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Biological Journal of the Linnean Society, 2013, , . With 1 gure

REVIEW ARTICLE A reection on biological thought: whatever happened to the organism?

20 Pickwick Road, London SE21 7JW, UK
Received 24 May 2013; revised 9 August 2013; accepted for publication 9 August 2013

Biological thought requires at least both a concept of organism and a concept of evolution to be causally efficient; this is above and beyond any materialistic cause or causes, however formulated. Although the concept of evolution has been much debated and developed over the last century and a half, the concept of organism has been neglected, despite promising modern beginnings with the works of W. E. Ritter and others. The independent efforts of ve biologists, including Ritter, to develop a concept of organism are outlined, key works are cited, and some similarities and differences of their approaches are outlined. E. S. Russells teleology of organism, far from being a failed worldview, is considered to be a major signicant step towards a more complete biology. 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, , .

ADDITIONAL KEYWORDS: Agnes Arber E. S. Russell J. H. Woodger Kurt Goldstein materialism

mechanism teleology W. E. Ritter.

The present essay was inspired by two meetings at the Linnean Society of London in 2011: Straininduced assembly hypothesis and the growth of form (24 March 2011), organized for David Knight by David Cutler and Andrew Packard, and The role of behaviour in evolution (8 September 2011). Both meetings addressed aspects of possible intrinsic exploration and explanation of problems associated with organisms and their organization, rather than the currently more usual approach of extrinsic explanation for organismal form, function, and existence. Both meetings concerned themselves with a welcome return to the organism after what seems a very long digression during which organisms have been relegated to the status of billiard balls set in motion solely by the action of external evolutionary forces. In point of fact, organisms are active, maintaining, growing, developing, and reproducing entities. Furthermore, organisms display an endless continuity and are capable, in part, of continually choosing paths

at the spatial and temporal edges to their own futures, albeit for brief durations only, and of uncertain consequences. What follows are some reections on the history of certain ideas in biology for the period 19191954, framed by the writings of W. E. Ritter, commencing with his organismal conception expounded in The Unity of the Organism (Ritter, 1919b), and concluding with the posthumous publication of Charles Darwin and the Golden Rule (Ritter, 1954). The latter includes a full bibliography of Ritters written works.


The main idea herein traced is that of the organismal conception but not, for example as Mayr (1997) categorized it, as organicism, an abstraction or extension, or as systems theory, another extension, which von Bertalanffy (1952) would ultimately fashion from a similar starting point to Ritters, or as holism as developed by Smuts (1926), yet another abstraction. This organismal conception, or to use Ritters phrase the unity of the organism is rather just deep reection on the concrete nature of the organism and of the


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R. W. BRUCE thesis. For example, the works of John Scott Haldane spanning the rst few decades of the 20th Century are replete with considerations of the nature of organisms (Haldane, 1929) but, in the key work of Huxley (1942), even the term organism, as a concept, is absent from the subject index. Moreover, the thoughts of J. S. Haldane are not cited therein but, instead, those of his son, J. B. S. Haldane, are now brought to the fore. I am not arguing for a simple cause and effect here, just noting that as evolution as a concept expanded, it appeared that organism as a concept contracted. I do not believe this to be a wholly intellectually balanced state of affairs. The ve biologists considered in this account are William Emerson Ritter, Kurt Goldstein, Agnes Arber, Edward Stuart Russell, and Joseph Henry Woodger. A naive classication would list them as three zoologists (Ritter, Russell, Woodger), a botanist (Arber), and a physician (Goldstein). Many others should also be included, although my lack of knowledge and space limitations require the present circumscription. J. S. Haldane and Ludwig von Bertalanffy already mentioned above, and Conwy Lloyd Morgan and Joseph Needham immediately come to mind as requiring further consideration within this arena of ideas. Indeed, Needham appears to manage to be both for and against the concept of organism in his writings of this period, an ambivalence that might reward close scrutiny. But it must be left to another essay or to others to pursue the efforts of these biologists. The largely Anglo-Saxon outlook is a result of my linguistic and cultural parochialisms, and my failure to embrace the world of micro-organisms is another signicant limitation. These shortcomings withstanding, I have come to the belief that, if one scratches the surface of biology in any culture or language during this period, approximately 19191954, evidence of a concept of organism would be quickly exposed. Thus, my chosen ve are in effect exemplars of a widespread approach to biology now largely in eclipse. As such, the ve do present service for the past efforts of the many. Their efforts are both real and substantial but now carelessly overlooked. All ve had more than sufficient ability to let their written works continue to speak for themselves. The diversity of outlooks that the ve had on organisms are, I believe, refreshing and stimulating, and serve as a balance against the current monopoly of evolutionary materialism.

