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bacterial cyclogeny
Prolegomena to Investigations
about structure, sexual and asexual reproduction
planting and development of bacteria

From

Professor Dr. Günther Enderlein

curator at the Zoological Museum of the University of Berlin

With 330 illustrations

Berlin and Leipzig 1925

Walter de Gruyter & Co .

formerly GJ Göschen'sche Verlagshandlung — J. Guttentag, Verlagsbuchhandlung
Georg Reimer — Karl J. Trübner — Veit & Comp.
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All rights, especially the right of

translation, are reserved.

Printed by Walter de Gruyter & Co., Berlin W. 10

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Foreword.

This treatise is unchanged in form and content from when it was written
in 1915—1916. Newer entries refer mostly to literature from later times.

In a series of preliminary reports in 1916 and later, all the new points of view
were formulated clearly and distinctly, so that they can be regarded as
previous reports. I welcome with satisfaction the fact that one of the most
outstanding bacteriologists, the American Löhnis, has also recently begun to
recognize the individual parts of the development cycle, in his latest work of
1923, almost seven years after this was done in my preliminary reports, so
that a more rapid naturalization of this approach can be expected.

The uniform magnification of most images to ten thousand times

according to precise micrometer measurements made it possible to show
even the finest differences that are still visible to the eye, thus offering even
the layman the opportunity to gain a comparative insight into the organization
of these smallest living creatures, something that has previously been
impossible with the help of the microphotographic images in many textbooks,
since the many optical phenomena of these are not able to create a clear
image.
Berlin , December 1924.
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table of contents.

Page

I. Introduction 1

II. Historical overview of morphology, cytology and development

Development of Bacteria A. History 4
of Cytology and Caryology of Bacteria B. Overview of the most important 4
literature on the manifestations of
bacteria 17
C. Historical aspects of comparative morphology and biology D. Historical 37
aspects of the question of sexuality 43

III. Elements of the comparative cytology of bacteria A. Generative organelles 45

46
1. The Mych (the primordial core) 46
2. The Centriolite 48
B. Somatic organelles 49
1. Trophic organelles a) 49
Cytoplasm b) 50
Trophoconia, trophosome and trophosomelles c) Sporitin grain 51
d) Plasmodesmata 60
2. Protective 61
organelles a) Membrane b) 61
Mucous 61
membrane 3. Motor 62
organelles a) Flagella b) 63
Undulating 63
membrane 4. Respiratory, 64
assimilatory and excretory organelles 64
a) vacuoles b) 64
dye formation 65

IV. Elements of the comparative morphology of bacteria 65

A. The morphological unit (mychit) 66
1. The Mychit (the primordial cell) 67
2. The Mychomitosis and the Diplomychite 3. The 68
Mychomer and the Mychomerite 4. The 72
Triplomychite B. The 72
Pliomychite 1. The 73
Dimychite 2. The 73
Mychomitosis of the Dimychites and the Dimychosis 76
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VI table of contents.

Page

3. The Didimychit 4. The 80

Syndimychit 83

C. The Symmychon and the Symmychit 86

V. Reproduction of bacteria A. Monogony and 88

arthrogony of bacteria 1. Monogony 2. Isomorphic 88

arthrogony 3. 89

Heteromorphic arthrogony a) 89

Fructifications ÿ) The gonidia ß) The 90

cystite ÿ) The cystoid b) 91

Embryonic 91

formations a) 97

Arthrothecite ß) 101

Endothecite c) Partial 101

formations ÿ) Oidia .10 1

formations ß) 104

Pseudothecite T) Oothecite 105

and sphaerothecite B. 105

Sexual reproduction : 106

of bacteria 1. The gonit 107

107
111

2. The spermite a) 117

The organization of the spermite b) On the 118

measurability of ultramicrometric quantities c) The centriolite 3. The 12Ó

oite 121
121

4. The Symplast 5. 122

Fertilization a) Search for 124

the Ç by the ¿ b) Copulation c) Union of the 124

mychomeres d) 126

Conjunction VI. Cyclogeny of bacteria VI. 128

ÿ) Causal factors of 129

cyclogeny A. The Cyclostadiums 1. The Basite (The 129

Probasite. The Anabasite) 135

135
137
2. The Basoit 140

3. The phytite a) 141

The prophytite b) The 141

anaphytite c) The 141

cataphytite d) The sporite 141

(disporite and didisporite) 142

4. Group of catatactous ascites a) Phytascite 149

b) Sporascite c) 154

Gonascite d) Cystascite 154

e) Thecascite f) 155

Catascite 155
156
156
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table of contents. VU

Page

5. Group of syntactic ascites a) The 157

synascite b) The 157

endothecite and the arthrothecite 6. The plastite 164

164

a) The proplastite b). 164

The cataplastite 7. The 164

zoite 164
8. The Pseudoscite 168
9. The Fructifications 170
10. The Coenobium 170
B. The formants 170

1. Gestalt designants (cyclostadials) 171

a) Mychostasis b) 172

Flagella 172

2. Physiological and biological designants a) Mucus formation 173

b) Dye excretion from 173

dye-containing culture media c) Dye formation d) Carbohydrate cleavage (sugar 175

fermentation) e) 176

Alkalogeny f) Acidogeny g) Protein cleavage h) Gelatin 178

liquefaction i) 181

Virulence and 181

pathogenicity k) 181

Luminosity 1) Agglutinability m) 181

Haemolysis n) Parasitism o) Oxygen 182

tolerance, anaerobiosis 183

183
185
186
186
) Thermoresistance q) 186

Colony form 186

C. The Colony 187
1. The primary colony 187
2. The secondary colony 192

D. The Mochlosè and Mochlolyse 195

1. The Mochlose 195

2. Mochlolysis 197
3. The Pseudomochlose 198

4. Pseudomochlolysis 199

E. The Cyclode (Cyclogenetic Basic Law) 199

VI. ß) Conditional factors of cyclogeny A. Trophomorphoses 1. Solid neutral 209

nutrient medium 210

2^0

2. Liquid neutral nutrient medium 3. Parasitism, 211

symbiosis 4. Syntrophosis B. 211

Chemomorphoses 212
212

1. Alkalis (alkalomorphoses) 212

2. Acids (oxymorphoses) 213

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VIII table of contents.

page
•3. Salts (halomorphoses) 213

4. Poisons (toxomorphoses) 214

5. Other influences C. 216
Thermomorphoses D. 216
Aëromorphoses E. 217

Photomorphoses F. 218
Electromorphoses 223

VII. Fundamentals for a comparative morphological classification of bacteria

fication 223

A. The organic relationships of the Protomychotes B. Systematic 232

overview C. Overview table of the 235

genera of bacteria D. The phylogeny of bacteria (Monomychota and 290

Dimychota) 292

VIII. Relationships of bacteria to other lower organisms ... . 295 295

A. Relationships to the fungi,,
B. .„ ,, Cyanophycea 297

IX. Relationships of the Mychotes to the Metamychotes 298

A. Relationships with the protozoa B. 299

" "" metazoans and metaphytes 305

C. Phylogeny of the nucleus and the cell 311

1. The morphological significance of the centriolite 2. Polismatology 311

316
D. Remarks on a Classification of the Protota .. 327

X. The significance of bacterial cyclogeny for medicine ... . 330

A. The cyclostatic moment of virulence (virostage). (Therapeutic
importance of cyclogeny) 330

B. The diagnostic significance of cyclogeny 335

1. On the diphtheria question 336

XI. Alphabetical list of the main newly introduced morphological and biological terms A. Bacteriology B. Cytology
349
349
:... . 355

XII. Bibliography 356

XIII. List of Illustrations 369

XIV. Appendix: List of publications of the author XV. Alphabetical table of contents 371

385
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I. Introduction.
Comparative morphology is the basis
of all biology.

The trend of ideas in recent decades has resulted in a certain

disproportion between physiological and biological research and comparative
morphology, particularly in bacteriology. Comparative morphology has been
viewed by some as something of secondary importance, and it was believed
that by pushing it aside, an achievement had been made for the benefit of
biology, although in reality it represents the basis of biology itself and is the
mother of all phylogenetic, ontogenetic and systematic knowledge of nature.
The foundation of knowledge for this field of research also lies in the recognition
of the rhythm of all organic processes.

The basic condition for a morphological developmental study is the organic continuity of
successive developmental states.

In the course of systematically continuing, comparative morphological

studies on bacteria, the realization forced itself upon me with convincing force
that bacteriology, which has been built up particularly from a practical point of
view, does not correspond at all to this basic condition and is founded on a
basis which is completely contrary to the actual conditions.
The main moments of this false direction of knowledge are three
words indicated:

Monomorphism — mutation theory — monocytism.

In the present study, an attempt is made to build bacteriology on a comparative morphological
basis. Since this attempt was not entirely unsuccessful, the results are by no means theoretical, but
are of a thoroughly comparative morphological nature, but nevertheless appear to be capable of
providing orientation for entire areas of individual biological and physiological facts.

E ÿ de r 1 ei ÿ, bacterial cyclogeny. 1
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2 Introduction.

It is a difficult matter, given the different interpretations of the un-

to meet all requirements of our observation material.
It is therefore recommended that this work be entrusted to the specialist.
When carrying out this sketch - it could not be otherwise, given the extent of the area - the main
goals had to be kept in mind as the direction that would indicate the direction, so that some aspects
may have been neglected. All of this is left to the development and verification of special research. I
am always grateful for suggestions, whatever they may be in.

Thus, from the dust of past decades, a new, comparatively morphologically clarified
pleomorphism may arise, which will displace the theory of mutation and monocytism in bacteriology
and replace them with cyclogeny and pleocytism, so that bacteriology may move from the stage of
theoretical attempts at explanation to the state of absolute biological-morphological knowledge.

Comparative morphologically sharply defined terms, which are indispensable for an

understandable discussion, elevate bacteriology to a comparative morphological discipline. This
naturally made it necessary to create a whole series of terms, which may initially deter some people
from attempting to penetrate this subject. But the essence of recognized terms crystallizes as a
name. Without a sharp comparative-morphological nomenclature, it is impossible to gain a clear idea
of the essence of organisms. Even biological terms such as mochiosis, for which a conclusive
explanation and understanding was not possible, are fixed by the name and can be used. "For
precisely where terms are lacking, a word appears at the right time."

But even this concept of the Mochlose, which at first seemed theoretical and vague to me,
has in the course of time taken on an ever more sharply defined form in my imagination, and the
comparative parallelism of this with the life of Meloë, which I drew on in the introductory pages of
Chapter VII (Classification) in a much later entry, has completely dispelled all remaining doubts and
also eliminated the remnants of theory for this concept.

How this nomenclature arose in the progress of knowledge

and has been recognized as indispensable for full understanding, so will anyone who has penetrated
into the understanding of the whole of cyclogeny become convinced of its necessity. The structure
of the nomenclature, which only developed into this conceivable possible simplification with the
completion of the whole work and was so carefully thought out
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Introduction. 3

has extensive organic relationships to its meaning in morphological and biological terms and results
from the synthesis of very few words. For this purpose, short, monosyllabic words had to be chosen
as terms for the basic morphological elements, so that the morphologically complicated terms did
not become too long. It should be mentioned that the term for the primal nucleus, Mych, was derived
from "tò ÿÿÿÿÿ = the innermost", which represents the starting point for most compound terms, the
derivation of which is usually easily recognizable. The information in section XI is sufficient for the
etymological structure.

The essence of all sciences is the organization of matter. Only in this way is the overview
necessary for a deeper understanding possible.
The main results are the following: 1. Discovery of the
bacterial nucleus: Mych (primordial nucleus).
2. Justification of a comparative bacterial morphology.
3. Proof of the universality of gonidia formation in all bacteria as a fundamental form of
asexual fructification (previously only established for some vaginal bacteria - Crenothrix
and some sphaerotilids).

4. Evidence of sexual reproduction of the bacteria.

5. Morphological evidence for the sporite (the so-called bacterial spore) as a special form
of oidia formation peculiar to a small side branch.

6. Determination of the cyclogeny of bacteria and its diagnostic and therapeutic significance.

7. Determination of mochlose and mochlolysis.

8. Detection of the cyclostatic moment of virulence and pathogenicity (virostage).

9. As a practical result of the cyclogenetic conception, the polyetiological assessment of

diphtheria-like infections -
tions, regardless of whether they are in the basite, phytite or cystascite stage.

In carrying out this work I was kindly supported with material and literature by Dr. C.
DORMEYER in Chorin, Privy Councillor Dr. HANAU (F) in Stettin, Dr. HENNEBERG in Berlin, Privy
Councillor Professor Dr. HEYMANN in Berlin, Mr. KLEINE in Stettin, Professor Dr. SIMROTH (f) in
Leipzig, and Director Dr.

STÜRMER in Stettin, Dr. SWELLENGREBEL in Amsterdam and Professor Dr. E. ZETTNOW in Berlin. My long-time

friend, the painter PAUL FLANDERKY in Berlin, also supported me by producing some illustrations .

l*
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4 Historical overview.

I owe the extensive literature above all to the Royal Library in Berlin, the
university libraries in Greifswald and Leipzig and, at times, to the friendly mediation
of the city library in Stettin.
I would also like to take this opportunity to express my sincerest thanks for all
of your support.
I would especially like to thank Director Dr.
STÜRMER for the hospitality he granted me in the most accommodating manner for cultures and
examinations in the rooms of the Chamber of Agriculture in Stettin, as well as my highly esteemed
boss for the duration*) of my two-year voluntary bacteriological activity in the Reserve Hospital I of the
2nd Army Corps in Stettin, and General Surgeon Dr. WERNER for the kind permission to use the library
of the Garrison Hospital of the 2nd Army Corps in Stettin.

Stettin, October 1916.

II. Historical overview of morphology, cytology

and development of bacteria.
Having an eye for the inconspicuous means seeing more
clearly.

Laotsze, Taoteking, chap. 52. (7th century BC).

The following historical sketch of the development of bacteriology is divided

into four groups.
The first group deals with all those ideas that are connected to the conception
of the bacterial individual as a cytological and caryological unit, as it has had
general validity up to the present day.
The second group includes the manifestations and their different
Interpretation.
The third group contains the development of comparative morphology, and
the reasons and errors of the monomorphists and pleomorphists as well as the
mutation theory are discussed.
The fourth group contains those moments which led a few authors, on the
basis of incorrect observations or interpretations, to attribute sexual reproduction to
bacteria.

A. History of the cytology and caryology of bacteria.

Parts of the cytological unit of the bacterium are already early on the
Attention has been drawn, but mostly without any idea
*) From August 7, 1914 to July 31, 1916.
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History of cytology and caryology. 5

of unity. But a connection between such facts and the view that certain bacteria do not
represent a unity also appears at the same time.

Thus, a distinction is quickly made between two views: 1. the bacterial individual is unicellular;
2. “multicellular.”

" " "

Closely related to this is the question of whether the bacterial individual is
mononuclear or multinuclear.
In general, the multicellularity was soon abandoned, and the multinucleus theory
was soon suppressed by the mononucleus theory, especially by the view of the "diffuse
nucleus", which is particularly represented by WEIGERT, MITROPHANOW, ZETTNOW,
SCHAUDINN, and DoFLEIN . The view that the whole or almost the whole bacterial cell
represents the nucleus has had fewer supporters.

Some of the most important points from the historical development of the idea of the
cell structure of bacteria are highlighted below.

BUCHNER (1885) considered Microspira to be "four-celled". But NÄGELI (1877)

also considered all rods, threads, spirals, etc. to be composed of individual, approximately
isodiametric units.
The bacteria mainly consisted of nuclear substance
HÜPPE (1886) as well as KLEB s (1887).
WEIGERT (1887) assumes that a differentiation of the protoplasm
from the caryoplasm in the bacteria has not yet occurred.
, morphological elements in the anthrax
LÖFFLER (1887) does note (page 250, plate II, Fig. 4 a) important

pathogen (e.g. trophosome, trophode, synascite); the passage: "The group of bacteria on the right shows a deposit

of the dye in the form of fine grains on the bacilli, in the middle of which a weakly colored central thread can also be

seen. The coarser and finer grains are due to the bacilli not being colored for long enough and being washed out in

alcohol for a long time" but proves that they are considered artifacts. In the same place, L. reports the observation

that individual members have not absorbed the gentian violet (i.e. atrophosis).

SCHOTTELIUS (1888) was the first to use the term "nucleus" for the structure that sometimes
appears thread-like in the longitudinal axis of the Bacteria (i.e., consists of trophosomes and trophodes),
but he himself says that "for the time being he only wanted to designate the central position of this
component". He also adds: "The disintegration of the Bacillus into two individuals is always preceded by a
division of the nuclear rod."
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6 History of Cytology.

He also speaks of a central spherical nucleus in the trophosomes of cocci.

BUCHNER (1888 ) considers the polar grains (i.e. telotrophosomes) to be a degeneration

with reduced resistance. MITROPHANOW (1889 ) assumes that the protoplasm and caryoplasm
in bacteria are still mixed.

The first detailed work on cytological elements was carried out by ERNST (1889); the
"sporogenic grains" observed by him comprised almost everything that is densely coloured by
trophoconia accumulations, such as gonidia (ÿ. B. Plate V, Fig. 7, Plate VI, Fig. 5), trophosomes
(numerous) [telotrophosomes and ascotrophosomes], trophosomelles (e.g.

Plate VI, Fig. 1, 4, 16 etc.), center of the sporitin grain (e.g. Plate V, Fig. 12,13), cystite (Plate VI,
Fig. 17). He interprets them as bacterial nuclei and explains some adjacent trophosomes (i.e. of
two different dimychoses that did not actually develop from each other by division) as division
phenomena. The name "Ernst's bodies" introduced later therefore has no claim to comparative
morphological justification.

BABES (1889) had already previously pointed out the same structures, assuming that they
play some role in spore formation, but failing to recognize that they are spores in the strictest sense
of the word.
BILLET (1890) describes the endogenous spore formation (i.e. the formation of gonidia) in
Bacterium osteophilum Bill. 1890, in Sfhaerotilus (Ciado-thrix) diehotoma (Cohn), etc. In the latter
form he also describes the germination of the same and a more strongly colourable spot in them,
which he calls "point germinatif" and which possibly represents the mych or at least a trophosome.

WAHRLICH (1890 ) considers the entire bacterial cell as the nucleus.

ALMQUIST (1890) treats 3 species of Streptothrix Cohn, which grow into long, unarticulated
threads, which he calls mycelium, and which show true branching. The elements into which these
threads later decompose, he relates to BREFELD's oidia or oidia spores, while these are gonidia
(ascogonidia), the germination of which he observed.

BÜTSCHLI (1890 and 1896) ascribes to bacteria a honeycomb central body, which he
regards as a nucleus, with only a very thin plasma layer on top. Since Ghromatium Okeni is a
synascite, it is immediately clear how the idea of the honeycomb structure arose. The net granules
are the trophosomes or trophosomelles, the so-called honeycomb structure is simulated by the
numerous trophods, perhaps also pseudotrophods, lying in all possible directions.
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History of Cytology. 7

WÄGER (1891 ) considers a rod in the middle of a bacillus to be the nucleus; during division it stretches, a

dumbbell-shaped form is created, and finally the two young nuclei are separated by a small, lighter space. There is

no doubt that in the first case Yerf. had before him a didimychite with a middle trophosome, in the latter one with two

mesotrophosomes, which thus did not arise from division at all, as is repeated again and again in the literature.

ZETTNOW (1891 ) supports the view already expressed earlier by (WEIGERT etc.) that in bacteria
the protoplasm is mixed with the nuclear substance.

SJÖBRING (1892) believes he can even recognize caryokinetic figures in syntactic syndimychites of the

anthrax pathogen, etc. I only need to indicate that these elements consist of trophosomes, trophosomelles and

trophodes. His ideas about mitotic division figures in micrococci, etc., are even less in line with the facts.

The views of TRAMBUSTI and GALEOTTI (1892) on nuclear division in a water bacterium
and its relationship to mitosis in higher cells are to be interpreted in a very similar way ; apparently
this is also a synascite. The idea of the presence of spores put forward in this context is highly
doubtful.

SCHEWIAKOFF (1893 ) follows the Bütschli interpretation.

According to ILKEWICZ (1894), the spores of the anthrax pathogen contain 1-2 nuclei; to
assume nuclei in bacteria for which no spores are known to exist seems incomprehensible to him.

BUNGE (1895) was the first to establish that the difficulty of staining the spore was due to the
substance, not the shell. He treated the same corpuscles as Ernst and Babes.

BABES (1895) made further observations on the same diverse structures that ERNST
considered to be sporogenic grains and called them "metachromatic bodies". The author had already
seen them in 1887 in the tubercle bacillus . Here too, they are trophosomes, trophosomelles, gonidia,
cystites, etc.

ZETTNOW (1896 and 1897) sees trophosomes and trophodes in Spirillum undula and depicts
them without going into detail. In 1897 he points out the similarity of the nucleus of algal cells,
especially of Oscillarla , with the images of spirilla due to the "scattered chromatin spheres".

The fact that in Gram's staining the same species can simultaneously produce individuals
with positive and negative staining
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8 History of Cytology.

CZAPLEWSKI (1896) already noted that this can be due to age differences.

LÖWIT (1896) , like BÜTSCHLI first, and later BUNGE (1894) and NOETZEL (1896),
considered the main part of the bacterium to be the central body and at the same time the
nucleus; in the fine cortical layer he observed fine granules (apparently morphologically
trophosomelles of a synascites), which he interpreted as centrosomes.

LAUTERBORN (1896) points out that the chromatin granules during

disappear during cell division.
In an extremely important and instructive experiment, FISCHER (1895 and 1897)
showed that the staining reactions used cannot be used for chemical purposes. He precipitated
3% and 0.3% aqueous albumose solution (deuteroalbumin); the former formed large granules,
the latter tiny coccus-like grains; he mixed both and applied double staining to a dried coverslip
smear. The result was surprising. When Flemming's Safranin-G-entiana was used, the large
granules turned red and the small ones violet. But when Gentiana was used first and then,
after differentiation with acid alcohol safranin, the large ones took on the violet coloration and
the small ones the red. The same body, which was regarded as a chemical unit, was thus
safranophilic or gentianophilic as desired. Exactly the same results were obtained with double
staining using Gram's stain, Ziehl's stain and Delafield's hematoxylin and eosin. This put an
end to the chemical theory of most histologists and the physical interpretation, which had
already been developed by GIERKE (Staining for microscopic purposes. Journal of Scientific
Microscopy.

II. Vol.) came to convincing dominance. However, this also eliminated the possibility of using
the staining reaction for the morphological identification of cell components.

In very specific terms, FISCHER (1897) opposes the idea of considering the more
strongly colourable granules of the bacterial cell as nuclei. The most important reason given
by FISCHER, which today appears to be entirely in line with the facts, is that he considers the
granules to be reserve substances.
He also considers the idea of considering the central body as the core to be incorrect.

Chromatin granules in tubercle bacilli (i.e. ascotrophosomes) are found

MUCH (1898) and demonstrates its viability and virulence.
WAGNER (1898) describes coli and typhoid bacteria as mononuclear cells, which
usually have a centrally located nucleus. Fig. a shows that this is a didimychite, in which only
one of the 4 mych in the middle has formed a trophosomelle, and it is equally certain that the
nuclear division assumed in Fig. d is nothing other than a didimychite.
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History of Cytology. 9

with 2 mesotrophosomelles; such a storage of two trophosomelles of a dimychites would be

impossible. Fig. 7 actually shows the beginning of division of a didimychites. The dimychite
of a B. coli or a typhoid pathogen is strongly stenostat. The author's statement about longer
threads (i.e. ascites) is also completely incorrect, namely that the above nuclei should form
large nuclei by coalescence, which in reality represent nothing other than trophosomes.

RUZICKA (1898) describes and depicts more strongly staining granules in the
bacterial cell. These are trophosomes and trophosomelles.
He states that they cannot be detected in all individuals and not at all times. For a cell with
only a single central granule, the author points out the similarity to a cell nucleus, but
cautiously rejects the interpretation of it as a cell nucleus. The author considers a connection
between the granules and degeneration processes to be impossible.

CATTERINA (1898) also discusses such dyeable grains.

As already mentioned, FISCHER (1897) was the first to object to the assessment of
the chromatin granules of the bacterial cell as cell nuclei on the grounds that they represent
reserve substances; he considers the bacterial cell to be nucleus-less. MIGULA (1897)
takes the same view ; he writes (page 72): "After the many investigations in this regard, the
discovery of real cell nuclei is still possible, but has become very unlikely."

A. MEYER (1899) interprets trophosomelles as nuclei and also ascribes a central

nucleus to the spore grain; however, he has to admit that not all bacterial individuals have
these nuclei. The picture relating to nuclear division (Fig. 38) again shows only two
trophosomes arranged next to one another.

ROWLAND (1899) does not explain all of the chromatin granules of the bacterial cell
as being of nuclear nature, but considers them partly as secretion products.

MARX and WOITHE (1900 ) consider the chromatin granules of the Bac -
teria cell - they call them Babes-Ernst bodies - for a homolog of centrosomes; here too,
bodies lying next to each other (i.e. trophosomes) are wrongly interpreted as division
phenomena. A core concept is not linked to them. The authors are of the opinion that the
bacilli of Babes-Ernst bodies can be lost far from all damaging influences (e.g. in pure
cultures) and only regain them in the animal body. They see the appearance of the granules
as the expression of the highest development of species and life and in this, in turn, the
equipment for the fight for existence. The observational finding is interesting that in pigment-
forming organisms (. pyocyaneus and
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10 History of Cytology.

B. prodigiosus) the number of Babes-Ernst bodies in individuals of a species is

directly proportional to the intensity of the characteristic pigment formation.

FEINBERG (1900) regards everything in the cell contents that is strongly

stained as the nucleus and thus comes to the conclusion that the nucleus takes up
the largest part of the bacterial cell. However, the author does not want to discuss
whether such a nuclear structure of bacteria meets all the requirements that have
been placed on the nuclei of animal and plant cells.

VEJDOVSKY (1900) observed a fairly constant chromatin grain in the middle

of the length of a bacterium from Gammarus, which grows to a greater or lesser
extent, and therefore considers this to be the nucleus. He stresses that these are
not artifacts, involutions, or plasmolytic phenomena. So here again we are dealing
with a trophosome, which evidently often occurs in the middle of the length of the
ascites in this species. However, the fact that trophosomes can also occur at other
locations can be seen in the later publication on the same subject (1904) .

