture

experiments and in the

human

organism.

In char-

bon, the bacteria never form spores.

They remain

always relatively short, even in the points where they form extra- vascular masses, and where conse-

DEVELOPMENT OE THE BACTERIA.


quently
liquid

133

we cannot invoke
order to
tissues

the movements of the


their
division.
little

in

explain

bacteria of charbon, then, take but

The oxygen

from the
in the

they do not vegetate luxuriantly organism; and certainly, if we judge by a


:

calculation necessarily approximative, their devel-

opment

is

seven or eight times

less rapid

than in

the strongly oxygenated serum of culture experi-

ments (Toussaint).
Polymorphism.

The spores

of

which we have

traced the genesis constitute those germs of which

the origin has for a long time been misunderstood,

those permanent spores or durable spores

(Dauersporen), thus called because of their re-

markable degree of resistance to temperature, desiccation, and all the agents which kill adult
bacteria or arrest their development. These " organs " are disseminated in great

numlittle

bers in various media under

the

form

of

rounded corpuscles absolutely similar to the micrococci

from which
them.

it

is

absolutely impossible
is,

to

differentiate

It
if

indeed, very probable


all

that the greater part,

not

of these organisms,

are the spores of filiform bacteria.

In the impossibility of recognizing these forms nearly related, of referring them to such or such a determined organism, the attempt has been made to cultivate them, in order to follow their development. We have just seen the results of this cultivation for the Bacillus ; but, in the hands
so of the greater

number

of experimenters, the

re-

134
suits

PHYSIOLOGY OF THE BACTERIA.

of such culture experiments are far from being so certain. Not having succeeded in re-

moving them completely from the invasion of foreign germs, the greater number have seen the most diverse forms develop themselves, and from
this

have inferred the most remarkable transfor-

mations.

Thus, Hallier pretends to have observed the


transformation of Micrococcus into various fungi,

such as Mucors, Ustilago, etc. The M. of vaccinia comes from Torula rufescens, which is itself a phase of development of Ustilago carbo ; the M.
of

human

variola

is

derived from a fungus having

sporangia and pycnidia, related to Sternphylium sporidesmium ; that of the variola of animals from Cladosporium (Pleospora) herbarum; the M. of the blood of scarlatina belongs to the those of glanders and of syphilis g. Tilletia from a Coniothecium, etc. In the same way Letzerich has referred the M. of diphtheria to another
;

Tilletia,

the T. diphtkerica.
" of bacteria into " levures

The transformation

(yeast fungi), and these into Penicillium, has been

admitted by Hallier, Tre"cul, and others. But the researches of Brefeld and de Seynes have shown
is far from being demonstrated innumerous cultivations, de Seynes has never been able to verify such an affiliation and Nageli in his turn has never been able to obtain a transformation of schizomycetes into budding

us that this
deed, in his

fungi.
It is the

same as regards the transformation

of

DEVELOPMENT OF THE BACTERIA.


bacteria into moulds and mildews.
cultivations of moulds,

135

In some recent
care, Nageli has

made with

never observed the formation of schizomycetes, and reciprocally. Are we not permitted to believe,

now

sporangia
scopists

that we know of the formation of among the bacteria, that the microwho sustain a polymorphism so extended,

have taken these organs, of which they have not been able to follow exactly the development, for This explanation is the sporangia of Mucorini?
the more admissible as Tre'cul has seen the bacteria " swell up, and transform themselves separately," a

phenomenon

quite identical to that ob-

served by Toussaint.

form well demonstrated in the present state of science, and the only one which can be compared to natural polymorphism, such as it exists in a great number
resume". of of fungi, consists in the transformation of spores

En

The only change

into Bacteria, Bacteridia, Vibrios, etc., and in the

teria take in

modes of grouping that the cells of bacbecoming zooglcea, mycoderma, leptothrix, etc. To go further would be to lack* prudence and scientific criticism.
different

136

PHYSIOLOGY OF THE BACTERIA.

CHAPTEE

II.

DEVELOPMENT OE THE BACTEEIA IN


DIFFERENT MEDIA.
In studying the conditions
of life

and of develop-

ment
and

of bacteria in

the different media, natural

which they are met, we will consider the actions which they determine (or that they accompany) as particular cases of their nutrition and of their reproduction. We will constantly take, then, their normal physiology as our point of departure and we will try to explain in this way the phenomena, so diverse, with which they
artificial,

in

are associated,

fermentations,

putrefactions, con-

tagion of infectious maladies, etc.


It is especially interesting to

study the role of

bacteria in non-nitrogenized chemical media,

where

they accompany the phenomena called fermentation, properly so called in nitrogenized media, vegetable or animal, which they transform, as a result of special fermentations, which constitute putrefaction in the human organism, where they accompany frequently, if not always, the development of certain affections having special characThis will be the object of so many paraters.
; ;

graphs.

THE BACTERIA

IN

DIFFERENT MEDIA.

137

1.

Hole

of Bacteria in Fermentations.
is

We
Two

say that a liquid


its

fermenting whenever

modifications occur in

chemical constitution, as

a result of the nutrition of organized beings.

kinds of fermentation are commonly distin-

guished.

In the

first

group

(false

fermentations)

are arranged those which are produced by soluble

quarternary substances (diastase, soluble ferments)


secreted

by

living cells,

from which they may be


is

separated in order to study their action upon fer-

mentable

liquids.

This action

comparable to that

of certain mineral acids, which operate in the same manner, either by the breaking up of molecules with addition of water or by the phenomena of

hydration.

Veritable

chemical

reagents,

when

these substances are once precipitated from their


solutions, purified

and

dried, they preserve their

properties indefinitely.

A sufficient elevation of tem-

perature seems to destroy the edifice of their molecule ; for they lose all their specific power after having been subjected to a temperature more or less elevated, but always inferior to 100 (212 Fah.). In the second group (true fermentations) are

joined
tion

all

the

phenomena

of chemical

modificaor fungi
kill-

which appear intimately united


of inferior

opment

organisms,

to the devel-

algae

(figured ferments).

Compressed oxygen by

ing these ferments, and chloroform by suspending their vital functions, arrest the progress of these
fermentations, while the same agents do not mod-

138

PHYSIOLOGY OF THE BACTERIA.


Accord-

ify at all the action of soluble ferments.

ing to Dumas, borax has, on the contrary, the

property of entirely destroying the activity of soluble ferments without absolutely preventing certain true fermentations, for example, the al-

coholic fermentation of glucose.

We

will see fur-

ther on that this property of borax has been


utilized in the

treatment of catarrh of the bladfirst

der and of certain virulent affections.

view these two groups of phedifferent, they may, however, be compared the one with the other. Without speaking of the ammoniacal fermentation of urine, which, as we shall shortly see, may be arranged in either of these groups, we may admit that the only difference between these two series of chemical modifications consists in the fact that in one case the true fermentations being the last term in the interior nutrition of the cell have their seat in

Although at

nomena seem very

the interior of the

cell itself

while in the other

the
lar

first

terms of nutrition are always extra-cellufor effect, as CI.

phenomena, having

Bernard

has shown, to render assimilable or diffusible in the interior of the organism the aliment necessary to
the development of every organized being (transformation of starch into glucose, of sugar into glucose, emulsion of fats, liquefaction of albuminoid substances).
these

The study, from a chemical point of view, of phenomena of nutrition, of these fermentasuch
is

tions, since

their
it

much

progress, and

rational classification

name, has not yet made would be difficult to make a of them in the present state

THE BACTERIA
of our knowledge.
sify

IN

DIFFERENT MEDIA.

139

I will not then seek to clas-

them, but will content myself with describI shall only speak of the fermentations

ing them successively, commencing with the best

known.

caused by tbe development of bacteria, leaving, consequently, the fermentation which has been
best studied,

ing order
1. 2. 3.

the

alcoholic.

I adopt the follow-

Acetic fermentation of alcohol. Ammoniacal fermentation of urine. Lactic, viscous, and butyric fermentations of sugar.
Putrefaction, or nitrification.

4.

Acetic fermentation.

The

transformation

of

wine into vinegar is a phenomenon long known and utilized. From a chemical point of view, this transformation is due to oxydation of the alcohol.

The following formula represents


C 2H 6

this reaction

+ O2

C 2 H*0 2 +

0.

called

The agent of this oxydation is a micro-organism Mycoderma aceti. It belongs to the group
and we have already given
it

of the microbacteria,

the botanical description of

(page 83)

but

its

development presents some interesting peculiarities which we think it proper to indicate in the language of M. Duclaux " These little beings reproduce themselves with such rapidity that by placing an imperceptible germ
:

upon the surface

of a liquid contained in a vat

having a surface of one square metre, we may see it covered, in from twenty-four to forty-eight hours, with a uniform velvety veil. If we suppose

140

PHYSIOLOGY OF THE BACTERIA.

that there are three thousand cells in a square mil-

below the truth, this will give for the vat three hundred milliards of cells produced in a very short time." " The Mycodermi aceti is not always the same. Usually it forms upon the surface of a liquid a soft-looking veil, smooth at first, then wrinkled, which is with difficulty submerged and moistened. If a glass rod is plunged into the liquid, it pierces this veil and when it is withdrawn, a portion remains attached to the rod ; and the opening made immediately disappears, being occupied by the veil which seems never to have room enough in which to extend itself. In some unpublished experiments I have frequently observed another form of veil, dryer, finer, and sometimes showing prismatic colors. This veil does not wrinkle, but is covered with crossed undulations, having sharp edges, which recall the surface of a honeycomb. Sowed
limetre,

which

is

upon the

surface of various liquids,

it

reproduces

and it is difficult not to consider it a different form of the preceding. Finally, I have also met a species of mycoderma producing well-developed veils, but having scarcely any acetifying power, and reproducing itself with this
itself identically,

character."
to distinguish these forms the one from the other, by the microscope, because of their minuteness. We may, however, say that the second which I have described is sensibly smaller than the first, and the third more attenuated than either of the others. However, the differences are feeble." This veil is called the mother of vinegar. The
is difficult

" It

THE BACTERIA IN DIFFERENT MEDIA.


liquid in

141

which

this

mycoderma is

cultivated should

be a
the

little

acid, containing

one-half per cent of


conditions

acetic acid, for example.

Under these

Mycoderma

vini (a species of Saccharomycete),

the formation of which should be avoided, finds


conditions unfavorable to
this
its

existence.

Indeed,

second organism, commonly called flowers of wine, has an action quite different from that of

consumes the alcohol and carbonic We must acid it also consumes the acetic acid. sow the M. aceti, if we do not wish to see the M. vini develop in its place, as the germs of the latter seem more widely diffused in the air. In order that the acetification may occur, the oxygen of the air is necessary. Once submerged, the M. aceti develops, but no longer produces It is even probable that it consumes acetic acid.
the
aceti.

Mycoderma
:

It

entirely, transforming it into water

the acetic acid already formed, reducing


state of

it

to the

water and carbonic acid. It is the same when, developing upon the surface, it has transformed all the alcohol. " In effect, it is not then arrested in its work ; and without changing form
or

mode
it

of action,

it

carries the
it

oxygen

of the air

to the acetic acid which

has produced, transformIf we add some phenomena change the


:

ing

into carbonic acid

and water.

alcohol to the liquid, the

'acid is respected, and the alcohol anew into acetic acid " (Duclaux).

is

transformed

According to
of aceti-

the experiments of Mayer, the

maximum

fying power
to 86 Fah.),

is

obtained between 20 and 30 (68

and this power is lost below 10 (50 Fah.) and above 35 (95 Fah.).