necessity of such a concept in any signicant greater biology. A large part of the biology of the last 150 years has centred, not surprisingly, on species and their origins (i.e. an evolutionary concept concerning the change of collectives through time); however, such an approach presupposes that organisms as individuals and their continuity are both real and actual and this too must be considered in any analysis and possible attempted synthesis. If the present review has any aim beyond exposing the thoughts and writings of an extraordinary collection of biologists to those as yet perhaps unfamiliar with them and their works, it would be that of stressing the necessity of the concept of organism in any framework of biology. This might seem a strange or even foolish aim; surely organisms are always understood to be the substrate and essence of biology? But are they? Or are we just confusing means and ends here? Is not the concept of the billiard ball more convenient and less demanding for our explorations and our explanations? The evolutionary metanarrative has, for better or worse, become our only intellectual explanation of life in its many forms, although the fact that life presents itself always as organisms remains curiously understated. Life is a property of organisms, and organisms alone, as far as we know, bear this property. Conversely, concrete organisms are not a property of abstract life but rather part of an unbroken continuity of unknown duration. Unless we wish to return to animism in some form or other, we just have to acknowledge that life, in general and in particular, always resides in organisms. The organism as a concept in biology was not always overlooked and understated, as the efforts of the subjects of this essay bear witness. Without a concept of organism, biology is much diminished, and fractures into competing cults of artefacts and abstractions. This is not to say that biology is sufficient as an intellectual pursuit with only the concept of organism but rather that, without this concept, there is nothing to hang disparate categories, threads, narratives, theories, and abstractions on, and they drift and recede into the reied ether and beyond. Without the organism, there are no knots in the nexus. Rather than a dialogue with reality, we content ourselves with embracing abstraction, and then praise our efforts in so doing; organisms alone allow our biology to be grounded in endless, renewing and continuous actuality and, as such, the organism is a category of nature above and beyond any material considerations of its parts. The concept of organism was most clearly developed in biological thought within this period by Ritter and many others, and then appears largely to have been forgotten. This loss of concept of organism appears coincidental with the development of the modern syn-


The beginnings of any continuity must be problematic; this is as true for organisms as it is for ideas. Perhaps Hans Drieschs Gifford Lectures 19061908, published as The Science and Philosophy of the Organism (Driesch, 1908), or indeed Henri Bergsons

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WHATEVER HAPPENED TO THE ORGANISM? Creative Evolution (Bergson, 1911; in English translation) might have warranted consideration as just such beginnings. These works certainly shaped the eld where vitalism met materialism. But I think that Ritter serves as the best exemplar for re-developing the concept of the organism as something more expansive than materialism and mechanism, and yet more concrete than the vitalism of Driesch and Bergson. Although Ritter was instrumental in moving biology past that limiting and limited antithesis, the fact that, by default, we continue to return to the antithesis suggests that we have paid scant attention to Ritters lead. The world we all live in is, proximally, distally and ultimately, a world of organisms. Perhaps this is just a self-evident truth, although it does reect the reality of our past, present and possible future existence, and we can only interact with the world by being organisms ourselves; something Ritter wrote about in detail during the later years of his life and formed the basis of Goldsteins biological method. As an aside, perhaps Descartes should be considered as the primordium for a modern conception of organism with cogito, ergo sum, at least with respect to humans; existence and organism were thereby fused. It remains an irony of both history and Descartes metaphysics that dualism remains his enduring biological contribution down to the present. Alas, there was man and then the rest of nature in his taxonomy of the world of beings (and regrettably Darwins insights lay unattainable in future time). Descartes divisions of nature thus still arrest biological development. Yet these divisions reveal the remarkable relationship between a false natural classication and the development of thought. It is an example of the problems that arise from embracing the fallacy of the bifurcation of nature (rather than a laudable bifurcation of thought) as exposed by Whitehead (1920). By 1960, the concept of organism had fallen from intellectual interest, with the molecule (we all know which one), the gene, the cell, the system, including the ecosystem, phylogenetics and, most recently, informatics, now vying to ll our biological gaze, all operating under some form of a concept of evolution. Perhaps it is little wonder that we have no time or space any more for a concept of organism in this crowded intellectual world. The practical world cannot allow itself such self-indulgence; ask the plant and animal breeders, the sherman, the pharmacologist, and the sewage engineer: in their real world, continuity of variety, breed, species, strain or culture must be maintained. Indeed any concept of evolution requires, as a necessary corollary, a concept of organism; the former concept is a theoretical postulate concerning time, space and the ultimate causes of change in collectives, the latter concerns the empiri-