According to MÜHLSCHLEGEL (1900), the spore (spore nucleus, which the author calls the luster body) is

formed from the material of the globules or a correspondingly finely distributed material ; the structure takes place

from the inside outwards (not the other way round as ERNST claims); the spore shell consists of two layers, the

endosporium and the ectosporium; the former is difficult to stain, but allows colours to pass through; the innermost

point in the spore, which is often easier or differently stainable than the rest of the spore, is not to be regarded as the
nucleus.

MARPMANN (1900) takes a completely isolated position in the morphological

interpretation of the entire bacterial cell as nucleus and the secreted mucous
membrane as plasma .
The distinction that KROMPECHER (1901) wants to make between Babes-
Ernst corpuscles and Bunge corpuscles is invalid, since they are partly due to
chemical and partly to purely technical color causes; it has no morphological
significance. The author's metachromatic corpuscles are said to only occur in spore-
forming organisms; it seems likely to him that they represent a precursor to the
spore. It actually seems that this is nothing other than the first stage of the sporitin
grain.

NAKANISHI (1901) makes the same error as he did several times before in
interpreting one mesotrophosome in a didimychite as a nucleus, two as a nuclear
division (cf. Fig. 3) and by attributing a centrally located nucleus to the spore. The
fact that the author has nevertheless already made the correct
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History of Cytology. 11

The fact that he had the idea that the rods were not units is demonstrated by the fact that
he speaks of "polynuclear rods" when referring to the diphtheria pathogen.
ERNST (1902) has become doubtful about the unity of the chromophilic particles and he cannot
give any compelling reasons for the nuclear nature, as he himself puts it. With regard to the chemical
investigation of the bacterial cell by NISHIMURA, the author considers the possibility that the chromatin
granules of the bacterial cell are composed of fats, cholesterol, lecithin or glycogen; then

these as containers of reserve substances or as secretion collectors or

can also be interpreted as excretory collectors; however, certain structural elements are assumed to
underlie the grains. This study is also based on trophosomes (ÿ. B. Fig. 5,13 a, 16,18,19, 25),
trophosomelles (ÿ. B. Fig. 3, 12 b), trophodes and pseudotrophodes.

The bacterial nuclei of FEDOROWITSCH (1902) again correspond to the trophosomes and
trophosomelles. He considers the latter to be nuclei that can pass into the former, and from these the
spores are then supposed to form.
But even in non-spore-forming bacteria, imperfect spores develop from these grains, which he calls
protospores (i.e. gonidia and mychite).

BÜTSCHLI (1902) again supports the idea of the central body as a nucleus; he was the first to
observe that the chromatin granules of Spirillum volutane are hollow.
He writes (page 50): "The precise examination of these
often quite impressive grains showed their hollowness very definitely; it could even be seen frequently
that the weaker refracting cavity was eccentrically positioned" (cf. Fig. 4 g, 4 h). Noteworthy in Fig. 4 a
are a series of tiny, somewhat more strongly colored plasma particles on the flagella of Splr. volutane .

The chromatin grains in Spirillum volutane are called GRIMME (1902)

Yolutans balls; like FISCHER, he sees them as reserve substances, but otherwise he is completely in
line with A. MEYERS' view. GUILLIER-MOND 1902) shows that these grains (he calls them corpuscules
meta-chromatiques) are used up in the formation of spores.

The idea , already put forward earlier by WEIGERT (1888), MITROPHANOW (1889), ZETTNOW
(1891) and others, that in bacteria protoplasm and nucleus are mixed, was expressed in sharp terms by
SCHAUDINN (1902) on the basis of studies on a very interesting, gigantic form of bacteria (i.e. synascite)
from the intestine of the cockroach, which the author calls Bac. Bütschlii , namely that in Bae. Bütschlii
"the nuclear substances, which in other cells are united to form a morphologically differentiated cell
nucleus, are here in the vegetative state still diffusely distributed through the plasma"; "only during spore
formation does a
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12 History of Cytology.

"a structure comparable to the true cell nuclei of higher organisms".

It should be particularly emphasized that SCHAUDINN does not generalize these
conditions to all bacteria, as was later attempted, but rather he cautiously speaks
only of the species at hand. He also leaves it undecided whether the conditions
should be viewed as primitive or derived, but tends to the view that the simplicity of
the bacterial cell is more likely to be brought about by regression and that they
originate from more highly organized beings (flagellates?)

He finds the idea tempting that the first living organisms did not yet show a
separation into nucleus and plasma. The fact that the spores of the form under
discussion are not morphologically spores at all is shown elsewhere. (The mesh
structure which the author attributes to Bae.
Bütschlii , corresponds to the sum of the trophodes, the more strongly staining
granules in the nodes of the meshes correspond to the trophosomes and
trophosomes.)
ASCOLI (1902) assumes three stages of development of the anthrax
pathogen : in the first stage there is no differentiation of the structure; in the second
stage, granules sometimes appear, united in groups and usually at the poles; in the
third stage, when spore formation begins, they are crowded together at the poles;
the spore, therefore, contrary to other views, does not arise from the granules.

DIETRICH and LIEBERMEISTER (1902) report on the occurrence of nucleus-

containing granules in the anthrax pathogen , and attribute to them the function of
oxygen transfer.
A. MEYER (1904) undertakes a detailed chemical investigation of these
chromatic bodies . He introduces the name volutin for the matter of these grains;
however, this should be considered a chemical term.
He calls the volutan spheres, or all corresponding chromatin granules, volutin
granules. In his opinion, when intensively stained with methylene blue, the volutin
always swells, and the volutin granules sometimes swell to form hollow spheres. In
view of the nucleohistone content of the lymphocytes of the thymus gland and the
nucleoproteins of automotive leucocytes and spermatozoids, which are likely to
serve essentially as reserve substances and energy sources, the author also
assumes that volutin, a body or nucleic acid compound that is extremely closely
related to nucleic acid, is an ergastic structure (= reserve substance). The reason
given is that the storage of volutin in bacteria that are about to form spores (he
calls them sporangia) reaches its maximum ; while during the formation of spores,
volutin is used up like fat and glycogen.

The previously discussed bacterium, gammari Vejd. 1904 is described by

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History of Cytology. 13

VEJDOVSKY (1904) further investigated the "nuclear relationships". The results are almost
the same as before (1900) ; in the author's opinion, the nucleus is always in the center and
only exceptionally at one pole of the cell. In filamentous bacteria, it assumes the
independence of the successive cells, each of which is provided with a nucleus. The author
interprets the centrally located trophosome as the nucleus, while he regards the
trophosomelles (called Babes-Ernst bodies or metachromatic bodies by the author) as
assimilation products.

The culmination of the dependence on the staining technique is represented by the work of
PREISZ (1904), who, in general and specifically (on the same organism), gives the same structures a
variety of names. PREISZ classifies the morphological elements, which are called trophosomes and
trophosomelles in the present study , in sequence, depending on the staining characteristics, as follows:

1. The acid-fast (Bunge's) corpuscles for orgasmic structures (nutrients) (page 539 etc.).

2. The Babes-Ernst granules for random components (page 430).

3. The grains of Sjöbring, Ruzicka, Zettnow, Feinberg, if not as artificial products, then
arose as a result of granular-lumpy degeneration of the plasma (page 430).

4. The nuclear rod and the central nucleus of Schottelius and the
Nakanishi's core is "a part of the plasma characterized by its strong colorability, a cell
organ in which the acid-fast bodies originate, but which subsequently disappears
completely or only in small amounts" (page 433).

5. Granules in young cells that can be intensively stained with diluted fuchsin
as real nuclei (Fig. 434, 537).
6. As spore plants and spores (this is especially
around telotrophosomes) (pages 420, 422).
The fact that the structures listed under 5 are only trophosomelles and not real nuclei
is proven by the fact that he only mentions "1 nucleus" in Didimychites ÿ. B. (cf. Fig. 3 and
4). The alleged division phenomena (in Fig. 9, 10 and 11) are in reality nothing other than
trophosomelles of completely different Dimychoses that have moved close together, and
which therefore could not have originated separately at all.

Since the centre of the spore grain is the easiest to colour, the author naturally comes
to the conclusion that this also represents a nucleus. Regarding the assumption that the
spore grain is formed from the forespores and spore primordia (i.e. from trophosomes and
trophosomelles), it must be added that the author himself admits that
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14 History of Cytology.

that this is a construction ; he says on page 538: "Since I did not observe the process of spore
formation directly, but reconstructed it from numerous pictures, I must stress, in order to avoid
justified objections, that I cannot directly follow the transition from the cell nucleus to the nucleus of
the spore primordium."

Using vital neutral red staining, OTTOLENGHI (1904) determined that the network structure
of the anthrax pathogen is initially composed of large granules connected by threads; after formation
of the light-refracting central body (i.e. sporitin granule), these granules are much smaller and are
crowded together, especially at the periphery.

Based on investigations of the anthrax pathogen, KUZICKA (1904) came to the conclusion
that these bacteria were cytodes in the sense of Haeckel, namely naked nuclei, since according to
his view only nuclein substances are involved in their structure, without having to refer to HERTWIG's
(1902) view of the chromidia scattered in the cytoplasm as representatives of the nucleus.

LEHMANN and NEUMANN (1904) also object to the constancy of the results of Gram's coloring .

MENCL (1904), as earlier authors had often done, interprets a mesotrophosome of an

ascites as a "cell nucleus", the telotrophosomes as "polar caps", and small wall-mounted, i.e.
syntactically ordered trophosomelles as "protoplasmic humps". Later (1906) the same author
interprets trophosomelles connected by trophodes as division phenomena, and the numerous
trophosomes of an ascites lead him to speak of "a clearly defined and lawfully arranged nuclear
substance and a real, formed nucleus". On the other hand, he also assumes that not all bacterial
cells, even within the same species, have a nucleus. A year later (1907) he uses the concept of
nucleus formulated in 1904 as an example of Bact. gammati Vejd. by attributing to the central
nucleus (i.e. a middle trophosome of an ascites) a membrane (spherical) that contains the chromatin
granules in the form of globules.

In Spirochaeta BalMani, PERRI N (1906 ) interprets the totality of the

Cross-filaments within a cell called the caryosome.
SWELLENGREBEL (1907) interprets the "zigzag band" (also called spiral band, granular
thread, central thread) consisting of chromatin as the nucleus based on his investigations on Bac.
maximus buccalis Miller, whereby he points out that the longitudinal division of the granules and
transverse threads (false observation) is reminiscent of the longitudinal division of the chromosomes
of higher cells. Later (1907) SWELLENGREBEL holds out the hand of Bacterium Unucleatum Swell.
1907 from his mouth—which I casually
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History of Cytology. 15

said to be a younger stage of the ascites of Bac. maximus buccalis Miller 1892 — the
cells with chromatin-containing transverse threads, granular threads or zigzag threads
are younger stages of a binucleate organism.
For a better understanding, I add here that this organism represents a syntactic ascite
(synascite). In Figures 1-18 each individual consists of about 6-8 dimychoses, in which
the possession of more or less number of trophoconia is visible in: Fig. 1: 1 trophosome,
2 trophodes; Fig. 2: 2 trophosomelles, 2 trophodes; Fig. 3: 2 trophosomelles and 2
trophodes; Fig. 4: 4 trophodes; Fig. 5: 1 trophosome, 3 trophosomelles, 2 trophodes;
Fig. 8: 3 trophosomelles and 4 trophodes; Fig. 9: about 5 trophodes; Fig. 10: 4
trophodes; Fig. 13: 2 trophosomelles and 4 trophodes; Fig. 14: 3 trophosomelles and
3 trophodes; Fig. 15: 3 trophosomelles ; Fig. 17 and 18: 2 trophosomes each, etc. The
author considers these 2 trophosomes to be the 2 nuclei. The discussion of this
"bacterial nucleus" is interesting on page 201: "It will be left to future research to
investigate whether it is perhaps possible, in addition to the previously usual
morphological differentiation of bacteria into rods, cocci, etc., to establish a bacterial
system on a purely morphological basis based on the shape of the nucleus." However,
the phylogenetic and ontogenetic grouping of the nucleus development is incorrect
according to SWELLEN-GREBEL's opinion . He explains the diffuse nucleus as the
lowest stage, the nuclear spiral as the higher stage, and the "true bacterial nucleus"
(of Bacterium hinucleatum) as the highest stage of development. In reality, the three
groups represent the same stage of development caryologically and cytologically,
namely the highest stage of development to syntactic ascite (synascite). A later
comparative study of spirochetes and spirilla by the same author (1907) is based on a
similar basis.

GUILLIERMOND (1907 etc.) opposes the view of MEYER, VEJDOVSKY,

ÿ,ÿÿÿÿÿÿ and KRUI S and MENCL that the bacilli possess a real nucleus and sees in
the fine, colourable granulations with which the cytoplasm is almost always permeated
an equivalent of the nucleus, a view which SCHAUDINN has already formulated more
sharply ; on the other hand, the author considers the organisation of B. maximus
luccalis, which SWEL-LENGREBEL interprets as a nuclear spiral or nuclear thread,
to be a higher level of organisation.

RXJZICKA (1907) considers the stainable parts of the bacterial cell (chromatin
etc.) to be "nuclear substance"; the fact that the spores cannot be stained leads to the
false conclusion that this is linin. He calls the chromatin grains (i.e. trophosomes and
gonidia) sporoid grains (or sporoid spheres) and describes them as abnormal forms of
development which, as soon as the moment arrives in which exclusive
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16 History of Cytology.

lently these forms (i.e. gonidia and gonites) present in a culture are no longer alive. As a
result of the investigations, the finding is that the bacterial individual (specifically of the
anthrax pathogen) corresponds to a naked nucleus.

MOLISCH (1907) found that purple bacteria contain a green pigment

(bacteriochlorin) in addition to bacteriopurpurin; he pointed out that these substances
possibly play a similar role to chlorophyll and carotenoids (red) in the assimilation of
carbonic acid in the chlorophyll grain.

The concept of the spiral filament nucleus is also represented by FANTHAM

(1908) in the genus Spirochaeta, and provides schematic illustrations of his idea
regarding the changes in the microscopic
behavior of this core thread.
BREDEMANN (1908/09) calls the "coccal forms and short rods" (i.e. mychite and
dimychite) "microoidia" in B. amylobaeter in contrast to the term "oidia" adopted by
MEYER in bacteriology, which encompasses larger elements.

As in numerous previous cases, HOELLING

(1910) in Fusiformis Hoell. considered the close association of two mesotrophosomes
of an ascites as a nuclear division, which he called caryosome-desmos e. Depending
on the number of trophosomes (within an ascite), he considered an ascite to be
mononuclear or multinucleate.
On the other hand, GUILLIERMOND (1910) is against the interpretation of
chromatin grains (i.e. trophosomes) as a nucleus and wants the concept of a kind of
diffuse nucleus or chromidial system to be applied only to the totality of these grains.
Later (1910) he is against the use of the term volute (MEYER 1904) and uses the name
metachromatin used by FANTHAM and PORTER (1909) for the same matter.

MENCL (1910) observed pale and dark specimens (i.e. atrophic and pliotrophic
mychites) in micrococci, etc.; he sometimes had the impression that the dark ones were
formed by budding from the pale ones. He considered the eccentric positioning or even
the wall position of its nucleus (alo trophosomelle) to be a temporary one.

The appearance of individuals in gram-positive forms of streptococci, which

decolorize after Gram, is interpreted by LEONE LO MONACO (1911) as a phenomenon
of death.
A change in the core concept cannot be found in MEYER's summary (1912)
compared to his earlier view.
The two trophosomes of heterogeneous dimychoses, which are frequently
observed in the middle of an ascites, contain double-chromatin
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Historical information about the growth forms. 17

GLAS and DISTASO (1912 ) . They also attribute longitudinal division to the "nucleus" and allow it
to disappear completely (when the chromatin disappears). They want the chromidial network of
earlier observers to be interpreted as a degenerative form of the nucleus.

Important results on the chemical composition of bacterial cells were obtained by TAMURA
(1913—1914) .
MARG. ZUELZER (1917 ) also holds the view that in the cell of bacteria, as well as in that of
spirochetes, there is no differentiation between nucleus and plasma.

DEUS SEN (1921 ) recently determined that the essence of the gram-positive phenomenon
is nuclein, especially nucleic acid; he releases it from gram-positive cells (e.g. yeast cells, Bae.
lulgaricus = yogurt producer) by means of acid, and these then appear gram-negative; while these,
by adding nuclein and nucleic acid in the form of their Na salts,

appear again as gram-positive. Recently, KIRCHEN-STEINS (1921 ) again considers the

trophosomes as core elements and falsely considers the bacteria to be cells analogous to those of
plants and animals.

B. Overview of the most important literature on the

manifestations of bacteria.
The controversy between monomorphism and pleomorphism relates exclusively to growth
forms. The question is : do bacteria have only one growth form or are there species with more than
one growth form ? The monomorphist considers any shape or appearance that differs within a
species to be an abnormality, degeneration, involution form or teratology, or a mutant, i.e. a
completely new formation that is more or less consistently inherited. The pleomorphist, on the other
hand, admits the possibility that a clearly defined species - if we ignore all extreme pleomorphists -
can have more than one growth form or appearance, that the growth form or appearance of a
species can be changed. The expression

Growth form is contrasted with development form. In pleomorphism, development forms are not
generally mentioned, and if, as in ZOPF (1883), a developmental idea is linked to it, this can only be
understood in the sense of ontogeny, that is, that a bacterial individual can go through more than
one growth form in the course of its life. Any representation in the sense of cyclogeny is not linked
to it. At this point , only NYBERG (1912 ) should be mentioned, who, on the basis of the determination
of two different colony forms that he observed in several Pseudomonas species,

E ÿ d e¡rLl'e i ÿ, bacterial cyologellia. 2

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18 Historical information about the growth forms.

respected, took the view that these two growth forms of the colonies, which he calls a- and ß-colonies
and which, from the standpoint of cyclogeny, comprise all possible stages, represent an original,
simple form of alternation of generations (page 33). The fact that this is to be understood purely
theoretically is demonstrated by the fact that in the same place (page 17) the author also considers
the possibility of interpretation through an expanded concept of mutation. Finally, FUHRMANN
(1907) comes closest to the cyclogenetic view, based on studies on Eucystia cerevisiae (Fuhrm.
1906) and some related forms, who stresses with certainty that "there can no longer be any talk of
an immutability of the bacteria in the sense of COHN ". PRELL (1910, cf. below page 35) should also
be mentioned here.

In the following, some of the most important works will be examined for their position on this
question. HALLIER (1867), JOH. LÜDERS (1866, 1867), POLOTEBNOW (1872), BILLROTH (1874),
NÄGEL ÿ (1877) and others will be passed over here.

The founder of bacterial systematics, COHN (1872), is often considered a monomorphist.

The fact that he is not one is proven by the fact that he was the first to identify gonidia in bacteria,
namely in Grenothrix Cohn in 1870. It is not clear from the publication whether COHN introduced the
term gonidia completely anew or whether he wanted to adopt the term conidia directly from mycology,
but it can be assumed that COHN created a name with the same name based on fungal science,
especially since it is biologically and morphologically identical.

LANKESTER (1873) occasionally cleverly avoids the nomenclatural error in the description
of his Bacterium rubescens Lank. 1873 by describing the various growth forms and appearances of
the bacterial individual, which he calls "plastid", in very general terms, according to their shape:
spherical, biscuit-shaped or bacteroid, thread-like, needle-like, bacillary, serpentine, spiral-shaped
and snail-shaped; according to their structure: naked or slimy, homogeneous or granular; according
to the form of aggregation: linear, star-shaped, spherical, massive, tree-shaped, chain-shaped, net-
shaped and cubic-shaped. LANKESTER does not go beyond a purely morphological nomenclature,
but nevertheless ascribes a number of different growth forms to the red organism described.

As far as LISTER (1873) is concerned in assessing the various growth forms of his Rad.
lactis List. 187 3 , the objections of LÖFFLER (1887, p. 137) against the applied research methods
may be partly justified, but there are still noteworthy morphological findings among them which show
that L., even if he perhaps
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Historical information about the growth forms. 19

did not work with sterile media, had a keen, comparative morphological power of observation ; thus,
figures a, b, d, e, h on plate 21, e, f, k on plate 20 can easily represent different cyclostadies of an
organism.

In 1876 (Beitr. Biol. Pfl. II [1877] 1876 Heft 2 pg. 249) COHN made the significant discovery of
the persistent form of bacteria, for which he mistakenly adopted the term spore from mycology,
although in reality it corresponded more closely to BREFELDS's term oidia in mycology. He determined
its germinability and its extraordinary resistance to very high temperatures in Bac. subtüis Cohn.

In his excellent study of the anthrax pathogen, KOCH (1876) also followed the development of
the bacteria from the spore; when asked about the possible reasons for the long survival of this
pathogen, he theoretically puts forward two possibilities: "1. by alternation of generations, 2. by
spores", and decides on the second, although he apparently did not consider the combination of the
two as a possibility. I have already mentioned the pleomorphist CIENKOWSKI (1877) . The work of
WoLFF (1880) in the same direction on micrococci appears doubtful. WERNICH (1880) observed a
certain instability within small limits in various liquids and resulted in a "labile form stability".
PRAZMOWSKI (1880) and BREFELD (1881) found a great variety of forms for Bac. suhtilis, and
PRAZMOWSKI also for the genus Mierospira (M, rugula) . KOCH (1877 ) also confirmed earlier
findings of COHN on Spirillum (S. undula).

GAFFKY (1881 ) , as a pronounced monomorphist, declared that no one could claim to have seen that a spirilla or a spirochaete arose

from a bacilli-like form or that a bacillus arose from a micrococcus. A wide range of forms as well as gonidia

KURTH (1883) demonstrated the formation of spherical bacteria from the chicken stomach for Bad
Zopfi Kurth in 1883 , but objected to the false conclusion of BILLROTH, NAEGELI and others that all
possible species and genera could merge into one another and that there were only a few species of
bacteria. This was carried out even more clearly by ZOPF (1883), who definitely occupies an
outstanding position in the development of bacteriology. He only made the mistake of considering all
such forms as only ontogenetic forms of development from the observation that the spherical
individuals (mychites!) frequently appear as ontogenetic development forms of morphologically higher
bacteria. The consequence was that he completely dropped the genus Micrococcus Cohn etc. and
used the name as a morphological name. COHN had, however, already earlier referred to these
spherical forms of higher bacteria as gonidia.

2·
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20 Historical information about the growth forms.

or at least in part, since he did not know the basite and could therefore include it in this term. BÜCHNER also

demonstrated various growth forms of a species in 188 2 .

HUEPPE (1884, 1886) firmly supports the view that there are relatively uniform
species as well as pleomorphic species; however, he also clearly recognizes that this
does not refute COHN's classification, but merely a significant addition and extension.
In an excellent overview of the state of knowledge at the time (The Forms of Bacteria),
HUEPPE (1886) repeats his view, expressed in 1884 , that the higher the cleavage
plants are in the system, the more likely it is that the phylogenetically lower form
species can also appear as mere growth forms in their ontogeny. As I have already
mentioned, this clearly states that any cyclo-genetic idea was completely alien to him.
It is very important for systematics that he was already firmly opposed to the use of
colonial forms, especially the Zoogioea, for the concept of genus.

DE BARY (1884, 1885 ) distinguished between endospores and arthrospores, which

completely coincide with the older term gonidia.
GRUBER (1885) considers all bacteria to be morphological units composed of
isodiametric cells ; in the case of the cholera pathogen he saw individuals consisting
of 2, 3 and 4 such units.
BUCHNER (1885) and GRUBER (1885) assessed the extraordinarily varied
body shape of the cholera pathogen as pathological growth forms, but without death,
while FINKLER and PRIOR (1885) in their excellent and detailed study on Micro-spira
comma Schrot. 1886 (= Vibrio cholerae) and Microsp. proteus Buchn. 1885 rejected
this. The latter recognized the germination capacity of the gonidia (which they called
spores) and observed the fact, important for the cyclogenetic concept, that the
microspires that develop from gonidia after 12-24 hours are extraordinarily small and
only reach their normal size in the course of further growth. They have therefore
already observed the prophytite stage. BUCHNER (1885) interprets the weak or
lacking stainability as evidence of the pathological conditions of the spindle, monad
and bottle forms etc., for example the neck of the bottle has not yet degenerated, as
it still stains well; he accordingly interprets the "polar grains" (= telotropho-somes) as
intact end rows of the "four-cell"

Microspira. BUCHNER already had a certain idea that many bacteria were not
cytologically units, although without cytological and caryological foundations. He also
noticed the differences between pleotrophites and atrophites. Later (pages 121—129)
BUCHNER recognized the nomenclatural error of the pleomorphists and opposed it.
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Historical information about the growth forms. 21

that "the designation of the microscopic growth form of the cleft pike is used as equivalent to the
designation of the biological species" [the last term must be replaced by genus]. He also notes the
"long threads" (i.e. ascites) and "spherical forms" (i.e. gonidia) under certain conditions in the
typhus pathogen and other forms and recommends "using German names for the growth forms
and reserving the Latin ones for the genus designation (he writes species designation)." The fact
that all these considerations are to be understood purely in terms of form and not in any way in a
comparative morphological way is already demonstrated by the nomenclature used; he uses the
following terms for growth forms: spherical, oval, short rods, long rods, thread-like, half-screw,
short screw, long screw, spindle-like, oval rods, club-like, for growth associations: double sphere,
spherical row, grape-like, double rods, articulated thread, tetrad-like, cube-like. Compared to
LANKESTER, this grouping of growth forms represents progress, and these terms have largely
been retained throughout the literature up to the present day. The fact that the sentence he
formulated on page 125: "The theory of monomorphism of the split fungi held by many
bacteriologists to date must be definitively abandoned as an outdated point of view" is being fought
against even more than ever after 30 years can be explained by the extreme development of
monomorphism.

As extraordinarily valuable as LÖFFLER's work is for the medical and practical scientist, the consequences

of a pronounced monomorphistic view cannot be overlooked. For example, he speaks (1887) of a cliff-edge
similarity in form between the gonidia of Crenothrix Cohn and bacteria, which can only be understood by considering
Crenothrix to be a "colorless alga" and rejecting the gonidia formation of bacteria.

According to GASPERINI (1885, 1887), involutionary forms are signs of the tendency to
assume the characteristics of higher species; in his opinion, they are not games of nature, but
rather indications of a return of original and therefore higher types in the service of the continuation
of the species. The author is therefore of the opinion that bacteria are degenerate mycetes.

FERRAN (Z. klin. Med. 9, 1885) already regards the involution forms of the cholera
pathogen as fructification organs , but concludes that it is therefore a P eronospora species.

