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Edmund B. Wilson's The Cell and cell theory between 1896 and 1925

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Hist. Phil. Life Sci., 24 (2002), 357-389 ---357

Edmund B. Wilson’s The Cell and Cell Theory

between 1896 and 1925
Ariane Dröscher
History of Science Unit and Historical Archives
Stazione Zoologica A. Dohrn
Villa Comunale, 80121 Naples, Italy

ABSTRACT – Edmund Beecher Wilson is generally celebrated for his contribution to

chromosome theory and genetics, whereas opinion concerning his cytological thinking is
often restricted to the idea that he provided evidence for the dominant role of the
nucleus. But Wilson’s cell theory was much more. It was a child of the German
Zellforschung, and its attempt to provide a comprehensive cellular answer to a wide
range of biological and physiological questions. Wilson developed a corpuscular,
micromeristic and preformistic concept, and treated the cell as an organism subject to
ontogenetic and phylogenetic processes. He defended his comprehensive theory even in
the 1920’s, when cytological research had become specialised and directed at more
practical goals. For many of his younger readers this concept might have seemed
antiquated, but today many of its features sound surprisingly modern.

From the first appearance of The Cell in Development and Inheritance

Wilson was praised as the worlds leading cytologist (see e.g. Conklin
1897; Driesch 1897), and historians today agree. Many Americans have
expressed the opinion that Wilson’s The Cell ‘has been the single most
influential treatise on cytology during the twentieth century’ (see e.g.
Allen 1976, 430). Jane Maienschein calls it the last monumental,
synthetic and all inclusive cytology textbook to have been written by an
individual author (Maienschein 1991a, 46-47). Moreover, it is generally
acknowledged that it prepared the way to the linking of cytology with
Mendelism, and Scott Gilbert suggests that Thomas H. Morgan was
induced to revise his opinion on the role of chromosomes in inheritance
by those students and collaborators of his who had used The Cell for
their theoretical cytological preparation (Gilbert 1978). No biographer
fails to underline the importance Wilson attributed to the cell.
Wilson himself stressed that he considered the cell the fundamental
unit of development and heredity, and indispensable for the solution
of any of the great questions of biology. But the simple declaration
that the study of the cell is central to biomedical inquiry cannot be
the only reason for Wilson’s outstanding position. Similar statements
are to be found in nearly all anatomical, histological and cytological

0308-7298/90 $ 3.00 © 2002 Taylor and Francis Ltd

358 ARIANE DRÖSCHER

textbooks. Further research is required about how he imagined that

the different branches of biology converged in the cell. It, therefore,
seems worthwhile to inquire what exactly Wilson’s conception of the
cell stands for, and how it fits into his general idea of vital
phenomena.
There is almost no aspect of cytological importance that was not
touched on by Wilson, and all his contributions to the various debates
would be worth further study (see e.g. Sapp 1998). It is not the aim
of this paper to analyse all of them, but to indicate the basic outlines
of his conception of the cell and to put forward three theses: 1.
Wilson’s cell theory is central to an understanding of his ideas of
organisation, development and inheritance. 2. In spite of the praise,
the value of Wilson’s contribution to cell theory was and still is
underestimated. 3. Wilson’s cell theory was undervalued because it
was based on a scientific philosophy that had become less and less
popular during the first half of the 20th century.
The paper will deal first with Wilson’s general approach to cell
theory, and then, more in detail, with two essential cytological
problems and the solutions Wilson proposed for them. The first refers
to the place cells (and their constituents) occupy within the hierarchy
of organic organisation. The second refers to the origin and
development of endocellular organisation. It will be shown that for
Wilson, both problems – organisation and development – were
inseparable. The present analysis is mainly based on the three editions
of Wilson’s The Cell in Development and Inheritance. The first edition
appeared in 1896, the second in 1900, and the third in 1925, covering
nearly three decades of Wilson’s scientific activity. Besides the
treatises, some research papers and general essays have been taken
into account. Three essays (Wilson 1923; Wilson 1925b; Wilson
1926) are of particular interest, because they gave Wilson the
opportunity to apply himself to natural philosophy and to make
speculations he could not express to this extent in his textbooks.

Cytology in Wilson’s period

By around 1900 it had become difficult to discern the state of the

art in cytology. To talk about cell theory in the 1920s is even more
complex. As John Baker has already emphasised, it is indispensable to
clarify which cell theory or which part of it one is talking about
because each investigator has a different idea in mind (Baker 1948,
104-106). The expression ‘cell theory’ will be used here in a rather
THE CELL AND CELL THEORY 359

wide sense, as a broad theoretical conception that joins together

various areas of biological research, and is based on the cell and its
vital expressions.
Cell theory is generally considered one of the two great synthetic
biological theories, a theory that established biology as an autonomous
scientific discipline and united the various branches of natural history,
microscopy and physiology. Moreover, it represented an important
step towards the integration of biology and the basic medical sciences.
The history of cell theory can be seen as subject to two different
movements: expansion and splitting. The more cytology expanded
into further research fields, the more cell research itself split into
various specialities, and the more it became difficult to formulate
conceptions of the cell that were acceptable to everyone.
In the course of its history cell theory underwent several radical
modifications. Until the 1840s cell research had been a domain of
botanists, investigating the cellular origin and constitution of plant
organs. Matthias Schleiden’s Zellenlehre united anatomy and ontology
(Schleiden 1838). Theodor Schwann showed that the same principles
were valid for zoology (Schwann 1839). During the 1850s and 1860s,
cell research was increasingly dominated by zoologists and
physiologists. Schleiden’s definition of the cell was superseded by that
of Max Schultze, the anatomical building-block cell theory replaced
by the protoplasm theory of the cell, Schleiden’s Zellenlehre by Ernst
von Brücke’s Elementarorganismen, and classical cell-research by
cytology. The physiological concept of the cell was more dynamic, and
offered promising approaches to embryological, developmental,
evolutionary, protozoological, and taxonomic questions. Again, each
branch had specific needs and standpoints.
In the 1890’s, by the time Wilson began his cell studies, another
major conceptual change had taken place, the transformation of
cytology into Zellforschung (experimental cell research). Its aims and
approaches were developed mainly by Oscar and Richard Hertwig,
Max Verworn and Theodor Boveri. In his book Die Zelle und die
Gewebe (Hertwig 1892) Oskar Hertwig announced his intention to
provide a new cell theory that regarded the cell as a ‘complicated
organism and a small universe’. In the same year appeared Otto
Bütschli’s Untersuchungen über mikroskopische Schäume (Bütschli
1892) and August Weismann’s germplasm-theory (Weismann 1892),
directing attention to the cell as the centre of the basic physico-
chemical and hereditary processes. Zellforschung elevated the cell to
the rank of universal key for all vital phenomena, and cell theory to
the central theory of the biological sciences (Jacobs 1989). The
360 ARIANE DRÖSCHER

consequences for biological research were twofold. First, cells were

treated as if they were small but autonomous ‘microcosms’ equipped
with the whole range of vital expressions, and, second, all vital
phenomena of multicellular organisms were explained in terms of
cellular mechanisms and properties. Cell theory had become a
synthetic theory that included all fields of biology.
The centre of Zellforschung was Germany,1 but even here not all
cytologists were in agreement. In Britain it was criticised as
morphological, preformationist and static. Thomas Huxley’s epigenetic
and ‘protoplasmic’ conception of the cell found greater acceptance,
and influenced British cytology throughout the 20th century
(Richmond 2001). Many French cytologists preferred the physiological
and cytochemical approach, whereas Italian cytologists practised
predominantly morphological cell research, but without formulating
the far-reaching conclusions of Zellforschung (Dröscher 1996). Apart
from certain national styles, more detailed analysis reveals a great
variety of conceptual and experimental approaches.
During the first decades of the 20th century the universal claim of
Zellforschung lost its attractions. It was not superseded by another cell
theory, for the general attitude of cytologists towards their enterprise
had altered. This change was reflected and reinforced by a series of
institutional phenomena: The ever increasing number of researchers
working with one or other aspect of cellular phenomena, the
continuing expansion towards new research fields, and the process of
specialisation within cytology, caused a splitting of the community of
cytologists into more and more confined and isolated circles. When
Wilson’s third edition of The Cell was published, Zellforschung and its
aspiration to furnish a universal answer to the secrets of life had been
given up, and synthetic cell theory split into a variegated research
field.
Cytology became too broad to be surveyed in its entirety, but
cytologists did not even feel much need of a comprehensive view. As
a research field, cytology has always been particularly plastic, involved
in a continuous process of disciplinary transcendence, and the
integration of techniques and approaches of adjacent disciplines. At
the beginning of the 20th century it had also become sufficiently
autonomous, stable and self-confident to permit internal diverging
opinions. Research was restricted to specific areas that promised