cal and proximal reality of individual continuity of living beings: the theory and the practice of life, respectively. The times under consideration are then the second to sixth decades of the 20th Century; less than half a century of concerted intellectual effort as evidenced by the works of Ritter, Goldstein, Arber, Russell, and Woodger. The degree of direct interaction between these individuals and their detailed interrelationships are not pursued to any extent here. Indeed, such an undertaking would be a huge task, although there was interaction. However, I do not suggest that there was a concerted and conscious unity of purpose in their efforts; they were merely explorers of organisms, which they found to be eternally fascinating. My purpose is to note that the concept of organism was something in the air during this period, to celebrate the efforts of these biologists to pin the concept down and to give it intellectual form, and to give an indication of works that can be consulted by any reader moved, like me, to acknowledge the extraordinary contributions of these ve during this period. Beyond their contributions, the loss of their ideas and with it a loss of continuity into modern intellectual culture is both disappointing and troubling. Disappointments can of course be reduced by replaying them through remembrance and nally dispelled by renaissance. But, if something as necessary, articulate, rational, reasonable, empirical, sane, and concrete as the concept of organism can disappear almost without trace and comment from mainstream biology, what does it say about our resolve, our science, and ultimately our values? Why did this happen? For the present, I only speculate; at an immediate level model organisms and shallow intellectual models impoverish biology, albeit unconsciously. Technicism, the appeal of novelty and specialization fracture biology. The title of Agnes Arbers nal book, The Manifold and the One, directs us to what biological thought ought to be but seldom is. Perhaps, at a deeper level, evolutionary materialism was allowed to gain a monopoly of the mind, a monopoly to which we have all contributed. In the nal analysis, a scientic theory which explains everything explains nothing. I therefore found it encouraging that the Linnean Society of London hosted two meetings in 2011 that placed organisms, once again, front of stage as individual concrete beings, and the metaphysical and abstract outlooks of much of current biology were challenged.


Ritters long life and career demonstrates a restless mind searching always for areas for further growth,

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once one attains a world view which truly strives to include, but makes no pretense to having already included, the whole world wholly in that view (Ritter, 1919b: xviii)

be it personal, institutional, professional, national or international. First and foremost, he was a general zoologist, reecting always on the organismal nature of life: shes, frogs, newts, snakes, lizards, birds, echinoderms, hemichordates, although mostly ascidians, colonial and solitary, were held in the gaze of his researches. Description, denition, and classication were the foundations of his knowledge but he refused to stop there; the organism in its environment was always part of his philosophy of zoology. Thus, the need for a marine laboratory on the Californian coast became a further quest, and the result grew into what would become Scripps Institution of Oceanography. The need for the proper dissemination of scientic results to a wide audience also animated him, and the result was the Science Service. Humans as part of the natural order of life caused him to reect on human nature from a zoological perspective, and he was a strong advocate of the League of Nations after the First World War. In a time of possible idealism, he was an American Idealist but with a thoroughgoing realistic streak born out of scientic scepticism and social pragmatism, especially when considering the nature of the human animal; progress was possible but not inevitable, and ideology, any ideology, was repugnant to him. The concept of the organism in its modern form belongs to Ritter (Russell, 1930); contra Descartes universe, all life forms were now a conjoined unity. An online bibliography (Ollhoff, 2013) and a brief biography (Day, 2006) are available. The titles of his books surely must intrigue biologists to at least browse his contributions, which are profound and humanitarian, in a way that few biologists of any age could match: War, Science and Civilization (Ritter, 1915); The Higher Usefulness of Science (Ritter, 1918a); The Probable Innity of Nature and Life (Ritter, 1918b); An Organismal Theory of Consciousness (Ritter, 1919a); The Unity of the Organism (Ritter, 1919b); The Natural History of Our Conduct (Ritter, 1927, with E. W. Bailey); The Californian Woodpecker and I (Ritter, 1938); and Charles Darwin and the Golden Rule (Ritter, 1954, with E. W. Bailey). Ritters ability to hold up a mirror to the human organism, especially himself, identies him as a remarkable zoologist. As a marine biologist, Ritter found much compatibility with Aristotles life and the latters world view, and especially with the concept of entelecheia, which Ritter considered best translated as complete reality. Two brief statements of Ritters are worth recalling. His political philosophy, if it may be so called, is summed up in the preface in The Unity of the Organism thus:
The reason why sincere humility and the spirit of democracy are alien to all forms of idealistic philosophy becomes clear

Some decades later, Karl Popper, echoing Ritter, would describe idealistic philosophy as the enemy of the open society. The second quote sums up Ritters biological philosophy, if it may be so called, again from the introductory section of The Unity:
. . . the organism in its totality is as essential to an explanation of its elements as its elements are to an explanation of the organism (Ritter, 1919b: 24)

For Ritter, no simple materialistic chain of temporal causality was therefore sufficient for the explanation of living things. What was necessary for Ritter was a reciprocation between the materials of the parts and the organism and the organism and the materials of the parts. Although Ritters views, by his own consent, can ultimately be located in the tensions between the concept of organism of Aristotle and the elementalism of Lucretius, more proximal inuences are to be found in the efforts of three American biologists, C. O. Whitman, E. B. Wilson, and F. R. Lillie to pursue and attempt to clarify the relationships between cells, embryos, and organisms. These relationships would be returned to again and again by Ritter, Russell, and Woodger with, I believe, penetrating insight.