NEISSER (ÿ. H. 1888) interprets the trophosomes in xerosis bacilli [Corynohacticerium

xerosis (Neisser et Kuschbert 1883)] etc. as endospores and the clubs as gonidia.

Significant experimental findings, which were later used for mutation-theoretical

purposes, were initially interpreted by WASSERZUG
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22 Historical information about the growth fornias.

(1888) still completely in the correct sense, i.e. pleomorphic. He found, using Bac.
prodigiosus, Bac. violaceus, a water bacterium, etc., that these organisms form
rods of very different sizes after 3-5 days in broth slightly acidified with tartaric
acid, while "micrococci" (i.e. mychites) appear in slightly alkaline broth. He
therefore clearly recognized that different growth forms can be promoted by
external influences (i.e. phytite, ascite, gonidia). Also very important is the
observation that the long threads (i.e. ascite) in the acidified broth form balls again
(i.e. gonidia) after 2 or 3 days, and that at the same time the soil gradually
becomes slightly alkaline. In a publication not long after, WASSERZUG (1888)
showed that similar results to those achieved with acid can be achieved by
applying heat (e.g. 50
0
for 5 minutes)
and that combined use produces an increase.
He even states that this resulted in a permanent variation in the form of a water
bacillus. He artificially produced the asporogenic form of the anthrax pathogen on
an acidic medium. In this work, WASSERZUG veers noticeably in the direction of
mutation theory; he believes that once a form has been acquired in a medium, it
begins to fix itself into a permanent form; if the medium is then modified, the form
does not vary or varies very little; and page 157: "In reality we are going to
intervene in the law of heredity". WASSERZUG has thus actually bred and
cultivated various cyclostadia of the same species, with a clear awareness of the
unity of both. WEIBEL (1888) connects the involution forms in the case of the
cholera pathogen in part with the spores and believes that in those "which in their
appearance are reminiscent of spores, the products of a failed fructification,
as it were dead spores" (page 291). He speaks of involution forms as deformities
and of degenerate dead products.

BUCHNER (1888) opposed GAFFKY's assumption of genuine endogenous

spores in the typhus pathogen. He cultivated long rods (i.e. ascites) on sour
potatoes and believed that the polar grains (i.e. telotrophosomes) were removed
from their polar position by colorless gaps (i.e. atrophic areas) and were then no
longer located at a point (here, ascotrophosomes). As a result, he completely
denied the ability of polar grains to germinate and declared them to be a
degenerative phenomenon with reduced resistance. He declared the characteristics
of genuine spores to be 1. resistance to the penetration of dyes, 2. resistance to
drying out, 3. germinability.

While DOWNES and BLUNT (1877) claimed a damaging, sometimes even

fatal influence of light and especially direct sunlight, RAUM (1889) can , on the
basis of a detailed literary
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Historical information about the growth forms. 23

The only thing that can be maintained by a rare study is that light has a damaging
effect on some pathogenic bacteria. The extensive literature can be found in RAUM
ZU . However, it is clear from Chapter VI on photomorphosis that there is no
damaging effect of sunlight at all.

In order to pave the way for his interpretation of the sulphur bacteria,
WINOGRADSKY (1889) had to take a stand on the basis of monomorphism. For
this purpose, a controversy against WASSER-ZUG and LANKESTER was
necessary. LANKESTER (1873) and WARNING (1876) had characterized the
Begiatoa roseo-persicina (Lank. 1873) as an extraordinary polymorphic
phenomenon. W. now believes that they had not followed the course of development
directly, at least with very few exceptions. W. could not confirm the pleomorphism
that ZOPF found in Cladothrix either , with the exception of the formation of gonidia
(he calls them athrospores). Not suspecting that the accusation of lack of consistent
and systematic observation would have to be directed at him, erpag says. 263:
"We have demonstrated that all new pleomorphic theory is based on observational
errors." As already mentioned, WINOGRADSKY was the first to divert all
morphological or biological deviations from the species type onto mutational
theoretical paths and thus gave the development of bacteriology a direction which
made the possibility of a healthy development of pleomorphism impossible.

In fact, pleomorphism, which had been established by outstanding researchers such as CIENKOWSKI,
ZOPF, METSCHNIKOFF, WASSERZUG, PRAZMOWSKI, HUEPPE, BREFELD and others and which
could have given rise to good hopes, was nipped in the bud. The study of sulphur bacteria by
WINOGRADSKY (1888) should be judged entirely from this perspective. From the observation of further
development and division in slide cultures, he concluded that there was constancy in the monomorphic
sense, although it was already striking that he was unable to obtain pure cultures. The cyclogenetic
approach clearly shows that such conclusions are unjustified. If we now consider all the morphologically
completely vague genera of red sulphur bacteria established by W. in comparison with the variety of
forms of CMamydothrix or of Syncrotis buccalis (Rob.), it seems quite possible that all these forms can
be cyclogenetic manifestations of a single species.

An analysis is possible on the basis of the excellent illustrations, which clearly

show the morphological composition and thus also the main cyclostadia.

Thus, Thiopolycocms with mychite, dimychite and didimychite, i.e. belonging

to the plastite and phytite stage, is found in a culture of
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24 Historical information about the growth forms.

Lamprocystis, i.e. synascite. Gonidia formation would have occurred here, and other stages are found
at the same time, which may have developed from this.

From these lower stages, one can trace two series, one ascending to ascite,
the other to synascite, with all possible transitions in between. All findings that cannot
be unusual in a synascot organism.

Thus, in the ascite series, TMopolycoccus is followed by Amoebobacter (A.

bacillosus) as dimychite and didimychite and Thiòdictyon as didimychite and
syndimychite.
It is noteworthy that the three Amoebobacter species represent three completely
different cyclostadia; A. granula probably corresponds to the gonidia or basite, A.
bacillosus to the phytite, and A. roseus corresponds to a short synascite.

In the synascite series, TMopolycoccus is followed by Thiopedia , which

presents a colourful picture morphologically; it apparently contains dimychites and
syntact didimychites, divided in 2 directions of space ; Thiocapsa occupies a similar
position with division in 3 directions of space. A short synascite represents Thiocystis,
as does Thiothece with zoogtfoea formation, larger synascites are found in Ghro-ma
tium, and finally long synascites are found in Ehabdochromatium and Lamprocystis ,
which are curved like spirillae in Thiospirillum .

A decision on this can naturally only be made on the basis of detailed breeding
experiments, but one thing can already be said: the probability is high that all of these
genera will emerge as developmental stages of one or only a very few species.

WINOGRADSKY's importance lies in the physiological field; he is particularly

responsible for proving that nitrite bacteria (1891) are able to feed without organic
substances .
That a comparative morphological study can achieve brilliant results even
without pure cultures is demonstrated by METSCHNIKOFF (1889), who, as an
outstanding morphologist, resolutely relied on pleomorphism in his description of the
spirobacillus (Spirob. Cienkowslcii Metschn. 1889) , which partly contained
bacteriopurpurin , from Daphnia magna on seaweed from Odessa; the illustrations
show a continuous whole up to the formation of gonidia (Fig. 14). Any objections to
the fact that certain stages were colourless are sufficiently refuted by the proof of
the dependence of the colouring matter on certain cyclostadia. In a later work
METSCHNIKOFF (1889, p. 265—267) turns particularly against WINOGRADSKY;
clearly keeping the goals in mind, he emphasises that the main thing is not to know
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Historical information about the growth forms. 25

whether the theories of earlier researchers are right or wrong, but to determine whether
there are species among the bacilli that can sometimes take the form of a bacillus,
sometimes that of a spirilla, and sometimes that of a coccus. He shows this using the
example of the cholera pathogen, B. prodigiosus, the pathogen of chicken cholera, etc.
He refutes WINOGRODSKY's postulate of a continuous series of observations on living
organisms to prove pleomorphism by referring to embryology and to studies on the
development of many parasites, which can never be followed on living objects.

BEHRING (1889) shows that the asporogenic form of the anthrax pathogen forms
"involution forms" very quickly at 37 degrees. He interprets the partial or total lack of spore
formation as a partial manifestation of degenerative processes. An interesting reference is the
observation of a weakened anthrax pathogen that no longer kills mice, was bred from a
completely normal form and cannot be distinguished from a normal one either in terms of
spore formation or in any other way.

BILLET (1890) opposes the idea of considering the short rods as bacteria and the
long ones as bacillus and cites a large number of species in which both forms of growth
occur.
For Sphaerotilus (Cladothrix) äichotoma (Cohn) and some other species he sets up
several stages of development. He groups the various forms of development into the
following stages: the 1st filamentous stage (état filamenteux); 2nd forked stage (état
bicladé); 3rd branching stage (état polycladé); 4th dissociated stage (état dissociacié); 5th
tangled stage (état enchevêtré) and 6th zoogloea stage (état zoogléique). Most of these
belong to the ascite stage, such as 1, 2, 3, 5; the dissociated stage (4. ) partly coincides
with the phenomenon that BRE -FELD 1889 called oidia formation (in fungi) and with the
catascite stage of the present study; However, BILLET also classifies completely different
morphological elements in this stage, such as synascite, thecite, etc. Finally, the zoogloea
stage (6.) corresponds to the appearance of the ability to form mucus (i.e. a formant).
BILLET understands all of these developmental stages only from the point of view of
ontogeny; cyclo-genetic concepts are missing.

VAN TIEGHEM calls the structures previously considered as gonidia

(1891) Cysts.
FRENZËL (1892) provides purely objective information of the greatest interest on a
green bacillus from Argentine tadpoles. The detailed description and the illustrations clearly
show that it belongs to the genus Schauäinnum Enderl., to which S. Bütsehlii (Schaud.
1902) belongs as the type and which I have designated in Chapter VII as 8. ranicolum
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26 Historical information about the growth forms.

n. sp.; it is a highly organized synascite with the highest fructification, which I have called endothecite.

A very important moment in the development of bacteriological knowledge is the introduction

of the term "secondary colony" by GÜNTHER ( 1895).

GOTSCHLICH and WEIGANG (1895) examine the fact first demonstrated by GRUBER and
WIENER (Arch. f. Hyg. 15) that cholera cultures "lose their virulence with astonishing speed" as
they age. FLÜGGE had already argued that the assumption of a qualitative change was not
necessary at all, but rather that the quantity of inoculated bacilli was lower. The authors believe that,
on the basis of their detailed investigations, they can assume that the number of living individuals in
a colony decreases rapidly with increasing age. The number of individuals capable of germinating
on solid nutrient medium from the material of a continuous culture is 7.43% after 44 hours of culture
according to Table I, 11.7% according to Tables V, VI and VII, and 0.8% after 68 hours of culture
according to Table I, and 2.76% according to Tables V, VI and VII. At the edge of the colony, the
"death" is less faster than in the middle, that this is only an apparent death, can be seen from the
section on Gonite (V ÿ 1).

ZETTNOW (1896) correctly observed the disintegration of Spirillum undula into spheres (i.e.
gonidia), but explained these as phenomena of death ; later (1897) he came to the same conclusion
on the basis of further test material, with particular emphasis on the fact that these are not
permanent forms. He interpreted the formation of ascogonidia in Spirillum undula minus and
Microspira rugula , which are forced out of the bacterial body, as forking, and those in Sp. serpens
as buds.

GRIXONI (1896) cultivated a polymorphic bacillus from pus, which

all forms of wax from coccus to long threads.
A very advantageous view of the differences in the growth forms of a species in a
pleomorphic sense can be found in LAFAR (1897, p. 36), which has fortunately found greater
acceptance among fermentation physiologists and has prepared the ground for the cyclo-genetic
view in a favorable way. Above all, this discipline has clearly recognized that external influences
have formative powers that can influence the life of these beings, namely through the temperature
and composition of the nutrient medium. However, here too, the more strongly deviating changes
in form are viewed as involution forms, i.e. as signs of death.

WAGNER (1897) observed in older colonies of B. coli and

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Historical information about the Wuehsformen. 27

Typhoid pathogens produce numerous "tubes and threads" (i.e. ascites)

and also note their disintegration into balls (catabiosis).
At the same time, GRASSBERGER (1898 ) noted the formation of "false
threads" (i.e. ascites), in which he sometimes claimed to have observed a bifurcation.
An extremely important work, not so much for the assessment of the
diphtheria question but especially for the cyclogenetic view, appeared in
1898 by HEWLETT and KNIGHT. They established that the typical growth
form of the diphtheria pathogen almost always occurs occasionally from time
to time in Gorynóbacterium pseudodiphtheriticum (Hoffm.-Wellenh. 1887),
and that pseudo diphtheria forms also occasionally occur in a single culture
of the true diphtheria pathogen. From a broth culture of the true diphtheria
pathogen heated to 450 °C for 17 hours , the authors grew a culture that had
completely assumed the type of a pseudo diphtheria strain ; polar body
coloration was absent, the rods were short and plump, and the animal
experiment with guinea pigs was negative. This would therefore be the
cultivation of the phytite of the diphtheria pathogen. Furthermore, they have
grown a typical diphtheria strain from a typical non-acid producing
pseudodiphtheria strain by frequent passages through serum culture media
and animal passages. The authors conclude that pseudodiphtheria forms are
sometimes modified forms of the true diphtheria pathogen, although not
always, since more than one species of the same morphology can exist.

In the "Lectures on Bacteria", in which A. FISCHER (1897) also discusses

his own earlier findings in detail, such as the permeability of salt solutions
through the bacterial membrane and the resulting strong plasmolysis, an
overview of the most important aspects of the state of bacteriology is given.
He himself takes the view of monomorphism and explains all the diverse
phenomena, including the gonidia, as random occurrences and as forms of
involution. The asporogenic form of the anthrax pathogen is regarded as a
general degeneration. According to their way of life, bacteria are divided into
3 groups : 1. Prototrophic bacteria, do not require organic food; the nitrate
bacteria are able to process nitrogen in its elementary form; the sulphur
and iron bacteria decompose organic compounds to obtain vital energy.

2. Metatrotrophic bacteria, with very different organic

Food.
3. Paratrophic bacteria, the parasites.
It is of great importance that FISCHER was the first to oppose the view of chromatin
granules as cell nuclei on the grounds that they represent reserve substances.
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28 Historical information about the growth forms.

The excellent work of MIGULA (1897) offers a detailed overview of bacteriology as a whole.
He considers all differences within cultures to be a simple development of the bacterial individual
and considers it incorrect to describe this as pleomorphism.

He thus stands on monomorphistic ground. At first, only the idea of regular states in cultures of
Bac. suUilis (page 233), "which, independent of the individuality, always recur under normal
conditions at a certain stage of development and represent a form in the life of the species", can at
least be interpreted as a certain idea of cyclogenetic processes. However, since (page 228) these
different forms of development are compared with the forms of development of the sun rose -
intended as an example - this may have been a mistake. This is stated even more clearly in the
sentence (page 228): "If the Bacillus subtilis, starting from the spore, first forms immobile germ
rods, then mobile swarmers, which after a while become immobile again, grow into long threads in
which spores are formed, and if the same stages develop again and again in the same order in
countless generations, then this is nothing other than the different stages of development - phases
of life of a form or species, but never pleomorphism." This actually makes it clear that he, considering
himself a monomorphist, is actually a pleomorphist, because most pleomorphists have not
understood it to mean anything else (apart from NAEGELI, BILLROTH and similar extreme
representatives, of course), and such a view has been fought against by the monomorphists. Any
cyclogenetic idea is thus ruled out.

MIGULA has made a great contribution to clarifying the term G o ni die.

He found that the gonidia had a much greater resistance to higher temperatures and drying out
than the vegetative states. A morphological fixation of this term was naturally not possible for him
in view of the lack of a comparable morphological basis. Likewise, Migula did not recognize the
general significance of the gonidia for bacteria, but only for the highest forms (chlamydobacteria).
Migula also already held the view that bacteria had not yet differentiated from animals and plants.

GÜNTHER (1898) considers the involution phenomena to be signs of death; according to

him, the cells swell up, the plasma is permeated by vacuoles, and malformations and curlicues
form.
MEYER (1899 ) transfers the BREFELD term Oidiu m (1899), which included permanent form-like structures and

for short sections of fungal

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Historical information about the growth forms. 29

threads (also called smut spores), in a completely different sense to bacteria; he thus describes as
oidia all dimychites, didimychites, short ascites, short synascites, sporites etc., in short everything
that has a shorter or longer shape that is remotely similar to an egg; this term has neither
morphological nor biological meaning in this form. MEYER pursues a very good idea, which could
easily have developed into a morphological term, by using the smallest cells of the Bacteriaceae,
which he describes as usually twice as long as the spores, as a unit of measurement for cell
length; cells twice as long he calls two-long, three-times as long three-long etc. This is the first hint
for the morphological terms dimychite, didimychite etc. formulated in this treatise, but they are only
formulated in terms of shape. MEYER also explains multinucleate cells in this sense. Threads with
spores or spore primordia are called sporangia, also a term used in mycology for something
morphologically quite different.

MACCHIATI (1899) describes two "Streptococcus" species that grow in the form of cocci
and rods, depending on the nutrient medium; he therefore believes that the habitus of the colony
must be regarded as a reliable identifying feature of the species. If the observation is correct, it
cannot of course be the genus Streptococcus .

An extraordinarily significant work is that of MATZU-SCHITA (1900), although he does not

make any observations on the results. He simply regards all the deviations that occur as
transformations and degenerations. He uses agar medium with the addition of 5-10% common
salt to culture numerous types of bacteria.

The results are surprising. There are such strong deviations in shape. Micrococcus rulefaciens (=
tardus Mig.) and N. flavus liquefaciens take on the shape of rods; Bact. Irunneum without Na Cl
short or long rods, with Na Cl spheres ; Bac. lactis innocuus without Na Cl fine short rods with Na
Cl spheres and rods. Bac. aeropMlus, liquefaciens, megatherium-simulans, implexus, proteus
Zopfii tend to form conspicuous spindle or spirilla shapes in addition to spherical formation. Bac.
typhi forms long threads, Bac. prodigiosus very diverse shapes. The diphtheriaerger forms strong
clubs. Bac. pestis, Bac. acidi ludici (Hüppe), Vibrio cholerae and Bac. pyo-cyaneus form large
spheres in addition to other shapes. These large spherical shapes had already been observed
earlier in Bac. pestis by HANKIN and LEUMANN (C. ÿ. 1,12, pag. 438).

SKSCHIVAN (1900 ) observed extraordinarily long threads, some of which even had
branches, in old cultures of Bac. pestis; he used the name bacterial mycelium for these and, since
they can transform into normal forms in a short time, he opposed the establishment of
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30 Historical information about the growth forms.

as degenerative phenomena. In contrast, GALLI-VALERIO (1900) describes similar forms of the same
organism as involutional forms. GAMALEJA (1900) opposes the earlier explanation of involutional
forms as degenerative phenomena and takes the view that this is a heteromorphism linked to the
chemical composition of the nutrient substrate ; he points out the possibility that the forms
obtained in this way could have evolutionary, systematic and phylogenetic significance.

MAAS SEN (1904) points out the importance of involutional forms (teratological forms) for
diagnostic purposes .
According to A. MEYER (C. ÿ. I, 31, 1902) "we must regard bacterial species of which we
do not know of spore formation, such as diphtheria bacteria, as incomplete species in comparison
with those whose complete development process is known, such as Bac. subtilis, whose juvenile
forms continue to multiply without going through their complete development process."
Cyclogeny teaches us, however, that the typical diphtheria bacilli are by no means juvenile
forms, and that the sporite is only a manifestation of the phytite stage and only occurs in a very
limited group of organisms.

The term "secondary colony" introduced by GÜNTHER (1895)

is unnecessarily split into two terms by PREISZ (1904) :
1. Secondary colony: small nodules that usually appear after a few days, usually hemispherical
and raised, give the impression "as if colonies of some foreign germs had developed on
the original cultivated lawn".

2. tertiary colony: further sculptural differences, usually with a raised center, a ring-shaped
depression around it, a denser wall on the outside ; sometimes warts or nodules after
weeks or months.
The author is of the opinion that secondary colonies are formed from spores. He considers
the abnormally formed forms in 2-3 day colonies to be degenerated bacteria that are doomed to
extinction. In anthrax colonies he observes that a number of cells do not reach sporulation, but
degenerate and decay. He comments on this (page 659): "But why it is that of two cells that are
next to each other under apparently similar conditions, one forms spores and the other does not,
is and remains a mystery that we cloak in the concept of individuality" (so this is a case of gonidia
formation of the asporogenic form of the anthrax pathogen). He considers such colonies that do
not form spores to be doomed to extinction (page 660).

ABOTT and GILDERSLEEVE (1904) wrongly object to the use of fork

formation in the diphtheria pathogen for systemic
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Historical information about the growth forms. 31

matic purposes, which has even been used for a separation of these and other forms
from the bacteria to the hyphomycetes (ÿ. B. LEHMANN and NEUMANN 1904); however,
this phenomenon is wrongly explained as involution or degeneration.

ALM QUI ST (1904) points to long mycelium-like threads, called "myceloids",

which occur in the pathogens that cause cholera and typhoid fever. He believes that
these have been ignored until now (they were already demonstrated by FINKLER and
PRIOR in 1885). In contaminated soil with a little broth or in fertilized soil without broth,
the author observes ball formations which he compares to the conidia of fungi and does
not consider them to be resting spores. "Visibly empty balls" etc. which he observes in
the pathogens that cause cholera, he interprets as a sign of degeneration. WALKER and
MURRAY (1904) grew similar long threads in Bacillus coli, the
pathogens that cause typhoid fever and cholera, on agar media which had been
mixed with methylene violet, methylene blue or fuchsin. PERRI N (1906) regards the
thinner specimens (i.e. ascites) of Spirochaeta (S. Balbiani) as male gametes, and the
thicker ones (i.e. synascites) as female gametes. Since he classifies this form as a
protozoa (as trypanosoma), he falsely applies the term cysts to structures of the
Spirochaeta (in reality they are arthrothecites).

In the Spirochaeta gallinarum Marchoux et Salimbeni 1903, VON PROWAZEK

(1906) interprets the forks observed as longitudinal division (this is to be understood as
bifurcation or branching). The spherical structures, which PERRIN interprets as cysts,
are called protoplasm buttons by the author.
MÜHLENS and HARTMANN (1906) interpret the aggregation of individuals of
Spirochaeta in Spirochaeta dentium , as assumed by PERRIN , following the example of
PROWAZEK with the help of longitudinal division; he also takes the erroneous view that
the spirochetes are protozoa and not bacteria.

HEI M (1906) also regards the round globules (i.e. gonidia) in the cholera pathogen
etc. as degenerative or dying forms; he rejects a relationship to arthrospores or persistent
forms.
Huss (1907) developed morphological comparisons using Bac. esterijicans Massen
1899 (grown from butter) and Bactrinium trifolii Huss 1907. In addition to sporite formation
(Fig. 17), the former form also appears to exhibit gonidia formation (Fig. 13) and
arthrocystite formation (Fig. 14); the figures show several cyclostadia and a fairly strong
pleomorphism in this species.

The first to introduce the concept of mutation in the sense of DE VRIES (1901 ) into bacteriology
were MASSINI (1907 ) and on the basis of this
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32 Historical information about the growth forms.

Work by MASSiNis, later by NEISSER (1906*)); it concerns the ability of the Bacillus coli mutabile
to excrete no fuchsin from endoagar in the primary colonies (whitish), but fuchsin in the secondary
colonies (red) ; in further breeding, the secondary colonies always give red colonies, the former
always white and red.

RUZICKA (1907) disputes the assumption that the so-called "involution forms" are death forms
and calls them hypertrophic forms and the phenomenon, since he associates them with an excessive
filling with chromatin substance,
, hyperchromasia.

In the case of purple bacteria, MOLISCH (1907) represents the same systematic views as
WINOGRADSKY for the sulphur bacteria; for these too, detailed investigations on the basis of cyclogeny
are necessary.

FUHRMANN (1907) came very close to recognizing the cyclode using Pseudomonas
cerevisiae Fuhrm. 1906 and Ps. myxogenes Fuhrm., which may be classified as belonging to the
genus Eucystia . He speaks of development cycles. The illustrations show that Fuhrmann's "granules
resistant to drought etc." (Fig. 5) are to be interpreted as gonidia, the "rods" shown in Fig. 1 as the
phytite stage, and the "elongated and partially thread-formed rods in Fig. 2" as ascites. The
"granules and spheres" in Fig. 3 and 4 are trophosomes, the "threads" in Fig. 3 are mycascites.
The "club-shaped swollen rods" in Fig. 6 and 9 and the end bulbs in Fig. 7 and 10 are cystites. The
terms all have only descriptive meaning and have no morphological fixation.

Much's granules (1907-08) are partly trophosomes and trophosomelles, partly gonidia and
cystites ; Much's investigations refer to the tuberculosis pathogen, which usually appears in sputum
only as a uniform long rod, and in which the formation of "granules" is not so frequently observed
without the use of special stains.

The work of WE IS ÿ (Vienna clinical research group) is moving in the same direction .
Wochenschr. 1899 No. 47), WIRTHS (Brauers Beitr. Bd. 11, Heft 1 and Münch, med. Wochenschr.
1908, V. 55, p. 1687—1690), WEGELIN (Korresp.-Bl. f. Schweizer Ärzte No. 29, 1910), ADAM, J.
(Inaug.-Diss. Leipzig 1910) and especially the work by KNOLL (1912) , which is provided with very
beautiful illustrations, based on very instructive staining of the tuberculosis pathogen using fuchsin-
methylene violet-resorcin.

The same morphological conditions as in Bac. Bütschlii Schaud. are found in the colon of
Bufo vulgaris by DOBELL (1908).

*) This work only appeared earlier.

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Historical information about the growth forms. 33

Bac. flexilis Dob. (both belong to the genus Schavdinnum Enderl .).

Like Prowazek, Mühlens and Hartmann, FANTHAM (1908) also

the bifurcation phenomena in Spirochaeta occur as longitudinal division.
KRYSTALOWICZ and SIEDECKI (1909 ) also classify the spirochetes as
protozoa, namely as mastigophores, and establish a special group on these, which
they call Spiroflagellata .
TADDEI (1909) observed strong vesicular swellings in streptococci , interpreted
them as atavistic characteristics, as the last effort of the organism for its survival,
and brought them into phyletic relationships with similar phenomena in the Nostococci
(algae); he opposed VINCENT 's (1902) view that similar changes in streptococci
were related to the reaction of the culture medium.