1 The concept of the central role of the cell was no exclusive achievement of German cytologists.
Jacyna has shown that, for example, John Goodsir (1814 - 1867) had expressed similar ideas long before
(Jacyna 1983).
THE CELL AND CELL THEORY 361

concrete and more immediately practical results and avoided excessive

theoretical deduction.2

Wilson’s general approach towards cells and cell theory

Even if it is better to speak in the 20th century of various, often

only unconsciously adopted concepts of the cell, it is nevertheless
acceptable to call Wilson’s conception of the cell a synthetic cell
theory. Since his time as an unknown student, and except for the
years between 1903 and 1912, when he ‘moved from cells and their
morphological characters to different, more immediately productive
research ventures, notably in cytology of the chromosomes and
genetics’ (Maienschein 1986, 42), the constitution of the cell was
Wilson’s favourite subject and leading interest. Until the end of his
scientific activity, his fundamental conception of the cell, as outlined
in the first edition of The Cell, did not undergo essential
modifications.
The definition of the cell Wilson proposed is characteristic of his
general approach. At first sight it seems rather traditional and, in the
course of time, it was slightly modified only to emphasise certain
aspects. In the first edition of The Cell he adopted the definition
formulated by Max Schultze and Franz von Leydig more than thirty-
five years earlier: the cell is ‘a mass of protoplasm containing a
nucleus’ (Wilson 1896, 19). In the 1860s this definition had been
rather revolutionary, because it had destroyed the static definition of
the Zellenlehre that had concentrated on the cell wall. In 1896,
however, when cell morphology had been enriched by an infinite
series of mononuclear and polynuclear, unicellular and polycellular,
syncytical and quasi-cellular forms, and by an extraordinary variety of
cell types with various endocellular structures, cytologists felt
increasing discomfort with Schultze’s definition (see e.g. Flemming
1882, 71-73). Wilson did not ignore their arguments, but continued to
defend the validity of the traditional definition.
In 1925 Wilson was still using it but stressing two specific features.
First, that protoplasm had to be understood with the modern
terminology of cytosome, and second, that ‘both, nucleus and

2 Evidently, in spite of the growing belief that purely technical issues were being dealt with, the
general ‘philosophical’ problems were still present. They reappeared at every step of research under the
mask of unconsciously made preliminary decisions concerning the experimental approach, the selection
of the experimental subject, the choice of suitable technique, and the interpretation of results (see e.g.
Fantini 1985).
362 ARIANE DRÖSCHER

cytosome, arise by division of the corresponding elements of a

preëxisting cell’ (Wilson 1925a, 21). Neither feature was innovative.
The first can be traced back to Eduard Strasburger,3 and the second
had been part of Schultze’s classical work of 1861.
Introducing his paper Protoplasmic Systems and Genetic Continuity,
published in the same year as the third edition of The Cell, Wilson
found no difficulty in expressing the same conception of the cell by
stressing its conformity to the basic ideas of colloid chemistry:
The cell is a complex heterogeneous system that arises only by division of a
preexisting system of the same general type, and with certain minor qualifications
the same is true of each of its two primary components, nucleus and cytosome –
each of these is itself a complex system that arises by division of a preexisting
system (Wilson 1925b, 481).

So, in 1896, and even more in 1925, Wilson’s definition of the cell
corresponded to a traditional one, and considering the fact that more
than sixty years had elapsed since 1861, one may even prefer
antiquated to traditional.
However, Wilson’s importance for cytology should not be sought
for in isolated statements. He himself did not attach much importance
to his definition of the cell. In the second edition of The Cell Wilson
added to the chapter ‘General Sketch of the Cell’ the introductory
phrase:
We are, it is true, still unable to specify all its essential features, and hence can
give no adequate brief definition of the cell. For practical purposes, however, no
such definition is needed, and we may be content with the simple type that has
been familiar to histologists since the time of Leydig and Max Schultze (Wilson
1900, 17).

Nor should Wilson’s persisting in his youthful beliefs be interpreted

as a diminishing intellectual engagement or as an obstinate incapacity
to assimilate new ideas. It is certainly true that by this time he was
able to incorporate into the third edition of The Cell, published
twenty-five years after the second edition, such a considerable
quantity of important new literature and information because the
basis had already been laid by the first two editions. These were much
less voluminous, but in their general conception not essentially

3 Strasburger had been the most authoritative voice on the terminological problem connected with
the use of ‘protoplasm’. He suggested a distinction between the term protoplasm intended as cytoplasm,
and hence the amorphous liquid surrounding the organised or solid parts of the cell, and protoplasm
meant as protoplast, and hence the whole body inside the cellular borders, including the formed parts.
THE CELL AND CELL THEORY 363

different. In her comparative analysis of all three editions, in fact,

Maienschein has noticed no major innovation in Wilson’s treatment of
the general sketch of the cell except the more detailed and more
confident exposition (Maienschein 1991b). But one does no justice to
Wilson by considering the third edition as nothing other than a simple
revision of an antiquated scheme.
The chapters dealing with the morphological and physiological
features of the cell were completely revised and notably enriched.
Reading them with the three papers on cellular constitution (Wilson
1923; Wilson 1925b; Wilson 1926) that were published around the
same year, it becomes evident that Wilson continued throughout his
life to compare his own ideas with the results of recent research and
with the explanations proposed by the new schools of cytology.
Three other features characterise Wilson’s general conception of the
cell: it was ‘philosophical’, ‘comprehensive’ and ‘synthetic’. In
opposition to the overwhelming majority of his younger colleagues,
Wilson did not give up thinking about universal conceptions. On the
contrary, although he was very careful to avoid an excessive exposition
of actual speculations, he felt pleased rather than uncomfortable over
illustrating the general implications of recent cell-research. All three
editions of The Cell are characterised by philosophical depth. In 1923,
probably feeling that this approach had made him increasingly
isolated, especially in the United States, he almost apologised for his
seeking for a unifying principle:
Perhaps there is no problem or none that we can formulate without talking
nonsense. Perhaps we should go no further than to record and analyze the existing
order of phenomena in living systems without losing sleep over the imaginary
problem of a unifying principle. Let us politely salute all these uncomfortable
possibilities and go our way. For my part, I find it more amusing to look forward
to a day when the great riddle may give up its secret (Wilson 1923, 286).