K. GOLDSTEIN, 18781965
Goldstein was principally a physician, neurologist and psychologist, and inuenced much of mid-20th Century psychology, although little of the biology of this period, which is truly unfortunate. Anyone who wrote a text entitled The Organism (rst German edition 1934) that still remains in print (Goldstein, 1995), deserves signicant recognition in the biological thought of any subsequent times. What current biological interest there is for Goldstein centres on holism (Holdrege, 1999a, b). But before expanding thought to holism, the empiricism of the organism, and all that that entails, must be acknowledged. In this writers view, Goldsteins life and career were about exploring the reality of the organism, especially with respect to human mental disabilities and what such clinical cases can reveal about the normal human organism. His studies on braindamaged World War I veterans at the outset might appear worthy but limited, although, in the hands of Goldstein, this work grew to be expansive, trenchant, and penetrating, in ways unmatched by any initial expectations. Goldstein demonstrated what nature can tell us if we are humble enough to listen to its quiet messages; the biological roots of humans as organisms and their potential for individual growth

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WHATEVER HAPPENED TO THE ORGANISM? as human beings, irrespective of the severity of their limitations, illuminate his writings. Goldsteins deant humanity is well displayed in his William James Lectures Human Nature in the Light of Psychopathology delivered at Harvard in 1940, and published thus (Goldstein, 1940). Russell refers to this work in the appendix to his nal presidential address, on the behaviour of animals, made to the Linnean Society in 1943 (Russell, 1944). The preface to the German edition of The Organism (reprinted in the 1995 English edition) outlines Goldsteins clinical philosophy:
The intention to write this book goes back many years. It dates to that time of the world war when it became my special task as a physician to take under my medical care a great many patients with lesions of the brain. As a director of a hospital for brain-injured soldiers, my experiences compelled me to broaden the medical frame of reference to a more biological orientation. It soon became evident that only the biological approach is adequate to evaluate the changes which these suffering fellow men have undergone; moreover, the facts taught me that there was no other procedure available which could render aid, however imperfect, to these patients (Goldstein, 1995: 15)

abruptly shattered, who had seen co-workers and colleagues killed, and who faced an uncertain future at an uncertain time in western civilization, this text is a remarkable combination of positive thought, unyielding humanitarianism and scientic virtuosity.

A. ARBER, 18791960
Of the ve, Arber is probably the most appreciated currently. She has had a signicant inuence on botanical thought, and this continues and indeed deepens (Barlow, Lck & Lck, 2001; Claen-Bockhoff, 2001; Rutishauser & Isler, 2001), and she has been well served by biographies and bibliographies (Thomas, 1960; Packer, 1997). Her knowledge of botanical history was extensive and provided her with deep foundations for her later researches and speculations. Artistic training by her father, a professional artist, greatly aided her botanical presentations and inuenced her later biological epistemology, reected in her chosen title for this work The Mind and the Eye. Working institutionally, then privately on botanical history, plant morphology, and growth, and nally by thought alone, she pondered the nature of botanical form in a way that few others have done, to a depth few could consider. The later publications on the natural philosophy of plant form, origins, and methods of biological knowledge, and, nally, the metaphysics of existence, are roads largely untravelled, even unconsidered by most modern biologists. Her continual awareness of the limits of language, especially when confronted with the germinative nature of life (in the bud) serves as an eternal caution to scientic aggrandizement, especially true now in the age of the algorithm, when information and language have become conated, and it has become easy to pretend that machines do our thinking and talking. Arber was a Linnean Society gold medallist in 1948. Her plant books remain part of the botanical canon; for the benet of nonbotanists, they include Herbals (Arber, 1912; and subsequent reprintings); Water Plants: A Study of Aquatic Angiosperms (Arber, 1920); Monocotyledons: A Morphological Study (Arber, 1925); The Gramineae: A Study of Cereal, Bamboo, and Grass (Arber, 1934); and The Natural Philosophy of Plant Form (Arber, 1950). Her later speculations into the nature of biological knowledge, The Mind and the Eye, A Study of the Biologists Standpoint (Arber, 1954), and metaphysics, The Manifold and the One (Arber, 1957), are products of a critical, reective, and mature mind drawing on vast biological knowledge gained over many decades. Arbers outlook on biology is demonstrated in her preface to The Gramineae:

Some three decades later Goldstein again stressed the nature of his biological method:
. . . this would not change the books [The Organism] essential character, which consists not so much in the communication of facts as in the clarication of the problem of method in biological research and in elucidating ways of conceptualizing the empirical material (Goldstein, in the authors preface to the 1963 edition, reprinted in Goldstein, 1995: 17)