MENCL (1910) observed in Sareina lutea a rapid "nuclear division" which the
protoplasm cannot follow ; he interpreted this as syncytia formation.
The iron and manganese content of the iron bacteria treats
MOLISCH (1910).
Biological and morphological changes in the cholera
pathogen in the human body is pointed out by ZLATOGOROFF (1911) .
In a varied and excellent summary, MEYER (1912) for the first time puts
forward the view that the sulphur and purple bacteria must be classified in the system
according to morphological principles. The earlier views on oidia, spores, etc. have
remained the same.

In contrast to KOCH and PETTENKOFER, EMMERICH and JUSBASCHIAN

(1912) support the view that the dejection cholera bacteria are relatively harmless
and have a weak toxic effect ; this is justified by the fact that when fed with nitrate-
rich food, the cholera pathogen produces large amounts of nitrous acid, thus severely
impairing its ability to reproduce and form nitrite; only after it has grown for a few
days on contaminated soil is it said to become stronger again.

HOROWITZ (1911) points out that in the case of symbiosis of the cholera
pathogen with a yellow Sareina, the vitality, virulence and agglutinability of the
former would be increased.
LUMBROSO and GERINI (1911) draw attention to the great variability and
instability of the morphological character of the cholera pathogen in bacilli carriers .

As interesting and useful for other purposes as the arrangement of

JENSEN (1909 ) may be on the establishment of a "natural bacterial system", nothing is further from a
natural bacterial system than
End of ricin, bacterial cyclogeny. 3
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34 Historical information about the growth forms.

this attempt. A natural system can only be based on a comparative morphological basis for
all organisms and not on a completely physiological one, as is the case here. If one judges
this system from the standpoint of cyclogeny (the phenomena of the formants, which were
previously considered mutations, are particularly important here), it cannot be avoided that
the same species belongs to one genus or another, or even to a family, depending on the
physiological phenomena associated with the respective stage. The systematic results are
therefore also not in line with the actual circumstances ; for example, the genera
Micrococcus and Sorcina , which belong to the most primitive forms , are considered to
be the most derived forms.

The question of the formation of individual granules (i.e. gonidia) of the tuberculosis
pathogen is dealt with by KNOLL (1912); the fact that the tuberculosis pathogens form
such granules or break down into such granules in the lymph gland is used by him to
conclude that the lymph gland is thus a biologically valuable factor in the organism's fight
against tuberculosis.
NYBERG’s work (1912) has already been discussed in more detail on page 17.
gone.
WALTER (1912) is convinced of the constancy of the appearance of the diphtheria
rod, especially of the presence of the so-called polar bodies (i.e. ascite with telotrophosome
or telocystite) ; however, this work contains important material on species differentiation
(Bac. mono-polaris, bipolaris and multipolaris). SCHÜRMANN (1915) takes a similar
view , although the possibility of different growth forms is recognized here, especially with
regard to the polar bodies (i.e. telotrophosome or telocystite or arthrothecite).

The isolated metachromatic bodies (here, gonidia) are considered by BABES (1914 )
to be the longest-viable parts of the tubercle bacillus (this was already established by
COHN 187 0 ), and he believes that this view must also be extended to all other bodies of
the remaining bacteria (i.e. to trophosomes).

BURCK-HARDT (1914) considers flagellation to be one of the most constant

properties of bacteria .
BAIL (1915) opposed the view that the variability of the colony forms of bacteria is a
result of mutation, using cultures of the cholera pathogen. He regarded them as merely
developments of the range of variability that belong to the character of the species. He
achieved a greater range of variation, particularly after treating material from the cholera
pathogen for 8 days at 420 in saline solution (cf. halomorphoses VI ÿ ÿ 3).
Machine Translated by Google

Historical information about the growth forms.

35

SCHMITZ (1916) is against the term mutation in bacteriology, for which the necessary
preconditions have not yet been fulfilled. He calls the variations that revert at fairly fixed times
"circular variations" and those that do not revert after a certain period of time "irreversible
variations".

In some preliminary communications (1916/17) I defined all comparative morphological and

biological concepts which are discussed in detail in the present treatise.

Prell (1917) recognizes the presence of two different growth forms in Bacterium coli ; one in
the form of cocci he calls the coccocytic phase, the other in the form of rods the rhabdocytic phase. He
also makes a distinction between, pleomorphic phenomena caused by internal conditions that have little
influence on the shape of the organism (called fluctuations), and those caused by external conditions
that often have a strong influence on the structure of the organism (called modifications). He calls firmly
induced modifications that cannot be reversed in experiments transformations. Apart from the fact that
PRELL 's publication appeared after my preliminary reports, the terms used in no way coincide with
those I have put forward.

First of all, they are comparatively not morphologically fixed and biologically too broad and too limited.
Thus, above all, the external conditions are just "external conditioning causes" which give the organism
the opportunity to transform itself into another "cyclostage" or another "formant". Both the fluctuations
and the modifications

in PRELL's sense, therefore, contain both "cyclostadia" and "formants". The PRELL term transformation
would therefore, in my opinion, correspond to: cyclostadia)

Mochlosc .
Formants J The

different size of the cells of the coccocytic and rhabdo -

PRELL compares the size of the haploid and diploid nuclei in echinoderm larvae from unfertilized and
fertilized eggs. The theoretical assumption of a generational alternation (cyclomorphosis) in a
continuous alternation of these two phases is based on this assumption of a different size of nuclei or

cells in the coccocytic and rhabdocytic phases. Therefore, even if these two phases are neither
morphologically nor biologically fixed and therefore the coccocytic phase cannot be recognized as
mychite on the one hand or as basite on the other, and the rhabdocytic phase cannot be recognized as
dimychite, didimychite, syndimychite, phytite, ascite, etc.

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Machine Translated by Google

Historical information about monomorphism and pleomorphism. 37

Jacobson (1910), Klein (1912), Lehmann and Neumann (1912), Langelsheim (1913), Mandelbaum
(1912), Markow (1912), Marks (1910), Massini (1907), Metschnikoff (1894 etc.), Müller (1912),
Neißer (1906), Neumann (1897), Nyberg (1912), Ornstein (1913), Penfold (1910—14), Pringsheim
(1910—14), Revis (1910—12), Römer (1914), Rosenow (1914), Sachs-Müke (1913), Salz-mann
(1914), Seiffert (1911—12), Schierbeck (1900), Thaysen (1912), Toeniessen (1913—15), Trautmann
and Gaehtgens (1913), Zupnik (1912) and many others.

C. Historical aspects of comparative morphology

and biology.
The extraordinarily unusual phenomena of bacterial cyclogeny were naturally
incomprehensible to primitive scientific methods of investigation. A completely natural consequence
of this was the early division of research into two camps, monomorphism and pleomorphism. The
first, as the primary, is the result of the simplest observation. The astonishing thing about this is the
fact that the battle between two schools of thought lasted for decades, both of which represented
apparently diametrically opposed viewpoints and yet both of which had a claim to legitimacy in all
essential points of the observation results. However, when the pleomorphic observation material
later accumulated so that it could no longer be denied by the monomorphists, people helped
themselves by assuming abnormalities (teratology) and coined the term involution forms for such
phenomena, which has long been valid and is even widely used today. But even this soon became
insufficient for the ever-increasing abundance of pleomorphic facts. Instead of considering the
possibility of a unification of both opposites, the monomorphists attempted to divert the explanations
to an area that was later called mutation since DE VRIES (1901) . The first person, to my knowledge,
to consider the possibility of such a path is WINOGRADSKY (1889). He says 1. c. pag. 264: “De ce
que les theories de M. NAEGELI et ZOPF sont mal fondées, faut-il conclure à l'impossibilité de faire
varier les espèces bacté-riennes ; de produire lentement par des moyens appropriés des variétés
con-stantes ? Certainement non; au contraire, l'experience des jardiniers et des agriculteurs, par
exemple, avec les plants de culture, rend this éventualité très probable." Now it was assumed that
the appearance of other shapes or other properties was a process of reformation. People believed
they could eavesdrop on nature in areas where it was producing new forms, even species.
Machine Translated by Google

38 Historical information about monomorphism and pleomorphism.

which inherit their acquired characteristics more or less consistently (the mutants of the theory of
mutation). It was only through NEISSER (1906 ) and MASSINI (1907 ) that the word M-utation was
transferred to bacteriology.

Strangely enough, this assessment of pleomorphic phenomena in the morphological,

biological and physiological fields from the point of view of mutation theory has become the generally
prevailing one and still is today. However, this has completely suppressed pleomorphism and given
monomorphism absolute dominance.

The idea that different forms of the same organism could appear alternately is so far outside the
realm of possibility for the monomorphist that he prefers to consider the assumption of completely
new phenomena in the morphological, biological or physiological fields.

The difficulty in mastering the matter lies in the cyclogeny itself. People were used to
correctly assessing pleomorphic moments in the ontogeny of plants and animals; they knew that a
frog develops from a tadpole, that a fly must first gradually pass through the egg stage, the various
larval stages, and the pupal stage before it becomes an individual capable of reproducing itself. But
the idea that such a metamorphosis, as is developed in this work, could be distributed over an
enormous number of generations and an unimaginable number of individuals, and that even each
of these stages could, under certain conditions, appear in endless generations with consistent
properties, was something for which no concept or comparison was possible.

But despite all this, this one-sided position is extremely surprising. If one compares the
factual material that DE VRIES and his followers used to prove mutation in higher organisms with
that used for the mutation-theoretical assessment of the variety of phenomena in bacterial species,
one immediately realizes that there is a strong discrepancy. While the former is an extremely

laboriously acquired and only observed in individual forms

material, the latter has accumulated to an ever-increasing extent, which not only suggests a certain
regularity, but even reveals a fundamental deviation from the theory of mutation in that the "acquired
characteristics" are lost again, even if they have proven to be hereditary for many generations.

EISENBERG (1914) believes that these clearly proven facts

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Historical information about methodological errors. 39

To be able to take into account that he is of the opinion that the requirements for heritability in relation
to mutation should not be set too high: "Hereditary does not necessarily have to mean absolutely fixed.
Even a temporally limited constancy can be considered heritability."

The constant heritability of the acquired characteristic is precisely the postulate on which the
theory of mutation is based. Thus, such attempts to interpret all phenomena of cyclogeny contained
the germ of death. The fact that this line of thought was nevertheless pursued further is one of those
mysteries of the human mind that are probably rooted in the law of persistence.

Thus, a whole series of objections raised against the interpretation of these phenomena with
the help of mutation have had no influence on the development of another interpretation. Among these
objections, that of SEIFFERT (1912) occupies a prominent position, since he rightly leaves open the
theoretical possibility of mutation in bacteria - he definitely denies that exact proof of any mutation in
bacteria has been provided, and proves with the help of a very remarkable example that new properties
can appear which have no connection whatsoever with external conditions.

I will come back to this example later.

The main errors of the two directions of explanation, which stood in the way of an agreement,
can be divided into the following three groups:
1. Methodological errors.
2. Errors of reasoning.
3. NounMatoric errors.

Viewed from this perspective, the numerous controversies are not without a certain humor. After
all, all the accusations that were intended to affect the other party also had to affect one's own party
with regard to cyclogeny. An exception to this are, of course, those points in the first group that relate to
working with impure cultures or similar inaccurate methods.

1. Methodological errors.

The most important methodological error that the pleomorphists were accused of was that
they did not work with pure cultures.
This accusation has been justified in numerous cases, such as in the case of HALLIER (1867 etc.), BILLROTH (1874), Jon .
LÜDERS (1866,1867) , and others, and these have unfortunately done extraordinary POLOTEBNOW (1872 )

harm to the healthy development of pleomorphism.

In many cases, however, such an accusation was unfounded.

The main methodological error of the monomorphists was initially the lack of methodical
observation with regard to cyclogeny
Machine Translated by Google

40 Historical information on errors of conclusions.

the changes in pure cultures over time; mostly only young cultures were used, often only one-day-
old ones. A large part of all older justifications of monomorphism are based on this error.

The same methodological error still dominates today

the whole monomorphism, since it is not possible for him to differentiate between the two concepts
of culmination and cyclostage. He is just ÿ. B.
"Probazite" and "culminate in the probazite stage" are exactly the same thing; he calls both ÿ. B.
Micrococcus. At first it seems as if there is at least a certain consistency of nomenclature in this;
all mychites are called micrococci, with the exception of the gonidia. In reality, however, this error
belongs to all three groups.

The above-mentioned interpretation of the two terms as Micrococcus is only correct for the
second term, while the first can include all other genera of bacteria except Micrococcus ; to take
an example, it can be a diphtheria pathogen in the probabilistic stage.

2. Errors of conclusions.
It was quite understandable that the pleomorphists attached too much importance to their observations.

The correct observation of the variety of forms in morphological, biological and physiological respects within a

species led to the false conclusion that all these differences in bacteria could therefore have no significance. The

result was a negation of the concept of genus and even of species. As an example of a very extreme representative
of this group I cite NÄGELI (1877); According to him, "the same species takes on different morphologically and

physiologically unequal forms in the course of generations, which in the course of years and decades sometimes

cause the souring of milk, sometimes the formation of butyric acid in sauerkraut, sometimes the aging of wine,

sometimes the putrefaction of proteins, sometimes the decomposition of urea, sometimes the discoloration of starchy

foods, and sometimes produce diphtheria, sometimes typhus, sometimes recurrent fever, sometimes cholera,

sometimes intermittent fever." It is really no wonder that such an over-the-top assessment of the situation brought the

whole of pleomorphism into complete disrepute.

The practitioner, such as the doctor, the fermentation technician, etc., needed a
differentiation of the concepts; he could not do anything with a view that hindered their further
development.
As an example of an even more far-reaching false conclusion based on apparently correct
observations, I would like to mention the recent work of DUNBAR : On the question of the position
of bacteria, yeasts and moulds in the system, 1907, although it is quite apart from the
Machine Translated by Google

Historical information on errors in reasoning. 41

The question of actual pleomorphism is at stake. DÜNBAR observed that bacteria, yeasts and moulds
grew out of algal cells and concluded from this that all three were merely manifestations of the algal
cell, and thus not independent organisms, but belonged to the developmental cycle of algae. It would
be unfair to deduce from the rejection of such a conclusion, which has naturally come from various
quarters, a rejection of DUNBAR's observations, which were apparently carried out with great love
and care ; if we consider parasitism or even symbiosis when trying to assess the findings, this
observation material can still achieve outstanding significance; I am just thinking of the symbiosis
between algae and fungi in lichens ; perhaps we are dealing with primitive precursors of such a
symbiosis, in which the symbiotic association of the two heterogeneous organisms is still loose, so
that an automatic separation of the two is still possible.

In contrast to pleomorphism, the possibility of the existence of monomorphism as a whole is

based entirely on false conclusions. The line of thought of the older school can be summed up briefly
in the following words: The extraordinarily numerous generations of a young, e.g. one-day-old, pure
culture have not resulted in any changes, and therefore there are no changes. When the monomorphist
then actually encountered undeniably occurring differences, he preferred to think of impure culture or
later he helped himself with teratology (interpretation as teratological phenomena, abnormalities or
forms of involution, which were usually denied viability). As a very characteristic example, I take from
the wealth of literature on this subject the view of WINOGRADSKY (1888), who considered numerous
cyclostadia of Beggiatoa roseopersicina (Lank. 1873) to be as many genera and species, because he
observed the isomorphic arthrogonia on living pieces of such cyclostadia on the microscope slide and
made the wrong conclusion from the agreement of the division products with the original form: "The
division products are constantly the same, and a change to other of the existing forms is impossible
and does not agree with the observed phenomenon (cf. also pages 23-24 and Chapter VII).

I have already discussed that the application of the theory of mutation to bacteria is to be
attributed to a new direction of monomorphism.
Apparently the whole range of observations pertaining to this topic belongs to the group of false
conclusions. This in no way diminishes the positive value of objective findings; new, clear points of
view can only increase it.

With what caution, by the way, KOCH, who in practice stood entirely on monomorphistic
ground, knew how to avoid the difficulties,
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42 Historical information on nomenclatural errors.

proves his opinion (Mitt. K. Ges.-Amt 11881 pag. 31): "that all those bacteria which,
on the same nutrient medium and under otherwise identical conditions, retain unchanged
the properties by which they differ from one another through several re-cultivations,
are also to be regarded as different, whether one calls them species, varieties, forms
or whatever one likes." From the cyclogenetic point of view, a special emphasis must
naturally be placed on "varieties and forms".

3. Nomenclatural errors.
The error of monomorphisms that belongs to this group is already listed in the
first group.
The exact same error, but here purely nomenclatural, only in the opposite
sense, had an immeasurably severe, paralyzing effect on the development of
pleomorphism . It is the fact that systematic genus terms, especially their names,
were used to designate biomorphological terms. Thus the name Micrococcus, which
is supposed to designate the genus M., was used for the probasal stage, i.e. for a
developmental stage of an organism with a higher culmination. The first person to
make this mistake of applying the genus terms, which had been established by COHN
only a few years earlier , to developmental stages of phylogenetically higher bacteria
was CIENKOWSKI (1877). He observed growth forms in a bacterial species that
corresponded to those of the genera Micrococcus, Bacillus and Leptothrix , and named
these growth forms with the genus names mentioned. This gives the impression that
C. regards these three genera as growth forms of a single one, as VAN TIGHEM
(1878) actually did, which earned C. a sharp controversy from this side and his view
was dismissed as "pas conforme à la vérité" .

But CIENKOWSKI had not had such an interpretation in mind; he only wanted to say
what the nomenclature of cyclogeny would mean: the observed organism grows as
baesite as well as phytite and ascite.

This example clearly shows that a whole series of research results that had
been considered completely irrelevant for decades, and were even mockingly
dismissed by contemporaries, are now regaining their importance as serious factors. I
would also like to mention names such as ZOPF (1883), WASSERZUG (1888), and
METSCHNIKOFF (1889), who are among the first names in the field of the forward
development of evolutionary bacteriology.

A renewed examination of the BiLLROTH (1874 ) investigations of the Coccobacteria

séptica Billr. 1874 will probably still provide some important
Machine Translated by Google

Historical information on the question of sexuality. 48

Findings in the much-maligned work, even if the methodology and conclusions are
not flawless.

D. Historical aspects of the question of sexuality.

All the considerations discussed so far about the possibility of sexual
reproduction of bacteria have been expressed with the greatest reserve ; however,
they are also invalid because they all refer to the highest cyclostage, the ascite.
As I have stated here, it is precisely the lowest cyclostage that comes into
consideration for a sexual process.

Nevertheless, all such notes found in the literature will be reported here
using the newly proposed morphological nomenclature. All relevant citations will
be cited.

The first person to regard certain processes in bacteria as a "primitive form

of self-fertilization" is SCHAUDINN (1902). In Bac. Bütschlii [= Schauäinnum
Bütschlii (Schaud. 1902)], the author observed that in the highly developed
synascite, before "spore formation" (i.e. endothecite formation), a median division
phenomenon occurs, but then soon disappears again, and he regards this as the
"most primitive form of copulation" (page 338).
In doing so , he draws on an observation by HERTWIG concerning a protozoa
(Actionosphaerium) , where the direct descendants of a cell copulate with one
another. Since the protozoa is a caryomone and the bacterium is a multiplicity of
primitive caryomones, such a comparison was not useful. Similar phenomena are
common in bacteria, particularly in the families Sclerothrichidae and Fusiformidae.
This phenomenon will be discussed in more detail elsewhere, and can at best be
explained as a primitive form of association or colony formation or stock formation.

DOBELL (1908) later referred to Schaudinn's view when describing a closely related
species.
PERRIN (1906) goes much further in the interpretation of certain
morphological conditions . In Gristispira Ballimi (Certes 1882), which is considered
a protozoa (Trypanosoma), he explains the finer ascites as "male gametes", the
synascites as "female gametes" and intermediate forms as "indifferent forms".
Thus, a phenomenon found in protozoa is transferred to an organism wrongly
considered to be a protozoa.
That there can be no talk of observed copulation is clear from what is said about
out of the , the conjugation: "This appears to take place but rearely"; "I have
only succeeded in obtaining a few doubtful
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44 Historical information on the question of sexuality.

examples in stained reparations" (page 147). The arthrothecites (here

incorrectly called cysts - a term valid for the protozoa) are considered to be
the results of the conjugation.
VON PROWAZEK (1906 ) , when mentioning the athro-thecites (which he called
plasma buttons and drops), adopts the Spiroch.
Obermeyeri by OBERMEYER, ERICHSEN and HEYDENREICH, in Sp.
hucealis [ by him, in Sp. gallinarum by SCHAUDINN and him, in Sp. Baïbiani
by PERRIN and in Sp. anatodontae by KEYSERLITZ , apparently supports
SCHAUDINN's view. He believes that they are not mere products of
degeneration, "but represent special resting stages in which the protoplasm
undergoes an intimate mixing and connection with the chromatin inhal+ and
which may have been preceded by peculiar sexual processes (autogamy of
two individuals resulting from division)" (page 565).

Also referring to Schaudinn's notes: MÜHLENS and HARTMANN (1906); they say
on page 104: "One could imagine that the long individuals described above represent special
growth forms which break down into a number of short, broad forms by transverse division
and that a kind of multiple reproduction takes place here to produce certain, possibly sexual
(?) development forms. In this context, small forms with a lighter-coloured swelling could
also be found which also showed two darker-coloured spots (i.e. arthrothecite). It is
conceivable that in this way (after possible fertilisation?) permanent forms would arise,
similar to those found by PERRIN for the SpirochaetaBALBiANi and, to a certain extent,
PROWAZEK for the chicken spirochaete." On the other hand, they reject PERRIN's view .

The views like PERRIN , on the other hand, are represented by

KRYSTALOWICZ and SIEDEOKI (1909); They say about the importance of
the small, delicate and short spirochetes (fine ascites)lc (pages 226-227):
“nous les avons définies comme individus males, c'est-à-dire microgamctes,
parce que nous avons aperçu plusieurs fois ces petites spirochetes attachées
aux côtés des grands individual à large corps. Toutefois, après avoir passé
soigneusement en revue diverses preparations où les grosses spirochètes
étaient abon-dantes, de même que les petits exemplaires, nous sommes
arrivés à la conclusion qu'une agglutination accidentelle des spirochètes est
bien difficile à distinguished from the union. Surtout , si sur la preparation ,
outre Spir. Les images d'accouplement des spirochètes provenaient des
preparations où en outre de Spir. pallida se trouvaient aussi Spir. refringens
ou Spir. dmtium;
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Historical information on the question of sexuality. 45

Nous constatons par conséquent, qu'une erreur dans leur interpretation n'est pas
exclue; c'est pourquoi nous les considérons présentement, sous toute réserve,
comme une manifestation de la fécondation.
Toute1 fois, en mettant en doute nos conclusions précédentes, nous
remarquons expressément que nous considérons les deux categories de formes,
les spirochetes courtes et les grosses, de même que les exemplaires petites et
minces, comme des individus tout à fait normaux, non dégénérés; It is likely that
our appearance in the syphilitic lesions is about to begin at a new stage in the
world of the spirochete. This new stadium pourrait être le stade de la reproduction
sexual où des forms décrites ci-dessus pourraient participer comme cellules
différenciées sexual-ment. This supposition nous paraît assez vraisemblable,
parce que chez d'autres spirochètes (Spir. laliiani et probablement aussi S fir.
gallinarum) les manifestations sexuales ont été constatées presque avec tous
leurs-détails. "Chez les spirochètes, la différenciation des individus sexués est
liée à un changement essential de leur structure et de leurs dimensions."

It should also be mentioned here that VICENTINI x) assumed sexual fertilization processes
in oral bacteria; however, this publication is of no importance because VICENTINI regarded all
bacteria of the oral cavity and sputum, including the tuberculosis pathogen, as developmental
phases of Syncrotis luccalis (Rob.) (= Leptotîirix iuccalis) . The extremely diverse appearance
of this organism led this author to such erroneous conclusions.

After my first observation of sexual reproduction in 1916 and 1921, LöHNis

(1922) has recently interpreted certain processes in the symplast as conjunction;
furthermore, POTTHOFF has also done so in Spirillum etc. (1921).

III. Elements of the comparative cytology of bacteria.

Omnia rerum principia parva sunt.

Cicero, de finibus. V 21.

With the exception of a few basic sections, only the

The most important thing is to be indicated, which is important for the understanding of the following
can serve as a basis for a non-expert to gain insight

VICENTINI, F. , Nuovi studii batteryologici sugli sputi sulla morphologia e biologia

de' microbi boccali. Memoria, presenta alla R. Accadenna medico-chirur-gica di Napolia 1892.
[Reference in: Baumgarten's annual report 1891.]
Machine Translated by Google

46 Myeh.

If these suggestions are a little too sketchy, I refer to the excellent basic works of
FISCHER (1897), MIGULA (1899), MEYER (1912), BENECKE (1912) and others.

A. Generative organelles.
The generative organelles are the essence of the bacterial cell and the
carriers of life. The most important organelle is the mych (the primordial nucleus).

1. The Mych (the primordial nucleus).

The mych, the primordial nucleus, is the caryological component and the
carrier of life in the primordial cell (mychite). In the mychite there is only one single
mych, which has a spherical shape and is always located in the spherical mychite
against the wall (Fig. 1). It can, especially if it is not too small in relation to the
mychite, protrude somewhat flatly like a wart over the spherical wall of the mychite
(Fig. 1). Of the bacteria

•O
Fig. 1. Micrococcus aureus (Rosenb. 1884). Two mychites, the left one a pliotrophite, the right one
an atrophite, in which the mural mych is recognizable. Magnification 10000:1.

only the spherical ones (e.g. Micrococcus) contain a single mych, all the others
contain two to numerous mych. It contains no chromatin, is only slightly more
strongly stained by fuchsin than the cytoplasm (somatic plasma) and is usually
completely unstainable by methylene blue, or at least the mych can rarely be
stained slightly more strongly with methylene blue than the surrounding cytoplasm.
The diameter of the mych in bacteria is Viooo o to 74000mm (0)1—0)25 u). If it is
clearly more strongly stained by methylene blue, it is probably always surrounded
by a very fine reserve substance shell or has dissolved nutrients evenly distributed
inside.
In the mychit, the primordial cell with only a single mych, the mych is only
recognizable if no reserve substances have accumulated in it.
These reserve substances, the yolk elements of the primordial cell, are either
distributed throughout the cell or are stored in the form of ultramicroscopic
granules, the trophoconia, around the mych and completely cover it. These
trophoconia are the chromatin elements of the mych and, particularly due to their
nucleic acid content, can be strongly stained with all dyes, and especially with
methylene blue, especially since - a lack of dye absorption is a sure reagent for
the lack of
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Mych. 47

The chemical components of trophoconia are very diverse ; in addition to

nucleic acid compounds, they usually also contain lipids and numerous other
substances.
The trophoconia sheath surrounding the mych, which in coloured
The trophosome always appears as a colored grain in preparations.
The matter of the mych is the mychin. The mychin is a substantive term,
not a chemical one. Since the mychin always comprises the pure mass of the
mych, it can naturally contain chemically different things in the different species.
The mychin appears colored with fuchsin (e.g. carbol fuchsin 1:10 H2 0) as a
completely homogeneous, structureless mass; the coloring is completely
uniform, without any trace of grains or other differently colored components.