Maybe Wilson was the last cytologist of his time to seriously

propose a comprehensive answer to all aspects of cellular life and
constitution. Maybe he was the last to develop a real cell theory.
Certainly, he was able to write The Cell only because had developed
a synthetic cell theory that served as a guide. His cell theory enabled
him not to be content with the explanation of individual phenomena,
but to show their intimate connection. The most enduring impression
in the reader is not provoked by the wording of his definition of the
cell, but by its omnipresence throughout the book. Every biological
problem was logically attached to cellular processes. Cells were the
key to the questions of vital phenomena in general.
364 ARIANE DRÖSCHER

Wilson’s relation to Zellforschung

Wilson’s attitude towards cell theory demonstrates what he owed to

the German Zellforschung. The enthusiasm with which the young
Wilson imbibed the new ideas, personally experienced during his time
at Heidelberg, Munich and Naples, emerges from his papers. The
most striking testimonies to his debt to the deans of German cytology
are in the dedication of The Cell to his friend Theodor Boveri, and in
the quotations taken from Rudolf Virchow’s Zellularpathologie, from
Schwann’s Untersuchungen, from Brücke’s Elementarorganismen and
from Ernst Haeckel’s Anthropogenie, placed at the beginning of the
chapters dealing with cell structure. The quotations were translated
into English in the third edition only.
During the years of Wilson’s second stay in Germany (1891-1892)
the debate about Zellforschung reached one of its climaxes. Two years
earlier Richard Altmann’s Die Elementarorganismen (Altmann 1890)
had been published. Discussion was heated and exciting. Bütschli’s
dynamic perspective, the stress he laid on causal-mechanical
explanation, and his interpretation of cells as physico-chemical
systems, a conception that was to initiate cell research in terms of
colloid chemistry, deeply impressed Wilson. Above all, Wilson
assimilated the concepts of Zellforschung through Boveri’s ideas about
cellular constitution and the intracellular processes of development.
The Entwicklungsmechanik (experimental embryology) of Hans
Driesch, and Curt Herbst, opened up new experimental approaches
to questions of development and determination.4
This influence was not limited to the purely intellectual sphere. At
Naples Wilson realised his ‘wildest most unreal dreams’ (Morgan
1941, 320). The unity of science and the arts as practised at the
Stazione Zoologica, the presence of young enthusiastic scientists from
all over the world, the concerts frequented at Leipzig and Berlin, the
discussions with Boveri while drinking, dining and playing billiards
together at the Café Heck (Morgan 1941, 321), made individual
scientific notions become part of a fundamental and lasting vision.
In Wilson the basic ideas of Zellforschung found a fertile terrain.
Allen describes him as a ‘meticulous and exhaustive encyclopedist’
speaking six languages (Allen 1976, 423). He stresses that Wilson was
a great lover of music and an excellent cellist. He was thus aware of

4 For the importance of the Naples Station and the Entwicklungsmechanik for the transformation of
biology into experimental science see Müller 1975; and 1976; for the impact it had on Wilson, see
Maienschein 1991c, 96-104.
THE CELL AND CELL THEORY 365

the existence of half-revealed meanings (Allen 1976, 435), and looking

for harmony, and for synthesis rather than the analysis of single notes.
This explains Wilson’s capacity to unite ideas that were actually quite
distant from each other, and often apparently contradictory. Two
research results could at first sight exclude each other, but within a
broader context it was possible to find an answer that would satisfy
both.
With regards to genetics, Jonathan Harwood and many others have
referred to the German ‘style’ as a research field guided by the broad
approach, aiming to connect the various questions of inheritance,
development and evolution, whereas the North American ‘style’ was
characterised by rapid specialisation, and priority given to small steps
aiming at practical results, rather than caring attention whether or not
they fitted into universal conceptions (Harwood 1987). If we accept
this as a general characterisation of biology, then Wilson can be seen
as a mediator. Intellectually he tended towards the German way, but
institutionally, and considering where his research school led
scientifically, he laid the bases for the American way of doing science.
Wilson personified transition: from biology as a branch of natural
history to biology as a science based on rigorous quantitative and
experimental analysis; from biological research dominated by German
personalities to the expanding American research centres; from the
Entwicklungsmechanik of the Naples Station to the experimental
developmental research of the Woods Hole Marine Biological
Laboratory; and from Zellforschung to the cell biology of the 1950s.5

The cell as basic morphological and physiological unit

Half a century after the formulation of the Zellenlehre, it had

become habitual to think of higher organisms as cellular constructs.
Nevertheless, the debate whether or not cells should be considered
the unique and fundamental unit of life was still far from being
settled. The central position that had been attributed to cells since the
time of Schwann and Schleiden was ‘menaced’ from two opposite
directions: on the one hand, by concepts of a superior, supracellular
organisation that considered single cells as secondary subdivisions of
an organism-wide substrate; on the other hand, by conceptions of

5 Wilson did not contribute directly to the birth of ‘cell biology’, founded as a distinct research
discipline during the 1950s, but he led cytology in this direction. For the institutionalisation of ‘cell
biology’ see Bechtel 1993.
366 ARIANE DRÖSCHER

subcellular units that regarded cells as secondary aggregates of still

smaller ‘carriers of life’. The dispute may seem a mere philosophical
problem of individuality, but it was also of considerable anatomical
and physiological interest. Whether a researcher regards the cell as the
producer or as the product of cellular substance, and hence, as either
the agent or the passive expression of vital phenomena, and likewise,
whether he concentrates his studies of vital phenomena on a universal
substance, on a cell, or on specific submicroscopic particles,
profoundly influences his experimental approach and interpretation.
The concepts of Zellforschung enforced the central position of the
cell for anatomical and physiological studies, and placed it in a central
position for biological inquiry in general. At the same time, however,
Altmann published his Die Elementarorganismen, in which he
described the cell as an assembly composed of smaller units, the
Bioplasten (Altmann 1890). Altmann was not the first to put forward
ideas of subcellular units. The conception of invisible but material
carriers of life was an old one, and revived during the 1860s with the
hypotheses of Herbert Spencer, Charles Darwin, Eduard Pflüger, and
many others, proposing ‘physiological units’, ‘gemmulae’, ‘living
protein’, and other agents. Altmann was probably the first to assert
that he had actually observed them.6 Supporting data came initially
from botany and embryology. In 1883 botanist Andreas Schimper
drew the conclusion that plastids, specific organelles of plant-cells,
were equipped with a special chemical constitution and the faculty of
persistent identity, and even self-perpetuating properties (Schimper
1883). Another heated debate began a few years later in embryology
about the nature of the centrioles (or centrosomes), that were
supposed to form the division-centre of the cell (Sapp 1998). Around
the turn of the century improved microscopes and more advanced
cytological techniques revealed a myriad of further cell structures –
mitochondria, chondriosomes, Zentralkapseln, chromidia, the
ergastoplasm, the apparato reticolare interno (Golgi apparatus), and
Throphospongien, to name a few – leaving the cytological community
puzzled over their significance (Dröscher 1999). The majority of the
cytologists were cautious about becoming too speculative, but some
went further, formulating far-reaching theories. Even symbiotic
theories were suggested maintaining that various intracellular bodies
had once been independent organisms (Sapp 1994). The crucial point
for cell theory was not whether there existed a cellular metastructure,

6 Later cytologists agreed that Altmann’s Bioplasten were mitochondria or chondriosomes (e.g.
Hughes 1959, 118-125).
THE CELL AND CELL THEORY 367

but to what extent those subcellular units were believed to be capable

of regular life and performance without being part of a complete cell.
On the ‘supracellular’ side, the position of the cell as the
elementary unit was under attack from many protozoologists and
physiologists, especially in Britain. Clifford Dobell, for example,
preferred the whole organism, whether or not it exhibited a cellular
subdivision, as the starting point for analysis, and drew the conclusion
that: ‘the cell must be defined in terms of the organism, and not the
organism in terms of the cell’ (Dobell 1911, 284).7 A similar argument
was put forward by Edward Stuart Russell, who maintained that the
‘fundamental organization precedes cell-formation in time and
regulates it, rather than the reverse’ and consequently that ‘the whole
cannot be completely explained from the parts into which conceptual
analysis resolves it’ (Russell 1930, 262-265).8 This interpretation had
already been adopted by Ray Lankester and Michael Foster, and
remained popular also among American developmental physiologists
like Charles Otis Whitman and Frank Rattray Lillie (Corliss 1989;
Richmond 1989).
Other supracellular opponents met around an advanced version of
the protoplasm theory, finding strong support from the ideas of
Thomas Huxley. Stressing the role of intercellular bridges and
plasmodesmata that connect neighbouring cells, and were frequently
observed after the 1860s, protoplasm was interpreted as the
fundamental and organism-wide unit of life,9 or, as Leonard Doncaster
put it:
Nowadays, however, opinion tends to the opposite direction – to regard the
organism as the individual, with common life running through it all, and the cells
not as units of which it is built up but rather as parts into which it is divided in
order to provide for the necessary division of labour involved in so complex a
process as life (Doncaster 1920, 3).