Goldsteins concern with the nature of the individual, whether it be normal or pathological, from a biological standpoint, nds little echo in most contemporary biology where the tness of the collective has become the ready explanation for all biological realities, and the adaptiveness and adaptability of the individual organism within its own lifetime has been conveniently overlooked. Goldstein (1940) attempts to root humans to their biological past as organisms and to consider their potential as human beings as organisms; of the former we can know about, of the latter we have the possibility of understanding. From understanding ourselves, we can then explore other organisms; from such a standpoint, from viewing all living forms as the organisms that they are, Goldsteins biology thus takes on a radical perspective. It is an enquiry into biology via the organism that one has the greatest possibility of understanding (ones self) and from there one can explore the rest of the world of life-forms. Written by someone who was hounded from his native soil, who had his career

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R. W. BRUCE Russell, President of the Linnean Society of London 19401943, wrote a series of books that have yet to be fully appreciated for their originality and clarity of thought. A biographer with the ability to take on his life and works is now needed to distil that originality and clarity for modern readers to appreciate. Russells works include, in addition to Form and Function, The Study of Living Things: Prolegomena to a Functional Biology (Russell, 1924); The Interpretation of Development and Heredity: A Study in Biological Method (Russell, 1930); The Behaviour of Animals: An Introduction to its Study (Russell, 1934); The Overshing Problem (Russell, 1942); The Directiveness of Organic Activities (Russell, 1945); and, posthumously, The Diversity of Animals: An Evolutionary Study (Russell, 1962). Ulett (2010) gives a brief but instructive biographic sketch. Russell is too easily dismissed as a teleologist by evolutionary biologists, although this is to misunderstand the nature of his teleology. For Russell, the teleology of the organism was a logical necessity born of the very nature of the organism and just had to be accepted as such; this perhaps is a classical view point that is now too far out of touch with modern material sensibilities. That materialism cannot survive with teleology was of little import for Russell but that the organism cannot survive without teleology was his lasting contribution to biological thought. For Russell, organisms do not only encapsulate material within their boundaries. Organisms also encapsulate time and space within themselves and self-generate directiveness, be it towards maintenance, adaptation or reproduction. Without signicant biographical reappraisal, such insights are likely to languish in the shadows of current evolutionary hegemony, and biology will remain but half a science and entirely a slave to materialism. The last book by Russell published during his lifetime was The Directiveness of Organic Activities. The preface outlines Russells intent, in the plainest of language:
This book is an experiment or adventure in biological thought. I have tried to work out the consequences of rejecting the mechanistic point of view in biology, and I have, not unnaturally, arrived at a conception of the living organism and a method for Biology which are entirely heterodox, and run counter to the ideas commonly accepted in the present theory and practice in biology. My rejection of mechanism is quite deliberate and for a good cause. The living thing can be treated as a physico-chemical system or mechanism of great complexity, and no one would dream of denying the validity and value of biochemical and biophysical research. But such an approach leaves out of account all that is distinctive of life, the directiveness, orderliness and creativeness of organic activities, and completely disregards its psychological aspect. I try to show that we cannot disregard these unique charac-

. . . I wish to stress the fact that this is only one of numberless possible books, for which, to different minds, the grasses might supply material. I have of the Grass, as Coleridge had of the Rose-tree, a distinct Thought, but I am continually conscious of his unanswered question what countless properties and goings-on of that plant are there, not included in my Thought of it? (Arber, 1934: x)

and in the closing lines of the same work:

What is the meaning of the differences that separate the Gramineae so delicately, yet so denitely, from any other order, and that so prevail that a grass remains a grass, however freely the type may vary? To attribute these differences to genic constitution is an explanation of a merely descriptive kind; it enables us, indeed, to assign a place to them in the mental framework which we impose upon reality, but in so doing we have shelved, not solved, the problem. The mystery abides (Arber, 1934: 409)

The exploration of the continuity of the rhymes and rhythms of organisms are Agnes Arbers enduring contribution to biological thought. The breadth and depth of her readings and writings are a sobering reminder of the nature of true scholarship in our age dominated by pervasive ideology and universal technicism. Her considerations of the reticulate nature of many of lifes relationships serve as a critical counterpoint to the current prevailing model of phylogenetic bifurcation, and strengthen and deepen any appreciation of organisms and their connections. With increasing evidence of lateral gene transfer between micro-organisms, Arbers insights into biological forms and their reticulations now appear fresh and felicitous when viewed against the linear model of bifurcation.

E. S. RUSSELL, 18871954
Russells perceptiveness is well displayed in his rst book, Form and Function, A Contribution to Animal Morphology (Russell, 1916), which is a critique and appraisal of biological thought from classical beginnings to the 20th Century. Russells own education was both classical and biological, and perhaps accounts for the success of this historical commentary. Lauder (1982) presents an interesting essay on this work and its continuing relevance to biology in the 1982 reprint, with a bibliography of Russells contributions. Russell worked as a sheries biologist but this practical vocation did not prevent his contemplation and pursuit of what was to be his lifes work: attempting to understand organisms as functioning wholes, and attempting to create a free biology, free from the strictures of materialistic thought, thereby raising biology to an appreciation and an interpretation of the organism.