Mychites with a distinct mych without a trophoconia cover are provided by,
for example, the micrococci, which in the vast majority of cases show more or less
numerous specimens without accumulation of reserve substances (cf. atrophites).
Here, however, the mych are particularly small. Gonidia from rod bacteria are
much better, as they also represent spherical mychites with a single mych.
Older gonidia from the cholera pathogen ÿ. B. often show a clear image of the
mych directly after brief staining with carbol fuchsin (1:10). However, if remains
of trophoconia are still found, which often only cover the mych like a veil, the
following method is used. After treating the cover glass with
1
On a absolute alcohol for 1 hour — often passing it twice
through the gas flame of a Bunsen burner is sufficient — a trace of gonidia, ÿ.
B. from Microspira comma Schrot., as they are usually found in large numbers
in cultures that have been around for 2 to 4 weeks, is placed in a small drop
of physiological saline solution and this is spread very thinly over the entire
cover glass using a square-bent platinum wire. The air-dry, unfixed preparation
is placed in absolute alcohol for 15 minutes to dissolve the lipids (phosphatides,
etc.) and to fix it at the same time. Since the nucleic acid dissolves in alkalis,
5% soda solution is used. This is heaped onto the cover glass and either
heated on a low flame for a while or brought to the boil 2-3 times over a high
flame. Then rinse well and stain with very diluted (1:10) carbol fuchsin for 2-5
seconds. Preparations that are more strongly stained are usually over-stained
and do not give clear images.

The same result of removing the trophoconia sheath of the mych in

unicellular forms in order to make the mych visible is achieved by enclosing
such forms in hanging drops in physiological saline solution on the hollow-
ground microscope slide (sealed with Vaseline); in this way, starvation forms
are produced at different lengths of time during which the trophoconia sheath
has been consumed.
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48 My ch.

In the association of several mychites to form a bacterial rod, the individual morphological
elements, the my chites, can be designated with my-choses.

A bacterial cell that contains more than 1 mych is a pliomychite.

In the pliomychite, the mych cannot be recognized even after dissolving the lipids (phosphatides,
etc.) using ether and alcohol and the nucleic acid compounds using alkalis (e.g. 5% hot soda
solution) for the following reasons : 1. insoluble residues of the yolk elements of the trophosomes
usually remain (in the case of the diphtheria pathogen, for example, very strong residues) ; if this is
not the case, at least 2. undissolved yolk residues of the trophoconia remain from the trophoconia,
which are evenly distributed over the entire cytoplasm; 3. the difference in the colorability of the
mych by fuchsin to that of the cytoplasm is too small to be recognized in the usually much thicker
layer of the same in the pliomychite.

The mych cannot be recognized in the pliomychite. If traces are sometimes recognized,
these traces are usually strongly stainable with methylene blue, and thus these parts belong to the
closely adjoining trophoconia shell and not to the mych itself.

2. The centriolite.

Just as the nucleus of the metazoan cell is opposed to the centriole (= centrosome) as a
caryological secondary component, and in the protozoa the main nucleus is opposed to the kinetic
or locomotor nucleus (= centrosome = blepharoplast = kinetonucleus = central grain of the heliozoa
cf. Hertwig 1902 = basal body of the trypanosomes etc.), so the centriolite in bacteria occupies the
same position as the mych.

Since both the centrosome of the metazoan cell and that of the protozoa are extremely tiny
nuclear derivatives, it was very unlikely that a corresponding organelle would be found in the
mychite. Just as in the protozoa the blepharoplast is most strongly developed in the forms with a
strong flagellum, so that it has actually been called the flagellum nucleus, the most favorable
precondition for finding a corresponding organelle in the bacteria was in those forms that have a
disproportionately large flagellum in relation to the body size of the mychite unit.

In fact, I have only been able to demonstrate this organelle, called centriolite, with certainty
for those morphological conditions that most clearly show the above-mentioned preconditions. It is
the spermite that "only a centriolite can be recognized with suitable staining. The demonstration of
this extremely tiny organelle is only possible in the case of spermite if the smear is as thin as
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Trophic organelles.
49

is prepared in such a way that it dries in the air in a few seconds and the individual
spermites are usually isolated so that the eye is not distracted by individuals lying nearby,
and with a staining process of approx. 2-5 seconds with carbol fuchsin 1:10. In the spermite
of the cholera pathogen, the size of the centriolite is between 0.01 and 0.02 ÿ (1 hundred
thousandth to 1 fifty thousandth of a millimeter). It has not been possible to prove whether
or not ordinary bacterial individuals have a centriolite. Either it is so tiny that it has not yet
been made visible, or a part of the mych has not split off. The former is probably the case;
One possible explanation is that in the much larger normal individuals of bacteria the
contrast is too great and therefore the picture is much less clear, and the separation of the
centriolite from the mych may also be smaller. In no case was anything similar even hinted
at.

I am unable to decide whether the observation by BÜTSCHLI (1902, Fig. 4 a) of

several tiny, more strongly coloured grains lying one behind the other in the flagellum of
Spirillum volutane can be related to the centriolite. I have not seen anything similar.

Since all knowledge of centriolites refers to bacterial spermite, these extraordinarily

difficult conditions in spermite (Chapter V ÿ 2) will be discussed in more detail.

B. Somatic organelles.

Even if not everything that is understood here as an organelle can be regarded as

such in the strictest sense, it should nevertheless be included here under this term. For
example, the fat body of the metazoa is the fat organ, and so the trophoconia and the fat
droplet, etc., are the most primitive organ structures and may well be referred to as
organelles.

According to their functions, organelles can be divided into: 1. nutritional organelles

(trophic), 2. protective, 3. locomotor, 4. respiratory,
assimilatory and excretory,,organelles.
"

1. Trophic organelles.
The trophic organelles represent the following groups:
1. Components of more liquid consistency (cytoplasm and substances
dissolved therein),
Enderlein, BacterialCyclogeny. 4
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50 cytoplasm.

2. solid components that are excreted in the cytoplasm

(trophoconia, trophosome, trophosomelles), 3.
particularly solid components of the sporitin grain, 4.
connecting bridges between loosely connected individuals (plasmodesmata).

a) Cytoplasm.
The cytoplasm is the somatic plasma that surrounds or surrounds the nuclear
component (Mych). According to MEYERS (1912), it is completely homogeneous and
only receives a structure (honeycomb structure, vacuoles, etc.) through the inclusion
of gaseous substances of a solid or liquid nature that are secreted in the cytoplasm.

Taking into account the microgonidia and microgonites described later, as well
as the spermite, in which the cytoplasm is greatly reduced or almost completely
absent, the assumption seems justified that the pure cytoplasm without inclusions (i.e.
structureless) ultimately represents an ergastic structure itself, which is responsible
for supplying the life-bearing organism (the nuclear part of the cell: Mych) with fluid
and nutrition, especially protein nutrition.

The cytoplasm stains alive with iodide potassium solution yellow to

brown, with very diluted methylene blue solution slightly blue.
The dead cytoplasm stains very differently with color solutions. A stronger stain
is not linked to the cytoplasm, but to the ergastic structures, which mainly represent
reserve substances.
The reserve substances are either more or less evenly distributed, dissolved or
undissolved, in the cytoplasm, which itself also represents a food supply as a
nutritional protein (e.g. lipids, phosphatides, alcohols, sugars), or they are deposited
in the form of extremely fine, tiny, solid or more liquid (e.g. oil droplets) granules
(trophoconia).
The former, depending on the amount present, cause a more or less strong,
uniform coloration of the entire bacterium, the latter form the strongly colorable
granules (cf. trophosomes, trophosomelles) by accumulation around individual or all
mycs. The lipoids are soluble in ether and also
white, waxy in alcohol and, according to TA - MURA (1914), represent a
mass ; the lipoid isolated from the diphtheria pathogen by means of ether extract had
a melting point of 180°, contained no phosphorus, was gram-positive, but not acid-
resistant. The snot pathogen is said to contain palmitin and olein (cf. CZAPEK) .

The phosphatides contained in the alcohol extract (there are no phosphatides

in the ether extract) vary in different forms. TAMURA (1913) isolated a
diaminophosphatide from the tuberculosis pathogen,
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Trophoconics. 51

which has also been found in other forms (e.g. Sclerothrix lacticola L. et N. ), from the
diphtheria pathogen a monoaminomonophosphatide related to lecithin, which represents
a yellow-brown, sticky mass.
TAMURA (1913 ) isolated a high molecular weight alcohol from the tuberculosis pathogen and
from Sclerothrix lacticola L. et N.

Formula C29 H5 6 0, which he calls mycol, but which is completely absent in other
species, e.g. in the diphtheria pathogen. The mycol is strongly gram-positive, acid- and
alkali-resistant. It is mostly contained in the bacterial body as an ester of a higher fatty
acid.
Of the carbohydrates, TAMURA (1914) demonstrated a pentose in several bacteria
(1. arabinose, partly as arabone, a hemicellulose), and in Sclerothrix lacticola L. et al.
also a hexose. But glycogen and logen (cf. A. MEYE R 1899) are also found in the
bacterial body.
The cytoplasm itself consists of protein building blocks, which are particularly
The following amino acids, as demonstrated by TAMURA (1913/14 ), represent :
Arginine, histidine, lysine, ru 1. proline, valine, tyrosine (= oxyphenylalanine) and
tryptophan.
In addition, there are other amino acids, which often vary between species.
Sclerothrix tuberculosis (Koch) and S. lacticola (L. et N.) also contain phenylalanine, while
Corynobacterium diphtheriae (Löffl.) also contains leucine and isoleucine, and perhaps
also adenine.
According to the findings of TAMURAS (1914), the inorganic components of the
same species can be subject to large quantitative variations depending on the living
conditions to which the bacteria are exposed.

In addition to these cell components, the colored bacteria also contain pigments
(cf. III ÿ 4 b).

b) Trophoconia, trophosome and trophosomelles.

With stained bacterial preparations, one occasionally has the opportunity to
observe a striking fact. Individual specimens are then found which only take on the color
very weakly or do not take on the opposite color in Gram's or similar staining, although
normal individuals of the same species are gram-positive. Often the outlines are then
only very weakly colored. Such preparations are easily capable of creating the illusion
that a number of the bacteria have fallen off the cover glass during treatment after staining
and only color fringes indicating their outlines remain.

However, when the tube is adjusted, it can always be seen that these bacterial individuals
are present in their entirety.
If such preparations come from pure cultures, then
4*
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52 trophoconia.

the idea easily arises that a stranger has crept in. However, cultural tests easily refute this suspicion.
It turns out that quite isolated colonies sometimes contain a large number of such specimens that
resist the assumption of dye. Moreover, careful comparative observation usually reveals all stages
of the transition from unstained to strongly colored individuals. Indeed, in the case of chain-forming
rods or spherical forms, such as in Streptococcus, one sometimes finds all these stages mixed
together in a very colorful series in a single chain.

A few such individuals are found in most species. There are also species in which about half
of the individuals show this phenomenon, but they are generally rare. Such species are most
commonly cultured from sputum, for example.

This phenomenon has received little attention so far.

BABES (1895 ) observed them in Micrococcus aureus Mig. 1900 . He noticed that
when stained with methylene blue, a greater or lesser number of individuals, in addition to the dark-
colored specimens, always took on a very pale reddish color, and explained them as transformation
products of specimens that form as outgrowths on normal individuals and have taken on the role of
carriers of the yellow dye of the colony.

He writes on page 424, 1. c.: "It has become clear to me that these structures contain the yellow
dissolved dye of the culture." That this view is incorrect is clear from the following ; however, it
should be mentioned here that the fact that young, strong yellow cultures can and usually contain
only a few pale (or gram-negative) individuals and that old cultures (of a strong yellow and also very
faint yellowish color) usually contain very numerous such individuals speaks against such an
assumption.

LEHMANN and NEUMANN (1904 ) rightly recognized the "common view" that every
bacterium is either gram-positive or gram-negative as incorrect, but that different colonies of the
same species can take on one or the other color. According to ZIMMERMANN, all species of the
genus Lamprella Enderl. retain the color in young colonies, although these are usually described
as gram-negative. HEIM (1906 ) also mentions on page 49 that in Gram's staining, some bacteria
only take on the color when they are young. Lo MONACO (1911 ) observed such differences in the
,
colorability of the same species in Mogalliapeumoniae (Weichselb. 1886); he says on page 117: "It
is evidently a matter of involution phenomena that can be traced back to the exhaustion of the soil,
and the pneumococci that no longer take on Gram are dead germs." DEUSSEN (1921 ) writes the
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trophoconia. 53

Gram-positive phenomena are attributed exclusively to the nuclein or nucleic acid ; by dissolving the latter
with acid, it makes cells (yeast, Bac. bulgaricus) gram-negative ; these become gram-positive again by the
addition of nuclein and nucleic acid in the form of their salts; as will be explained below, however, all
possible reserve substances are possible, including lipids.

As will be seen from the following sections, the phenomenon that both gram-positive and gram-
negative individuals can be present in the same cultures can be observed in a large number of different
forms from all groups of bacteria and in all possible stages of development. It now remains to attempt to
gain an understanding of this.

If such bacteria are treated with Gram's stain, it can be seen that mostly the weakly stainable
individuals are gram-negative (pale red) and the strongly stainable ones are gram-positive (blue).

If one follows the different stages of the disappearance of the colourable substances, e.g. in short
rods (cf. dimychite), it becomes apparent that first the colourable matter disappears in the middle zone and
in the two

Poles each have a spherical (granular) accumulation.

In the case of micrococci, there is only one point around which the stainable substance is grouped. This
can be observed not only in stained individuals, but also in living, unstained specimens; the more stainable
parts appear here as more strongly refractive grains. It is noticeable that, particularly in specimens that are
in the early stages of cell division, the stainable substances gradually disappear. After

or with the division a new accumulation begins

the same.

The suspicion is therefore justified that the colourable substances are reserve substances that are
used up during the process of nuclear division, firstly because a strong energy consumption sets in for the
formation of two nuclei from one and then simultaneous food intake from outside is probably excluded.

Since in metazoans and metaphytes cell division there is often a strong consumption of yolk, which
can lead to the complete exhaustion of the yolk supply, a parallel physiological process is probably at work
here. However, these yolk particles in the somatic cells of animals and plants are always microscopically
recognizable as threads, granular threads or larger or smaller grains (called chondriocontes, chondriomites
and chondriosomes).

This does not seem to be the case with bacteria.

the matter in question appears as a fine haze in which no physical structures can be discovered even at the
highest magnification.
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54 trophoconia.

However, if one compares particularly large bacteria or similarly constructed oidia (so-called smut
spores) of lower fungi (see Section VIIIA), one can actually see a fine to extremely fine granular
structure, which is all the more clearly made up of extremely fine grains or rods.

composed, the more the reduction has occurred.

In the Oidias, these grains are often much larger and more distinct.
When the colourable substances disappear, the last remnants of these finest granules are
always found near and especially close to the boundary wall of the mych (primordial nucleus), a fact
which is entirely in harmony with the manner in which the yolk structures mentioned above appear
in higher cells.
There seems to be no doubt that these are structures that belong to the broad area of mitochondria.
According to the general usage, according to which all morphological elements in the cell that
intensively absorb pigments are called chromatin, the mitochodrines and corresponding structures
could also be called chromatin. But this would be a much too broad definition.

Term because in 1879 FLEMMING called the entirety of the strongly stainable part of the cell
nucleus of higher organisms, which is composed of chromosomes, chromatin, and the term should
be kept limited to this purpose, especially since yolk elements (food reserves) are probably
combined with true nuclear elements in the chromosome. The former are strongly stainable, the
latter (linin or achromatin) are not.

If the bacteria here really are yolk-like reserve substances, then it should be possible to
artificially produce such specimens with a reduced or even completely used up yolk content by
exposing gram-positive bacteria rich in yolk, i.e. highly stainable bacteria, to a state of starvation.
This has been achieved many times in numerous experiments. For this purpose, individuals from a
young culture were placed in a hanging drop of physiological saline solution and hermetically sealed
with Vaseline on the hollow-ground slide. This small chamber was kept in the dark for 3-6 days.
This assumption was indeed confirmed; the consumption of the stainable mass could be determined
both in living individuals and in the stained state.

Even in the case of micrococci (ÿ. ÿ. M. aureus), the presence of gram-negative individuals
increased significantly as a result of this treatment.
When examining the individuals in the hanging drop without food step by step, it can even
be determined that mostly the reserve substances in the dissolved state are used up first, such as
the mycol, the lipids, etc. Then, in forms that have a very intensive and uniform coloration and thus
no
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Trophosome, TrophosoraoUe. 55

Color grains (trophosomes) can be seen, these color grains stand out clearly, while the cytoplasm
becomes lighter. The same result is achieved by chemically dissolving these substances with ether and
alcohol. Here too, the trophosomes remain undissolved.

The above results in the following. ' The tiny, usually ultra-microscopic
particle of solid reserve material is called trophoconie.

The trophoconia is the structural unit of the strongly colourable food reserve substance
accumulated in the cytoplasm in solid or liquid form or at least in a more solid or dense form than the
surrounding cytoplasm, a tiny dust-like grain. This is therefore a morphological and not a chemical term,
since the chemical composition certainly varies between different species and also under different

external circumstances are extremely different. A strongly represented

The main component is nucleic acid, mostly as a nucleprotein, but other components of a yolk-like
nature (cf. below), oils, fats, etc. are also involved.
The trophoconia are often arranged densely around the mych like a sheath. If this sheath is
thin, the structure appears as a tiny, strongly staining granule, the trophosomelle; if it is thick and dense,
larger, strongly staining granules, the trophosomes, appear.

The trophosome is the more or less dense and thick trophoconia shell (i.e. a hollow sphere)
around the mych. The trophosome has often been wrongly considered to be the bacterial nucleus
(mostly the two united middle trophosomes of a Didimychites), furthermore also the Much granules, at
least in part (besides the gonidia).

"
The “sporogenic grains” of ERNSTS (1889) and the “metachromatic
ÿ. T. belong to the "chromatin granules" of BABES (1895), later called Ernst-Babes granules (another
part are gonidia, cystites, etc.); often the trophosomes are also simply called chromatin granules.
Further details can be found in Section II A.

The trophosomes and trophosomelles are particularly clearly visible in forms which contain
fewer lipids and mycoles, such as in the plague pathogen (Fig. 2—7) or in the diphtheria pathogen (cf.
IV ÿ 4).
In forms that contain more dissolved reserve substances, such as the cholera pathogen, etc.,
the trophosomes, which are admittedly less dense, are not visible with normal staining. In this case,
starvation treatment in a hanging drop or chemical elicitation with ether and alcohol, absolute, must be
carried out beforehand; if one then stains, the trophoconia usually become clearly visible. They can
also often be more clearly distinguished by Gram staining in some species, such as the tubercle
bacillus, in which the trophoconia are rarely visible with the usual Ziehi staining.
Machine Translated by Google

56 Atrophosis, miotrophosis, pliotrophosis.

because the majority of the sticks are not acid-resistant (cf.

MUCH etc.).
As in metazoans, eggs with missing yolk are called "alecithale"
If eggs are called, then the phenomenon of the absence of trophoconia and dissolved reserve
substances in the bacterial cell is called atrophosis (lack of yolk). Accordingly, the phenomenon
of the abundant occurrence of trophoconia or of dissolved reserve substances is called
pliotrophosis (abundance of yolk), and the sparse presence of one or the other is called
miotrophosis (lack of yolk).

2nd 3. 4th 5th 6th 7th

Fig. 2—7. Ewcystia pestis (Yersin 1894). Plague pathogen. Magnification 10,000:1. Specimens from human blood of
a plague victim.
Fig. 2.1 Dimychit. The two mych wrapped in trophosomes. Fig. 3. Didimychit. Three of the mych wrapped in trophosomes,
the fourth mych without a trophoconia sheath and therefore nothing visible in its place. Fig. 4. One of the two mych
divided, the daughter mych not yet moved further apart; the other undivided or, if already divided, the daughter mych
without a trophoconia sheath. Fig . 5. Didimychit; both end mych shortly after mychomitosis. Fig. 6. Didimychit; three
mych in trophosomes, one wrapped in a trophosome. Fig. 7.

The two young Dimychites resulting from the division of Didimychites ; the upper one with
only one trophosome, the lower one with one trophosome and one trophosomelle.

Finally, it is advisable to name bacterial cells (be it a simple mychite or a pliomychite),

the cells of the closest relatives and the cells in general, according to their content of trophoconia
and dissolved reserves, etc.

1. the atrophy = the yolk-less cell, 2. the miotrophy =

the yolk-poor cell, 3. the pliotrophy = the yolk-rich cell.

Depending on its location in the bacterial cell, the trophosome can be classified as: 1.
Telotrophosome, which is always located at the end of a bacterial rod or thread; 2.
Ascotrophosome,
which is located at any point on the thread,
except for the two ends.
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Chemistry of the trophosome. 57

Telotrophosomes include, for example, the structures formerly often referred to as polar
caps or polar grains, as long as they are not telogonidia, etc.

The trophosomes, trophosomelles and mychites without trophosomes (i.e. with uncovered
and therefore invisible mych) can appear mixed in a colorful row in the bacterial rod, as shown
in Fig. 2—7 of the plague pathogen. The accompanying Fig. 8—10 show some of the many
possibilities of a rod composed of 4 mychoses of Corynobacterium pseudodiphtheriticum (Löffl.).

8th 9th 10th

Fig. 8—10. Corynobacterium pseudodiphtheriticum (Lehm, et Neum.). Rods composed of four

mychoses. Magnification 10,000:1.
Fig. 8. Rod with one telotrophosome and three trophosomelles. — Fig. 9. Rod with two
telotrophosomes and two trophosomelles. — Fig. 10. Rod with one telotrophosome, two
trophosomelles and an atrophic mychosis.

It is easy to prove that the trophosomes and the substance that composes them, the
trophoconia, are by no means chemically uniform in different bacteria. Their main content, nucleic
acid (not exclusively, as SCHUMACHER claimed in 1922 for the polar granules of the diphtheria
bacillus1 )), is easily dissolved with dilute acids or alkalis. Thus, a 5% sodium carbonate solution
(soda) easily dissolves the nucleic acid, especially when slightly warmed2 ). In some bacteria,
the trophoconia are completely dissolved in this way, proof that only nucleic acid or related
substances were present (e.g. in Spirillum volutans Ehrenb.).

In others, such as the diphtheria pathogen, some trophosomes are completely different and
partially dissolved, so that completely different substances are certainly present in addition to
nucleic acid. Other bacteria, such as the diphtheria pathogen,

T) The proof of this is easy; if these objects are treated with alcohol and ether, the granules become
considerably thinner after the lipids have been dissolved. '

2 ) It is best to place the coverslip smear in absolute alcohol, which fixes it and at the
same time dissolves the phosphates, some of the other lipids, etc.; then, place the 1Va2 C03
solution on the coverslip and heat it several times. Finally, the remaining lipids can be dissolved
with ether. Only then should the sample be stained. Very nice contrast images are obtained from
preparations in which only half of the coverslip has been treated with soda.
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58 Trophosome and throphosomelle as hollow sphere.

The sulfur bacteria, for example, excrete sulfur as trophoconia, so that the trophosomes of
these organisms contain a lot of sulfur. In the case of iron bacteria, they can absorb iron
and manganese.
This extremely different behavior of the trophosomes in the face of chemical
influences thus clearly proves that a uniform substance that composes the trophosomes
of all bacteria cannot be thought of. When A. MEYER (1904) therefore introduced the
chemical term volutin (also recognized as a reserve substance) for the material of the
trophosomes of Spirillum volutane Ehrenb., which incidentally can certainly be considered
to be the closest relative of nucleic acids, the name volutin must under no circumstances
be used for the material of the trophosomes of other bacteria, but must be used specifically
for the specific nucleic acid form of Spirillum volutane Ehrenb., since it was fixed as a
chemical term and not as a morphological term, even though volutin may be involved in
the composition of the trophosomes of some other bacterial species. For this reason, the
name volutin grain, which MEYER would like to see applied to all corresponding chromatin
grains of other bacteria, is obsolete and falls entirely within the terms trophosome and
trophosomelle.

The trophosome and the trophosomelle always include a Mych.

Since the latter absorbs only very little or almost no staining, the trophosome and the
trophosomelle are always a strongly stainable hollow sphere.
However, this finding is only the result of long comparative morphological observations,
based on the conviction that every trophosome and every trophosomelle has a central
mych. I have never been able to directly observe this tiny, unstained central spot in the
pliomychite. The trophosome and the trophosomelle, when isolated, always appear as a
grain, which never shows any noticeable lightening in the center. Objects have often been
observed that appear to show trophosomes with a clear cavity (e.g. in the case of cholera,
diphtheria pathogens, etc.); however, precise comparisons have shown in all cases that
these were shrinkages caused by the cover glass being heated too much during fixation.

Even if the possibility of making the tiniest unstainable centre of the trophosomes and
trophosomelles visible cannot be completely dismissed, however extraordinarily small it
may be in view of the intensity of the stainability of the trophoconia, there is nevertheless
an extremely strong suspicion that the two observations of such cavities in the interior of
the trophosomes, which were found in the literature, are also deceptions due to shrinkage
phenomena.