142

PHYSIOLOGY OF THE BACTERIA.


Urine.

Ammoniacal Fermentation of
urine
is

When
:

freely exposed to the air,


it

we

perceive at

the end of a short time that

ammoniacal.
ate of

The urea is ammonia by the addition

has become strongly transformed into carbonof

water

CO(NH
Miiller urine, of

+H

i=

CO 2 + 2NH 3

suspected that the

deposit of altered

which Jacquemart had already recognized the particular activity, was an organized ferment, but this was only an induction drawn from the analogy with beer yeast. Pasteur showed that this sediment is formed of a mass of spherical globules, united in chaplets, which he considers the agent of ammoniacal fermentation. These globules are Micrococcus urece, Cohn, which we have
already described (page 75). This bacterium lives in the interior of the liquid,

and not on the surface like the Mycoderma aceti. alkalinis an obstacle to its development ity, on the contrary, favors it within certain limits. Van Tieghem has even seen the fermentation conAcidity
;

tinue until the liquid contained thirteen per cent

ammonia. mechanism of this fermentation ? M. Musculus has shown that we may obtain from altered urine a soluble ferment upon adding
of carbonate of

What

is

the

to

it

highly-concentrated alcohol

a precipitate
dried.

is

formed, which
into

may

be

filtered

and

This

precipitate, not at all organized, transforms urea

carbonate of ammonia.
it.

temperature of

80 (176 Pah.) destroys

This diastase appears,

THE BACTERIA

IN

DIFFERENT MEDIA.

143

then, to be a secretion of the Micrococcus urece;

and perhaps the role of the bacteria is limited, in the phenomena of fermentation, to the formation of this secretion alone. The ammoniacal transformawould consequently enter into the group of fermentations by the varieties of diastase. According to Arnold Hiller, if carbolic acid be added to urine, it does not become alkaline on the contrary, the acidity is even augmented, and
tion of urine
;

that notwithstanding

a considerable
it.

bacteria which develop in


free

number of Has the carbolic

acid killed the Micrococcus urece, leaving the field

an producing other transformations of the constituents of the urine ? In the memoir which we here cite, the author, resuscitating the ancient opinion of Liebig, wishes to demonstrate that the decomposition of dead organic matters, and putrefaction in general, are phenomena purely chemical, these decompositions being determined by the presence of organic substances, themselves undergoing transformations. We will not stop to consider these views, long since refuted the experiments upon which they
to other organisms capable of living in
of

acid

medium, and

are founded are easily criticised.


for

It is

sufficient

me

to say that they are in formal opposition

with

all

the

observations

contained in modern

works upon
It is

this question.

ammoniacal fermentation that the question of spontaneous generation has been discussed. We have already seen the results arrived at, and will not return to
especially in relation to

144
this

PHYSIOLOGY OF THE BACTERIA.


subject.

Let us, however, mention before an interesting work by MM. Cazeneuve and Livon, in which are reported some experiments which prove that urine never ferments in a healthy
closing

bladder.
Lactic, Butyric,

Sugars.

Saccharine

and Viscous Fermentations of


liquids, left

to

themselves,

are susceptible of divers fermentations, which

may

occur separately or simultaneously.


butyric,

have been best studied are three, the lactic, the and the viscous fermentations. We will
describe
1.

Those which

them

successively.

Lactic Fermentation.

Under

the probable

influence of a bacterium {ferment lactique of Pas-

teur) glucose

and the substances susceptible


such as mannite, malic acid,
etc.,

of

furnishing

it,

are

transformed into lactic acid. From a chemical point of view, there is in this nothing more than a molecular change, lactic acid

having the same composition as glucose. Taken in mass, the lactic ferment resembles beer-yeast ; its consistence is, however, a little more viscous, and its color more gray. But under the microscope, the aspect is very different, as we have seen in describing Bacterium lineola. An interesting point concerning this fermentation is the action of acids upon the bacteria which produce it (presumably). As soon as the medium becomes acid, even by the lactic acid produced, the transformation is arrested. It resumes its course, if chalk or carbonate of soda is added to the
liquid.

THE BACTERIA IN DIFFERENT MEDIA.

145

The most
(95 Pah.).

suitable temperature

seems to be 35

little about this fermentation. however, to be better studied. It is this which causes the spontaneous coagulation of milk: sugar of milk is transformed into lactic acid, which coagulates the caseine. We often see it occur in beef juice or in sour starch water; it must play a part in the formation of sour krout, and intervenes very certainly, and perhaps more than

We

know but

" It

merits,

the alcoholic fermentation, in the preparation of


bread.
of
its

Finally, it very easily invades beer, which our domestic drinks is most exposed, because of
slight acidity, to

become the
it is

seat of this ferit

mentation.
ing, so

All of these facts render

interest-

much

the more as
is

rarely

exempt from

complication, and

frequently accompanied, for

tion, far

example, by a commencement of butyric fermentamore disagreeable in its products " (Du2.

claux).

This is, in fact, alButyric Fermentation. ways preceded by a lactic transformation, and it is

by an

ulterior modification that the

lactic

acid

produces the butyric acid. The organism which accompanies it is a bacterium very nearly allied
to Bacillus subtilis,

Cohn.

The

reaction

represented

by the phenomena,
is

from a chemical point of view,

the

follow-

ing:
2C 3H6
lactic ac.
3

C4H8

2C02

+H

4
.

butyric ac.

146

PHYSIOLOGY OF THE BACTERIA.

According to Pasteur the butyric ferment belongs to his class of anaerobies.

great

This fermentation resembles putrefaction in a many particulars. Indeed some authors init

clude
3.

under the same head.

change so that they contain a mucilaginous substance and mannite. This viscous matter has the same com10 position as gum or dextrine (C 6 O5 ) at the
Viscous Fermentation.

Wines often
H
;

same time

it

disengages carbonic acid.

In the fermenting liquid, we find an organism which is not yet sufficiently studied. " There are chaplets of little spherical bodies, of which the diameter varies sensibly, according to the kind of wine attacked by this malady (Pasteur).
Pasteur has proposed the following formula
:

25(C 12 H 22 O n )

+ 25H

12(C 12 H20 O 10)


gum.
2
2

24(C 6 H"0 6 )
mannite.

+ 12C0 + 12H 0.

which represents the phenomena well enough as it usually occurs. There is produced in the vicinity of 51.09 of mannite and 45.5 of gum for one hundred parts of sugar. But sometimes the gum exIn this case, ceeds the mannite in quantity.
according to Pasteur,
ent nature
results

we can always

verify in the
differ-

liquid the presence of a larger


;

ferment of a

and the same author adds


of this

that, per-

haps, in this case the increased production of

gum

from the presence

second ferment,

which transforms the sugar only into gum, without

THE BACTERIA

IN DIFFERENT MEDIA.

147

any

correlative formation of mannite.


isolated.

ferment has never been


in another

But this M. Monoyer has

explained the variation in the proportion of

gum

manner

(see his thesis for the doctorate

in medicine, Strasburg, 1862).

White wines are more subject than red wines


this fermentation, called graisse des vins.

to

Accord-

ing to M. Francois, the absence of tannin in the

white wines is the cause of this disease, and it may be prevented by adding this substance. This remedy is even very highly appreciated in champagne, according to Pasteur. What is the exact action of the tannin upon the gummy ferment? The only means of knowing is by cultivating this ferment in a state of purity and treating it with
this agent.

We
water,

have united together the


all

viscous ferments, because


selves in the
etc.
;

beer, sweetened and because they have for effect the We ought to say a transformation of glucose.
liquids,

same

wines,

and three manifest themlactic, butyric,

word here

of

some other

inferior organisms, per-

haps bacteria, observed also in the same liquids, Not but which have not been as well studied. but we only are they not known systematically,

do not know precisely what is their chemical action upon the elements of the medium which nourishes them. I shall only enumerate them. "These 1. Ferment of Turned Beer (Pasteur).

are rods or filaments, simple or articulated into chains of variable length, of about 1 /i, diameter.

148

PHYSIOLOGY OF THE BACTERIA.

A high

power shows them divided

into a series of

shorter rods, scarcely born, not yet mobile at the

which are scarcely indicated." 2. Micrococcus of a beer, having a particular acidity, distinct from that of beer pique, having an acetic odor. " It consists of grains resembling
articulations,
little

spherical points jointed square " (Pasteur), etc.

by

pairs or in fours

2.

Role

of the Bacteria in Putrefaction

and Nitrification.
which we have just passed rapidly in review, we have always been

While

in the fermentations

able to study, at least summarily, the chemical


action of

the

different

organisms,

we

are

now

about to find ourselves in presence of phenomena


far

great
its

more complex. We will have to consider a number of these vegetables at work, without

being possible to assign to each its role, or to say what is its function. The agent of the nitric
fermentation has not as yet even been seen, and
is
it

only by analogy that


tlje
is

we

class this nitrification

with
It
still

true fermentations.

not only because of the obscurity which exists in regard to a great number of peculiarthese two phenomena, that

ities of

we have

united

them
of

in the

of the circulation

same study. Prom the point of view upon the surface of our globe
constitution of

the elements essential to the

organisms, they play an analogous rSle, although opposite the one to the other.

THE BACTERIA

IN DIFFERENT MEDIA.

149

Let us consider, for example, nitrogen in plants. This element, of which the atmosphere is the reservoir, does not enter directly into combination, as
does oxygen, with the other elements which with
it

are to constitute the immediate principles of the

The chemical properties of nitrogen may be characterized in two words, great resistance to entering into combination when it is free, and great facility, on the contrary, in passing from one combination to another when once it has associated itself with other elements. The circulation of nitrogen in a state of combination upon the surface of the globe is also an
tissues.

interesting question of general physics, as well as

the circulation of carbonic acid, of water, and of


the
air.

Let us seek to sketch the march of


culation.

this cir-

Whence comes
torial regions, in

the

ammonia which
?

is

found in

the sea, in the clouds which come to us from equathe dust of the air

The only
It is

known

source

is

the fermentation of organic mat-

ters out of reach of the

oxygen

of the air.

to this sort of fermentation that

we owe

the for-

mation
all

and the immense masses of combustible minerals which have formed during nearly
of peat

the geological periods.


itself
air,

We

see this sort of feror-

mentation develop
ganic liquid to the

when we expose an
is

but only in the inferior


dissolved

part of the liquid, the oxygen which

near the surface being arrested in the superficial zone, where a very different fermentation occurs.

150

PHYSIOLOGY OF THE BACTEEIA.


latter is essentially oxidizing
;

The

the material

is

almost completely burnt, forming water and carbonic acid at the inferior part, on the contrary, a
;

reduction
is

is

produced so energetic that hydrogen

disengaged.

The

metallic sulphates are there

transformed into sulphites, and even crystals of sulphur are sometimes found (see the history of the Beggiatoa, page 91). We see then the source of the ammonia, which, distributed upon the soil by the winds and the rains, becomes a powerful fertilizer. Now, vegetables do not absorb nitrogen under the form of ammonia, but

under the form


ation of

of nitric acid.