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teristics of life without losing all hope of building up a unied, coherent and independent biology (Russell, 1945: viii)

This book concludes:

Our conclusion that life processes are essentially and fundamentally directive and creative may be rejected as metaphysical or mystical. It is of course nothing of the sort. I make no hypothesis as to the philosophical basis or ground of directiveness or creativeness. I merely accept the patent evidence that they are characteristic of living things and of them alone. Nor do I suggest anything strange in the way of method. I suggest simply that, instead of making continual and vain efforts to squeeze biological facts within the materialistic frame, and attempting analysis without end, we accept them as biological, that we deal with the problems of development, maintenance and reproduction in terms of the observable activities of the organic agents concerned, without making the gratuitous hypothesis that these activities are mechanistic. Only in this way can we hope to establish the laws of organic activity (Russell, 1945: 192)

organisms and their intrinsic and extrinsic connections immaterial, material and organismal, and include Elementary Morphology and Physiology for Medical Students (Woodger, 1924); Biological Principles: A Critical Study (Woodger, 1929); The Axiomatic Method in Biology (Woodger, 1937); Biology and Language (Woodger, 1952); and Physics, Psychology and Medicine: A Methodological Essay (Woodger, 1956). Woodger concluded Biological Principles with a chapter titled The Future of Biology. One arresting sentence reads:
Clearly one of the most interesting and important problems for the biology of the future is that of the relation of characters to parts, of genetic factors to developmental processes, of the persistent racial immanent endowment to the process in which it is displayed in the course of temporal passage (Woodger, 1929: 484)

Russells quest for a free biology has but barely started, although he outlined the route on the clearest of maps. Esposito (2013) gives an extensive commentary and analysis of Russells synthesis, an unmodern synthesis in Espositos terms, which requires careful reading but from which I gain succour for my appreciations of Russells unique contributions to biological thought.

These problems still seem to lie in the future. But Woodger also gave the means by which substantial progress in biology will accrue. The lines are again to be found in the nal chapter of Biological Principles:
What biology requires is a better ventilation of its thought and a more critical scrutiny of its concepts; more openmindedness and more careful consideration of the relation between investigation and theoretical interpretation. But above all we need a wider recognition of the value of thought itself . . . In biology we require to think primarily about biological facts, not about hypothetical billiard balls. But even thinking about biological facts is not enough. We must scrutinize our ways of thinking too, in order to try to overcome the limitations of those two grooves into which thought has been conned since Descartes, from which it has resulted that a specically biological way of thinking has hardly been so much as considered (Woodger, 1929: 486)

J. H. WOODGER, 18941981
Woodgers lifes work could be summarized as the pursuit and capture of the logic of the organism. That life embraced teaching biology especially to medical students, descriptive and experimental embryological research, biological philosophy, philosophical biology, language and biology, and their interrelationships. In my view, his attempts to create a core of biological principles (Woodger, 1929) have yet to be superseded. Gregg & Harris (1964) edited a volume of studies dedicated to Woodger on the occasion of his seventieth birthday, including his curriculum vitae and a full bibliography of his published writings. The breadth and depth of the contributions to this volume by colleagues and friends is testament to the regard in which Woodger was held. Hull (1988) also adds interesting comments on Woodgers contributions to biology, as does Hall (1992). Woodgers paradox is reviewed and resolved (though not to Halls or the present writers satisfaction) by Medawar & Medawar (1983: 281282); the paradox concerns how one taxon becomes another taxon (i.e. how is the baton of lifes continuity passed on via individual organisms, and how do we demarcate the taxa so created). Woodgers books represent a remarkably successful attempt to wrestle logic from the cacophony of

Central to any biological way of thinking must be a concept of organism. In Biological Principles, Woodger cleared a path towards that concept.


It appears to me that the ve shared a profound contemplation of the nature of the organism, as shown in their many and varied writings, from many sources, viewpoints, and perspectives. Their common inspiring principle was the belief that the organism must ultimately be understood as a living, functioning whole. They all acknowledged that, although reductionism as a method is necessary, productive, and unavoidable, the sterility of a reductionist philosophy and the resultant totalitarian expansion to a world view must be guarded against. The ve demonstrate that a variety of views and perspectives are necessary if the study of organisms is not to be