BÜTSCHLI (1902) says of the chromatin granules of Spirillum volutans

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trophosomes and trophosomelles. 59

Ehrenb. pag. 50 “The precise examination of these often quite impressive grains showed
their hollowness very definitely; it could even be recognized frequently that the weaker
refracting cavity was eccentric (cf.
Fig. 4 g, 4 h)." Likewise, according to findings made by A. MEYER (1904) on the same
species, swelling of the volutin always occurs when intensively stained with methylene blue
and the trophosomes (volutin grains) sometimes swell up to form hollow spheres.
Furthermore, DOFLEIN (1911) says of Spirociiaeta plicatüis Ehrenb. 1838: "In regular
arrangement on both sides of this axial thread lie grains which are hollow and consist of
volutin."
The fact that trophosomes with cavities can be simulated even without shrinkage is
shown in Figure 11 of the diphtheria pathogen. To the left of the middle are two trophosomes
secondary

Fig. 11. Corynohaeterium diphtheriae (LöffL). From an old, very large, isolated colony.
Magnification 10000:1. Ascite with 11 trophosomes and trophosomelles; the 4th and 5th from the
left are connected by lateral pseudozyges, simulating a single trophosome that appears hollow.

connected by lateral strands of trophoconia (pseudozyges), so that the impression of a

hollow sphere can easily be created.
Confirmation of the view that trophoconia, trophosomes and trophosomelles are
reserve substances can be found in findings in the case of distinctly parasitic bacteria.
Distinctly parasitic bacteria, provided they are completely surrounded by food, are not
exposed to a lack of food or are not juvenile stages of higher forms (e.g. insect larvae), do
not need to accumulate reserve substances. Taeniids (tapeworms), for example, do not
produce fat. Accordingly, in the case of distinctly parasitic bacteria, particularly those which
present great difficulties to artificial breeding, atrophites and miotrophites should predominate,
i.e. forms which are difficult or weak to stain or which are gram-negative. This is actually the
case. Just think of the stunted nuchal pathogen (Diplococcus intracellularis Weichselb.) and
Treponema pallidum (Schaud.); The former* is gram-negative, only weakly stainable and
without trophoconia accumulations, the latter is extremely difficult to stain. Less pronounced
parasites, such as the cholera pathogen, whose highest developmental stages do not
develop in the host but outside it, can still be gram-negative, but here there are clear
trophosomes that only
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60 sporitin grain.

by lipids etc. which are also highly stainable and can be easily removed with alcohol and ether.

The fact that different formants of the same species can also prove to be different with regard to
the accumulation of trophoconia is shown by the experiments of CHRISTELLER (1914) on B. lulgaricus;
after passage through milk agar, colonies were formed on agar (the normal form does not grow on agar )
which showed poor growth in milk and lack of coagulation; these were gram-negative on agar, gram-
positive in milk. This B. lulgaricus formant is thus able to store reserve substances in milk, but not on agar.

c) sporitin grain.

As explained in more detail in section VI a A 4 d, the so-called bacterial spore is morphologically

not a spore, but a dimychite or didimychite. Therefore, the term sporite is used for this structure.

During the formation of sporites, the reserve substances of the bacterial cell are dissolved away,
especially the trophosomes etc. gradually disappear completely, and other components are deposited
as grains between the two mych of the dimychites. This is the sporitin grain. Its substance is sporitin,
which, however, is not a chemical but a substantial term. In the didisporite, the two central mych are
included in the sporitin grain.

The sporitin grain begins to deposit in the middle between the two mych, eventually filling the
entire space between the two mych and almost the entire cell. This is where the idea of an endogenous
formation of the sporite comes from, because the sporitin grain was considered to be the essential part
of the sporite. The sporitin is a homogeneous, very solid, highly light-refracting mass, which is usually
very difficult to stain, cannot be stained with many dyes, and represents a reserve substance.

In the interior of the sporitin grain, in its center, there is a tiny point in the didisporite (cf. page 142) which
is often more accessible to certain dyes and is therefore different in structure or composition. A. MEYER
(1912) describes several such stains on page 70 and shows some of these pieces in Figs. 8, 9 and 10 ;
he interprets these sporitin grain centers as bacterial nuclei on the basis of their colorability, which is
due to the trophoconia shell, and without taking into account the nuclear nature of the polar grains.

During germination, the sporitin is completely dissolved or transformed into another state of
aggregation. From its strong light refraction, one could conclude that it is a solid, perhaps very solid
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Protective organelles. 61

Body. One could think of dried egg white in particular. The extraordinary hardness
of dried egg white (such as Chinese dried chicken egg white) would also explain
the extraordinarily long viability of disporites and didisporites, which are a
completely isolated phenomenon in nature. The sporites of the original cultures
of Migulanum anthracis (Koch 1876)

[Anthrax pathogen] Koch's has proven to be viable even today. The hardness
and impermeability of the dry protein should then retain the last necessary
residue of organic water in the innermost mych in order to make such a long life
possible. d) Plasmodesmata.

If two or more individuals of bacteria remain attached to one another in a

chain-like manner (desma formation, cf. VA 2), two at a time are connected to
one another by a fine connection, for which A. MEYER (1912) uses the term
plasmodesma, which was initially proposed by STRASBURGER (1910) for
plasma connections between cells of higher plants and animals . Noticeably long
plasmodesmata are shown in Fig. 175.
Whether, as MEYER believes, plasma strands are always contained in the
fine connection, or whether they are sometimes only membrane connections,
does not seem to me to have been clarified with certainty.

2. Protective organelles.
The protective organelles consist of a solid membrane that surrounds the
cell (membrane) and an adjoining gelatinous mass, which is usually only a very
thin layer but can sometimes reach a considerable thickness (mucous membrane).

Both the membrane and the mucous membrane are secretion products
from the cytoplasm.
a) Membrane.
The bacterium is, as ZOPF (1883), DE BARY (1884), A. FISCHER (1895) and others have
already established, always covered by a membrane. This is probably always the result of
secretions of the cytoplasm, as is already shown by the fact that a bacterial individual hatching
from a sporite leaves the old membrane behind and forms a new one or already within

the old sheath has formed. Even in cases of frequent gonidia formation, as is
usually the case with Crenothrix Cohn 1870, the old filamentous sheath remains
behind and the gonidia, covered with a new membrane, push out of it.
According to MEYER, the membrane is not dissolved by dilute potassium
hydroxide solution and dilute hydrochloric acid, but is dissolved by 5% sulfuric
acid at 85° in 372 hours.
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62 mucous membrane.

A reliable detection of the membrane is possible with the help of A.

FISCHER (1897) discovered plasmolysis. Even in 1% saline solution and in 2.5% potassium nitrate,
the membrane stands out clearly from the shrinking cytoplasm. Even when the sporites, gonidia, etc.
hatch (germinate), the remaining membranous shells are sometimes

visible (if they are still attached and being towed).

In some bacteria the membrane attains a considerable thickness, as in Chlamydothrix Mig.,

Sphaerotilus Kütz. etc., which has led some systematists to regard these as a special group (vaginal
bacteria). The justification of this view has not been confirmed on a comparative morphological basis.

b) mucous membrane.

A number of bacteria have a thick shell of slimy, sometimes highly refractable, highly water-
containing substance, which is attached to the membrane and emerges from it or is secreted. A.
MEYER (1912) already recognized: "Few or probably no vegetating bacterial cells are covered only by
the described delicate membrane at their stage of development." In most or perhaps all

There is a more or less thin, sometimes very thick layer of gelatinous substance (e.g. Bact. pneumoniae
Friedl.), which often looks like a capsule.
-like and has therefore often been referred to as capsule formation.

The mucous membrane is usually uncolourable or very difficult to colour and

therefore can only be recognized in differently colored substances. For example, the mucous
membrane of Mogallia pneumoniae (Mogallia pneumoniae) is easily recognized in sputum.
Those of Bact. pneumoniae Friedl. or of Bacillus tumescens Zopf etc. are easily visible to the eye by
spreading the bacterial mass on a cover glass in a drop of physiological saline solution mixed with a
little liquid Chinese ink; these smears can then be stained.

According to ZETTNOW (1907), the mucous layer of Streptococcus alla Zettn. dissolves in
sulphuric acid and in chloride of zinc iodine. According to MEYER (1912), the mucous membrane
(especially of B. tumescens Zopf) is soluble in copper oxyammonia, as shown by smears mixed with
India ink.

A. MEYER was able to stain the mucous membranes by prolonged application of Delafield's
hematoxylin; also by using Magdala red in 30% alcohol (several hours) and subsequent transfer to
glycerine.

Since bacteriologists have often referred to drying zones as capsules, A. MEYER (1912)
recommends using the term capsule
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Motor Organelles 63

to drop it altogether, especially since the term capsule completely coincides with that of the mucous
layer.
The substance of the mucous layer is composed differently in different species, as Meyer
(1912) pointed out on the basis of microchemical findings. More detailed investigations into this have
recently been carried out by Kramák (1921); according to him, the capsule of Calymmato-, bacterium
pneumonicum (Fried. 1882) (Friedländer's pneumonia bacillus) consists of galactane, a polymeric
carbohydrate which, after inversion, yields galactose; that of the anthrax pathogen (Migulanum
anthracis [Koch 1876]) is proteinaceous and sulphurous, but phosphorus-free; the mucous substance
of Bac. radicicola consists of dextran, a polymeric carbohydrate.

,
hydrate, which upon hydrolysis yields glucose.
For all these reasons alone, the objections recently raised by Plasa j (cf. ÿ. I. Orig. 91, 1924,
pages 353—355) against the existence of the mucous membranes, which he describes as artifacts,
are invalid ; according to experiments carried out by myself, there is an even simpler direct proof of
this: if one spreads out mucous membranes using the ink method (i.e., mixed with Chinese ink before
drying) and stains these smears after they have dried, one obtains extremely beautiful and instructive

preparations.

In the Chondromycids, the mucous membrane of the synascites has increased phylogenetic
and systematic importance, since in these forms it mediates the adhesion of the synascites to form
complicated higher associations, which represent growth forms that are partly reminiscent of growth
forms of higher fungi.

3. Motor organelles.

The movement organelles of bacteria are the flagella. Only in a few forms is the occurrence of
an undulating membrane possible.

a) Flagellum.
The movement organs of bacteria, the flagella, are thread-like, plasmatic processes. According
to their arrangement on the bacterial individual, Messea (1890) first divided bacteria into monotrichous
,
lophotrich and peritrich forms are distinguished, names which have now become generally accepted.
The former have only one terminal flagellum, the lophotrich a tuft of 2 or more flagellum at the end and
the pefi-trich have more or less numerous flagellums distributed over the whole surface.

While the seat of the kinetic energy of the flagellum in flagellates is in a nuclear section at the
base of the flagellum, the blepharoplast,
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64 Respiratory, assimilatory and excretory organelles.

is located, nothing similar has been found in the bacterial flagellum. Only in the flagellum of the
bacterial spermite (cf. III A 2 and ÿ ÿ 2) has a tiny granule been found that is probably an equivalent
of the blepharoplast and has been called a centriolite.

As already mentioned on page 49 (III A 2), it is also uncertain whether the grains (sometimes
several in one flagellum) observed by BÜTSCHLI (1902, Fig. 4 a) in the flagellum of Spirillum
volutane Ehrenb. can be related to the centriolite.

There is no doubt that there are flagellated forms of bacteria which do not possess flagella
in all stages of development. FISCHER (1895, p. 49) already points to a flagellated form of
Bacillus subtilis . MEYER (1912, p. 107) also considers this to be probable.

If FISCHER, MIGÜLÄ and others nevertheless point out the absence of scourges
to be a general difference, this must of course be taken into account with the restriction that all
developmental stages (apart from spermite, for example) must be flagellated in such a case. The
genus Bacterium therefore only includes forms with the properties mentioned above.

As MEYER (1912, p. 108) points out, the decision in detail is naturally often very difficult,
as, for example, it is often not easy to determine whether flagella are present or absent in
representatives of the genus Micrococcus Hall and Planococcus Mig. MEYER (1912, p. 105—141)
provides a detailed section on flagella .

b) Undulating membrane.

An undulating membrane as a movement organelle is found only in some spirochetes. It

is very clear in Cristispira Ballianii (Certes) and even broader in Cristispira anodontae Keyss.,
where it extends over almost the whole length of the rod. DOFFLEIN (1911) and KEYSSELITZ
(1907) depict it.

4. Respiratory, assimilatory and excretory organelles. a) Vacuoles.

As far as the bacterial cell actually contains cell sap vacuoles

They probably serve partly to absorb reserve substances and partly to perform an excretory
function. According to MEYER and others, fat vacuoles also occur in bacterial cells.

MEYER (1912) also counts volutin vacuoles among the vacuoles.

However, as shown in section III ÿ 1 b, the trophosome represents a deposit of tiny solid particles
in the cytoplasm (tropho-conia), which encloses a mych in the center, so that one cannot speak
of vacuoles here.
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dye formation 65

For the present implementation, a closer look at the

Vacuoles not required.

b) Dye formation.
In higher organisms, even in unicellular ones, the pigment is bound to special grains, the
chromatophores; such as the brown diatomine of the diatoms, the blue-green phycocyanine of
the cyanophyceae, the red phycerythrin of the rhodophyceae, the brown phycophaein of the
phaeophyceae, the yellow phycopyrin of the peridineae, the red haematochrome of Euglend and
Haematococcus , etc., which seems to be close to carotene, and also the chlorophyll, xanthophyll,
carotene, etc. of the higher plants.

These grains and their pigments are often associated with respiratory and assimilatory activity,
particularly the breakdown of carbonic acid.
It has not yet been proven with certainty whether the pigments of bacteria have a similar
function. This was claimed by ENGELMANN (1883) , but WINOGRODSKY (1888) disagreed
with his statements. MOLISCH (1907) stated that oxygen secretion does not occur in purple
bacteria, but pointed out the possibility that they could assimilate CO2 without expelling oxygen.
MEYER (1912), however, rejected such a possibility.

The pigments of colored bacteria are partly soluble substances. For example, the red
pigment of Bact. prodigiosan Ehrenb. dissolves in alcohol to form an orange-yellow color, which
according to FLÜGGE (1886) turns into carmine red when acid is added and into yellow when
alkali is added. According to the findings of MOLISCH (1907) , purple bacteria have two pigments,
namely a green one that is soluble in absolute alcohol, bacteriochlorin, and a red one that is
soluble in chloroform or carbon disulfide, bacteriopurpurin, as the latter was already called by
LANKESTER (1873) . These substances are formed primarily in the light, like some of the other
bacterial colors.

IV. Elements of the comparative morphology of bacteria.

Natura non facit saltus.

Linnaeus, philosophia botanica.

It is impossible to obtain a clear idea of the nature of organisms without the establishment
of clear and precisely defined morphological units and without a sharp morphological
nomenclature.
Even for bacteriological research on a comparative morphological basis, the first
requirement is the introduction of a strict morphological nomenclature, the complete absence of
which would hamper the development of
Bnderl e in , Bacterial Cyclogeny.
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66 Mychit.

comparative bacterial morphology beyond the most primitive beginnings.

We have seen that the reason why CoHN's correct ideas of a morphological and systematic
nature, even though they were far removed from cyclogenetic knowledge, gave rise to an endless dispute
between pleomorphists and monomorphists, lay in the complete lack of comparative morphological
training of the disputants, who, with few exceptions, used CoHN's classificatory genus names
(Micrococcus Bacterium, Bacillus, Vibrio) as morphological names - a usage that has unfortunately
become completely established to this day and has thereby brought hopeless confusion into the entire
morphology and systematics of bacteria. However, these names were used for morphological purposes
because no morphological names were found in bacteriology. Strangely enough, this need for
—,
morphological names, which soon became apparent following the CoHN conception, especially since
the pure form concepts of LANKESTER and BUCHNER were not taken into account for comparative
morphology, has not been recognized by anyone in almost half a century since COHN up to the present
day , and while systematics has experienced significant development in recent times through the
influence of outstanding botanists, I mention only FISCHER, MIGULA, MEYER, morphology has
remained almost entirely at COHN's standpoint . The main reason is that the outstanding later advances,
which were mainly in the field of biological morphology (development) and are linked to the names of
important morphological researchers, I recall only ZOPF, METSCHNIKOFF, WASSERZUG, were
crushed by the monomorphists, the main contingent of whom are practitioners and physiologists, and
have been completely suppressed to this day. Monomorphism still prevails in bacteriology to the extreme
that anything that shows any kind of morphological difference is regarded as degeneration or death, or is
attributed to teratology (involution, deformity) and mutation.

It is therefore no wonder that even recent, outstanding individual results - I only need recall the
diphtheria results of HEWLETT and KNIGHT - were ignored simply because they contradicted the
phantom "monomorphism".

A. The morphological unit (mychit).

The mychite is the morphological unit and building block of which all bacteria are composed, with
the exception of the Mono-mychota, which represent only a single mych.
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Mychit. 67

1. The mychit (the primordial cell).

A cell with only a single mych is the mychite (see Fig. 1).
As will be explained elsewhere, the mychit is the primordial cell.
The shape of the mychite is usually that of a sphere with a wall-mounted or nearly
wall-mounted mych (see Fig. 1).
Examples of mychites are all individuals of the Monomychota with the exception
of those in division, e.g. Micrococcus. The basites and gonidia of the dimychotes also
belong here.
The mychite may be completely without a trophoconia sheath (in atrophosis) or
it may have a weak trophoconia sheath in myotrophosis, thus possessing a
trophosomelle, or it may have a thicker trophoconia sheath and be trophosomes, or
the whole mychite may be filled with trophoconia in pliotrophosis (Fig. 1).

The atrophite is very weakly stainable and is usually gram-negative when

stained with Gram. The pliotrophite is gram-positive and usually strongly stainable.
The same species can exhibit both gram-positive and gram-negative mychites.
All so-called gram-positive micrococci always have, in addition to pliotrophs, more or
less atrophies in cultures, which are then gram-negative. The number of atrophies
increases particularly in older cultures (e.g. Micrococcus aureus).

The mych of the monomychotes (e.g. Micrococcus) is very small, usually about
0.1 ÿ in diameter, that of the mychites of higher forms (Dimychota) is usually much
larger, about 0.2—0.25 ¡ÿ. Thus, in general, the assumption seems justified that the
presence of a larger amount of nuclear material (mychin) in a mych enables the
organism to develop more phylogenetically.

The proof that the mych in mychite is actually always attached to the wall can only be
established by long comparative observation, since naturally in the microscopic preparation the majority
of mychites do not appear to be attached to the wall at all. The mych of an atrophite usually appears
optically to be in all possible positions away from the edge, which is due to the transparency and
spherical shape of the mychite and can easily be confirmed by scattering glass balls each painted
with a black spot. In the present work, in order to always make the wall attachment of the mych clear,
the mych have been drawn almost consistently on the edge, although the microscopic image often
shows other images.

5*
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68 Mycomitosis.

2. Mychomitosis and diplomamychit.

The division process of a mychite is initiated by the division phenomenon in
the mych itself, the mychomitosis or mychokinesis. Due to the exclusive possibility
of reliably observing the mych in the colored state in mychites without trophoconia
accumulation, certainty about mychomitosis can only be obtained in those individuals
which, due to atrophosis, allow easy insight.

A form that naturally has a strong tendency towards myotrophosis would be

particularly suitable for the detection of mychomitosis. Such a form is found, for
example, in Diplococcus intracellularis (Weichselb* 1887), which, due to its extremely
low ability to deposit reserve substances in the form of trophoconia, shows a very
special sensitivity to less favorable conditions and, as a result, easily dies off, since
it is a pronounced parasite.

12th 13th 14th 15th 16th 17th 18th 19th

Figs. 12-19. Diplococcus mtracellularis (Weichselb. 1887). Veigr. 10,000:1.

Fig. 12. Mychite with simple mych. — Fig. 13. Mychite with enlarged mych. — Fig. 14.
Mychite with a mychozyg, consisting of 2 daughter mychites and a connecting mychomite. — Fig. 15. The two daughter mychites are
separated by reduction of the mychomite. — Fig. 16. The two daughter mychites move away. — Fig. 17. The two daughter mychites
move further away. — Fig. 18. The two daughter mychites approach the poles while the cell simultaneously elongates. — Fig. 19. After
reaching the poles, cell division occurs. The two daughter mychites are still flattened against each other (diplomychite).

The Mychites growth form is more common in cultures that have been in
existence for about two days. The last remnants of trophoconia can be dissolved
away by heating the coverslip smear with a 5% soda solution or by subsequent
treatment with alcohol and ether.
The division of the mych now takes place as follows, as will first be shown
using Diplococcus intracellularis (Weichs. 1887), the pathogen that causes stiff neck,
in the basal stage.
A normal mychite is shown in Figure 12. Figure 13 shows an ellipsoidal
widening of the mych. A further stage is shown in Figure 14; here, at the ends of the
rod-like, slightly curved, attached to the cell wall, two button-like swellings have
developed, the daughter mych. The rod connecting the daughter mych is the
mychomite. The mychomite including the button-like daughter mych is to be called
the mychozyg (= arch of the nucleus). Still in quite
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DiplomycHit. 69

At a relatively high distance, the mychomite in this species is reduced, as shown in Mg. 15.

This picture shows us for the first time a mychite which contains not just one mych, but
two, which, as always in division, are exactly the same size. In strict morphological terms, the
simple cell of the mychite has already progressed to a double structure, the dimychite. The
dimychite as a further unit will be discussed in more detail later. Since the dimychite is only a
transitional form here, it is not even considered in more detail.

As the process continues, the distance between the two daughter

mych, the mychostasis, more and more, as can be seen in Figures 16 and 17. This often results
in deviations from the spherical shape, which can appear very differently in the various species
and not infrequently give the appearance of a widening of the transverse diameter (perpendicular
to the mychostasis). Only then, with ever further lengthening of the mychostasis, does the cell
stretch in the axial direction and at the same time narrow (Fig. 18). The beginning of the
constriction and separation of the two cells from each other can take place before, with and
after the daughter mych moves into the poles of the cell. In the present case, this only occurs
after this process. The result is that in this, as in many other species, clear dimychites can be
found in large numbers, albeit with a short mychostasis (stenostat).

The resulting daughter mychites, as long as they are still flattened against each other,
represent a diplomamychite (Fig. 19). An example of a diplomamychite is Diplococcus gonorrhoeae
(Flügg).
This state can either be replaced by further development of both to a spherical shape,
but it can also persist. Both can occur in the present species.

Such a development in the mychite can be demonstrated for all bacteria in all stages of
the forms. It is found in Micrococcus as well as in Bacillus, Corynobaäerium, Microspira etc.

Very nice pictures of this kind are provided, for example, by the cholera pathogen, also by the
typhoid pathogen, Syncrotis buccalis (Rob.), many species of bacteria, etc. In many species, a
large amount of material must be examined in order to find suitable atrophy of the mychite, or
the disturbing trophoconia must be dissolved using soda (5%) and alcohol + ether. Interesting
pictures can be found in those individuals or species that show few trophoconia accumulations
without showing complete atrophy, especially when a more pronounced prolongation
of the mychostasis (eurystat) occurs.
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70 trophode.

Such a case is shown in Figs. 20 to 26 in the case of the cholera pathogen.

These pictures, which can easily be found, for example, in the germination
of gonidia among miotrophites and pliotrophites, show significant deviations
from the processes just discussed. Fig. 20 shows the mychite with typical
mych.
Here, a fine trophoconia sheath is found around the mych. In îiig. 21,
the daughter mych has already moved far away, although a connecting
thread is still formed. According to color and structure, however, this thread
is not a mychomite, but a fine trace of trophoconia accumulations, the
trophode. The formation of the trophode can be understood as follows:

20th, 23rd, 26th21st 22nd 24th 25th

Figs. 20-26. Microspira comma shot. 1886. Gonidal mychomitosis. Enlarge 10,000:1.
Fig. 20. Mychite with a simple mych. — Fig. 21. Pseudozyg (two daughter mych with trophoconia sheaths
and a trophode connecting them). — Fig. 22. The same, the daughter mych further away. — Fig. 23. The same,
the daughter mych reaching the pole. — Fig. 24.
Dim., the trophode fading in the middle. — Fig. 25. Dim., the trophode disappearing in the middle. — Fig. 26.
Dimychite without trophode.

If the mychomite has a trophoconia sheath, this will continue to exist for a
while after the reduction of the mychomite. The trophode is also always
attached to the wall, even if it often appears to be positioned differently, as
shown in some of the above illustrations. The resulting dumbbell-like arch
consisting of two daughter mychs with the connecting trophode is therefore
not a mychozyg, but is to be described as a pseudozyg. The mych moving
away from the original mych appears smaller than the first in the inserted
illustration ; this is, as morphological comparisons show, a phenomenon
that can be traced back to the difference in the thickness of the trophoconia
sheath; in reality, the two daughter mychs are completely identical after a
mychomitosis. The differences in the sheath are perhaps also due to the
greater consumption of nutrients by the moving mych.

In dividing Mych, the trophoconia may be evenly or very unevenly

distributed, so that one side of the dimychite may be provided with
trophosomes while the other is atrophic.
The pseudozygous has been observed several times in recent literature and is usually
referred to as a dumbbell or barbell-shaped structure.

As the cell continues to stretch, the daughter mych approaches the

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Mychit. 71

Pole in Fig. 22 more and more, to reach it in Fig. 23. The trophode also persists up to
this stage, in different individuals and species to different degrees. Fig. 24 represents
a subsequent stage in which the trophode begins to fade, especially in the middle.

In Fig. 25 the reduction of the trophode by further consumption of the trophoconia has
progressed even further, so that only two fine remnants are inserted at the two mych,
while the middle is broadly interrupted.
Finally, Fig. 26 shows the dimychite completely free of a trophode.
This example shows a longer mychostasis. This stage is followed by a constriction that
precedes the disintegration into two mychites.

In individuals that are heavily filled with trophoconia, none of these morphological
processes can be observed. In some species, too, no individuals can be found that
contain few trophoconia. In these, all such phenomena are completely concealed by
the trophoconia.
In such cases, individuals with few trophoconia are often preserved by creating
unfavorable conditions. This is also the case with starvation forms, which can be
produced in a hanging drop in physiological saline solution after a greater or lesser
number of days, but which usually no longer show any signs of division.

Especially in forms whose highest development lies in the alternation between

mychite and dimychite, i.e. in organisms that consist of only one mychite, such as the
genus Micrococcus, Streptococcus (here mychites in chains), etc., most species are
characterized by a strong ability to accumulate trophoconia. But even here, atrophites
are always found in older colonies.

Now and then there are species that are an exception to this, such as the Diplococcus
intracellularis mentioned above . But even among members of the genus Streptococcus one
sometimes finds species whose individual individuals in the chains take on very different colors
when stained with ordinary dyes. Some are very dark and full of trophoconia, others are almost
completely uncolored. With a little attention, all transitions are easy to detect. The chains, which
contain individuals with strong and pale colors, mixed in many different ways, usually take on color
in part when stained with Gram, and in part they do not, and a colorful picture is created.

Such a species was cultured several times from tonsil swabs, a Streptococcus
spec., of which two chain pieces are shown in Fig. 27. The same pictures were seen
in this species as those described for Microspira . All stages are mixed in a colorful
series. In the two chain pieces shown, one can see both
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72 Mychomerite, triplomychite.

Fig. 27. Streptococcus spec., from tonsil smear. Two chain pieces. Magnification 10,000:1.
Mychite (pliotrophic and miotrophic), mychite with pseudozyges, dimychite.