How is this transform-

ammonia into nitric acid effected ? The Erdmann, Mensel, and T. Phipson show that in the phenomena of destructive putreobservations of
faction, nitric acid, far

from being produced,

is

on

the contrary reduced to the state of nitrous acid

on the other hand, Th. Schlcesing and A. Miintz conclude from their experiments that in the putrefactions essentially oxidizing produced by Penicillum glaucum, Aspergillus niger, Mucor mucedo, etc., there is no formation of nitric acid. But,
according to these authors, nitrification
cial
is

a spe-

phenomenon which takes place

in every soil

permit a free circulation of air, and of which the agent is a micro-organism. This organism has not yet been perceived, it is true;
sufficiently loose to

and it is evident that it would be difficult to seek and observe, because of its peculiar situation.

But the action

of chloroform

upon

nitrification
is

tends to prove that the agent of this process


THE .BACTEKIA
IN

DIFFERENT MEDIA.

151

truly an organized ferment.


this

Indeed, chloroform,

anaesthetic, suspends nitrification,


kill

and seems

even to

the ferment.

then, this phenomenon, but little known, we may distinguish in the agents of putrefaction, or more generally of fermentation, two

Leaving,

groups of micro-organisms,
other reducing.

one

oxidizing,

the

observed upon the surface of liquids undergoing putrefaction. "We may distinguish a great number of forms, Bacterium termo,
first

The

are

Monas
also to

crepusculum, Spirillum,
include

etc.

We

ought

Mycoderma

aceti,

which, like the

others, vegetates on

the surface

a great

number

of organisms of

of liquids, and which we cannot

speak here. The second are met, on the contrary, in the interior of liquids or of fermentable bodies they are analogous to the butyric and lactic ferments, and perhaps to the other agents of diseases of wine and beer previously enumerated.
;

En

resume, the

little

beings which

we have been
:

considering have an important role

they cause

the return of dead organic matter to the atmos-

phere and to water.


" Without them, organic matter, even exposed

would not be destroyed or would be transformed with extreme slowness, in consequence


to the air,
of a slow

combustion produced by oxygen.

With

them, on the contrary, its destruction takes a rapid march and becomes complete. If, then, the
equilibrium
is

maintained between living nature

152

PHYSIOLOGY OF THE BACTERIA.


if

and dead nature,


composition,
tilizing, it is

the air has always the same

if the waters are always equally ferthanks to the infinitely minute agents of fermentation and putrefaction" (Duclaux). But the rftle of bacteria is not limited to this. " They invade also the living organism," says Duclaux, " and bring in their attack this double char-

acter of infinite smallness in the apparent

means

and powerful destructive energy in the results. From this source come diseases of which medicine, not long since, did not know the cause, and which she only commences to refer to their veritable origin. For those who are au courant with the first steps which she has made in this new line of
research, with the fecundity of her
first

glimpses,

with the richness of her first results, it is not doubtful that she will soon succeed in demonstrating the parasitic nature of the gravest epidemic
maladies."

3.

bole
in

of the bacteria in contagious Maladies and Virulent Affections.


shall first pass in

We

review the different

af-

fections

which the presence of bacteria has been indicated, whether they have been given as the cause of the malady or considered as simple epiphenomena.
Septicemia.

According

to

the

hypothesis of

upon the nature of septic blood, Sedillot demonstrated by some very conclusive experiments that the infective power is due to formed elements (des eUments figures).
Borsieri and of Gaspard

Plate IX.

$<&>
r-
\r*,

^>r"

-V

i
?fM
/ ~^

iv

vr-'

iU

W3*

s ;-

<

PLATE

IX.

Copied from photographs by Koch in Cohn's " Beitrdge zur Biohgie der Pjlanzen," Bd. II., Heft 3.

X500. Spirochete from human mouth, resembling the SpirochFig. X 500. ete Obermeieri of relapsing Fig. with spores from rotten onion. X 500. Fig. with spores from surface of rotten potato.
Fig

1. Zooglosa ramigera from fluid containing rotting algae. 200 by Seibert's immersion objective No. 7. Spirillum undula. Fig. 2. X 500. Bacillus (subtilisf), from hay infusion, showing flagella. Fig. 3.
1

4.

fever.

5.

Bacilli,
Bacilli,

6.

X
1

500.

A higher power (500) shows that these branches are made up of oval bacteria

154

PHYSIOLOGY OF THE BACTERIA.


first

experiments of Davaine brought him " The effects of putrefying substances do not go beyond the animal

The

to the following conclusions

into

toxic

which these substances are injected. The agent of putrid matters does not regenitself.

erate

Putrefaction acts

upon the animal


opinion should

economy
have for

as a poison."
it

The

first

the authority of Robin.

Already in

1864, Leplat and Jaillart, after a series of inoculations made with septic blood, arrived at deductions analogous to those
of

Robin.

At

the

same epoch,

Billroth

and Weber, having injected

the gases of putrefaction, expressed the opinion


that the septic agent was of a molecular nature
(particulate).

Bergmann,
alcohol

of Dorpat, admitted as

contagious agent an azotized substance, not organ-

and ether at a temperature of 100, and passing through filters. This theory was identical with that of Panum. It is to Pasteur that the honor belongs of having first
ized, resisting

affirmed the parasitic nature of septicemia.

This

communication was followed by confirmatory ex-' periments by Coze and Feltz. These experimenters also proved that the bacteria of putrid blood do not possess the property of traversing the epithelium, and that " the infectious element gains in passing through similar organisms." In 1868, changing his first Davaine, opinion, admitted the presence of bacteria in the blood of animals which Hallier of Jena and Be'champ die of septicemia.
micro-organism, Micrococcus
of Montpelier also believe in the

presence of a

for Hallier, Micro-

zyma

for Be'champ.

THE BACTERIA

IN CONTAGIOUS MALADIES.

155

Coze and Feltz, in their work, have demonstrated the constant presence of bacteria in the

blood of animals dead from septicemia.


relation has driven
is

This cor-

them

to

admit that " there


the infections acci-

a direct relation between


little

dents and the

foreign organisms which play,


researches confirmatory of the

in the blood, the role of ferments, and reproduce

themselves."
first

New

have been communicated to the Academy by the same authors. Opposed to these conclusions upon the bacterial origin of septicemia, some experiments have recently appeared in England, then in Berlin, which weaken them. Zuelzer, struck by the analogy which exists between septicemia and
intoxication
testinal

by atropine

(dilatation of pupils, inof

paralysis, acceleration

heart's action),

has sought for the presence of an alkaloid by the

method
teria

of

Stas.

In collaboration with Sonnenit.

schein, he has succeeded in discovering

Bac-

and introduced in considerable quantity into the mouth, under the skin, and into the vessels of various animals, have never seemed to him to produce septic accidents. But the scene changes as soon as an addition is made to the injected matters of two to five centigr.
cultivated
artificially,

of neutral sulphate of atropia.

The period

of in-

cubation always lasted from nine to twelve days.

The same

year, Riemschneider deduced from his

observations the same result, and confirmed thus

the similarity between atropine and sepsine.

The

bacterial origin of septicemia received, however, a

new

support from the experiments of Vulpian.

156

PHYSIOLOGY OF THE BACTEEIA.

Having injected into a rabbit two drops of blood from a man who died of pulmonary gangrene, the Nuanimal died at the end of twenty hours. merous bacteria were found in its vessels. The
liquid extract of these, injected into other rabbits

produced death in

twenty-four hours,

and the

were observed in their blood. Henrot the bacteria penetrate by the pulmonary mucus membrane, and only act in contact with blood rendered " phlogogene " by pus. There is then a necessity for two producing

same

parasites

According

to

In support of his opinion the author cites experiment. He injects into the jugular of two rabbits a mixture of distilled water and pus, for the purpose of rendering the blood "phlogogene;" in two others he injects pulverized
causes.

the

following

coral suspended in water.

One

of the first rabbits

and one

of the second, placed in a

pure

air, resist

perfectly.

The two others are subjected

to

ema-

nations from putrefying " anatomical " fragments.

The
of a

rabbit into which pus

in three days, the second

had been injected died was still living at the end

month. Cavafy admits the presence of bacteria in septic liquids, but does not regard them as the efficient cause of the thromboses following inoculations. According to Moritz Traube and Gschleiden, living organisms into which one injects

blood containing a great quantity of the

bacteria of putrefaction, do not suffer able


1

any dur-

harm from the


and

injections.

At

the end of

I have injected various liquids containing bacteria into the circulainto the cellular tissue of rabbits,

tion of dogs

any

serious results follow such injections.

G. M.

but have never seen


S.

THE BACTERIA

IS CONTAGIOUS MALADIES.

157

twenty-four hours, the arterial blood drawn from a rabbit into which one and one half centimeters
.

of

liquid containing

bacteria had been injected,

can be preserved (by protecting it from exterior germs) during several months, without presenting traces of putrefaction. The bacteria then are dead in the living organism. Nevertheless, the living blood is powerless to resist beyond a cerThese results seem difficult to rectain point. oncile with those which Feltz has reached as the result of new researches with the toxic principle Compressed air passed through a of putrid blood. septic liquid has no influence upon its toxic properties or upon the minute beings contained in
it:

there

is

the vibrios.

simply a diminished movement of In a vacuum the toxic power is


the Cocco-bacteria and bacteria be-

diminished
lose

vibrios and spiral bacteria but the smallest forms are not killed. W. Moxon and J. F. Goodhart have recognized the presence of bacteria in the blood

come motionless, the


their
activity,

and

in

the

inflamed

tissues

of

septicemic pa-

According to Virchow, also, the active agent is a bacterium which, injected, with or without putrid liquid, produces death by septic
tients.

intoxication.

Livon and Zuelzer have never observed any

symptom

of putrid infection to follow injections

of these micro-organisms into the blood.

In presence of so many divergent opinions, each supported by scientific authorities, we do not feel justified in adopting any definite conclusion.

158

PHYSIOLOGY OF THE BACTERIA.


of Coze and Feltz, however, as which confirm them, lead us to

The experiments
well
as

those

consider the constant presence of bacteria in putrid

blood as a great probability in favor of the


is

parasitic genesis of septicemia.

There
just

a variety of septicemia which presents

we have spoken namely, puerperal septicemia. The preceding researches, with their consequences, are all applicable to this form of septicemia, and exthe closest resemblance to that of which
;

plain

to

us

its

nature.

This manner of seeing

seems to us justified by the labors of Orth of Bonn, according to which the lymph and the blood contain Micrococci in considerable numbers. Klebs has verified the presence of the same
parasites
in

the

putrid

infection

consecutive

to

Like the preceding authors, in the liquids of septicemia the presence of Micrococci, and does not admit any other parasites.
Birsch-Hirschfeld
recognizes

gun-shot wounds.

Charbon.
charbon.

A malady

inferior organisms has

in which the influence been especially sought

of
is

We

will

examine successively the

results

fur-

nished by experimental pathology and by clinical


observation, and will finish
of the nature of charbon.

by

a general discussion

Although
scribed

this

affection

has been

known and
de-

studied from the highest antiquity, and was

by Chabert (1782),

Gilbert (1795), and

many

others, its parasitic nature has not long

been

THE BACTERIA

IN CONTAGIOUS MALADIES.

159

known.
ell,

Let us note, however, that Fuchs, Brau-

Pollender, and Delafond, had remarked


Prof. Delafond

some

corpuscles in the blood of animals attacked with

made, twenty years ago, some researches, which he communicated to the


charbon.

Central Society of Veterinary Medicine, upon the


rods of sang du rate.