In the course of the 20th century, explicit attacks against cell theory,
like those of Doncaster, became rather rare, but colloidal chemists still
treated protoplasm as something not necessarily having a cellular
organisation, stressing the common properties of all protoplasms, and
7 It is significant that Dobell’s arguments show strong analogies with the 18th century theory of the
fibre as outlined by Berg (1942).
8 Without penetrating too deep into the question raised by Russell, it is necessary to underline that
supporters of the idea of a cell-state could likewise be convinced that the whole was more than the sum.
For the origin, significance and variety of cell-state metaphors see Mazzolini 1988.
9 Marsha Richmond calls this standpoint the ‘holistic view typical for the epigenetic conception of
development’ (Richmond 2001, 58).
368 ARIANE DRÖSCHER

making statements on vital physiological phenomena without any

reference to cells or the particular cellular organisation (e.g. Loeb
1922; Bayliss 1923; Bělař 1928; Frey-Wyssling 1938). Some, like
Gottfried Berthold in Germany or Luigi Buscalioni in Italy, even tried
to create artificial cells by mixing several colloidal substances
(Buscalioni 1920).
In the 1920s, however, many young colloid chemists of the cell
began to dissociate themselves from ideas of supracellular
protoplasmic organisation. The Austrian physiologist Armin von
Tschermak-Seysenegg, for example, defended the cell as an
ontogenetic and functional unit (Tschermak 1924, 416). Robert
Chambers, through his micro-dissection experiments, found strong
evidence in favour of the autonomy of the single cell. He concluded
that ‘protoplasm is a cellular unit which cannot exist without its
nucleus and its cortex and, therefore, must be regarded not as a ‘stuff’
but as a mechanism consisting of visibly differentiated and essentially
interrelated parts’ (Chambers 1924, 238). Even more explicit was
Victor Heilbrunn, who insisted that the colloid chemistry of
protoplasm should always study protoplasm itself and not gelatine or
other inanimate colloidal substances. He warned against treating
protoplasm as one single universal substance, just as there was no one
single universal type of cell: ‘Generally if one is to obtain information
concerning the physical properties of protoplasm one must study the
individual cellular units. This is a broad statement and is not to be
taken too exactly. Nevertheless, in the main, it is true’ (Heilbrunn
1928, 6).

Wilson and the hierarchy of organic units

Like many supporters of Zellforschung, Wilson had been

particularly impressed by von Brückes’s idea of the cell as an
Elementarorganismus (Brücke 1861). He himself treated the cell both:
as an elementary unit and as an organism. More than his German
counterparts he was aware that the first aspect required a clear
definition of the role of the cell within the hierarchy of organic
organisation. With regard to the multicellular context, Wilson stated
in the first edition of The Cell: ‘As far as structure and origin are
concerned, the tissue-cell is unquestionably of the same morphological
value as the one-celled plant or animal’ (Wilson 1896, 41). Though he
never lost sight of the necessary restrictions and exceptions to be
taken into account while dealing with much less independent tissue-
THE CELL AND CELL THEORY 369

cells, he considered the cell to be mainly an autonomous entity;

autonomous, but not isolated. The functionality of the cell could not
be understood without the intercellular context. Thus, to say that the
cell represents only a part of a higher organism, and to say that it
represents a small but autonomous microcosm, was no contradiction.
On the contrary, the study of just this double role was the key to the
understanding of functionality:
There is at present no biological question of greater moment than the means by
which the individual cell-activities are co-ordinated, and the organic unity of the
body maintained; for upon this questions hangs not only the problem of the
transmission of acquired characters, and the nature of development, but our
conception of life itself (Wilson 1896, 41).

Wilson was used to thinking in more than one category, so that

inserting the cell in more than one context came easily to him. In the
same way he approached the question of cellular metabolism, for
example, or the question of inheritance. The part and the whole, the
single morphological unit and the role it played within broader
physiological contexts were not opposites, but complementary
counterparts. At the beginning of the chapter on the problems of cell
organisation, Wilson wrote:
Whether structure or function is the primary determining factor in vital
phenomena is a question that has been a subject of debate for many generations
of biological philosophers. As thus stated, however, the question has proved
barren, for all students of the problem have in the end had to admit that
structure and function are inseparable. It is certain that vital action is not known
to us apart from an organized material basis, and equally certain that vital
structures exist only as products of protoplasmic activity (Wilson 1925a, 670).

The solution to the understanding of vital phenomena had to be

sought within the structure as well as in their reciprocal relation, or,
in short, in the interaction of formed structures. Wilson’s line of
thought was hence characterised by some sort of dialectic process,
integrating two apparently opposing aspects, the more static
morphological and the more dynamic physiological approach.
With a similar attitude Wilson treated the problem of subcellular
constitution. In the same way that the cell represented an autonomous
unit and a part of a multicellular organism, it was also a whole and
made up of subcellular parts. And in the same way that the
functionality of a tissue or an organism could be understood only by
studying the interaction of its constituents – the cells, subcellular
constitution was the key to the questions of cell-life.
370 ARIANE DRÖSCHER

For Wilson the cell consisted of two main systems, the cytosome and
the nucleus. Both represented autonomous interacting entities. Wilson
stressed that ‘the differentiation of the protoplasmic substance into
nucleus and cytoplasm is a fundamental character of the cell, both in a
morphological and in a physiological sense’ (Wilson 1896, 17). Besides
these two, in 1896 he admitted a third distinct element, the centrosome,
arguing that also the centrosome probably possessed functional and
morphological specificity as well as persistency (Wilson 1896, 17). In
plant cells the same was valid for the plastids. In the second edition,
however, the importance of the centrosome was diminished by his calling
it now only ‘a subordinate part of the general apparatus of mitosis’.
Wilson still considered the centrosome a constant feature of the cell, but
it now ranked as a subordinate part of the cytosome (Wilson 1900, 52).
Wilson did not stop at the level of the dualistic constitution of the
cell. He was firmly convinced that neither nucleus nor cytosome were
homogeneous mixtures, but different in their specific local
composition. Their difference was not purely chemical. With regards to
the nucleus, Wilson enthusiastically spoke of ‘the existence of an
organization of the nuclear region of the cell-system that is as complex
and wonderful as any pictured by the fantasy of the speculative nature-
philosophers’ (Wilson 1923, 280). Since 1896 Wilson included in his
textbook a diagram of the typical cell possessing a definite constitution
(fig. 1). In all three editions the nucleus was made up of four

Fig. 1. Diagramm of a cell in the first edition of The Cell (Wilson 1896, 14).
THE CELL AND CELL THEORY 371

component parts: the nucleolus, the chromatin-network, the linin-

network, and the karyosome. In the third edition, only the terminology
was slightly changed (fig. 2). The cytosome, too, consisted of various

Fig. 2. Diagramm of a cell in the third edition of The Cell (Wilson 1925a, 23).

formed components: the attraction sphere with its two centrosomes,

passive bodies, and vacuoles and plastids, the two latter only in plant
cells. In the third edition of The Cell, chondriosomes and Golgi bodies
had been integrated into a diagram that otherwise showed no
substantial modification. All subcellular structures, too, were treated
like distinct morphological entities.
Wilson’s conception of the cell did not stop at the level of visible
formed cell-organs. It went still further into the dimension of invisible
bodies. Since 1896, Wilson expressed the opinion that: ‘in many if not
in all cases the division of cell-organs is in the last analysis brought
about by the division of more elementary masses of which they are
made up’ (Wilson 1896, 236). He had been particularly impressed by
Strasburger’s remark, that it would be quite by chance if microscopes
captured the lowest possible visible unit of the cell. Wilson, for his
part, drew the analogous conclusion that ‘no hard and fast line is to
be drawn between matter in pieces visible to the naked eye, down
through ultra-microscopic particles to molecules’, and regarded it as
certain that ‘the diminishing series does not end at the purely artificial
372 ARIANE DRÖSCHER

boundaries determined by the working limit of our instruments’