Trophodes, simple mychites without division phenomena, miotrophic and also

pliotrophic, dimychites etc.

3. The Mychotner and the Mychomerit.

During the formation of the sex cells, the mych is reduced to a half-size,
the mychomere. The cell with one mychomere is a mychomerite. .

This includes gonit, spermite and oit, which are found in

V ÿ 1, V ÿ 2 and ÿ ÿ 3 are treated.

4. The Triplomychite.
The triplomychite is a growth form in which three mychites are arranged
radially, close to each other and flattened against each other. They do not form
a morphological unit.
The three mychites can be of equal size, but usually one of them is slightly
larger, as shown in Fig. 28. Its formation probably takes place mostly from a
diplomamychite, with one of the two mychites dividing perpendicular to the axis
between the two mychites before the other mychite becomes isolated or rounded
off to a spherical shape.
Triplomychite does not have any special significance.
It is generally rare. It also occurs temporarily in some Micrococcus species, but
also occasionally in morphologically lower developmental forms of higher bacilli.

That the triplomychite may also sometimes arise from a dimychite, in which
one mych is perpendicular to the mychostasis

28th 29th

Fig. 28. Barcina spec. Triplomychite. Enlarge 1:10,000. — Fig. 29. Dipbcoccus intraeellu-
(Weichselb. 1887). Dimychite, by nuclear division of one mych across the mychostasis
with 3 mych arranged in a triangle. Magnification 1:10,000.
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Pliomychit. 78

Occasional findings, such as that shown in Fig. 29 for the stunted neck pathogen,
seem to support the idea that the cell has doubled in size after the division, i.e.
from a single cell with 3 Mych.

B. The Pliomychite.
Any bacterial individual that contains more than one mychite, i.e. deviates
from the usual spherical shape of the mychite, is a pliomychite.

As we have already seen in the previous section, the first step of the majority
is the union of two mychites into a single unit, the di-mychite. However, while the
dimychite discussed was a labile entity and was only a very temporary
manifestation of the mychite during the division into two mychites, the dimychite
appears as a solid organic unit during the phylogenetic ascent as the first stage
of the association of two mychites. This unit is the building block from which all
higher bacterial forms are constructed and composed.

The union of two dimychites to form one individual is the didimychite, of

more than two such building blocks the syndimychite.
Examples are all those bacteria that can develop beyond the spherical
shape as their main growth form.
The two or more mychites united to form a single bacterial individual have
no boundary walls between them, but only a common shell. The fact that the
uniting of mychites in this tiny dimension involves completely different mechanical
conditions than in the case of the much more highly organized cells of the protozoa
and metazoa, etc., should fully explain this fact.

1. The Dimychit.
The dimychite is the union of two mychites into a single cell. The two
mychites are located close to the two poles of the elongated cell.
The dimychite is a morphological unit and the building block for all higher levels of
bacterial organization. In the association of dimychites, each of these building
blocks is appropriately designated by a special name: dimychosis. Likewise, a
mychite in the association within a dimychite or dimychosis is designated by
mychosis.
While in mychite the mych is wall-aligned, in dimychite it is apparently not
always wall-aligned, but is often more or less offset from the pole. Fig. 39 would
show this, but such images are not absolutely conclusive, since the mych does
not have to be exactly polar, and in fact often is not polar, and this is therefore
only an optical phenomenon. In contrast, such images sometimes appear so
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74 Dimychit

regularly and exclusively, so that in such cases a wall attachment is excluded.

Since the pure mych without a trophoconia sheath can no longer be determined
microscopically in the dimychite, and also not in the atrophite, a fine trophoconia
sheath is necessary and most useful for determining its position. The trophosomelles
thus indicate the position of a mych with considerable accuracy. But the
trophosomes also localize at least the location of a mychite, so that experience
shows that the location where the mych is found can be fixed at least approximately
precisely. In the case of the pure atrophite (Fig. 40), one has to rely entirely on
comparative morphological aspects.

The distance between the centers of the mych in a dimychite or in a

dimychosis is the mychostasis. This is of the greatest importance for comparative
morphological investigations and must be compared again and again for these
purposes in order to arrive at a reliable idea of the structure and the morphological
relationships, especially to the unit (the dimychite

Fig. 30 and 31. Microspira comma shot. Two dimychites with trophosomelles.
Magnification 1:10,000.
Fig. 30. Stenostates dimychit (during division of gonidia and basites). — Fig. 3L Eury-states
dimychit (during germination of gonidia or phytite).

and dimychosis). Even in dimychites with delicate trophosomes, mychostasis can

still be detected with certainty.
The mychostasis can be short (stenostat), as shown in Fig. 30 for a
myotrophite of a dimychite of the cholera pathogen and as occurs in the division of
gonidia and basites, or it can be long (eury-stat). One such is also shown in Fig.
31 for the cholera pathogen as a di-mychite, which is common in phytite and also
occurs in gonidia germination. In general, one can say that the shorter the
mychostasis, the more the mych are mural and vice versa.

The axial length of a bacterial rod is the median length of the entire rod
without regard to the position and number of its mycocytes.
At this point, attention should be drawn to phenomena which can very easily
give rise to gross deception, although these are higher orders which will be dealt
with in a later section. In order to avoid these pitfalls, this should be mentioned
here.
These are long rods, which at first glance appear to be dimychites due to their
terminal trophosomes (cf. Fig. 33), but
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myochostasis. 75

are noticeable by their greater length. Since there are indeed sometimes
very strongly stenostatic dimychites and dimychoses, a reliable
morphological interpretation is only possible by detailed comparison with
other individuals of the same culture, combined with comparative
measurements. Thus, Fig. 32 represents an ordinary dimychite with two
trophosomes from the typhus pathogen, while Fig. 33 probably simulates a
dimychite, but belongs to a longer group, namely the group of two
dimychoses (Didimychite), in which, however, the two middle mychs do not
betray their presence due to the lack of a trophoconia sheath and are even
completely invisible in the stronger and more rich cytoplasm.

32nd 33rd

Fig. 32 and 33. Acystia typhi (Eberth 1880). Enlarge 1:10,000.

Fig. 32. Dimychite with 2 trophosomes from a young culture. ·— Fig. 33. Didimychite with 2 telotrophosomes
from a four-day culture on agar with 654% sodium chloride.

The mychostasis of Dimychites is only slightly shorter than its axial

length.
In every dimychite, every mych can be uncovered, i.e. invisible, or it
can be concealed by trophosomelles or a trophosome, thus making its
presence noticeable. There are all transitions here. The extreme on the
strongly stainable side is the short rod (dimychite) completely filled with
trophoconia, which also no longer allows any trophosomes to be seen,
since everything is uniformly dark and takes on the color strongly, and on
the weakly stainable side is the completely atrophic dimychite.

Regarding the distribution of trophosomes and trophosomelles on the dimychite, there are
only the following possibilities: 1. 2 trophosomes (Fig. 37);
2. 1 trophosome + 0 trophosomelle
(Fig. 38); 3
.1 " +1
.0 " +1 " (Fig. 39);
4 5. 2 trophosomelles; 6.
without trophosomes and without trophosomelles (Fig. 40).
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76 Mychomitosis of Dimychites.

Some of these forms in the distribution of the reserve apparatus are illustrated using
developmental forms of Corynöbacterium pseudodiphtheriticum (Lehm, et Neum. 1899) (Fig. 34 to 40).

We see that even within a dimychite a division of labour can take place with regard to the storage
of reserve substances. However, these differences can also be attributed in part to differences in the
life activity within the dimychite, as a result of which the reserve substances are used up faster in one
place than in another.

other places.

That this division of labour can acquire even greater significance in higher associations, we will
see later in the Syndimychit and especially

#
34th 35th, 36th 37. 38. 39. 40.
Figs. 34-40. Corynóbacterium, pseudodvphtheriticum (Lehm, et Neum. 1899). Enlarge 1:1 0 000.
Fig. 34. Mychite pliotrophic. — Fig. 35. Mychite miotrophic. — Fig. 36. Dimychite formation from the gonidia.
— Fig. 37. Dimychite with 2 trophosomes. — Fig. 38. Dimychite with 1 trophosome. — Fig. 39. Dimychite with 1
trophosomelle. — Fig. 40. Dimychite, atrophic, without trophosome and without trophosomelle.

in the formation of gonidia, cystites, sporites and oidia. Then certain dimychoses take over the nutrition
of these reproductive and permanent forms etc. These dimychoses are the trophodimychoses. Whether,
alongside this nutrition of the reproductive and permanent forms etc. by trophodimychoses, a separate
nutrition also takes place at the same time, could not be found either for or against.

2. The mychomitosis of dimychit and dimychosis.

Since in dimychite the mych is not recognizable as a morphological unit (not as a division
phenomenon of the mychite) even in atrophites due to the denser cytoplasm and its thicker layer,
because the color difference is too insignificant compared to the size of the mass, the mychomitosis in
its actual processes in dimychite can no longer be directly recognized with the tools available to date.

In contrast, the subsequent phenomena, especially the formation of pseudozyges (dumbbells)

and the formation of trophosomes, are excellent moments for the morphological understanding of growth
phenomena and division phenomena. Very small trophosomes, which are formed by only a very thin
layer of trophoconia, are particularly suitable for assessing these questions, firstly because
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Mychomitosis of Dimychites. 77

they fix the presence and location of the mych sufficiently through their delicacy and often even
present a size only slightly larger than the mych itself, and secondly because their extraordinarily
strong colorability means they represent a happy mediation of the morphological conditions for the
eye. Since experience shows that the formation of trophodes and pseudozyges follows mychomitosis
and only ever occurs during division phenomena, this phenomenon provides a good standard for
the morphological analysis of growth and arthrogony. Only in higher dimychosis associations is one
phenomenon to be observed, namely the formation of pseudotrophodes, which are trophode-like
trophoconia accumulations between two trophosomes or trophosomelles, which belong to different
dimychoses. One such pseudotrophode is ÿ. This is illustrated in Fig. 179 (section VI a A 5 a) in the
syn-ascite of Spirillum undula (Müll. 1786) between trophosome 9 and 10. In more primitive
dimychosis associations, the danger of confusion with the trophode is not to be taken into account
because the neighboring mych of different dimychosis are usually much closer together than the
mych belonging to one dimychosis, and if a larger accumulation of trophoconia occurs at this point,
a thread-like connection never develops, but both mych belonging to heterogeneous dimychosis are
then enclosed by an oval or roundish trophoconia accumulation, which, especially in the latter case,
can be confused with a trophosome, although in reality it represents two fused trophosomes.

To recognize all the morphological stages that lie between a dimychite and the formation of
two dimychites from it, species that are naturally prone to atrophosis or myotrophosis are naturally
particularly suitable, unless one wants to resort to chemical interventions. Since many species, in
addition to rich trophoconia material, which often fills the entire cell to such an extent that even the
trophosomes cannot be recognized, also have lipoid substances evenly distributed in the cytoplasm,
which also take on the color strongly, all such bacterial species that are richly equipped with reserve
substances and are usually very vigorous even outside the host body are not suitable for this purpose.

Species prone to atrophosis or myotrophosis are not common.

However, such species are sometimes found in sputum and tonsil smears. The findings shown in
Fig. 41 to 48 come from such a species cultured from sputum. Fig. 41 shows a pliotrophic dimychite,
i.e. a cell individual which is so richly filled with reserve substances (trophoconia, lipids, etc.) that it
has a similar coloration.
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78 Mychomitosis of Dimychites.

and neither trophosomes nor trophosomelles can be seen. In mychomitosis,

which seems to begin in this state, food intake is probably completely eliminated,
since individuals in the process of dividing use up their reserves and so
myotrophosis and finally atrophosis occur following the division processes and
as a result of the associated energy consumption. Fig. 42 shows the beginning
of the consumption of reserves, Fig. 43 even more so. Here, the denser and
chemically often different inner layers of the trophoconia shells stand out clearly
in the fading trophosomes, so that trophosomelles can already be spoken of here.

41st 42nd 43rd 44th 45th minute 46th minute 47th minute 48th

Fig. 41—48. Bacterium spec., from sputum. From agar culture. Magnification 10,000:1.
Fig. 41. Dimychite, pliotrophic. Densely filled with reserve substances, so that the trophosomes are not recognizable. — Fig. 42. Dimychite. The 2 trophosomes
become clear. — Fig. 43. Dimychite. The 2 trophosomes reduce their size and become less dense, so that trophosomelles can already be seen. — Fig. 44.
Dimychite, miotrophic. The 2 trophosomelles stand out sharply; the upper one forms a pseudozygous. — Fig. 45. Cell individual, miotrophic. The upper
pseudozygous fully developed, the lower one forming. — Fig. 46. Dimychite, miotrophic. Both pseudozygous fully developed. — Fig. 47. Dimychite, miotrophic,
again with somewhat more reserve substances. The upper pseudozygous fading in the middle of the trophode; beginning of the separation of the two dimychites
from each other. — Fig. 48. The 2 resulting dimychites at the beginning of pliotrophosis, both still connected, but shortly before separation. Each with 2
trophosomes.

Fig. 44 shows the formation of a pseudozygous, starting from the upper mych,
after the entire cytoplasm has faded considerably as a result of continued
consumption of reserves.
In Fig. 45 the upper pseudozygous is complete, the lower one does not appear
to be complete yet. The trophodes are strongly curved in this species and thus
allow the path of the daughter dimychi to be recognized. Fig. 46 shows the two
pseudozygous completely, now the entire cell individual gradually begins to
stretch. A fading of the trophode through consumption or distribution of the
trophoconia can be seen in Fig. 47 in the upper pseudozygous ; the fading begins
most strongly in the middle. At the same time a gradual constriction of the two
daughter dimychites that have been created begins and a renewed accumulation
of reserve substances begins. In Fig. 48 this is even further advanced, the two
daughter dimychites that are still attached to one another but are already strongly
constricted from one another have
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Isozygy, protozygy. 79

the trophodes are completely lost, and the four mych that are now present are represented by four
trophosomelles. Then even more reserve material accumulates as a result of the full ingestion of
food, the pliotrophites fall apart completely, and the process described repeats itself again.

Such processes can easily be observed in other species in a similar way through findings of
all these stages and their transitions.
In the species under discussion, the processes described are also usually simultaneous in
both mych. Such a simultaneous occurrence of mychomitosis and the subsequent processes in
both mych of a dimychite or a dimychosis is called isozygy. The non-simultaneous course described
above, which is not so common — at least in the dimychite—but is a quite common phenomenon in
higher groups, is protozygy.

If the separation of two dimychites, which have arisen from a dimychite by mychomitosis of
the two mych, does not occur, but rather remains firmly united, then the result is a di dimychite. The
two morphological building blocks corresponding to the dimychites, from which the didimychite
arose as a phylogenetically higher association, are to be referred to as dimychoses in this
association, as has already been suggested. If one wants to make visible in species which, as a
result of their plio-trophic predisposition, conceal all these morphological features under rich
accumulations of reserves, there are two ways of removing them.

One way is the artificial creation of starvation forms, which reveal many morphologically
important findings. For this purpose, bacterial material is placed in a hanging drop of physiological
saline solution and placed in a hollow slide under a tight seal for a greater or lesser number of days
in the dark. When such material is stained, it shows that a large part of the reserves have been
used up and the obstacle to the penetration of the eye has been removed. Such preparations give
very beautiful pictures, e.g. of cholera.

pathogens.
The other method is based on chemical interventions. The best way to release the lipids is
to use a mixture of alcohol and ether, into which the air-dried or alcohol-hardened coverslip smear
is placed. Here too, the cholera pathogen can be cited as an example, which then clearly shows
the trophosomes, which would otherwise be hidden by the abundant lipids.

A large part of the components of the trophosomes, especially the nucleic acid compounds,
can be removed by a 5% soda solution.
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80 Didimychit.

To do this, the soda solution is heaped onto the air-dried and fixed cover glass smear
and then heated several times over a small or large gas flame*, either slightly or
strongly. A few longer breaks are advisable in between. In individual cases, only the
experiment and a comparison of the then colored preparations will give the correct
result. ÿ. For example, this method is very useful for morphological diphtheria
investigations, since many morphological details can only be recognized in this way,
especially since the diphtheria trophosomes are often very large and the gonidia and
cystites otherwise conceal everything.

For some purposes, especially for the diphtheria pathogen, a combined

procedure is recommended. In this case it is advisable to first use soda and then
dissolve the lipids, because the bacterial layer easily peels off the cover glass.

3. The Didimychit.
The didimychit is the union of two dimychoses to form a higher association
and an individual. "As will be explained later, the German name Zelle can certainly
be used for the my chosen of all lower and higher associations, as well as for the
latter themselves, since scientifically this cannot be regarded as a uniform
morphological term, and this collective name is also used for protozoan and
metazoan cells.

Right at the beginning we should point out the possibility of differences in the
arrangement, which are already established in the association of two Dimychoses
and which become of outstanding importance in the construction of higher associations.
Fig. 46 (IV ÿ 2) shows that the two middle mychs are not located one behind the
other in the longitudinal axis of the cell, but that a considerable deviation from this is
noticeable. However, while in the division process of the dimychite this deviation in
the orientation of the mychostoses from the cell axis is quickly compensated for by
the fact that, as Fig. 47 clearly shows, these two middle mychs move into the axis,
in the dimychite differences of fundamental importance arise through the fixation of
one or the other state. According to this, the arrangement of the dimychoses in the
didimychite can be: 1. catatact, 2. syntact.

The catatactous arrangement of the dimychoses in a didimychite or a higher

association (syndimychite) is the arrangement of these in a row one behind the
other, so that the mych and the mychostases lie in the longitudinal axis of the cell.

The syntact storage of the dimychoses in a didimychite or a higher association

(syndimychite) is such an arrangement that the mych and the mychostases are not
arranged one behind the other in the longitudinal axis of the
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Didimychit. 81

Cells are not arranged in the same way, but rather they are arranged irregularly,
obliquely or transversely to the longitudinal axis, either some of the dimychoses or
all of them, or two or more mychostases are arranged next to each other in the
longitudinal direction, either parallel to the longitudinal axis of the cell or mostly in a
disordered manner. Here, compare Section VI ÿ A 5, ÿ. B. also Fig. 182.
While in the dimychite the possibility of the arrangement of trophosomes
and trophosomelles was quite limited, in the catactous didimychite it reaches
a quite significant number, even if all transitions and intermediate forms are
disregarded.
First , the possibilities for the arrangement of trophosomes are
summarized using some Corynobacterium pseudocliphtheriticum (Lehm, et
Neum.) 1899 isolated from a tonsil swab.

49. 50. 51. 52. 53. 54.· 55. 56. 57.

Fig. 49—57. Corynobacterium pseudodipMheriticum (Lehm, et Neum. 1899). Various possibilities
of the arrangement of the trophosomes in the didimychite. Magnification 1 :10 000.
Fig. 49. Atrophy. No trophosomes. — Fig. 50. 1 telotrophosome. — Fig. 51. 2 telotrophosomes.
— Fig. 52. 1 ascotrophosome. — Fig. 53. 2 ascotrophosomes. ·— Fig. 54.
1 telotrophosome and 1 ascotrophosome. •— Fig. 55. 1 telotrophosome and 2 ascotropho-somes.
— Fig. 56. 2 telotrophosomes and 1 ascotrophosome. — Fig. 57. 4 trophosomes (2 telotrophosomes
and 2 ascotrophosomes).

can occur. In Fig. 49 to 57 these are shown in consistently observed findings ;

only one possibility is not shown here.
There are a total of 10 possibilities. The atrophy shown in Fig. 49 lacks any
trophosome. A terminal trophosome (telotrophosome) is noted in Fig. 50, two
telotrophosomes in Fig. 51.
In the middle part there may be one (Fig. 52) or two trophosomes
(ascotrophosomes) (Fig. 53). One telotrophosome and one ascotrophosome
may be distributed, as shown in Fig. 54, or both may lie close together (not
shown). Three trophosomes can only be arranged in the two possibilities
shown in Fig. 55 (1 telotrophosome and 2 ascotrophosomes) and 56 (2
telotrophosomes and 1 ascotrophosome), and the presence of all four
trophosomes is shown in Fig. 57.
In the same way, trophosomes can be distributed and finally trophosomes
and trophosomes can be found mixed together in the most diverse ways. In
the case of Didimychit, there are already 46 possible combinations of the
latter. With a little patience, one can easily
Enderlein, Bacterial Cyologeny. 6
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82 Didimychit.

All these possibilities can be observed in the material and countless transitions
can be found which often appear quite different in the different species but
nevertheless have the same morphological basis.
Some further examples of this kind are taken from a species of the
genus Bacterium isolated from a tonsil swab and chosen at random from
numerous other combinations. In Figs. 58 to 62 they are shown exactly as they
were in the original. Fig. 58 shows 1 telotrophosome and a terminal
trophosomelle, Fig. 59 2 terminal trophosomelles. In Fig. 60 the presence of
all 4 Mych is made visible to the eye by trophosomelles, in Fig. 61 by
trophosomes, which are finally replaced in Fig. 62 by richer accumulations of
reserve substances.

psr·· ; .·•-"
k'iOi* . . I »·

58th 59th minute 60th 61st 62nd

Figs. 58-62. Bacterium spec, from tonsil swab. Didimychite. Enlarge 1:10,000.
Fig. 58. 1 telotrophosome, 1 trophosomell. — Fig. 59. 2 trophosomells. — Fig. 60.
4 trophosomelles. — Fig. 61. 4 trophosomes. — Fig. 62. Pliotrophite, which just shows the 4 trophosomes.

are quite heavily concealed. Of course, all possible combinations can be found in this species too,
including atrophites, pliotrophites with even stronger accumulation of reserves, etc. Pliotrophites are
naturally present in the majority, as is usually the case.

The findings to date already confirm the knowledge that the extraordinarily
diverse appearance of the rods of a single species, as they so frequently occur
within morphologically similar structures, here of Didimychites, does not
indicate any comparative morphological differences, but that they are only an
expression of the different accumulations of reserve substances in the various
morphological units of a higher association.

As with dimychite, with didimychite too, after some practice and with
comparative measurements, it is easy to determine in most cases, using the
trophosomes and trophosomelles as an aid, that didimychite is composed of
4 mychites and thus must contain 4 mych.
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syndimychit. 83

4. The Syndimychit.

Syndimychi t is the union of more than two dimychoses

to a higher organization and to an individual.

As with the Didimychit, the difference in the arrangement of the di-mychoses is even more
apparent here. The tactual and syntactical arrangement is much more obvious and of even greater
importance.

The catactous syndimychit arises from the didimychit by further division of the mych of one or
both dimychoses and simultaneous arrangement of the resulting new mych in the longitudinal axis of
the rod. Further divisions of this kind can lengthen the rod more and more, and it can expand to form
extraordinarily long threads.

As we have already learned from the simple association of two Dimychosen, in the case of
Syndimychit the two associated

Fig. 63. Bacillus Solmsi Klein. Piece of a Syndimychites with 4 Dimychoses.

Magnification 1:10,000.

Mych of one dimychosis are often further apart than two mych of neighboring dimychosis. Again,
forms with numerous trophosomelles offer suitable preparations for making such a structure visible.
Since a favorable stain for trophoconia is methylene blue, it is advisable to use this stain for this
purpose. Fig. 63 shows a selected piece from the syndi-mychite of Bacillus Solmsi Klein 1889, which
forms rather long threads. The entire thread consisted of 10 dimychosis.

In many forms, however, the dimychoses are arranged in such a way that the mych are
approximately equally spaced. In some, the two mych of the dimychoses appear to be closer to each
other than to the neighboring dimychose. Species whose catactous dimychite threads are not of
uniform thickness also show a certain difference in the arrangement of the dimychoses of a
syndimychite.

Such formations are frequently found, for example, in the diphtheria pathogen (Fig. 64 and 65).
However, it must always be remembered that atrophic mychosis can occur within every dimychosis.

The mychoses can, as in lower associations, also in the syn-dimychit, depending on the
amount of stored reserve substances, have a very
6*
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84 Catatactous syndimychit.

64th

65th minute

,Fig. 64. Corynobacterium dipMheriae (Loffi.). Syndimychite. Enlarge 1:10,000. — Fig. 65.
'The same syndimychite, composed of 12 mychoses, i.e. 6 dimychites. Magnification 1:10,000.

They can present a variety of appearances and be arranged in the most varied ways. Atrophic
mychoses, mychoses with trophosomelles and with trophosomes, etc. can follow one another in
any order. Fig. 64 and 65 show such arrangements. For example, 4 and 10 in Fig. 65 are atrophic
mychoses, so that this diphtheria rod is made up of 12 mychoses. The first and eleventh, both more
strongly thickened mychoses, are structures that will only be described as cystites in the following
section.

Fig. 66. Treponema Vincenti (Blanch. 1906). Syndimychit with trophosomes and trophoso-
mellen. Magnification 1:10,000.

Another example from the area of catactous syndimychites is taken from the spirally coiled
bacteria. Fig. 66 shows a specimen of a Treponema Vincenti (Blanch. 1906) taken from a tonsil
swab taken during a case of Angina Vincenti, a disease caused by the above-mentioned spirochete
in association with Fusijormis hastilis (Seitz). This rod is also morphologically a catactous
syndimychite. In favorably colored, more trophic specimens, trophosomes and trophosomelles can
also be found.

In the catactous syndimychite, the terminal trophosomes are also called telotrophosomes
and the others ascotrophosomes.
The biomorphological term ascite mainly includes catatact syndimychites. Further information
on catatact syndimychites can be found under VI a A 4.

The syntactic syndimychite comprises associations of more than two dimychoses to one
bacterial individual, in which the mychostases are all or partly arranged obliquely or transversely to
the longitudinal axis of the rod. The formation of such forms is due to the fact that the
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Syntactic syndimychit. 85

Fig. 67. Migulanum anfhracis (Koch 1876). Piece from the middle of a syntactic syndimychites (Synascites)
from a three-day culture on agar with 6% Na Cl. From the border between catactic and syntactic syndimychites.
Magnification 1 :10 000.

Mychomitosis of the individual Mych is then not arranged along the longitudinal axis of the rod, but
obliquely or perpendicular to it.
As an example, a piece from a thread of the anthrax pathogen is shown here (Fig. 67), which
comes from a three-day culture on agar with 6% sodium chloride. On the left is another piece with a
catatact arrangement.