Davaine,

who had

observed,

with Eoger, the presence of rods in the blood of charbon as early as 1850, did not attach any importance to the fact. After the work of Pasteur

resumed his researches and the results which he obtained were communicated to the Academy of Sciences the 27th of July and the 10th of August, 1863, then the 22d of August, 1864. His
in 1861, he

experiments established the fact that the blood of animals attacked with charbon contains organisms (elements figures), and that, injected into a healthy animal, it kills it by reproducing the same symptoms. There remained a step to make, to prove
that

the

bacteria

alone

possessed
cases.

the

infective

power, even in epidemic


discoveries,

Notwithstandto find

ing the labors of Signol (1864) corroborating his

Davaine did not

fail

oppo-

nents.
sults

Leplat and Jaillard made


of

known
to

the re-

their

experiments, according

which

the bacteria were not the cause of sang de rate.

In 1867, Bouley and Sanson, and in 1870, Bailstudied the nature of the malady known under the name of mal de montagne. Klebs, in Switzerland, having, with Tiegel of the Pathological Institute of Berne, made some
let,

negative injections with filtered blood

(of

char-

160

PHYSIOLOGY OF THE BACTERIA.

bon) demonstrated thus that the disease was truly due to the solid particles ; but he could not, as he did, affirm that the bacteria alone were endowed with virulent power, for he included at the same time all the other solid elements (fibrine, globules), and could not therefore eliminate the granulations Klebs does not of a virus other than the bacteria. believe that the bacteria cause death by asphyxia. This view is also sustained by Recklinghausen and

Waldeyer, who believe that death results from embolism according to Burdon-Sanderson, on the
:

contrary, this

is

not the case.

The

observations and experiments up to this


disease.

time demonstrated that the blood of charbon would


transmit the
the

Davaine had said that the

bacteria constituted the condition sine qua non of

development of these diseases, but he had him the experiments of Leplat and Jaillard. Besides, as he injected at the same time other corpuscles figurees, it was difficult to prove that they went for nothing in the production of
against
Finally, this theory could not explain endemics (pastures in Auvergne). One could truly say, with Burden-Sanderson, that the virulent element can exist in two forms, one fugitive (bacteria), one permanent, unknown. The point was to demonstrate it. This is what Koch has done. Having taken some bacteria, he cultivated them in urine, or the aqueous humor of the eye of a horse, and remarked that they underwent a certain elongation, then presented brilliant points of condensation which became free; injected

charbon.
certain

THE BACTERIA

IN

CONTAGIOUS MALADIES.

161

into the blood of sheep

and of

rabbits, these cor-

puscles produced

death with the

symptoms

of

charbon, and the blood of the animals presented

numerous

bacteria.

The appearance

of spores in

the liquid under cultivation, containing bacteria, occurs in twenty-four hours at 35 (95 Fah.), in three days at 18 above 45 and below 12 it is
;

no longer

possible.

Once produced these spores

and alternation of humidity and dryness, during several years. On the contrary, the adult form of the Bacillus anthracis dies under the influence of putrefaction and of oscillations of temperature. It has not seemed to develop itself in the dog, the cat, birds, and cold-blooded animals. The immunity which these enjoy has recently been the object of a study by Pasteur, Joubert, and Chamberlain. Believing that it might be attributed to their temperature, incompatible with the life of the bacteria, they have refrigerated a fowl, and have ascertained that Besides, by placing the it lost this immunity. infected animal in an oven at 30 (86 Fah.) they have seen the temperature come back rapidly to the normal and the symptoms of charbon to reresist putrefaction, desiccation,

cede.

The

labors,

then, of

Koch add an

additional

element of probability in favor of the parasitic They show us the existence of an organtheory. ism which we would be able to invoke as the cause
of spontaneous epizootics.

All these results

Cohn
11

of Breslau has obtained

upon repeating the experiments of his compatriot.

162

PHYSIOLOGY OF THE BACTERIA.

In France, Toussaint commenced, March 21, 3875, a series of successful inoculations upon rabbits, with blood obtained from the spleen and an abdominal tumor of a sheep which had died of the
rate. These specimens had been sent to Chauveau by Joly, veterinary surgeon at Gien. Having preserved some blood in the air, Toussaint remarked, as Davaine had done and as Koch had observed, that putrefaction kills the Bacillus enclosed in a close vessel, it succumbs as soon as oxygen is wanting, which occurs sooner when the

mal du

temperature
It

is

elevated.
of

was upon the presentation

these results

to the

Academy

of Sciences that
is

Cohn expressed

the opinion that charbon

but not

to a special virus.
act, it is

If the filtered
filter,,

not due to a bacterium, blood does


at the

because the

same time,

retains the, virus.

Pasteur, in his letter of Aug. 18, 1877, replies

that a virus would be impotent to resist the numer-

ous cultivations endured by the liquids in his experiments, and that the bacteria alone remaining,
it

tious

was very logical to attribute to them the infecpower possessed by the liquid of the last
Paul Bert had at
first

cultivation.

believed, with Cohn, in

the existence of a virulent agent other than the

Indeed, after having treated blood of charbon with compressed air and alcohol, which kill bacteria, he had been able to transmit charbon. But, abandoning this first idea, he expressed himbacteria.
self as of

the same opinion as Pasteur and Joubert

THE BACTERIA

IN

CONTAGIOUS MALADIES.

163

(July 30 of the same year), recognizing that the


persistence of the virulence
is

due to spores (corthe causes of de-

puscles germes) which resist


struction.

all

Quite recently, finally, Toussaint, while studying


the mechanism by which the bacteria cause the

death of rabbits, horses, and sheep, arrives at the conclusion that it is because of asphyxia of mechanical origin,
pillaries.

embolism

of the

pulmonary
is

ca-

The phlogogene

action of the vibrionien

some-

is a rupture of the capillaries, and even lesions graver


still.

times such that, in addition to embolism, there

The phlogogene material is more active, according to the subjects from which the bacteria are
obtained.

"

The animals which


:

I have studied

may

be arranged as follows the rabbit, guinea-pig, sheep, ass, horse, dog." As to the hog, it is not
at all susceptible.

In the

last place,

Toussaint has presented, through

Bouley, a note upon a form of charbon caused by a Vibrion aerobie. This affection was already recognized as contagious, but the agent of contagion was not known.
is

Toussaint has found that

it

caused by a Bacterium, differing in certain char-

from Bacillus ; he has cultivated this microbe, and has seen it reproduce itself under the microscope, in an apparatus invented by Ranvier. The malady has been transmitted to rabbits in the same burrow without inoculation.
acters

By

taking the excrement, reduced to powder, of

164

PHYSIOLOGY OF THE BACTERIA.

an infected animal, and sowing it upon the food given to an animal in good health, the latter has
contracted the disease.

Besides Malignant Pustule without Bacteria. the numerous facts concerning charbon, in which the presence of a bacterium has been verified, it is proper to cite those cases in which none has been discovered. Some authorities, such as Toussaint, Mannoury, and Salmon, who have given
these instances, consider this absence of bacteria

a favorable prognostic sign.

The following
:

marked example has recently been observed Louis Donin, a tanner, aged forty, was admitted to the Hotel-Dieu of Lyons, June 15th, 1876, service of Fochier. On the morning of June 13, he had noticed three large flies, eagerly attacking the skins upon which he was working. One of bit him in them the face. The same day his cheek
during the night a large vesicle formed, surrounded by an areola of other smaller vesicles; the skin having become pruriginous, Donin rubbed
swelled
;

well-

the central vesicle.

Upon

his arrival his general

condition was satisfactory.

Upon

his

left

cheek

was seen an areola


meter.

of little vesicles surrounding

a slough having a diameter of less than a centiThe periphery was oedematous and hard,

trembling and extended downwards as far as the xiphoid appendix. The eyelids and the oedematous lips were opened with difficulty.

In order to

verify the diagnosis, Toussaint inoculated a rabbit

with the debris of a pustule; a second, with blood;

THE BACTERIA

IN

CONTAGIOUS MALADIES.

165

and a

third,

with serum.

The

result of these three

inoculations

was completely negative. The microscopic examination by Charpy and Colrat did not reveal the presence of any vibrionien. On the
16th of July, the patient
left the

hospital cured.

Let us add that the interne of the service, having punctured his finger with a syringe employed in making injections of carbolic acid (twenty per
cent), did not experience

any

ill

effects.

Darreau, veterinarian at Courtalain, attributes to bad food this variety of charbon, in animals,

without the presence


scribed an epidemic

of

bacteria.

He

has de-

on a farm where the forage quality. bad Charbon is then due, acwas of cording to him, to impoverishment of the blood. Decroix, veterinarian in the army, has examined
with the microscope the blood of horses submitted to his observation, from the jugular vein, and also the tumors. He has never found any bacteria. The These experimental and horses have recovered. permitted Bouley to estabclinical results have lish the unity of the charbonneuse malady, contrary to the opinion of Prof. Bouillaud, who renews
the hypothesis of a multiplicity of charbonneuse
affections.

In effect we see the same bacterium everywhere producing the same disorders. In the very rare and generally favorable cases of charbon which do
not seem to be of bacterial origin, we may say with Pasteur, " When the parasite has not been perceived, it is probably because sufficiently high

powers have not been used.

The phlogogene

ac-

166

PHYSIOLOGY OF THE BACTERIA.

tion of the bacteria, brought to light


is

by Toussaint,

necessary in order to explain the production of

the tumors of charbon."

Davaine, in consideration of the immobility of the Bacillus anthracis, admits that every tumor
results

from a

local inoculation.

We

have called

attention to the fact that the virulent agents of


variola

and of rugeola, although motionless, proIn the duce nevertheless local manifestations. second place, the charbon of the horse is often accompanied by internal tumors of which the orFiigin evidently cannot be an external cause. nally we have pointed out by Bouley some horses of La Plata in which the local manifestations did not appear until after the general symptoms. This view had already been sustained by Chabert. However, notwithstanding all the proofs furnished turn about by experimental pathology and clinical study, one desideratum still remains. It is necessary to verify the presence of the spores of Bacillus anthracis in the lands where charbon prevails as an epizootic and to discover its means of
transportation.

we think it proper to examine the efficacy of antiseptic treatment. Carbolic acid, studied by Koch, has been employed with success boracic acid, which acts
Before abandoning this question,
;

upon the

bacteria

by depriving them

of oxygen,

has been utilized by Decroix, veterinarian of the

army
acid,
rise

by
to

tincture of iodine, employed, like carbolic

subcutaneous injections, has given grave accidents. Beside these general

THE BACTERIA

IN

CONTAGIOUS MALADIES.

167

proceedings based upon the idea of an infection of the entire organism, are placed the local treatment
destined
of entry
:

to

destroy the bacteria at their point

thesfe are, caustic potash, sublimate,

the

paste of Canqouin, the hot iron.

These two categories of means employed sometimes alone, sometimes simultaneously, show that the idea of the clinicians has always been the destruction of an
infectious organism.

Variola.

The

partisans

of

the

parasitic

na-

ture of variola
1.

may

be divided into two groups

Those who, with Coze

and

Peltz,

attribute

the virulence to a Bacterium; 2. Those who, with Luginbiihl and Weigert, attribute it to a Coze and Feltz have indeed disMicrococcus. covered Bacteria in the blood Of variola, and this

blood injected into the veins of a rabbit has given


it

a mortal malady, which these observers consider

variola.

But Chauveau has shown

that the af-

fection which proved fatal to the subjects of the

experiment was not and could not be variola. Another objection is that Bacteria are not found in all those who suffer from variola. However, Coze and Feltz and Baudouin affirm that there are in variolous blood numerous rods, of which the appearance is similar to that of Bacterium bacillus and Bacterium iermo of Miiller. These elements do not at all resemble those found in other infections, and when inoculated possess the power of
reproducing variola.