(Wilson 1925a, 635). Once again, Wilson had been convinced by one
of von Brücke’s ideas, that cells were congeries of self-propagating
units of a lower order (e.g. Wilson 1896, 237; Wilson 1923, 283).
Not at home with the ideas of ‘gemmulae’, ‘physiological units’,
‘Bioplasten’, ‘Biophoren’, and the like, Wilson strictly avoided assigning a
definite term to these units. In the first edition of The Cell Wilson
introduced the semantically broad term ‘microsomes’ (small bodies) to
indicate the ultimate units of the linin-network of the nucleus and of the
cytoreticulum of the cytoplasm (Wilson 1896, 223-224). Later he also
spoke of ‘hypothetical units’, ‘particles’, ‘ultra-microscopical bodies’,
‘primary units’, ‘stuffs’, ‘not-splittable masses’ or ‘larger molecular
aggregates’, and his use of the term ‘microsome’ remained vague on
purpose. Sometimes it indicated all visible endocellular bodies (Wilson
1926, 113-115), sometimes a visible but non classifiable structure besides
those known as chondriosomes, fat droplets etc., and still other times it
indicated granules ‘lying at the furthest limits of microscopic vision’
(Wilson 1925a, 33).
Yet although they continued to be neither seen nor exactly definable,
Wilson asserted his corpuscular conception with increasing certainty.
The idea of their existence opened up great opportunities. In the
second edition of The Cell Wilson emphasised how the conception of
‘ultimate protoplasmic units’ helped the understanding of many
problems of cellular constitution and of structural persistency during
cellular and nuclear division (Wilson 1900, 292-293). In the third
edition, Wilson was more careful to point out the hypothetical character
of his view, showing himself not less firmly convinced that ‘particulate
conceptions of cell-structure, sometimes offer the simplest and most
effective means of formulating the observed facts’ (Wilson 1925a, 719).
Finally, in 1923, he stressed that also disease-germs or genes still could
not be seen but were, nevertheless, supposed to be real (Wilson 1923,
282-283). Hence he felt justified in imagining that something of this
kind must exist, and in treating these particles like units.
Wilson was aware that the majority of cytologists and physiologists,
especially in the 1920s, met corpuscular conceptions with great
scepticism, but he defended its usefulness and its consistency with
other cell conceptions, like those of colloid chemistry:
And lastly, I would remark that I am not here attempting to resuscitate the old
conception of the cell as an assemblage or colony of elementary organisms or
primary vital units – perhaps it is such, perhaps not – nor am I able to see how
the possibilities here considered are in any manner out of harmony with the
conception of the cell as a colloidal system (Wilson 1923, 284).
THE CELL AND CELL THEORY 373

Though a convinced morphologist, Wilson was likewise convinced

that the study of structure alone was not sufficient to explain the
processes occurring inside the cell. He praised the colloid chemistry of
the cell as the best approach to investigating cellular mechanisms and
metabolic processes (Wilson 1925a, 633). Yet he evaded emphasising
that he had a quite different idea of what the principles of colloid
chemistry were. Conceptions that were popular among the physiologists
around the turn of the century and that interpreted cells as diphasic
and structureless systems, and all seats of cellular action as necessarily
liquid, had absolutely nothing in common with Wilson’s point of view.
Wilson’s conception of cell chemistry continued to be linked to the
views expressed by Otto Bütschli and Carl von Naegeli. Thus in 1925
he welcomed the younger generation of cell physiologists who began to
dissociate themselves from the extreme positions of the old colloid
chemistry of the cell, and to develop concepts that regarded the cell-
substance as a polyphasic-differentiated system. Wilson took for granted
that the so-called ‘disperse phases’ were represented by suspended
particles and that cytological interest had to be directed just to these
particles.
On introducing his concept, Wilson adopted Naegeli’s term
‘micellae’, imaginary particles that lie below the horizon of
microscopic vision, but later he preferred to talk about ‘larger
molecular aggregates’. These particles ‘form the original source of
many still larger bodies that emerge into microscopical view as the
smallest visible formed components of protoplasm’ (Wilson 1925a,
634-635). On this ultimate level, the constitution of the nucleus and
the constitution of the cytosome, the structural basis of the
chromosomes and the structural basis of the cell-organelles, the
constituents of the cytoreticulum and the constituents of the
chromatin-grains were all essentially the same.
Wilson did not supply any details on how he imagined that cell-
organelles fulfilled their task. By this time, this would have hardly
been possible. Cytologists even disagreed about what exactly the
function of the various organelles was. Concerning nuclear units, on
the contrary, Wilson’s ideas were plain:
Genetic evidence is now opening far-reaching horizons of future discovery by the
accumulating demonstration that no one of the nuclear units plays an exclusive
rôle in the determination of any single character. It has been made clear that the
individual unit may affect the production not merely of one character but of
many. Conversely the probability is shaping itself that the production of any
single character requires the cooperation of several or many units, possibly of all.
I believe it is not a great overstatement when I say that every unit may affect the
374 ARIANE DRÖSCHER

whole organism, and that all units may affect each character. We begin to see
more clearly that the whole cell-system may be involved in the production of
every character. How then are hereditary traits woven together in a typical order
of space and time? It is the same old puzzle made larger and more insistent but
not yet, so far as I can see, brought nearer to its solution. We are ready with the
time-honored replies: it is the ‘organism as a whole’; it is a ‘property of the
system as such’; it is ‘organization’ (Wilson 1923, 285).

In short, Wilson imagined the physical constitution of the cell as a

hierarchy in which each organisational level possessed a certain degree
of individuality and genetic constancy (Wilson 1923, 283). He imagined
‘as if viewed through an inverted telescope’ an infinite series of units of
various entities, beginning with larger molecular aggregates, passing to
smaller and larger visible endocellular structures, endocellular systems,
and cells (Wilson 1923, 282-283). In this way, Wilson managed to link
cell morphology with the colloidal chemistry of the cell, anticipating
basic ideas of molecular biology or, at least, preparing his readers for
future conceptions of a macromolecular constitution of the cell. In this
way, he also managed to link the subcellular with the supracellular
dimension. The cell was not an isolated unit, but an open system,
embedded in a hierarchy of organic units.
The question that immediately arises is: supposing a hierarchy of
units of different degrees exists, which level represents the truly vital
one? Wilson gave an intriguing answer: chondriosomes, Golgi bodies
and centrosomes probably possessed morphological identity and
permanence, and the power of growth and division, but only the cell
represented the vital unit, because ‘life can only be properly regarded
as a property of the cell-system as a whole; and the separate elements
of the system would, with Sachs, better be designed as “active” and
“passive”, rather than as “living” and “lifeless”’ (Wilson 1900, 29).
Within the hierarchy of organic units, the cell represented only one
unit of a series, but it was the one that enabled the treatment of
important biological questions in a way that promised a good chance
of scientific progress. Inheritance was linked to nuclear units, and
cellular metabolism probably to the ultramicroscopic units of the
cell-organelles but all processes could be fully understood only within
the framework of cellular organisation and interaction.