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R. W. BRUCE (2005). These works reect on the relationships between the observer and the observed, as ways of seeing. But this is surely just part of the universe of perception of the organism, as would have been recognized, I believe, by all ve. In this regard, Russells comments (1916: 4551) on Goethe require careful consideration, especially the relations between Gestalt, whole form, and Bildung, form change: a world of being and a world of becoming, respectively. Living organisms only exist in the world of becoming; perhaps this is just another way of expressing Russells teleology of organism. Driesch too requires further consideration (Driesch, 1908), partly as the originator of the structured thought that the ve would shape their world views against. But Drieschs views need also to be considered as a awed attempt to create a greater biology beyond the restraints of materialism. Rdls (1930) chapter on Driesch and his thought is both sympathetic and critical and worthy of re-reading at the present time, especially if organisms are once again to be allowed to become central in biological thought. A slim volume by the theologian Thomas F. Torrance (1969) might appear to be a strange place to try to nd support for the ve and their efforts. However, Torrance, citing the physicist Jean Chardon and the chemist Michael Polanyi, points out that biology will not make the advances that physics has made if it remains stuck in mechanistic concepts and language but that it must adopt a notion of a eld with its own characteristic structure to make use of physicists space-time. What is needed is a transition to the level of organismic thought. The ve were all engaged in trying to go beyond the mechanistic, to an organismic conception with, I believe, some degree of success. Roll-Hansen (1984) felt that the efforts of Russell and Woodger were a failure. Perhaps rather than a failure, the rst round was just not yet successful, and the issue remaining is just one of needing to try again, and harder. Russells desire for a free biology, free of materialistic restraint, was not just a result of the times (given as November 1943, in the preface to Russell, 1945) of his writing about it, but Russell knew that freedom must be won. The irony of a theologian, a chemist, and a physicist supporting the direction of the efforts of the ve, whereas the biological community largely fell silent and remains so, is indeed poignant. What holds biologists back from trying again? Fear of ridicule, fear of failure or just fear of fear? Has the seductive embrace of the meta-narrative numbed our empiricism or has the gangrene of specialization entered our collective blood stream? Or have organisms become irrelevant to modern biology and can evolutionary materialism now answer all the questions that we are capable of asking? If this is answered in the affirmative, then we

hollowed out and reduced to the banal pointlessness of studying the abstraction without the being. All these biologists recognized organisms, including humans, as ends in themselves, and thus worthy of contemplation and study, and not just as means to justify theoretical fashions, intellectual whims and a current meta-narrative. Another characteristic shared by the ve was a resistance to move the organism from the foreground of their thought. It is important to stress they are not really organicists or holists but rather explorers of the organism, in the full sense of the terms. What was to be understood were organisms, not abstractions about organisms. In von Bertalanffys later writing, he used organisms to develop his theory of general systems, with general systems becoming the foreground of his thought; however much merit there might be in general systems theory, the organism has been displaced. All were also generalists as well as specialists, as any consideration of their published works will show; they were biologists with a breadth of outlook that would be unusual, if not unknown today, with open and expansive philosophies but with critical and focused practical outlooks. They were all philosophical omnivores rather than obligate parasites of a single philosophy. None of the ve displayed any evidence of developing what Jacques Barzun described as the gangrene of specialization apparent in so much of western contemporaneous thought, which so distressed him (Anonymous, 2012). All ve were engaged in the enterprise of growing and expanding biological thought not only by factual accretion, but also by fundamental research, review and criticism of the very basis of biology. They explored the nature of time and space and how these relate to organisms, and how organisms interact with time and space, intrinsically and extrinsically, and they tried to frame a language which can begin to address these concepts. Russell, especially, realized latterly that biologists just have to accept the teleology inherent in the organism, and adapt to that reality, rather than succumb to mechanistic strictures, sophistry and superstition. Ultimately, for Russell and the other four, the world of life was a world of connection among organisms, not just a world of connections among materials. To root the world views of the ve to any single source is of course a fruitless task. Their sources of inspiration are as many as their readings and experiences allowed. However, it does appear worthwhile to ponder where their persistent contrariness issued from, when compared with consensual contemporary materialism. Recently, the contributions to science by Goethes thoughts are receiving overdue and worthy review; for example, see Wahl (2005) and Ebach

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WHATEVER HAPPENED TO THE ORGANISM? have reached the limits of our mental evolution. At a time when physicists are willing to ponder the possibility of multiple universes, to this writer, biology appears happy to die by its own hand through material strangulation.

I do not think it is necessary to erect another abstraction such as holism to categorize the thinking of the ve; they all seemed to recognize, perhaps intuitively, the place and nature of the organism in biology. Ritter largely used ascidians to explore and

Figure 1. Willi Hennigs (1966: 31) scheme of hologenetic relationships. The original caption reads, somewhat cryptically: Figure 6. The total structure of hologenetic relationships and the differences in form associated with its individual parts. Note the diagram for an individual, bottom right. This shows successive stages (semaphoronts) in the life of an organism as it undergoes metamorphosis (e.g. from fertilized egg until death), the successive semaphoronts being linked by their ontogenetic relationships (ontogeny). Hennig (1966: 6) dened a semaphoront as the organism or the individual at a particular point of time, or even better, during a certain, theoretically innitely small, period of its life. The individual organisms making up a species typically exhibit polymorphism, notably sexual dimorphism, and have tokogenetic (nonhierarchical, reproductive) relationships. Division of the stream of tokogenetic relationships (Hennig, 1966: 58) leads to loss of reciprocal tokogeny between daughter lineages (streams) but establishes phylogenetic (hierarchical, sister-group) relationships between species. From Phylogenetic Systematics. Copyright 1979 by the Board of Trustees of the University of Illinois. Used with the permission of the University of Illinois Press.
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coming relevance in the science to which that society is dedicated (Rosen, Nelson & Patterson, 1979: x)