As

to

the Micrococcus of variola, they have

PLATE

X.

Copied from photographs by Koch in Cohn's " Beitrage zur Biologie der Pfianzen," Bd. II., Heft 3.
Fig.
1.

Bacilli of Miltzbrand from the substance of the spleen.


col-

X700.
Fig.
lected
2. Bacilli of Miltzbrand in blood of basilar artery two days after death. X 700.

Fig. 3. Fig. 4.

Spirillum Obermeieri.
Bacilli

X
x

700.

of Miltzbrand cultivated in aqueous

humor,

showing development of spores,

700.

1 The lithographer has not succeeded in making a satisfactory copy of Koch's photo-micrograph, in this figure.

THE BACTERIA

IN

CONTAGIOUS MALADIES.

169

been studied by Luginbiihl, Weigert, Hallier, These micro-organisms possess the and Cohn. characters of all the spherical bacteria, and are
found in the variolous pustules, the
rete

Malphiyii,

the liver, the spleen, the kidneys, and the lymphatic ganglia. "We can only insist upon the fact
of the concomitance of the variola

and the pres-

ence of Micrococcus, since experiment cannot be resorted to in this disease, of which the complete
evolution only occurs in man.

We

also in

find

in

vaccine

point of view, to those of variola.

lymph analogous Micrococci, Cohn


species,

every

considers

them

both, not as distinct species, but as two races

of the

same

the Micrococcus

vaccinae.

and Feltz have found in the blood of scarlet-fever, taken from patients, living or recently dead, some rods as well as moScarlatina.
bile points.

Coze

This blood injected into the cellular


death,

tissue

of

rabbits has sometimes produced

and the blood of the animals experimented upon


has presented the same bacteria as
of scarlatina
:

human

blood

they are simply a little larger and longer. As to the mobile points, they appear to correspond to the Micrococcus of scarlatina
described

by

Hallier.

Rugeola.
sles

The examination

of the blood of

mea-

has shown to the same experimenters, bac-

extreme minuteness and great mobility. The inoculation of this blood has not produced the death of rabbits ; however, these animals have
teria of

170

PHYSIOLOGY OF THE BACTERIA.


sick for

two or three days and have presented in their blood very slender and active rods. In the period of invasion the nasal mucus already
been
contains small bacteriform elements.

Diphtheria.
Tigri,

All

the labors undertaken since


CErtel, Letzerich,

by Trendelenburg, masi, and Hueter, from a


Micrococcus.

Tom-

parasitic point of view,

have attributed diphtheria to the presence of a There is however a single exception, Eberth of Zurich, in a work published in 1872, regards a Bacterium as the agent of the diphtheritic contagion.

According to new researches, which appeared in 1873, the pus of pyaemia, or of purulent peritonitis, inoculated, produces diphtheria because of the
presence in
it

of bacteria.

Laboulbene had already pointed out the presence of Bacterium, accompanied by Vibrios and Micrococcus in pseudo-membranous affections. Some
researches

made

in

collaboration with

Robin had

but these savants did not admit a relation of cause to effect between the micro-organisms observed and diphtheria. According to Duchamp, as shown by experiments
result,

given the same

cited in his inaugural thesis (1875), there are in


false

membranes

Bacteria,

Vibrios,

and

granu-

lations.

Taken alone, these micro-organisms appear to possess a very injurious action, but their inoculation does not produce diphtheria. The demonstration of a causal relation

between the Micrococcus

THE BACTEBIA and diphtheria


last
is

IN

CONTAGIOUS MALADIES.

171

not, then, yet established

by these

experiments.

found bacteria in the blood of a man dead with typhoid fever. These organisms were also found by Signol (1863) and Megnin (1866) in the blood of horses attacked by a disease called by the veterinarians

Typhoid Fever.

Tigri

first

typhoid fever. This blood, by inoculation, produced the death of some rabbits, with the same
alterations in the blood.

rabbits

Coze and Feltz (1866), having inoculated some with the blood of typhoid fever, have produced results which they consider analogous and as accompanied by the same pathological localizations in the glands of Peyer.

The blood of be used upon a second rabbit, with positive results, as in variola and scaran injected rabbit

may

latina.

The

species of

Bacterium which

is

case recalls the Bacterium catenula, but


sions are less.

found in this its dimen-

Glanders and Farcy.


tion of the contagious

The
power

universal recogniof a liquid

coming

from an animal with glanders had, a priori, led to the supposition of an element of special contagion. The first indication was given by ChrisThese experimenters tot and Kiener (1868). discovered in the secretions and vascular glands of animals attacked with glanders, bacteria of two sorts 1. Rods, sometimes having a vibratory motion without changing place, sometimes having
:

172

PHYSIOLOGY OF THE BACTERIA.

a rectilinear or eccentric motion of translation 2. Some spherical granules of variable diameter,

homogeneous, animated by a rapid gyratory movement and a movement of translation in various


directions.

The
in

latter are

certainly Micrococci.

an experiment designed to demonstrate that the organisms alone are active, took ten grammes of pus from a pulmonary abscess of a horse attacked with acute glanders the virulent elements were so numerous that the water became opalescent. This pus was washed four or five times in five hundred grammes of distilled water, was then collected and dried, and finally was inoculated, and the inoculated animal perished with glanders. Another, on the contrary, into which the filtered liquid was injected presented nothing abnormal. The particulate elements are, then,
Chauveau,
:

alone active.
tagion
is

But

in glanders, as in charbon, con-

not always demonstrated. There are which spontaneous origin appears incontestable (case of M. Boulay d'Avesnes, three cases
cases in
cited in the " Eecueil de Mtjdecine Ve'te'rinaire,"

June

15, 1877).

Finally, this opinion has recently

been supported

by Delamotte,

who

accords in this with Tabourin,

might perhaps see minute germ, playing, in regard to the bacterium of glanders, the same role as the germ of charbon does with regard to the Bacillus anihracis. This is an hypothesis which no researches have yet confirmed. As proof of the functional analogy which may
in these cases the action of a

Bonnaud, and Chenier.

We

THE BACTEEIA
exist

IN CONTAGIOUS MALADIES.
of

173
of

between the bacteria

'

glanders and

charbon,
the

we

recall

the fact that Bedoin, having

mixed some powdered borax (two grammes) with


blood of a glandered horse, has found the

two hours. These results correspond with those which Decroix, veterinarian of the army, has obtained by treating horses attacked with charbon with
bacteria completely motionless at the end of

boracic acid.

Ulcerative Endocarditis.

In

this affection,

it is

well

settled
all,

to-day that the


that the

cardiac walls

and,

above
masses.

the valves are covered with

parasitic

Some think
;

malady

is

due to the

introduction of these

parasites into the interior

on the contrary, like Hiller, deny that the bacteria bear any causal relation with the lesions of ulcerative endocarditis. Gerber and Birsch-Hirschfeld have recently made an observation which is a complete refutation of the ideas supported by Hiller. They have found at the autopsy some hemorrhagic foci disseminated in various organs, the greater number of which
of the tissues
others,

contained some particular corpuscles belonging to


the class of the bacteria.

Belapsing Fever.

In

1868,

Otto Obermeier

discovered in the blood of the sick attacked with

recurrent fever certain bacteria, called by

Cohn

Spirochete Obermeieri.

These organisms are only

found during the


cess of fever

febrile

paroxysm

after the ac-

they disappear, but often they are

174

PHYSIOLOGY OF THE BACTERIA.

again found a few hours before the new access. They are no longer found when convalescence is
established.
feld,

Heidenreich, Weigert, Birsch-Hirsch-

and Cohn have confirmed these observations.

Intermittent Fever.

In

all

the analyses of air

made
found.

in

the vicinity of lands

where

this

fever

prevails,

numerous
1

inferior organisms

have been

I refer to a previous

work
to

for additional

information,

and limit myself

pointing out

the observations of Griffini,

who

has found in the


etc.
2

dew

of places
lineola,

subject to this fever, Vibrio bacil-

lus, V.
1

Bacterium termo, B. catenula,

Ant. Magnin, Rech. Geol., Bot. et Stat, sur lTmpaludisme dans les
et le

Dombes,
2

Miasme

Palude"en.

Paris, 1876.

who have recently spent with the intention of investigating the cause of malarial fevers, have published an account of their researches. Eroni an abstract of their report, published in a recent
" Professors Klebs and Tommassi-Crudelli,
in the

some time

neighborhood of

Rome

number
task.
.
.

of the

'

tigators followed a very deliberate plan in the


.

Medical Times and Gazette,' we learn that the invesperformance of their

"Professors Klebs and Tommassi-Crudelli first succeeded in producing the symptoms of malarial poisoning in animals by injection of watery extracts from the marshy soil. They then proceeded, by the process called fractional cultivation,' to isolate the active material, that is, the true generator of the disease, supposed to be a living organism. Lastly,
'

they isolated the organisms by filtration and, comparing the results obtained in injections of the filtrate with those produced by the residue containing the organisms, they proved that the poison of malaria resides
;

The fungi obtained appeared as small rods of 0.002 to 0.007 millimeter in length, growing into long, twisted threads. The fungus is
in these.

markedly
rise to
etc.

aerobiotic.

these fungi

true

If air is excluded, it dies out.

bacilli

malaria.

into' healthy

The

injection of

animals always gives

symptoms

of intermittent fever, with enlargement of the spleen,

Rome, was able to demonstrate spores and bacilli in the spleen, the marrow, and the blood of three persons who had died of pernicious fever, showing the same characters as those observed by Klebs and Crudelli. In summarizing the results of their investigations, the authors consider the following facts as proved 1. That
Later, Dr. Marchiafava, at
:

THE BACTERIA

IN SURGICAL LESIONS.

175

4.

Of

the Role of the Bacteria in


Surgical Lesions.

Existence of Bacteria in

Since the day when theraby Surgical Lesions. peutics has entered upon a road truly scientific,
the study of the liquids secreted upon the surface
of divided tissues has occupied in

the

Liquids Secreted

an important place

the observations and researches of surgeons.

Little

by

little

the discussion of operative methods


societies,

has fallen to the second place, so that to-day, both


in the press
is

and in the

but

little

attention

given to the
;

mode

of

proceeding or to the form


is

of the flaps
all

but the greatest interest

taken in

questions touching the pathological physiology


solutions of
to

continuity. Thirty years ago it chemistry that we looked for an explanation of the complex phenomena which favor or
of

was

prevent the cicatrization of wounds to-day it is above all to the microscope or rather it is to that part of chemistry which is the most particularly indebted to the microscope for the progress which
:

it

has accomplished, that

is

to say, to the science

of ferments, to

zymotic chemistry.

To show by

what labors this tendency has been brought about, to what facts they have led, and what progress has been realized, is the object which we propose to
ourselves in this paragraph.
it is

possible to reproduce malarial infection in every


it is

which

known

in

men

2,

form That the malaria produced

in rabbits in
artificially in

is generated by organisms existing in the malarial soil at a time the outbreak of the fever has not yet taken place." Extract from Leading Article in " Philadelphia Medical Times," March 13, 1880.

animals

when

176

PHYSIOLOGY OF THE BACTERIA.