The cell as an organism

The second implication of Brücke’s Elementarorganismen adopted

by Wilson referred to cells seen as organisms. Though morphologically
THE CELL AND CELL THEORY 375

and functionally interconnected with the lower as well as with the

higher levels of biological organisation, the cell showed a certain
degree of self-sufficiency and completeness, so it could be subject to
evolutionary and developmental categories. This cell-organism-concept
enabled Wilson to see cells as beings equipped with a specific internal
organisation, and to apply to cytology the patterns of variability and
development in which he had been trained to think as a young student
at Baltimore and at Cambridge. The cell was not an abstract
philosophical unit, but, to use Boveri’s expression, ‘a microcosm, a
living system’ (Wilson 1923, 278). In his textbook he became even
more explicit: ‘The cell, like the whole organism, thus appears to us as
a delicately balanced moving equilibrium, offering a picture that is
continually changing yet always remains within the boundaries of a
specific type’ (Wilson 1925a, 637).
Like a higher organism, the cell had undergone ontological and
phylogenetic processes, reproducing, metabolising, synthesising, and
interacting with its environment and the surrounding organisms, and
differentiating in relation with all these internal and external factors.
This conception was useful to Wilson, in order to explain the
plenitude of cellular organisation. It enabled him to interpret
plasmodesmata, intercellular bridges, syncytia, polyenergids, and the
great variety of visible endocellular structures, not as counter-proofs
to the theory of cellular individuality but as variations of the same
fundamental cell-typus that had undergone different histories,
adapting to a variety of demands. It also enabled him to overcome the
contradiction between the idea that the cell potentially possessed ‘in
itself the complete apparatus of life’, and the admission that highly
integrated tissue-cells could be regarded only as ‘localized areas of
specific activity’ (Wilson 1925a, 101-104). And, in the end, it also
enabled him to make do with a rather simple cell-definition like
Schultze’s, and to present in his textbook not only figures of specific
cell-types, but also a diagram of the cell. Few cytological textbooks
ventured to do this, preferring to avoid pictorial representation, or to
furnish a series of figures of specific cell-types.
The way Wilson treats the cell as an organism reveals that his
approach to cell theory is not so much physiological or morphological
or embryological, but biological, based on what Bernardino Fantini
has called the dialectical process between the search for common
data and the multitude of concrete mainfestations (Fantini 1988, 111-
112).
376 ARIANE DRÖSCHER

Epigenesis, preformism and the physicochemical constitution of

the cell

Wilson’s ideas about the constitution of the cell cannot be separated

from his ideas about organellar genesis. Like other cytologists, he
attributed morphological value to the various constituents depending
mainly on whether or not they were permanent, and on whether or not
they possessed the power of growth and division. The concept that a
body is formed out of an amorphous substance implies that this body
remains subject to this substance. If, on the other hand, it is capable of
self-perpetuation, it is considered an autonomous entity. Wilson was
aware that also his ‘larger molecular aggregates’ had to be defined in
this regard. He did not avoid the question about the origin of living
matter, and in the course of time he became increasingly conscious that
he was in the middle of the dispute between epigenesis and
preformism.
During the preceding centuries, more than once, the dispute
between epigenesis and preformism had been declared settled. In
reality it had only shifted down to a smaller scale. In the 19th century,
no serious scientist still thought of a homunculus sitting inside the egg
cell or the spermatozoon waiting to develop, nor did he assume that
worms grew out of decaying organic substance. But the controversy
found fertile terrain in the discussion about the formation of cells, and
in the 1860s, in the idea of protoplasm as living matter in its simplest
and most primordial state (Geison 1969). Around the turn of the
century, the debate took another step downward to still smaller
entities: the genetic and the morphological continuity of chromosomes
and cell-organelles.
During the first decades of the 20th century it became increasingly
difficult to completely reject the idea of the existence of formed bodies
within cells. Even the supporters of the colloid chemistry of the cell that
accused cell morphologists of studying mere artefacts could not deny
that there had to be something inside the cell that caused the specific
staining reactions. The debate concentrated on two aspects: 1. Whether
or not these ‘structures’, namely chromosomes, chromidia, centrosomes,
Nissl bodies, chondriosomes, mitochondria, and Golgi bodies, were
permanent, and hence distinct morphological entities. 2. If they were
permanent, whether they had derived from identical pre-existing
structures of the mother-cell or whether they had been built up ex novo
from the homogeneous protoplasm of the undifferentiated cell.
Those who considered all visible particles mere passive ‘lifeless’
products of the cell-metabolism, or of the gelification of fluid phases,
THE CELL AND CELL THEORY 377

had no difficulty in explaining their progressive formation due to

spontaneous, simple and usually reversible physico-chemical processes
within the cytoplasm. The processes had to be investigated, not the
structures (see e.g. Berthold 1888; Verne 1923; Lepeschkin 1924, 54-
57; Bottazzi 1925, 32; Pensa 1925, 67). But those who regarded at
least some of the ‘bodies’ as permanent and physiologically active
‘living’ cell-organelles, were faced with the dilemma of admitting
spontaneous generation in the case of their formation ex novo, or
preformation in the case of self-perpetuation.
It is not easy to decide to which of the two factions Wilson belongs.
On the one hand, he was a convinced morphologist emphasising the
structural features of the cell. But he regarded the whole cell as an
interacting system, and in his developmental studies attached greater
importance not to the nucleus itself, but to its position within the cell.
According to Alice Baxter, he divided germ cells ideally into two
separate elements: an epigenetic cytoplasm responsible for development,
and a preformed nucleus responsible for the transmission of the
hereditary qualities. Thus, he was at the same time a preformationist
who admitted an underlying structure of organisation, and an
epigeneticist who argued that the ‘formative stuff’ is not necessarily pre-
existent in the egg cell but is formed by progressive formation. In this
way, Baxter argues, Wilson separated the processes of development and
transmission, and thus resolved the epigenesis – preformation –
controversy (Baxter 1976). Hermann Joseph Muller underlines that
Wilson advocated only chromosomal continuity, disproving any
evidence that indicated cytoplasmic heredity (Muller 1943).
Both are only partly right. As we will see, Muller’s view is simply
erroneous. Baxter’s interpretation is mainly due to her definition of
epigenesis and preformation, that follows Russell’s. In 1930 Russell
had defined the epigenetic approach as dynamic, vitalistic and
physiological, stressing time and process, and emphasising function,
and the preformistic approach as static, mechanistic, deterministic and
morphological, stressing space and momentary state and emphasising
form (Russell 1930, 29). Applying these criteria to the question of
cellular constitution, it would follow that an epigeneticist tends to
presume a homogeneous substance as a starting-point of development,
at least with regard to the very first step, the germ cells. Hence he is
not likely to be seeking any specific material or structure within the
cell. His approach will probably be more physiological, treating the
cell as an ‘unbreakable whole’ and as more than the sum of its parts.
During the experiment he will be anxious to maintain the cell intact
so as not to impair its functionality. A preformationist, on the other
378 ARIANE DRÖSCHER

hand, will follow a morphological approach. He will search for

specific, permanent bodies or carriers of determined functions, and
tolerate the use of aggressive cytological techniques in order to reveal
the latent, invisible structures. Preferably these special stuffs are
imagined as handed over from past cell-generations.
These categories are intriguing but not very helpful, except to give
a very superficial perspective. By applying Russell’s definition there is
a very good chance of finding innumerable intermediate positions.
There were scientists that could be called epigeneticists for certain
features but not for others, or preformationists in one period of their
career and afterwards not. Ernst von Brücke, for example, was a
physiologist, but he understood cells as structures of a far more
complex morphological organisation than that revealed by
microscopic analysis. The comparative anatomist Thomas Huxley, on
the other hand, considered his protoplasmic theory a physiological
and epigenetic view but also a triumph over vitalism (Geison 1969,
279). Schleiden’s and Schwann’s cells blocked epigenetically out of
amorphous blastem. Nevertheless, the emphasis they laid on cells as
developmentally autonomous units was criticised by Huxley as a new
form of preformationism (Richmond 2000, 250).
More than an overall division into morphological preformationists
and physiological epigeneticists, it is necessary to differentiate exactly
which structural level represents the point of debate, and which
criteria were applied to distinguish their own position from those of
the others. Peter Bowler has shown that in the 17th and 18th century
the difference between epigenesis and pre-existence-theories lay in
whether an organism was supposed to be formed before or after
conception (Bowler 1971). Hence definition has to take into account
the temporary border-line, the exact moment at which one may decide
whether or not a structure already exists. The second aspect concerns
the operating level. Jan Sapp, for example, has shown that the
distinction between genetic continuity and the formation ex novo of
centrosomes became less clear with the transition from light to
electron microscopy (Sapp 1998). At the macro-anatomical level the
answer is different from the one given at the microscopic, at the ultra-
structural or at the biochemical level.