expand his biological thought; Goldstein used his brain-damaged patients; Arber used the growth and form of plants; Russell saw the teleology inherent in the organism; and Woodger reviewed and rened the logic of the organism. Down very different byways of nature, the ve arrived at the same destination: an appreciation of the unity of the organism and a biological way of thought. Despite largely having been written out of the contemporary biological narrative, the efforts of the ve have left and will continue to leave their marks in many elds for those willing to look for them. For example, Phylogenetic Systematics (Hennig, 1966) remains one of the central texts of the last half century in addressing and trying to resolve the complex, confused, and contentious relationships between natural classications and evolutionary theories. In some way or other, this book launched many other texts and thousands of papers. It is worth stressing that, in his work, Hennig laid great value on Woodgers efforts to develop and clarify the nature of hierarchy and its internal logic and its relevance to organisms, time and space (Hennig, 1966: chapter 1), even though Knox (1998) has suggested that Hennig misunderstood or misrepresented Woodgers logic of hierarchy and its relevance to biological systematics (see also Hull, 1988: 214). Consider Hennigs well-known gure of hologenetic relationships (Hennig, 1966: 31, g. 6; reproduced here as Fig. 1), an attempt to transform tokogenetic relations into phylogenetic relations by way of speciation. The underlying logic of this gure concerns space, time, and organisms and their continuity, together with their interrelations; these were Woodgers perpetual concerns and the substance of his paradox. Recently, Rieppel (2010) has re-tilled Woodgers hierarchies with respect to Hennigs interpretations. This suggests that Woodgers presence still remains clear and actual, even in current taxonomy, and the possible consequences of this are as yet difficult to determine. The larger question, as to whether hierarchy is a necessary and sufficient model on which to base lifes reticulations as perceived for example by Arber, still remains largely unvoiced.

In the spirit of forthcoming relevance, and indeed with Woodgers prevision (see Biological Principles; Woodger, 1929), the efforts of the ve, and other named and unnamed colleagues, require reappraisal, and their results must again become part of the core of biological principles founded on the concept of the organism and its relationships with space and time. I believe that the two meetings of the Linnean Society that inspired this essay have already started this necessary and long overdue reappraisal. A biology that cannot nd space for the efforts and insights of Ritter, Goldstein, Arber, Russell, and Woodger would not only be poor in body of fact, but such a science would also demonstrate a failure of application of disciplined and creative imagination. If, after all this, we choose to remain trapped as underlings by the metaphysics of materialism, so be it, although it is a choice and as Cassius pointed out, the fault is not in our stars.

The many and various contributions of the Linnean Society of London are duly recognized, not least in maintaining a library that not only stores books, but also forges conversations. Earlier drafts of the manuscript were much improved during the editorial process, and a later draft by the comments of three anonymous referees and further editorial contributions. All remaining errors and misconceptions are of course entirely my responsibility.

Anon. 2012. Jacques Barzun: obituary. London: The Daily Telegraph, 27 October. Arber A. 1912. Herbals, their origin and evolution. Cambridge: Cambridge University Press. Arber A. 1920. Water plants: a study of aquatic angiosperms. Cambridge: Cambridge University Press. Arber A. 1925. Monocotyledons. a morphological study. Cambridge: Cambridge University Press. Arber A. 1934. The Gramineae: a study of cereal, bamboo, and grass. Cambridge: Cambridge University Press. Arber A. 1950. The natural philosophy of plant form. Cambridge: Cambridge University Press. Arber A. 1954. The mind and the eye. Cambridge: Cambridge University Press. Arber A. 1957. The manifold and the one. London: John Murray. Barlow PW, Lck HB, Lck J. 2001. The natural philosophy of plant form: cellular autoreproduction as a component of a structural explanation of plant form. Annals of Botany 88: 11411152. doi:10.1006/anbo.2001.1397.

Returning to Hennig, some comments in the foreword to the 1979 reprint of Hennig (1966) are worth recalling:
This reprinting of Phylogenetic Systematics has been initiated through the action of the Fellows of the Linnean Society of London a venerable organization whose members have somehow managed repeatedly to penetrate the mists of their own traditions, and maintain a sense of current and forth-

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Woodger JH. 1929. Biological principles. London: Kegan Paul, Trench, Trubner & Co. Woodger JH. 1937. The axiomatic method in biology. Cambridge: Cambridge University Press. Woodger JH. 1952. Biology and language. Cambridge: Cambridge University Press. Woodger JH. 1956. Physics, psychology and medicine, a methodological essay. Cambridge: Cambridge University Press.

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