In Bacteria in Liquids Exposed to the Air. order to comprehend the importance of bacteria

from a surgical point of view, it is sufficient to put a drop of pus from an open wound under the microscope.

bacteria will be demonstrated.


is

In the majority of cases the presence of And, truly, there

nothing in this which should astonish us, if we remember what has been said above, since we know the facility with which these vegetables develop in all the liquids of the organism, and the

which they offer to all but the most However, all pus is not equally suitable for the development of these organisms. It is easy to remark that they are more abundant in pus of a bad character, in that which smells bad and exhales an odor of butyric acid. They are also more commonly found in pus which has remained a long time in wounds having hidden siresistance

powerful agents.

nuses.

But while recognizing these


confess that there
is

differences,

we must

nothing absolutely fixed about them, and above all that they do not bear a constant relation with the conditions which govern the genesis of putridity.
It is certain, also, that the conditions surrounding the sick person, the quality of the air in which his wound is bathed, are conditions which it is

necessary to

And this results from what we have said relative to atmospheric germs. For example, when we submit to maceration in distilled water, dust gathered in a hospital, on one hand, and on the other, dust from a different locality, taken in the country, for instance. The
consider.

THE BACTERIA IN SURGICAL LESIONS.


first

177
etc.,

will

much

sooner develop Micrococcus,


(L.

and in far greater numbers


it is

Julien).

Now

evident that this result

is

exactly applicable to

the subject which occupies us, since the pus of

wounds exposed

to the air can be compared, in

a certain measure, to the infusions of dust.

elements found in pus, they are Most frequently the cocco-bacthemselves present in the form of little chains teria

As

to

the

equally variable.

They rarely exhibit any moveThe appearance of bacteria upon the surface of wounds occurs at the end of sixteen to twenty-four hours. During the first hours following the division of the tissues, as is well known,
(strepto-bacteria).

ment.

the only exudation which appears

is

a yellowish-

pink serum. In this inflammatory exudation, which

always contains a considerable quantity of blood


globules, the parasites do not at once appear.
as I

But,

have

said, at

the end of a day or two Micrococci

and Bacteria in chains, very small, and of average size, make their appearance. It seems, then, as has been remarked by G. Nepveu, to whom we owe an excellent work on " The Role of Inferior

Organisms in Surgical Lesions," that


cretion of

this first se-

wounds

is

quite propitious for the de;

velopment
able to

of bacteria

and, perhaps,

we might be

draw from

this fact

of their formation

an argument in favor and their existence in the blood.

Later, with free suppuration, the inferior organ-

isms increase

still more. It is, however, to be remarked that they are never very abundant upon the surface of healthy wounds. If we gather this

12

178

PHYSIOLOGY OF THE BACTERIA.


it

open air for some time, we may follow the increasing development of these germs. An important remark, due to Dr. Molliere
pus, and preserve
in the
of Lyons, is

that the

phenomena

of putrefaction

are hastened by the presence of blood in the puruA quantity of pus drawn by aspiralent liquid. tion from a deep abscess, and not mixed with blood,

was exposed to the air for fifteen days, without any bad odor being developed.
Bacteria in Purulent Collections not Exposed
1875, Dr. Albert Bergeron communicated to the Academy of Sciences (Seance of
to

the Air.

In

Feb.

26)
in

the
the

results

of of

numerous observations
Prof.
if

made

service

Gosselin, for the

purpose of ascertaining
contains bacteria.
sions of his

the

pus of abscesses
are the conclu-

The following memoir


:

1. Vibrios are found in the pus of abscesses, without any contact with the external air and

without, usually, any indication that the organism


is

seriously infected by their presence ; 2. We cannot admit that in these cases the vibrios have pen-

etrated into the interior of the abscess through the

lymphatic system, or through the circulatory system, both being intact. The pus of warm abscesses
in adults often contains vibrios
;

if

the case of infants the fact has not


3.

they occur in been observed


vibrios

The pus

of cold abscesses in the adult, as in the


;

infant,

never contains them

4.

The

may

be considered as indicating a serious inflammatory state, and a certain tendency to decomposition of

THE BACTERIA
the

IN SURGICAL LESIONS.

179

humors which contain them, without, however, exercising any toxic action upon the organism 5. The author is far from rejecting the
as a rule,
;

possible intervention of vibrios in the pathology

and he explains the happy exemption of infants from septicemia, in a majority


of purulent infection
;

of cases,

by the

fact that these organisms are not

found in the pus of abscesses in young children.

The conclusions

of the

memoir

of

M. Bergeron

were the object of earnest discussions. According to the accepted theory, there ought never to be a development of organisms, unless the germs had been introduced from the air; if, then, we admit
the correctness of these observations, the explanation given

that

many

by Pasteur breaks down. Let us add, times, by the bedside of the patient,
has furnished results
absolutely
-

the microscope
contradictory.

Sometime before

M. Bouloumie

had formally established, as the result of long continued researches, that pus coming from any collection whatever, not in communication, directly or indirectly, with an open wound, never contained organized elements, mobile or motionless, which
can be considered as microzoa, or microphytes,
except

some highly-refractive moving points,

often

joined together in pairs.

We dare not say that the long discussions to which these communications have given rise have thrown any light upon the grave problems which they have attempted to resolve. Let it suffice for us to have pointed out these different points
of view.

180

PHYSIOLOGY OF THE BACTERIA.


the Effects

It would be rash of Bacteria. to attempt to-day to give a definite verdict upon the greater or less noxiousness of the bacteria.

Of

At

the outset of the studies which disclosed their presence in many pathological liquids, it was believed that we had finally found tbe true explanation of the obscure phenomena which retard or

complicate the normal repair of wounds.

But

what

shall

we

think to-day of the redoubtable role


first

which was at

attributed to a microbe,
its

when

we

ascertain that
clinical

by the
praised

development is not fettered means which are most highly

? These are the difficulties of the problem upon which the future will, without doubt, shed some light, but of which I repeat it notwithstanding the amount of labor which they call

forth every day,

it

is

ulate the solution to-day.

not possible for us to formLet us try, however, to

indicate

that which

experiment has taught us

up

to the present time.

Upon

the surface of a granulating wound, not

any vascular or lymphatic be believed that the microbe constitutes only a very contingent danger. It may
offering to absorption
orifice, it is to

multiply in greater or

less

degree, and will absorb


birth, perhaps, as

more or

less

oxygen, giving

result of chemical decompositions, to a virus

upon

the surface of the wound, which, notwithstanding,


will arrive at cicatrization.

But, if a solution of continuity happens to break through the thick

by the spontaneous movements of the patient or by those


layer of fleshy granulations, either

THE BACTERIA IN SURGICAL LESIONS.

181

which occur in dressing the wound (the pressure and manipulations which it was formerly customary to resort to), a rent being produced, the infectious agent will be able to penetrate into the blood current or into the lymphatics, and the door will thus be opened to local or general complications.

As a
all,

local complication,

we

should

cite,

above

Vabces de voisinage. If the presence of bacteria in the pus of spontaneous abscesses is still under discussion, all observers are agreed as to

secondary purulent collections. Bouloumie has taught us that in pus coming from abscesses developed in the parts in the vicinity of a wound, whatever may be its extent and depth,
their presence in

we may

verify the

presence of

all

the

micro-

organisms found in the pus of the wound, or cer-

developed with or contiguous in parts continuous to the tissues of the wound. There is then no doubt i^iat,
tain ones only, according as the abscess is
in this case, these
find

elements are absorbed, or at least


the veins or lymphatics.
of micro-organisms,

their

way through

The inflammatory action

thrown

thus into the midst of healthy tissues, even of those

which become inflamed with the most


easy to demonstrate.

difficulty, is

this

The following unpublished experiments upon point have been communicated to us by our
:

friend Dr. Louis Jullien

Experiment
surgical

1.

Having gathered some dust


it

in the

wards

at Lyons, I put

in a flask of distilled

water.

At

the end of some days I had thus given rise

182

PHYSIOLOGY OF THE BACTERIA.

to the development of numerous micro-organisms, the most common form being Bacterium. The 20th of May, 1873, with a Pravaz syringe, I

injected four drops of this liquid into the right ocular Immediately afterward, the iris globe of a rabbit.

bulged forward, became clouded, and the pupil dilated irregularly. As a result of this injection a terrible inflammation was developed. On the 25th I note an intense conjunctivitis and a roughness of the cornea, from which the epithelium has disappeared at certain
points.

Upon

the posterior surface of the cornea


in certain points
;

is

a white cloud occupying the inferior part of the pupillary opening,

more opaque

and present-

ing the appearance of a hypopyon the iris was still bulged forward, tomentous, of a violet red color ; pupil
contracted.

June
to

4,

the

iris is still
;

touch the cornea forward.

the crystalline lens

bulged forward, so as nearly is also pushed

June

5.

By

aspiration I

a sort of

membrane having

withdrew from the vitreous the form of a whitish fila-

ment. Upon examining this with the microscope, I recognized the presence of pus globules, cryptococcus,

and bacteria. June 10. The ocular inflammation has disappeared, but a cataract remains.

Exp. II. The same as the preceding. When the syringe was withdrawn from the vitreous, into which four drops were injected, a chemosis occurred, resulting no doubt from the fact that a
little fluid

was extrav-

asated into the sub-conjunctival tissue.

Operation made

May

25.

ocular inflammation, conjunctiva of a uniform red color cornea clouded, of a pearl gray tint iris very red, tumefied. The animal seems to suffer much when it is examined.
;

29. Intense

THE BACTERIA
30. Still

IN SURGICAL LESIONS.

183

much

inflammation, the ear

is

very warm,
white,

fever intense.

June

4.

capsular cataract

is

recognized,

milky, and glistening.


5. Aspiration by means of Pravaz syringe. A white membrane was obtained and examined under the microscope. It contained a great number of pus corpuscles,

a very great quantity of highly refractive yellowish


bodies, resembling cryptococcus,
bacteria.
10.

and a small number of

The inflammation
III.

is still

considerable.

Exp.
injection

As a control

experiment I made an in-

jection of distilled water into the eye of a rabbit.

The

was well made, and,

as in the preceding case,

a chemosis occurred.

There has been no reaction of any sort, either and it is impossible to distinguish the which the injection was made. eye into

June

10.

local or general,

The inflammatory properties


isms being demonstrated,

of the micro-organ-

we

can easily understand

the pathology of an abces de voisinage, and


ceive also the consecutive formation of
cesses as a result of the
first.

we connew ab-

This

is

one of the

modes derived from a


rulent infection
is

local process

by which pu-

accomplished.

As a general complication, we must mention the penetration of the microbe into the blood, and the possibility, thanks to this liquid, of the infectious
agent being transported to
tion,
all

the organs, where

it

will form, either centres of softening

and suppura-

which have been called metastatic abcesses,

184
or,

PHYSIOLOGY OF THE BACTERIA.


a
little

by

different

mechanism, veritable emsince,


is

bolisms.

A very curious
of a

fact

was published not long


This observation

which supports this view. to Hjalmar Helberg (of Christiania).

due

In the case

woman

in

whom

uterus was found to

mucus membrane of the be covered with a great numthe

ber of micro-germs, the tissues of the eye presented a series of lesions, which microscopic ex-

amination showed to be due to the presence of parasites. In the cells of the cornea, in the vessels

of the choroid and of the retina, were seen

veritable plugs containing bacteria identical with

the micro-organisms found in the genital parts.

The author admits


I will

that the ocular inflammation


septic embolisms.

had been produced by


cess,

not insist further upon the embolic prois

which

studied in another chapter of this

work.