Wilson, a reluctant preformationist

For most of his life, Wilson was convinced he was an epigeneticist.

Indeed, in the first edition of The Cell he wrote:
THE CELL AND CELL THEORY 379

The facts point rather to the conclusion that all cell-organs arise as differentiated
areas in the common structural basis of the cell, and that their morphological
character is the outward expression of localized and specific forms of metabolic
activity (Wilson 1896, 212).

But to call Wilson an epigeneticist becomes problematic even in the

early pages of the same book:
We are compelled by the most stringent evidence to admit that the ultimate basis
of living matter is not a single chemical substance, but a mixture of many
substances that are self-perpetuating without loss of their specific character. The
open question is whether these substances are localized in discrete morphological
bodies aggregated to form the cell somewhat as cells are aggregated to form
tissues and organs, and whether such bodies, if they exist, lie within the reach of
the microscope. Altmann’s identification of the “granulum” as such a body is
undoubtedly premature; […] It is nevertheless certain […] that at least one part
of the cell, namely the nucleus, actually consists of self-propagating units of a
lower order than itself, and there is some ground for regarding the cyto-
microsomes in the same light (Wilson 1896, 22). (Emphasis his)

Later, in 1923, he became even more explicit:

For my part, I am disposed to accept the probability that many of these particles,
as if they were submicroscopical plastids, may have a persistent identity,
perpetuating themselves by growth and multiplication without loss of their
specific individual type (Wilson 1923, 283).

Wilson’s idea of organellar genesis looks different if read in the

context of his corpuscular cell theory. It becomes evident that he
tended very early to a preformistic conception of inheritance, as well
as of development. Becoming more and more aware that, in the last
instance, it was impossible to decide whether or not cell-organelles
were permanent, Wilson passed to the underlying level of his
hierarchy. At this point, again inspired by an intuition of von
Brücke’s, he made an important decision: On the microscopic level it
was quite possible that the visible structures were formed ex novo, but
on the sub-microscopic level something definite and self-perpetuating
had to exist that secured the identity of the nuclear, as well as of the
cytosomic structures throughout the ontogenetic and phylogenetic
history of the cell.
For several years the centrosome, and above all the question
whether it persisted or was formed ex novo, captured Wilson’s
interest. He did not regard the division of the cytoplasm as a simple
mass-division in succession of karyokinesis, but rather as an
autonomous process. He hoped that the study of the centrosome,
380 ARIANE DRÖSCHER

about the year 1900 regarded as the smallest visible structure, could
provide the necessary information on how the cytosome perpetuated
itself from one cell-generation to the next (Wilson 1895; Wilson
1901). Later, Wilson directed some of his students to study other
components of the cytosome, namely the mitochondria and the Golgi
apparatus (Bowen 1924; Bowen 1929; Wilson 1925b; Wilson 1931;
Wilson and Pollister 1937; Pollister 1932).
Wilson himself had a definite opinion on the harsh polemics
surrounding endocellular structures. He admitted the real existence of
plastids, centrioles, chondriosomes and even of the suspicious Golgi
bodies, and assigned them persistency, almost omnipresence, metabolic
functions, and the power of growth and division. In 1926 he went even
further, and stated that the Golgi bodies were received from the last
generation of oogonia, and hence from the mother-organism (Wilson
1926, 118). Thus Muller is certainly wrong when he says that Wilson
disproved any evidence that indicated cytoplasmic heredity (Muller
1943). But at the same time, Wilson accepted the testimonies of other
researchers that stated to have observed cytoplasmic bodies to be built
ex novo. The resulting dilemma considerably puzzled him. His
corpuscular hypothesis offered the aspired synthesis in so far as:
[…] by ascribing to these hypothetical units the power of growth and division,
in accordance with the pangen theory, we are enabled to get a certain amount of
light upon some of the most puzzling questions of cytology, such, for example,
as the ultimate nature and origin of dividing cell-organs like the nucleus or the
plastids, and especially such a contradiction as that presented by the centrosome
which may apparently arise either ex novo or by division of a preëxisting body of
the same kind (Wilson 1899, 19).

Likewise, his micromeristic conception resolved the question

whether the protoplasmic organisation was alveolar, fibrillar or
granular. The visible image was ‘of only secondary significance for the
problem of protoplasmic structure’ because all these pictures ‘are but
varying aspects, kaleidoscopic pictures, of heterogeneous systems that
comprise a great variety of components, both formed and unformed’
(Wilson 1926, 119).
Wilson had harboured ideas of hypothetical units that self-
perpetuated, and thus represented the basis of genetic continuity, at
least since the 1890’s. In the third edition of The Cell he repeated an
identical concept using nearly the same words (Wilson 1925a, 721). In
his study on cytokinesis of the same year, Wilson responded to the
question whether the components of the cytosome arose epigenetically
or from pre-existing material:
THE CELL AND CELL THEORY 381

The possibility nevertheless remains that cytoplasmic systems generally may

include many different stuffs that are self-perpetuating by some kind of
autocatalytical process. We may venture the further surmise that such stuffs may
undergo individuation in various degrees to produce visible formed bodies which
in their fullest development become definitely organized as such and capable of
self-perpetuation (Wilson 1925b, 494).

Quite some time passed before Wilson became aware that his ideas
corresponded to a preformistic view. This reluctance can be explained
by our reading the introduction of the third edition of The Cell. In his
sketch on the history of embryology and cytology, Wilson presented
preformation as the theory of encasement, hence in the sense of
Charles Bonnet’s emboîtement, and epigenesis as the theory of
progressive new-formation. In the following evaluation of both
theories Wilson left no doubt that the first represented a ‘logical tour
de force’, whereas it was the merit of the second to have provided the
foundation for modern embryology (Wilson 1925a, 6-7). Thus
considering Wilson’s prejudice towards preformation, in addition to
the fact that he indeed intended development as a process of
progressive formation, it is comprehensible that he felt rather
uncomfortable when, in the end, he was constrained to admit that, in
certain respects, he himself held preformistic ideas:
Admittedly, the foregoing suggestions involve a somewhat preformistic
conception of cytoplasmic systems; but it is one that differs widely, I think, from
the intracellular pangenesis of Weissmann, Altmann and similar micromeristic
hypotheses of other earlier writers. Should it prove well founded it might offer a
more acceptable alternative to those speculative constructions – one that is more
in accordance with modern views of protoplasm and comes more closely into
touch with the problem of development. Experimental studies in embryology
have clearly demonstrated the correctness of the early conclusion of cellular
embryologists that the cytoplasm of the unsegmented egg contains specifically
different, but not visibly formed, materials that are segregated in definite manner
by the process of cleavage. Whether any or many of these are self-perpetuating
is a question of far-reaching interest alike to the embryologist, the cytologist and
the geneticist, and hardly less so to the biophysicist and the biochemist (Wilson
1925b, 495).