The formation of the mychoses is visible, while the syntactic storage of the dimychoses follows. The storage of the mychoses

is characterized by the trophosomelles. Incidentally, some specimens of the anthrax pathogen shown by LÖFFLEE (1887 ,

Plate II , Fig. 4 a) also show a similar organization; however, LÖFFLEE considered the trophosomelles to be "a deposit of

the coloring agent in the form of fine grains"; LÖFFLER believes that insufficient coloring is to blame for these specimens

being considered abnormally colored. He says : "The coarser and finer graining is due to the bacilli not being colored for

long enough and being washed out in alcohol for a long time."

The fact that catatact and syntact syndimychit can be found simultaneously in one specimen is a
more or less frequent or rare occurrence in all Synascota. The transition is either gradual or sudden.
During division, one half of the part may occasionally appear catatact and the other syntact. PROWAZEK
1906 depicted such a specimen of Treponema gallinarum Bl. 1905, which is reproduced here in Fig. 68.

The syntactic syndimychite can also accumulate such a large number of dimychoses across the
width that gigantic bacterial individuals can arise, such as those found in Syncrotis Enderl.,
Phragmidiothrix Engl, and, in the most extreme case, Schaudinnum Enderl. A series of illustrations

Fig. 68. Treponema gallinarum R. Bl. 1905. On the left, syntactous, on the right, catatactous syndimychite
shortly before division. (After: PROWAZEK, Arb. Kais. Gesundheitsamt 23. 1906, Taf. II, Fig. 10 b.)
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86 Syntactic syndimychit.

genes of syntactic syndimychites can be found in the section on syn-ascite

(VI ÿ A 5 a).
In such huge syntactic syndimychites, individuals with very numerous
trophosomelles are not uncommon. Such specimens have a granular
appearance. They gave Schaudinn (1902) and others the reason to attribute
a "diffuse" nucleus to the bacterial cell, due to the chromatin substance
distributed over the entire cell, which according to the prevailing view
represents the nuclear substance.
That in higher dimychotic associations, such as the catactous and
syntactous syndimychites, here and there, in addition to the typical di-
mychoses, individual mychoses can also be interspersed between the
structure is understandable if one considers that isozygy does not always
occur in the two mychs of a dimychosis, but that individual mychoses can
remain through protozygy, which sometimes have a longer existence than
mychoses. Thus, ÿ. For example, in the syndimychite of Spirillum undula
(Müll., 1786) in Fig. 181, which apparently consists of 3 dimychoses, there
are 2 individual mychoses in the middle of the rod, which apparently remained
because the two end mychoses of the dimychite shown in Fig. 180 developed
into dimychoses by protozygy, while the two middle mychoses are likely to
represent dimychoses, even if they are capable of giving the appearance of a
dimychosis.

C. The Symmych and the Symmychit.

The symmych is a polydynamic (multi-valued) mych. Its content
corresponds not just to one, but to several mychs united into a single
homogeneous body. Its diameter is larger than that of a mych.

The symmychit is a mychit with a symmychon.

All the types of bacterial cells discussed so far, whether composed of
one or more mychoses, were always monovalent mychs. Division of the mych
only ever occurred when it was able to grow through external nutrition or by
means of reserve substances from the cytoplasm. Observing a simple division
of one dimychite into two under the microscope, even under favorable
nutritional conditions, is a task that requires patience.

However, if living material is placed without any nutrients, for example in a

hanging drop of physiological saline solution, all growth is quickly stopped.
Even all those phenomena that otherwise occur very quickly and easily under
unfavourable conditions, such as the formation of gonidia, are completely
eliminated, and it can easily also
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Symmychon, Symmychit. 87

After a few days it can be determined that growth has not progressed, but that each
bacterial individual has used up some or all of its reserves. Such forms of starvation
often provide good objects for comparative morphological investigations.

The first form in which I noticed a striking, completely different behavior was
the ÿ oit of the cholera pathogen. A more detailed description of the zoite and
pseudascite follows in section VI a A 7 and 8.
Elsewhere it is pointed out that zoites are only found in cultures that originate
from cholera stools or that have not been cultivated for very long. If such material
with zoites is placed in hanging drops of physiological saline solution, which excludes
any nourishment of the material, and the zoites, which only have a single nucleus,
are observed, it can be seen that they grow into long threads (pseudascites) in less
than 2 hours, while the dimychites and di-dimychites, which contain 2 and 4 mych
respectively, do not elongate. If the one nucleus of the zoite were equivalent to the 2
and 4 nuclei of the cholera rod, there would be a much greater probability of the rod
stretching out into a long thread than with the zoite.

However, as mentioned, this is by no means the case.

However, if you continue to observe the pseudascite, you will soon see that it
splits into about 10-15 or more didimychites, which remain attached to one another
in a chain-like manner. This process is completed after another 2 hours.

How is it possible that under completely identical conditions 40—60 or even more nuclei
of the resulting

formed didimychite chain can develop in 3-4 hours, and at the same time the 2-4
nuclei (mych) of the normal cholera rod do not experience any proliferation?

The only possible explanation for such findings is the assumption of a polyvalent mychia, an
assumption which gains even more ground in the fact that very similar phenomena can be
demonstrated in certain protozoa, as has been proven beyond doubt by the excellent investigations
of HARTMANN (1911) . HARTMANN calls such nuclei polyenergid nuclei.
(1909 ,

For reasons of comparative morphological nature, which will be further

developed in Section IX C, it is not appropriate to transfer the concept of polyenergid
nucleus to bacteria, and the term polydynamic Mych or Sym-mychon shall therefore
be used for the polyvalent Mych.

When it was later discovered that another symmychon, namely that of the
cystites of the diphtheria pathogen, had a volume
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88 Monogony, arthrogony.

is significantly larger than that of the Mych, significant concerns could no longer be raised.

A mychite with a symmychon as its nucleus is a symmychite . The

symmychites include the cyst it and the ÿ o it, which will be discussed in more detail
in sections VA 3 a ÿ and VI ÿ A 7.

V. Reproduction of bacteria.
Omne vivum ex ovo.
Harvey 1651.

The reproduction of bacteria is divided into asexual (monogony and

arthrogony) and sexual reproduction.
Since sexual reproduction occurs only once within a complete cyclode (cf. VI),
it is not very conspicuous and is not easily recognized. It seems certain that sexual
reproduction occurs in all bacteria.

Asexual reproduction is so extraordinarily prominent due to its enormous

productivity that sexual reproduction has so far been completely overlooked.

A. Monogony and arthrogony of bacteria.

The asexual reproduction of organisms is divided into monogony and
vegetative reproduction by budding followed by division.

Monogony can be divided into two groups: 1. primary monogony, 2.

secondary monogony.
Primary monogony is based on simple cell division following nuclear division,
as occurs in protozoa, for example. A subgroup of this is frequent nuclear division
and subsequent disintegration into numerous cells, which SCHAUDINN (1899)
called schizogony (for example in sporozoa).

Secondary monogony includes the phenomena that arise through the

regression of fertilization processes (the second directional body is reabsorbed by
the egg cell and acts like a spermatozoon by replenishing the number of
chromosomes) and includes parthenogenesis and paedogenesis.

The latter is not an option for bacteria, but primarily monogony is. However,
this only occurs in mychites,

!) cf. Haecker, Fertilization. Handb. Natural Sciences, Vol. I, 1912, page 907.
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Isomorphic arthrogonia, 89

because only here does a division into two individuals occur following nuclear division.

This is different for all higher associations and also for the dimychite. Here, a division of the

mych into two is by no means followed by a separation of the two newly formed mychoses (mychites),
but a subsequent disintegration always takes place in a completely different place, as was clear from
section IV ÿ. For example, in a dimychite (after the division of the two mych into four), the two newly
formed dimychoses are separated from each other. This division and multiplication is therefore not
monogony at all, but multiplication by disintegration into individual higher or lower cell associations.
This phenomenon can therefore be classified in the group of vegetative reproduction. For this type of
reproduction and multiplication I use the term: arthrogony ]).

The products of arthrogonie can be of equal size, similar shape and morphologically
equivalent, or they can be of unequal size, different shape and morphologically different
value.
The former phenomenon is the isomorphic arthrogonia, latter
called the heteromorphic arthrogoni e.
Every individual created by arthrogony is a species hr o gone.
The contrast between monogony and arthrogony clearly highlights that the
division and multiplication of spherical bacteria (ÿ. B. of Micro-coccus) has,
comparatively morphologically, a completely different significance than the multiplication
of rod-shaped bacteria (ÿ. B. Bacterium).

1. Monogony.
Reproduction by primary monogony occurs among bacteria in all monomychotes
and in the basal stage of all dimychotes
instead of.

More detailed information on mycomitotic processes can be found in Section IV A 2

Information to which reference is made here.
The products of primary monogony can remain loosely attached to one another,
so that more or less long (2 to numerous) chains (desma) can be formed. For further
desma formation, see VA 2.
As an example of the desms created by monogony, the
genus Streptococcus should be mentioned.

2. Isomorphic arthrogony.
The isomorphic arthrogony of bacteria is the asexual reproduction by
disintegration into two cells or cell groups, in which both partial products are of the
same size, shape and morphologically similar.
cf. Enderlein (3) 1916, page 404.
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90 Heteromorphic arthrogonia.

The decay never separates mychoses that are products of one of the
mychomitoses of a single mychosis preceding the decay; the only exception is
the decay of dimychite to 2 mychites.
The section between two manifestations of arthrogonia
is a generation of the asexual form of the bacteria.
Earlier authors mistakenly used the term "generation" in bacteria to refer to
the arbitrary term for a passage through agar (or other nutrient medium) of one or
more days. The term "genode" is suggested for this term.

The most common reproduction in short (1 or a few

days) genodes is the isomorphic arthrogonia.
If an organism ÿ. B. grows on agar as a dimychite (ÿ. B. pathogen of typhus,
cholera), after mychomitosis a didimychite is formed, which then decomposes
again into two dimychites.
If a longer rod develops, e.g. a didimychite, then after mychomitosis of all 4
mych it forms a syndimychite with 4 dimychoses relative to 8 mych, which then
breaks down again into 2 didimychites.
A long thread-like syndi-mychite consisting of many dimychoses can,
whether only some or all of the mych are multiplied simultaneously by mycho-
mitosis, disintegrate into two equal-length fragments.
Isomorphic arthrogony also occurs when the resulting arthrogony does not
completely fall apart but remains loosely attached to one another. Long chains
can be formed by repeated arthrogony of this kind. The term desme should be
used for these chains, whether they are chains of two or longer, but it should also
be used for the chain formation of the globules formed by monogony (cf. page
180). The formation of desmes is part of the specific character of some species in
certain stages of development, and in the case of monomychotes it is also partly
part of the generic character (eg Streptococcus).

3. Heteromorphic arthrogonia.
The heteromorphic arthrogony of bacteria is the asexual reproduction by disintegration into
two or more parts that are of unequal size, shape and morphologically unequal value. Here too,
disintegration never separates mychoses, which are products of a mychomitosis of a single mychosis
preceding the disintegration, apart from gonidia.

Among these phenomena there is a surprising variety

diversity of shape, size and morphological structure.
The colorful abundance is initially divided into three groups: 1. Fruit-
r>ÿ · ÿ TT* ÿ ÿ·ÿ· ÿ .. ÿ ÿ ·ÿ ÿ
j ÿmM<
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gonidia. 93

a) Fructifications.
Only resting or persistent forms that represent the morphological building block, the mychite,
or are derived from it without mychomitosis, as is the case with the symmychite, are to be regarded
as fructifications of bacteria.

This definition excludes the so-called bacterial spore, which is a dimychite filled with a
peculiar grain of reserve substance (cf. sporite VI a A 3 d).

ÿ) The gonidia.
The gonidia is a spherical mychite, usually covered with a firmer membrane. It is formed by
the rejection of the supernumerary mychite after protozygy in the dimychite or in the dimychosis, or
by the disintegration of a dimychite (plastite). Like the mychite of the monomychotes, it can
reproduce itself by division by entering the basite stage (cf. basite stage).

The gonidia can be formed by all dimychotes and is the most common fruit form of bacteria.
It is formed in all ripening phenomena. All factors that favor ripening are listed in section VI b.
These are primarily nutritional, chemical, heat, sunlight, etc. influences.

There is no dimychote bacterium that does not readily tend to form gonidia.

Only in sporite (incorrectly called spores, cf. VI A 3 d) forming'

In most species, this phenomenon must first be eliminated before the true and original fruit form is
formed. I would just like to remind you of the so-called asporogenic form of the anthrax pathogen,
which can easily be produced by cultivation in sunlight or by frequent re-cultivation (frequent
change of the nutrient medium), etc.

The term gonidia comes from COHN (1870).

The gonidia is the actual and original fruit form of the bacteria.
It represents the shedding of individual morphological basic elements, the building blocks of all
higher bacteria.
HÜPIE, BLIESENER and others call the gonidia arthrospores.
NEISSER (1888) calls them spores (but also trophosomes etc.). Morphologically, the gonidia is the true
spore, which is homologous to the spores of fungi (conidia and ascospores).

The so-called bacterial spore, on the other hand, is morphologically similar to that of
1
What BREFELD calls Oidie and for which the name S ÿ o rit is applied here (cf. Y 3 c and VI a 3 )
d). Also BETECH (On a

cf. Enderlein (3) 1916, page 405.

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92 gonidia.

New method for the preparation of tubercle bacillus spores, C. ÿ. I. 1909 pag. 461—463) still
calls the gonidia spores.
The granules MUCHS (1907/1908 ) are ÿ. T. gonidia, partly cystites and mostly
trophosomes and trophosomelles (cf. also KNOLL WEGELI N 191 0 [Korrespondenzbl. f.
1912 , WIETHS 1908 , Schweizer. Ärzte
No. 29], ADAM [Inaug.-Dissert. Leipzig 1910 ] etc.).
Among all gonidia there are larger and smaller specimens, which COHN 187 0 already
distinguished as macrogonidia and microgonidia.
There is no significant morphological difference; there are usually all transitions. In terms of
color, there are sometimes differences within the gonidia, for example, some of the gonidia,
particularly the microgonidia of the tuberculosis pathogen, are not acid-fast, so they do not
stain according to Ziehl, but only by Gram staining; however, this also depends a lot on the
length and intensity of the acid effect ; with careful application of acid, a Ziehl stain is also
achieved for the most part; so, strictly speaking, they are only less acid-fast; MUCH calls these
gonidia (but also trophosomes) of the tuberculosis pathogen granules.

After their formation, the gonidia are divided into dichogonidia and
Arthrogonidia n to distinguish.
Dichogonidia are formed by the disintegration of a dimychite into two spherical mychites;
this is therefore a case of isomorphic arthrogonia. As an example, the cholera pathogen may
be mentioned, which forms dichogonidia in large quantities in older cultures. Fig. 75 shows
fairly small dichogonidia.

The arthrogonidia are shed by didimychites or syndimychites (ascites), rarely by

dimychites. Particularly in very short unions of dimychoses (2 or 3), it is easy to observe on
suitable objects that the shed of the gonidia is initiated by protozygy occurring in one dimychose
and the other mychose then becoming free, the union is loosened and finally the completed
gonidia is shed. Evidence that gonidia formation can also occur in unions of dimychoses
through simple disintegration of dimychoses can be seen in the formation of cataplastites (cf.
VI ÿ A 6 b).

The gonidia can be formed on a syndimychite (ascite) either at the end (telogonidia ) or
along the entire longer or shorter thread or rod (ascogonidia). If the gonidia is located at the
inner end of one of the two halves of a diplascite (see this), then one can speak of a
mesogonidia. —

Telogonidia of the cholera pathogen, as they occur in older cultures, are shown in
Figures 69, 72 and 76. The rods
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gonidia. 93

69

Fig. 69—76. Microspira comma shot. Forms from older cultures. Magnification 1 :5000.
Fig. 69. Syndimychite (ascite) in the middle in the beginning of division, at each end with 1 telogonidia (left beginning of gonidia
formation, right shortly before the gonidia is expelled). — Fig. 70. Cystite with ascite residue. — Fig. 71. Cystite with ascite residue.
— Fig. 72.
Ascite residue with macrogonidia and microgonidia. — Fig. 73. Cystite and microgonidia still connected. — Fig. 74. Ascite residue
with cystite and microgonidia. — Fig. 75. Two dichogonidia formed by the decay of a dimychite. — Fig. 76. Syndimychite (ascite)
with 1 telogonidia.

77th minute 78th, 79th 80th 81st

Fig. 77 Microspira comma shot. Syndimychite (ascite) below with telotrophosome, above with telogonidia. Magn. 1 :10 000. Fig.
78. Same, formation of trophosomes. Magn.
1 :10 000. — Fig. 79. Desg., formation of gonidia (ascogonidia) and 1 telocystite. Magn.
1 :10 000. — Fig. 80. Acystia typhi (Eberth 1880). Syndimychite (ascite) with trophosomes, above with beginning formation of a
telogonidia from a telotrophosome. From a four-day culture on agar with 6% Na Cl. Magn. 1 :10 000. — Fig. 81. Same, above with
telogonidia pinching off. From a four-day culture on agar with 61 /, % Na Cl.

Magnification 1:10,000.
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94 gonidia.

Some of the cells are long and some are short. The latter often represent rods, which are already
largely used up by the shedding of several gonidia.

Fig. 77 represents a syndimychite (ascite) from the cholera pathogen, which has a
telotrophosome at one end (below) and has formed a gonidia at the other end (above), which is soon
ready for rejection.
A syndimychite (ascite), also from the cholera pathogen, is shown in Fig. 78 at half the
magnification, which allows numerous trophosomes to be seen even with ordinary staining; at the
upper end the telotrophosome is already somewhat swollen; it is on the way to developing into a
gonidia due to a greater accumulation of reserve substances.

Four ascogonidia are distributed over the syndi-mychite (ascite) shown in Fig. 79, which
shows the beginning of cystite formation at the upper end adjacent to trophosome accumulations
(see next section).

Likewise, the formation of a telogonidia on a syndimychite (ascite) with trophosomes and

trophosomelles of the typhoid pathogen is in

Pig. 82. Treponema dentium (Koch 1877). Syndimychite (ascite) with trophosomes and a
telogonidia. Vergi. 1 :10 000.

Figure 80 shows a picture of this, as does a developed telogonidia, which is about to be shed, in Fig.
81. Both syndimychites represent only the end segments of even longer threads (cf. mycascite).

The formation of gonidia in a spirochete, Treponema dentium (Koch 1877) from human teeth,
is shown in Fig. 82. While the syndimychite in this species is usually without trophosomes and
gonidia, such pictures are rarer. After one slightly swollen end there are a few clear trophosomes
and at the end a telogonidia. Sometimes these gonidia are found already strongly constricted, but
this is rare. The "many small protoplasmic bodies the size of small cocci in the spirochete of Weil's
disease" mentioned by REITER and RAMME (1916) are the same gonidia.

In higher bacteria, especially in the synascites of some sphaero-tilids and syncrotids, such
as Phragmiäiothrix Engl., Crenothrix •Cohn etc., large quantities of gonidia are formed
simultaneously and then shed in large numbers, as COHN described and illustrated as early as 1870.

The gonidia owes its origin partly to a stronger accumulation

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gonidia. 95

formation of reserve substances. These are gradually used up during the resting state as isolated gonidia,
which is intended to survive unfavourable conditions such as food shortages, etc. As this state approaches,
the gonidia becomes increasingly lighter and when coloured (methylene blue, etc.) a pole that is only
very faintly colourable gradually appears (Fig. 84). Fig. 83 shows an isolated pliotrophic gonidia. As the
reserve substances (trophoconia) are used up, the remains of these are finally only collected at the pole
that contains the mych and grouped around the mych (Fig. 85). When the trophoconia are completely
consumed, the cell (Fig. 86) which can only be stained very weakly contains only very faintly stainable
cytoplasm and the myc, which can usually only be stained more sharply with carbol fuchsin (1:10 water)
and is usually unstainable or hardly stainable with methylene blue. The further fate of the

83rd 84th 85th minute 86th minute

Fig. 83—86. Microspira comma shot. Isolated gonidia. Magnification 1:10,000.

Fig. 83. Pliotrophic gonidia. — Fig. 84. Onset of miotrophosis. — Fig. 85. Miotrophic gonidia. — Fig. 86. Atrophic gonidia.

atrophic gonidia are to be further investigated in section ÿ 1 (gonite formation).

While in miotrophic gonidia the limitation of the tropho-

•conia collection is blurred and a gradual transition from the pale to the strongly colored areas can be
seen, whereby the trophoconia (reserve substances) are usually very strongly stainable, the border of
the mych is very sharp and clear and completely circular despite its extraordinarily much paler coloration
(ÿ. B. when stained with carbol fuchsin 1 ; 10).

Completely identical or extremely similar pictures can be observed in the gonidia of all bacteria
and in very numerous species of my

It is always possible to detect these processes.

We will not go into this in more detail here, but reference will be made to it in other places, either in words
or in pictures. For example, gonidia are shown from the tuberculosis pathogen (Fig. 222, 226), from
Syncrotis huccalis (Rob.) (Fig. 207), etc.

Just as the formation of gonidia was observed in the vast majority of bacteria examined, I have
also observed with certainty the development of gonidia into the typical rod in numerous species made
from pure gonidia material. I only mention syn-
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gonidia.
96

erotis luccalis (Bob.), Microspira comma Schrot., Acystia typhi (Eberth)

etc. etc.
Mychomitosis occurs in gonidia that find themselves in more or less favorable
living conditions. Depending on the nutritional conditions and the amount of reserve
substances, the resulting dividing dimychite breaks down into two mychites (cf.
basite stage), whereby mychostasis is very low.

Pig. 87—90. Microspira comma shot. Germination of macrogonidia. Magnification 1:10,000.

Fig. 87. Equatorial germination. — Fig. 88. Same. — Fig. 89. Polar germination. —
Fig. 90. Fully hatched dimychite.

91st 92nd 93rd 94th

Fig. 91 Corynobacterium diphtheriae (LöffL). Miotrophic didimychite. From a culture exposed to sunlight
for a long time. Magnification 1:10,000.
Fig. 92. Desgl., miotrophic gonidia. — Fig. 93. Desgl., mychomitosis of the gonidia. —
Fig. 94. Desgl., Microgonit.

95th minute 96th minute 97th minute

Fig. 95 Acystia typhi (Eberth 1880). From culture exposed to strong sunlight for 20 days. Pliotrophic
dividing dimychite. Magnification 1:10,000.
Fig. 96. Desg., atrophic dividing dimychitis. — Pig. 97. Desg., gonidia.

remains or, as is the case with macrogonidia that are particularly filled with reserve
substances (trophoconia), the phytite stage (cf. VI ÿ A 3) can follow immediately or
soon. The germination of a macrogonidia - again from our example used here, the
cholera pathogen - is shown in Fig. 87 to 90. As can be seen here, germination can
take place equatorially (Fig. 87 and 88), whereby
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cystitis. 97

At the same time, the two newly formed mych emerge from the old gonidia membrane, or only one
of the two mych appears outside the gonidia membrane, while the other remains stuck in the
membrane, as shown in Fig. 89 (polar germination). Here the mychostasis is immediately of
considerable length. A dimychite that has completely rid itself of the gonidia membrane is shown in
Fig. 90.

Some other gonidia, which come from cultures kept for a long time in sunlight, are shown in
the above figures, namely from the diphtheria pathogen and the typhoid pathogen.

That some gonidia, even of very large species, pass through the chamber-land filter was recently
established by MIEHE ; but similar observations had already been made earlier, for example by LOURENS for
the swine fever bacillus (C. ÿ. I. 44 1907 pag. 420—427, 504—512, 630 to 648), ALMQUIST for typhus
pathogens (C. ÿ. I. 60, 1911 pag. 167), by DOERR, R. (Über filtrierbares Virus; CB 50, 1911 and Beih. pag. 12
—23) , by WOLBACH (1912—1915) for spirochetes and by LÖHNIS (1921).

Microgonidia are particularly suitable here, and in some cases perhaps also gonites (see this).

ÿ) The cystite.

The cystite is a fructification form consisting of a sym-mychite (mychite with polydynamic

mych). It therefore corresponds to a gonidia with a symmychon as the nucleus.

The multivalence of the symmychon enables the organism to form a syndimychite (ascite or
development pseudascite) directly from the cystite, skipping a large part of the normal
(cyclogeny, see this).

The cystite has only been proven with certainty in some phylogenetically higher dimychotes.
Cystite formation has so far only been firmly established in spirochetidae, microspiridae and the
sclerothrichidae, and among the bacteriidae only in the genus Eucystia (E. pestis).

The cystite is formed either at the end of a syndimychite from a trophosome or a gonidia
primordium (telocystite) or at any point on the rod or filament, except at the end (ascocystite).

In the case of phylogenetically very high-ranking dimychotes, which attain the growth form of
the syntactic syndimychites, the assumption seems justified that endogenous cystites can form
inside the thick rods. This would be the case in the endothecite-forming schaudinnids.

The term endocystitis should be introduced for this endogenous cystitis.

After completion of the development of the cystite, it is often removed.
Enderlein, BacterialCyologeny. ^
Machine Translated by Google

98 cystitis.

The cystite thus isolated has, as was usually the case before, predominantly the shape
of a sphere (Fig. 98).
The cystite is usually much larger than the gonidia and often contains a much
larger amount of trophoconia and other reserve substances, which fill it up and make it
very easily stained.

98th 99th

Fig. 98. Microspira comma shot. Cystite. Enlarge 1:10,000.

Fig. 99. Microspira comma shot. Cystitis. Specimen during bursting of the membrane. Magnification 1:10,000.

The cystite usually has a firmer membrane than the gonidia, which bursts in
several places during germination (Fig. 99). However, pictures like the one shown in
Fig. 99 have only been observed sporadically in the cholera pathogen; usually the firmer
skin appears less clearly.
The diameter of a cystite in the cholera pathogen can increase to ÿ·2 ÿ and more
21(Fig. 98). Additional cystites of the same parasite

Fig. 100. Pseudostreptus puoqenes (Rosenb. 1884). Specimen from two-day culture.
S
Magnification 1:5000.
1 cystite (diameter 8.4 ÿ), symmychon apparently lateral. — 2. Pliotrophic cystite or gonidia. — 3. Arthrothecit. — 4. Desg. —
5.-7. Pliotrophic cystite or gonidia. — 8. Didimychite with 2 trophosomelles and 1 trophosome. — 9. Didimychite with
trophosomelles and 1 trophosome.

are already shown in Fig. 70, 71, 73 and 74. Here it can be seen that in many cases
the syndimychite is used up by the cystite through withdrawal of reserve substances
and shrinks completely.
The diameter of the cystite in the genus Pseudostreptus increases to over 8 ÿ-
(Fig. 100). Here too, the cystite can be isolated and