To sum
of bacteria

up, I will say that, so far as the influence

on wounds themselves is concerned, we know as yet nothing positive since, as I have said, we find these parasites upon the surface of solutions
;

which progress most rapidly and surely to a cure. Better informed in regard to the generalization of the process and the infection of the organism by products formed on the surface of the wound, we cannot dissimulate the fact that
of continuity

much

of our information

line of hypothesis,

and that

to accept

it

yet as

upon the borderwould be imprudent invariably settled. According


is
still

it

to all probability,

we

will find

the

key

to these

THE BACTERIA. IN SURGICAL LESIONS.


obscurities

185

when we know

better

how

to distin-

guish the different kinds of microphytes.

Indeed,

while Billroth only admits two invariable funda-

mental types, Coccos and Bacteria, Weigert has


recently expressed views quite opposed to these in

speaking of the bacteria of fermentations and of

chromogenes, which according to Cohn and Pasteur constitute as

many

physiological species as

there are different fermentations and colorations. According to Weigert, there are an infinite number of sorts of bacteria, arising from the fact that each Micrococcus assumes special vital properties according to the medium in which it finds itself, and consequently gives birth, as the result of decompositions which it effects in taking possession of oxygen, to various chemical products acting as morbid
viruses.

Influence of the Preceding


peutics.

We have

Notions upon Therafor

just

shown that
it

many

au-

thors -germs are the origin of the greater part of

the complications of wounds,

was then natural

that they should seek to prevent their develop-

ment.

In order to attain this result, the clinicians

have used means either physical or chemical. M. Alphonse Gue"rin, founding his practice upon
the ideas of Pasteur regarding the possibility of
filtering

and purifying the

air

by passing

it

through

cotton-wadding,
realized

conceived the idea of covering


of cotton,

wounds with a considerable quantity

the

fact cannot be denied

an

and

immense

progress in the treatment of severe traumatisms.

186

PHYSIOLOGT OF THE BACTERIA.


attributed his success to
all

But while the surgeon

the absence of germs, and gave

the credit to

M. Pasteur, the microscopists had no difficulty in proving that, far from being exempt from them, the pus of the wounds kept under these dressings swarmed with micro-organisms. It was, then, to some very different conditions that the real progress The as realized by M. Gue"rin should be ascribed.
constant temperature, the absolute immobility, the

continued pressure, and consequent absence of any


rent of the tissues, as well as the absolute want of

absorbent openings upon the surface of the wound,


are probably the circumstances to which so

many
is

happy cures have been due

since 1870.

Among

the chemical agents to which recourse


in the front

had we must place


extolled especially

rank carbolic

acid,

by Lister. Not more than for M. Guerin can we deny the fact that- the number
of cures after severe surgical operations has

been

considerably increased by operating under a cloud


of pulverized carbolic acid solution, and applying upon the wound nothing but dressings which have

been for a long time submitted to the action of


this agent.

But

in this instance also the interpre-

was a mistaken one, in seeking the secret of success in the exclusion of every microbe for, in a great majority of cases, Virchow has not been able to find any appreciable difference between the pus treated by the old methods and that of wounds submitted to that of Lister. And, nevertation
;

theless,

tions of

purulent infection disappeared, complicaall kinds diminished singularly in fre-

THE BACTERIA
;

IN SURGICAL LESIONS.

187

quency and union by first intention is attempted and often obtained in very extensive wounds, where formerly it could not have been hoped for, even in the cases which presented the most favorable
appearance.
results

Let us rejoice that we can record such favorable and, however cloudy the present theories concerning bacteria may be, let us recognize that the labors of Pasteur and Cohn have at least had the merit of inspiring great reforms, which are
;

subjects of just pride in the operative surgery of

the present day.

Many other substances have also been praised as being an obstacle to the development of germs. We will only mention the permanganate of potash,
the hyposulphites, chlorine water ,and tincture of
eucalyptus, of which the action
is

doubtful.

We

must

still

mention glycerine, which from

its affinity

for water has the property of fettering the

move-

ments

of bacteria,

and determining

at their ex-

pense a considerable exosmosis. We will not return here to the subject of the origin of micro-organisms, but refer the reader to the chapter in which this has already been
treated.

CONCLUSIONS.

"We may sum up as follows the actual knowledge upon the bacteria
:

state of

our

1.

The

bacteria are cellular organisms of vege-

table nature.

Their organism is more complicated than The principal was for a long time believed. their structure, the points brought to light are presence of cilia, the nature of the substances
2.
:

contained in their protoplasm,


grains of sulphur, etc.
3.

colored granules,

The forms
etc.

of torula, zooglcea, leptothrix,

my-

coderma,
4.

The multiple affinities of the bacteria, on the one hand with the algae, on the other with the fungi, differently understood by authors, and their development, still unknown for the greater number of species, make it impossible to classify these beings except in a provisional manner. 5. This development, well studied in several species of Bacillus, has proved that bacteria may multiply not only by fission, but also by formation of spores, and even by veritable sporangia.
6.

These spores or permanent germs are the

PHYSIOLOGY OP THE BACTERIA.


principal

189

means by which these

inferior organisms

are disseminated.
7.

As

to their role in fermentations, in putrefac-

tions, in

contagious diseases, and in surgical


considerable

le-

sions,

notwithstanding the

number
it is
1

of

labors of

which the bacteria have been the


not
it

object in these different points of view,

yet possible to define

in a certain manner.

1 The candid statement made in the last paragraph by a savant who has shown himself so familiar with the whole subject, and by a Frenchman who evidently has a high regard for his distinguished countryman,

Pasteur, shows us that there


ful

is still much to be done in the way of careand conscientious work in the study of the life-histories and physiological functions of the micro-organisms known under the general name

of bacteria.

6. M.

S.

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C.

t.

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le

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p. 69.

causae par un nouveau vibrion aerobic.

LXXXVII.

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Quar. Journ. Micr.


e

Sci.

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aerie,

t.

p.

elec-

trischen Stromes auf die

Vermehrung von Bacterien.


1,

Beitrage zur Biologie der Pflanzen., Bd. III. Heft


p. 141.

Dallinger.

Recent Researches into the Origin and Development of Minute and Lowly Forms of Life. Proc. of Roy. Inst. G. B. Vol. VIII, Part 4, No. 67.

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Friedrich.
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Pathogenetic
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No.

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Bacteria

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Harz.
mal.
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218

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Fourth Replies of M. Berthelot. C. R. Ac. des Sc. LXXXVIII, nos. 2, 3, 4, and 6. C. R. Observations on M. Felz's Paper. 1879. Pasteur.

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C. R. Ac. des

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Identity of Bacillus Amylobacter and

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M.

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LXXXIX,

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On the Nature of Tommasi-Crudelli and Klebs. the Specific Agent which produces Malarial Fever.
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Arndt.

Observations

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pp. 51-83.

on Spirochete denticola.
pp. 76-86.

The

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Mucus

Arch. Path. Anat.

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Ein Beitrag zur Dorpat, 1880, Bkantlecht. Algues d'eau douce, genique. Ann. d'Hyg. 2 Dallinger. Experiments
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Public,

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determine the
etc.

Thermal Death-point of known Monad Germs,

Journ. E. Micr. Soc. Ill, pp. 1-16. Declat. On the Analogies which seem to exist be-

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LXXIX,
Hansen, E.

p. 374.

C.

Contributions

the

Knowledge of

Organisms which live in Beer and the Wort of Beer. 1. Besearches on the Organisms which at Different Periods of the Year are found in the Air at Carlsberg and Neighborhood, and which develop in Beerwort, I, pp. 49-67.
2. 3.

On

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in

Compte Rendu Trav. Lab. Carlsberg 49-75. (French Resume), Oidium Red-colored Saccharomyces, 75-80. Hansen, E. exercised by Hansen, E. C. On the
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C.
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Organisms observed

Beer and Wort, pp. 72-75.


pp.

C.

pp.

Influence

the In-

troduction of Air on the Fermentation of Wort, pp.

81-88.

Hansen, E. C. Hansen, E. C. Hansen, G. A.


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Hypothesis Mycoderma Bacillus

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Aceti, pp. 94-96.

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225-240.

Hansen, G. A.
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Bacillus leprce.
pp. 32-42.

Arch. path. Anat.

Phys.

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220
1880. King-sett.
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BIBLIOGRAPHY.
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88 pp.

Kobert.

Naturw. (Gieble), IV, pp. 884-886. Lassar, O. Die Micrococcen der Phosphorescenz. Arch. f. d. Ges. Physiol., Bonn, 1880, XXI, 104-109. Loomis. The Relation of Bacteria to Pyaemia. Med. Gaz. N. Y., VII, 24. The Schizomycetes and their R61e in Luerssen, C.
Diseases and Fermentations.
pp. 242-260.

Bacillus
Beitr. Biol.
e

Anthracis.

Zeitsch.

Gesammt.

Rev. Internat.

Sci.

V,

Researches on the Bacteria suspended the 119-140. Pflanzen Miguel. Poussieres organisers de l'atmosphere. Ann. 226, 333. 2 sene, d'Hyg. Moss, E. L. Experiments on Septic Organisms
Miflet.
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in
III. pp.

Public,

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in

Rep. Brit. Ass. Adv. Sci. Meeting, 1879, pp. 416-18. Existence of Bacteria or Nencki, Giacosa.
Living
Tissues.

49 th
their

Germs
Nencki.
zoa.

in

the Healthy Organs of Animals.

Journ.

Relation of Oxygen the Life the Micro337-358. Journ. Prakt. Chem. XIX. the Nencki, M. Contributions
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Prakt. Chem.

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of

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to

Life-history of the
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Schizomycetes.
Leipsic, 1880.)

62 pp. 2 plates and 7

(8vo,

Ollive

C.

On the
with
p.

Resistance of Barbirine Sheep to

Inoculation

Charbon.

C.

R.

Ac.

des

Sc.

LXXXIX,
Pasteur.

Observations

792.

the Bacteridium of

(verbal) on the Cold which Anthrax and other Microscopic

Organisms can support without losing their Virulence. C. R. Ac. des Sc. LXXXIX, p. 1015. On Virulent Diseases, and particularly that Pasteur.

BIBLIOGRAPHY.
popularly called " Cholera des Pouhs."

221
C. R. Ac.

1880. Pasteur.

Pasteur. On

des Sc.

XC,

pp. 239-248.

Studies on Fermentation (trans, by F. Faulkner and D. C. Roff ). 8vo, London, 1879. Pasteur. On the Cholera of Fowls. C. R. Ac. des Sc. XC, pp. 952-958, 1030-1033.
the Extension of the
to the Etiology of

Theory of Germs some Common Diseases. C. R. XC,


the

pp. 1033-1044.

Pragmowski.

On

Development of some Species

of Bacteria and the Fermentation which they cause.


(Transl. from "Bot. Zeitung.") Bev. Internat. Sci. IV, pp. 546-554. Pragmowski. Researches on the Developmental History and Ferment-action of some Species of Bacteria, 58 pp. 2 plates (8vo., Leipzig, 1880). Romueguere. The Micrqscopic Organisms of the

Blood.

Rev. Mycol.

II, pp.

71-73.
Nitrification.

Schloesing and Munz.


C. R. Ac. des Sc.

Researches on
"Wiss.,

Soyka, G.
Air.

On the Passage
M.

LXXXIX, pp.

891-894, 1074-1077.

of Schizomycetes into the

S. B.

K. Bayer. Akad.

Munchen, 1879,

pp. 140-147.