Initially, for Wilson ‘preformism’ and ‘epigenesis’ were historical

expressions that referred to debates of preceding centuries. But, the
more he meditated upon his own ideas on genetic continuity, the
more he became aware that they still had a value in the discussion of
present-day problems.
Considering the various operative levels, the answer to the question
whether Wilson was a preformationist or an epigeneticist has to be a
382 ARIANE DRÖSCHER

differentiated one: On the cellular level Wilson was clearly a

preformationist and, around the turn of the century, only very esoteric
scientists would have been in dissension. On the level of cellular
systems, namely the nucleus and the cytosome, he was likewise firmly
convinced he was dealing with systems that were entirely passed on to
the next cell-generation, and this was still a point of harsh debate. On
the subcellular level he became increasingly convinced of the power of
self-perpetuation of certain cell-organs, but admitted also contrary
cases, and here he stood in opposition to nearly all leading cell-
physiologists and cell-chemists and even many cell-morphologists. On
the sub-microscopic level, finally, Wilson cautiously expressed a
preformistic vision too, apologising for its very speculative nature. On
this ultimate level Wilson found the smallest audience.

Final remarks

The three editions of The Cell in Development and Inheritance

reflect Wilson’s personal approach to biological questions. Though he
made considerable use of ideas or concepts, and even the words of
other cytologists, he never made use of them without having
thoroughly thought them through. Cell theory stands at the basis of
Wilson’s biological theories in general. He comprehended vital
phenomena not as a series of separate questions, but as a whole. In
the last instance, all biological questions converged in his conception
of the cell and its expressions, and, vice versa, his conception of the
cell was the essence of his biological thought. Through his
corpuscular and micromeristic conceptions Wilson held together his
ideas about constitution, organisation, functionality, development and
inheritance (Dröscher 2003).
Despite all the admiration and the universal praise that was
showered upon it, Wilson’s cell theory was and still is undervalued.
This is not the case with the value of his research contributions, even
if these, when compared with his contribution to chromosome theory,
was certainly minor. It is generally recognised that he gave significant
answers to a series of important cytological problems, like the debate
around the centrosome, the physical constitution of the cytoplasm, or
organellar research.10 These alone have ensured Wilson a place in the

10 For example, it is owing in great part to Wilson’s student Robert H. Bowen, prematurely deceased,
that the Golgi apparatus research definitely broke with its early static concepts and underwent
considerable progress in the 1930s (Dröscher 1998).
THE CELL AND CELL THEORY 383

history of cytology. Likewise, the underestimation does not concern

The Cell in Development and Inheritance as a textbook in general. Yet
the general attitude is well represented by Garland Allen’s remark that
Wilson was the one who had best raised the essential questions (Allen
1976, 430). The answers were evidently sought for elsewhere. Many
aspects of his conception of the cell were not grasped. It is
acknowledged that he produced an extraordinarily dynamic, plastic
and comprehensive conception of the cell, yet the unity he propagated
between cytology, heredity and development soon fell apart. His
preformationist, micromeristic, and corpuscular cell-organism-theory
found no direct adherent.
Today it is surprising how modern many of Wilson’s concepts
sound. Probably his contribution can be truly appreciated only
posthumously. Wilson adapted and transferred some of the most far-
reaching conceptions of Zellforschung into the new era of
experimental biology. Of course, his microsomes cannot be seen as
the direct precursors of Albert Claude’s microsomes. Claude referred
to definite morphological particles, provoking a renewed polemic in
the 1940’s and 1950’s about the actual existence of such structures
(Claude 1943; Brachet 1957, 40-45; Rheinberger 1995), whereas
Wilson’s conception referred to a more general and still smaller level.
But many other aspects – such as the unity of cytology, development
and inheritance, the cell as a complex but unbreakable ‘microcosm’,
the cell as subject to evolution, the cell as the principal unit of
biological investigation, the value of morphological research in
cytology, the concept of the interaction between the single parts and
the whole, the concept of the interaction between nucleus and
cytosome during cell differentiation – had been popular among the
supporters of Zellforschung, were propagated and adapted by Wilson
to the cytology of the 1920s, were increasingly abandoned by younger
cytologists, and revived several decades later. Even Wilson’s
corpuscular theory resembles today’s concept of the molecular biology
of the cell. The idea of ‘higher molecular aggregates’ originated in
Naegeli’s hypothesis of ‘micellae’, were transformed by Wilson into
the concept of self-perpetuating stuffs that ‘may undergo
individuation in various degrees’ and that constitute the structural
basis of the cell, and revive in cytological textbooks of today that
advocate very similar opinions (see e.g. Alberts et al. 2002, 11).
But when the time was ripe, and Wilson’s speculations became less
unimaginable, other models were adopted. In 1938, for example,
plant physiologist and colloid chemist Albrecht Frey-Wyssling
developed ideas that aimed in the same direction. Thirteen years after
384 ARIANE DRÖSCHER

the appearance of the third edition of The Cell, he considered the

structure of the protoplasm a ‘coherent molecular scaffold’
(kohärentes Molekulargerüst) made up of macromolecules. Frey-
Wyssling regarded cell-components as well as genes as ‘specific
proteic groups’ that ‘cytoplasm is not able to create ex novo’. He even
went so far as to declare that the single macromolecules had been
elaborated through the long process of phylogenesis (Frey-Wyssling
1938). Yet he made no reference to Wilson. Nor did Tschermak,
Chambers or Heilbrunn.
One of the reasons for his disregard is probably Wilson’s explicit and
firm adherence to the discussion of the end of the 19th century. Wilson’s
cell theory was a child of German Zellforschung. It was formed around
the year 1892, and remained basically the same, though adapted in the
course of the years to later discoveries and conceptions. But personalities
like Schwann, Virchow, von Brücke, Boveri, and Bütschli, as well as the
questions that had moved them, became increasingly unknown and
incomprehensible to the young generation of cytologists. In the 1920s,
Wilson’s comprehensive intellectual approach, and his pleasure in
drawing logical conclusions that went beyond the apparent experimental
results, were definitely out of fashion. During the 1920s and 1930s, the
scientific enterprise tended towards rigorous experimental
argumentation, physico-chemical approaches, and restricted research-
problems. Moreover, historians of science generally hold the opinion that
molecular biology arose on new methodological techniques like electron
microscopy, chromatography and ultracentrifugation, and on new
research subjects (see e.g. Rheinberger 2000). The influence of
theoretical reflections was minor, and hence Wilson cannot be seen as a
forerunner. Experimentally, in the 1920s, Wilson’s morphological
approach to cytology had reached its limits, and biochemical and ultra-
structural research still did not have the instruments necessary to
transform his intuitions into experimental research-projects.
Wilson was very aware of the very speculative character of his
theory. Indeed, as far as his presentation of cell theory is concerned,
there is a strikingly frequent use of expressions of uncertainty, or at
least of unwillingness to be too explicit, like ‘if it exists’, ‘in some
sense or other’, ‘perhaps it is, perhaps not’. Moreover, he never forgot
to make it clear that the explanation he was putting forward was
‘valid as a working-hypothesis’ or ‘aids practically in investigation’.
More than real uncertainty with regard to the very delicate questions
concerning the essence of life, these expressions indicate caution.
Especially in the United States, which set high value on their being so
different from ‘speculative European science’, as they called it, highly
THE CELL AND CELL THEORY 385

stimulating but purely theoretical concepts like those circulating in

Germany at the end of the 19th century found a very limited
reception. Wilson, on the other hand, was not willing to give them up
but used them to overcome experimental inadequacies or
contradictions. He aimed at a complete and synthetic cell theory, and
in order to fill the evident gaps in direct experimental results, he
needed ‘metaphysical speculations’. He did not claim he had seen the
‘formative stuffs’, but he was convinced they must exist.
Although Wilson’s cell theory did not receive the attention it
deserved, it is possible that it influenced general cytology indirectly. In
the same way as Muller emphasised that Wilson’s contribution to
chromosome theory had become wholly evident only several decades
later (Muller 1943, 30-31), his cell theory probably helped many
cytologists to retain a general concept of the cell compatible with
those molecular biology later outlined.

Acknowledgements
I would like to thank Desirée Sanges and Charles Hindley for the
revision of the manuscript, and my referees for their kind and helpful
suggestions.

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