Stapff, F.
Zeitschr.

Bacteria

in

the

Gotthard Tunnel.
to exist

Talmt.

On

Gesammt. Naturw.

(Giebel), IV, pp. 848-853.

the Analogies which

seem

between

the Cholera of Fowls and the Malady of Sleep (Ne-

On the Butyric Ferment the Coal LXXXIX, 1002-1004. Epoch. C. R. Ac. des Tieghem, P. Van. On the pretended Flagella of BacTieghem, P. Van.
in

lavan).

C. R. Ac. des Sc.

XC,

pp. 1104-1117.

Sc.

pp.

teria.

C. R. Ac. des Sc. Sp. Nov. C. R.

Tieghem, P. Van.
ferum.
C. R.

Development
On
pp. 141, 142.

LXXXJX,

pp. 37-45.

of Spirillum Amylipp. 65-68.

LXXXIX,

Tieghem, P. Van.
Tieghem, P. Van.
lose.

the Spores of some Bacteria.

On the Fermentation CelluBot. Fr. XXVI, 25-30. Malaria? Tommasi-Crudelli, C. Selinunte


of
Bull. Soc.

LXXXIX,

pp.

Bacillus

of

222

BIBLIOGRAPHY.

and Campobello. Atti R. Aocad. Lincei (Transunt.), IV, pp. 1 10-113; 653-696. 1880. Vallin. Sur la resistance des bacteries a la chaleur. Hun. d'Hyg. Public, 2 e sene XLIX, p. 259. The Diphtheria Fungus. Midi. Nat. II, Vizb, J. E.

E. Micro-fungi Britannici Cents. 1-4. Vize, "Waldstein. Contribution the Biology of the BacQuart. Journ. Micr. XX, pp. 190-201. Wernich. Experiments on with Micrococpp. 289-291.
J.

to

teria.

Sci.

Infection

cus prodigiosus.

Beit.

BioL

j^flanz. Ill, pp.

105-118.

INDEX.

B. genus established by Davaine,

A.
Abscesses, bacteria
Acetic ferment, 83.
Aerobies, 116.in, 178.

18.

B. luteum, 73. B. prodigiosum, 73.


B. violaceum, 74.

Algae, bacteria classed with, 56.

Bacterio-purpurine, 38.

Ammonia, does not


source

dissolve bac149.

Bacterium,
B.

80.

teria, 53.
of,

B. seruginosum, 85.

brunneum,

85.

Anaerobies, 116.
Antiseptic treatment, in charbon,
166.

B. catenula, 82. B. griseum, 81. B. lineola, 81.

Ascococcus^e. A. BiUrothi, 96.

B. littoreum, 81.
B.

'

punctum,

82.

B. rubescens, 85. B. sulphuratum, 86. B. syncyanum, 85.

B.
Bacillus, 87.
B. amylobacter, 88.

B. termo, 81. B. triloculare, 81. B. xanthinum, 85. Bastian, views and experiments Beggiatoa, 91. [of, 103. B. alba, 91.
B. arachnoidea, 91.

B. anthracis, 88.
B. ruber, 89.
B. subtilis, 87.

B. ulna, 89.
Bact^ridie charbonneuse, 88.
B. des infusions, 90.

B. leptomitiformis, 91.
B. minima, 91. B. mirabilis, 91. B. nivea, 91.
Billroth, views
of,

B.
B.

du levain, 89. du vin tourn6,

90.

B. glaireuse, 90. B. intestinal, 89.

22.
? 108.
of,

Blood, bacteria in
bacteria, 122.

Bacteridium auriantiacum, 73. B. cyaneum, 73.

Boracic acid, action

upon the

224
C.

INDEX.
Conclusions, 188.
of,

Carbolic acid, action


bacteria, 122.

upon

Coze and Eeltz, experiments


155.

of,

Carbonic acid, action


teria, 122.

of,

upon bac-

Culture-fluid of Cohn, 113. Pasteur, 112.

Carbon,

how

obtained by bacbacteria, 35.

Mayer, 113.

teria, 113.

Cell-membrane of
60.

D.

Characters, generic

and

specific,

Charbon, bacteria in, 138. Chloroform, action of, upon bacteria, 122.

Dallinser and Drysdale, extract from paper of, on " The Existence of Flagella in B. Termo,"
41.

Chlorophyll, bacteria destitute


56.

Davaine, classification
of,

of, 18.

Definition of bacteria, 13.


72.

Chromogenes,

Desmobacteria, 86.
39.

Cilia, described

by Ehrenberg,

extract from paper of Dallinger

Dimensions of the bacteria, 29. Diphtheria, bacteria, in, 170.


Dissemination of the bacteria, 103. in air, 103. in the human organism, 107. in water, 106. Distinction between animals and
vegetables, 53.

and Drysdale de-

scribing, 41.

seen by various authors, 39.


Cladrothrix, 97.
CI. dichotoma, 97.

Classification of Billroth, 23.

Bory de SaintVincent, 15.

Cohn, 65. Davaine, 18.


Dujardin, 17. Ehrenberg, 16.

of bacteria from inorganic substances,


49.

Dujardin, classification

of, 17.

Hoffmann,
Nageli, 57. Sachs, 57.

20.

E.

O. F. Miiller, 14.

Ehrenberg,

classification of, 16.

generic and specific


characters, 59.

Fat-qlobules, resemblance
to micrococci, 51.

of,

Coccobacteria septica of Billroth,


22.

Fermentation
classification of, 66.
of,

acetic, 139.

Cohn,
120.

ammoniacal of
urine, 142.

Cold, effects

upon the bacteria,

butyric, 145.
lactic, 144.

Color of the bacteria, 31. Colored bacteria, where found, 32.

viscous, 146.

Compressed

air,

action

of,

upon

Fermentations, role of bacteria


in, 137.

the bacteria, 121.

INDEX.
Forms
of the bacteria, 29. Fungi, bacteria classed with, 56.

225
aurantiacus, 73.

G.

Glanders,

171.

Gliabacteria, 45.

Gliacoccos, 45.

Grouping, different modes

of, 43.

H.

Heterogenesis, 102. Hoffmann, memoir of,

20.

I.

M. M. M. M. M. M. M. M. M. M. M. M. M. M. M. M. M.

bombycis, 76.
candidus, 74.
chlorinus, 73.

crepusculum, 75.
cyaneus, 73.
diphtheriticus, 76.
fulvus, 74.
luteus, 73.

of epidemic diarrhoea, 77. of exanthematous typhus, 78. of glanders, 78.

of intestinal typhus, 78. of rugeola, 77.


of scarlatina, 77. of stringy wine, 75. of syphilis, 78.
77.

Intermittent fever,

174.

K.

Koch, experiments
charbon, 101.

of,

relating to

M. of the variola of animals, M. prodigiosus, 73. M. septicus, 76. M. urea?, 75. M. vaccinae, 76. M. violaceus, 74.
Microbacteria, 65, 80.

Microspha^ra vaccinae, 76.


L.

Leptothrix,

90.

L. brevissima, 90.

Microsporon septicus, 76. Microzyma bombycis, 76. Miquel, experiments of, 104. Monas crepusculum, 75.

L. croespitosa, 90.
L. parasitica, 90. L. pusilla, 90.

L. radians, 90.
L. rigidula, 90. L. spissa, 90.

M. gracilis, 79. M. Okenii, 79. M. prodigiosa, 73. M. termo, 81.

M.
first

vinosa, 79.

Leeuwenboeck,
bacteria, 14.

to

observe

M. Warmingii, 79. Movement, brownien,


cause
of
of,

33.

34.
32.

Leptothrix, form of grouping of

two kinds,

the bacteria, 43.

Muller, O.

F., classification of, 14.


of,

Multiplication, rapidity

125.

M.

Malignant pustule, without


bacteria, 164.

Mycoderma aceti, M. form of, 45. M. vini, 141.


Myconostoc, 96. M. gregarium, 97.

83, 140.

Micrococcus, 72.
\

I;,

226

INDEX.
Respiration of the bacteria, 111. Reproduction of the bacteria, 123.

N.
Nageli,
149.
classification of, 57.

Nitrification, role of bacteria in,

by by

fission, 123.

spores, 126.

Rhabdomonas

rosea, 80.
of,

Nitrogen,

how

obtained

by the

Rugeola, bacteria in blood

169.

bacteria, 112.

Nutrition of the bacteria, 111.

S.

Saccharomycetes,
0.
Sachs, classification
Sarcina, 96.

57.
of,

57.

Ophidomonas sanguinea, 80. Origin of the bacteria, 101.


Oxygen, role
teria, 121.
of,

Scarlatina, bacteria in blood

of,

169.

115.

Ozone, action

of^

upon the bac-

Schizomycetes, 57. Schizophytes (Cohn), 66. Septicemia, role of bacteria in,


152.
Species, physiological of Pasteur,

Palmella

63.

prodigiosa, 73.

Pathogenes, 75.
Petalobacteria, 46.

value of, 61. Spherobacteria, 71.


Spirobacteria, 91.

Petalococcos, 46.

Pigmentary
series, 55.

bacteria, 72.

Place of the bacteria in vegetable

Spiromonas Cohnii, 80. Spirillum, 94. S. attenuatum, 96.


S. Rosenbergii, 96. S.

Polymorphism, 133.
Position of the bacteria, 48.

rufum, 94.
undula, 94. violaceum, 96.

S. tenue, 94.
S. S.

Protoplasm, currents granules


Pseudobacteria, 50.

in, 37. in, 37.

of the bacteria, 36.

S. volutans, 94.

Spirochete, 93.
S. gigantea, 93.
S.

Pulverulent precipitate, consisting of bacteria, 46.

Obermeieri, 93.

Pus, bacteria

in, 111, 177.

S. plicatilis, 93.

Putrefaction, r61e of bacteria in, 148.

Sporangia, 130.
Spores, development
of,

128.
131.

germination
R.

of,

Starch, in Bacillus amylobacter, 39.


of,

Ray-Lankester, researches
38.

Streptobacteria, 44.

relating to bacterio-purpurine,

Streptococcos, 44.
Streptothrix, 97.
S. Foersteri, 97.

Relapsing fever, bacteria

in, 173.

INDEX.
Structure of the bacteria, 35.
Surgical lesions, role of bacteria
in, 173.

227

Sulphate of Quinine, action of, upon the bacteria, 122. Sulphur, contained in bacteria,
38.

V. lactic, 83. V. lineola, 81. V. prolit'er, 94. V. rugula, 92.


V. V. V. V. V. V.
serpens, 93.
subtilis, 87.

Swarms, 46.

syncyanus, 85. synxanthus, 85.


tartaric right, 83.

T.

tremulans, 81. Vibrioniens, definition of Ehrenberg, 16.

Temperature,

action

of,

upon

the bacteria, 118.

Thermal death point


119.

of bacteria,

W.
Wounds,
180.
effects of bacteria

Torula form of bacteria, 43.

upon,

Typhoid

fever, bacteria in, 171.

Y. U.

Ulcerative endocarditis,
teria in, 173.

bac-

Yellow-eevek blood, no bacteria, 108.

contains

Ulvina

aceti, 83.

Z.

Variola, micrococci Vibrio, 92.


V.
bacillus, 89.

in, 167.

Zooglcea, genus established by Cohn, 21. Zooglcea form of grouping of the


bacteria, 44;

Zymogenes,

75.

University Press

John Wilson and Son, Cambridge.

wsm&^mmz l

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