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In Context #22 (Fall 2009, pp. 3-6); copyright 2009 by The Nature Institute

When Holism Was the Future
Stephen L. Talbott

Back in the autumn of 1999, writing in In Context #2, Craig Holdrege described our discovery of the work of Kurt Goldstein: "It is hard to express the joy and gratitude I experienced in finding a like-minded thinker and scientist, from whom I can learn so much." Born in 1878, Goldstein became a topflight neurosurgeon, and his 1933 book, Der Aufbau des Organismus (published in America in 1939 as The Organism) is a masterwork of rigorous, holistic thought. One testimony to its continuing vital relevance is the 1995 Zone Books reprint, with a foreword by Oliver Sacks.

But Goldstein was not the only such figure. The first half of the twentieth century produced a number of outstanding proponents of biological holism. Those who spoke out against mechanistic and one-sidedly analytic approaches to the living organism included, beside Goldstein, such well-known scientists as D'Arcy Thompson, Sir Charles Sherrington, and J. B. S. Haldane. But one exceptionally far-seeing luminary of this holistic movement in biology is rarely heard of today outside certain specialist circles, and my own meeting with his work has proved as unexpected and inspiring as that earlier encounter with Goldstein. His name is E. S. Russell.

Cells Do Not Make Organisms

Russell was a British naturalist pursuing fisheries research while also interesting himself in the entire history of biological thought, beginning with Aristotle. His 1916 work, Form and Function: A Contribution to the History of Animal Morphology, was deeply influential and continues to be regarded as a primary resource for students of the history of morphology. Robert J. Richards, author of the recent, highly acclaimed The Tragic Sense of Life: Ernst Haeckel and the Struggle over Evolutionary Thought, has written that

Russell's book has yet to be surpassed as a history of morphology ... One can feel the force that Russell gives to criticisms of the "mechanistic" approach of Darwin and Weismann. He gives compelling reassertion of a teleological view, which permits one to experience the real texture of the history of science.

The book that has most captivated me, however, is one Russell wrote in 1930 as a strikingly prescient plea for attention to the whole organism: The Interpretation of Development and Heredity: A Study in Biological Method. In it he fully granted the necessity for a use of analytic methods by the biologist, but he also showed how these methods, left to themselves, yield a habit of abstraction whereby the organism itself is lost from view:

The initial step which leads to abstractness of treatment is of course the isolation and definition of parts and part-processes as such. To define is to separate, and to separate is to ignore or to disregard in some measure the relations with other parts and with the whole. In the living thing there are in actuality no separate parts, no separate processes, for no part can be adequately characterized save in terms of its relations to the whole. (p. 145)

Russell recognized that the usual analysis of the organism into separate, distinct traits - the kind of analysis upon which Mendelian genetics was based - resulted in a falsification of the organism. We can, he pointed out, distinguish an infinite number of traits, "but yet none of them is in reality separate from the rest" (p. 151). This point is borne out by every study of the organism conducted with due respect to the whole organism. Many readers will recall Holdrege's study of the giraffe, where it is found, for example, that the long neck is reflected in all the other bones of the animal's body as well as in many other features (2005a).

So while it may be necessary to analyze and abstract as one step along our way toward understanding, this leads to severe distortion unless we simultaneously insist on recalling the indissoluble unity from which we are abstracting. Only this continual reckoning with unity can enable us to counter the falsification entering our science through abstraction.

Russell developed this line of thought with considerable energy and subtlety. However, such efforts themselves remain rather abstract, sterile, and ineffectual if they are not brought to bear in a practical way upon real biological problems. One is therefore happy to find Russell looking in detail at such topics as the nature of the cell and theory of the gene.

By Russell's day the cell was widely viewed as a largely independent and self-subsistent unit. This made it easy to imagine the organism as "built up" from cells in the way one assembles a structure with building blocks. The organism, instead of being a prior unity, tends to become some vague sort of "emergent reality" resulting from a successful assembly of parts. All explanation then resides in the understanding of the parts, there being no unitary organism as such to be studied. However, Russell's analysis of what was already known about the cell in his day led him to a much more balanced view, whereby he could conclude that "it is the organism that makes the cell, not the cells that make the organism" (p. 196). And "the cells taken separately no more constitute the organism than words and letters by themselves make a sentence"; the unity and the meaning "are prior to the components":

We accept the simple facts of observation that the organism acts as a whole, and that the activities of its parts are subordinated to, and co-operate in, whatever the organism as a whole is doing at the moment of observation. (p. 235)

Refusal of the Gene-Centered Approach

Just as Russell refused to reduce the organism to its cells, so, too, he refused to reduce the cell to its chromosomes or genes. He wrote before the discovery of the structure of DNA, at a time when genes were still only hypothesized "factors" designed to account for the results of Mendelian experiments. However, the genes were already being reified within scientific thought as concrete particles or unit substances in the chromosomes that, when finally identified, would explain all the various features of the organism and of inheritance.

Russell strenuously opposed this excessive reliance upon the gene as bearer of the organism's essential identity and as sole carrier of heredity. He approvingly quotes the botanist F. Noll, who wrote in 1903 (before the term "gene" was coined): "If the egg-cell of a lime tree is already a young lime tree, there is no need of any idioplasm, germ-plasm, pangens, or heredity-substance to render possible its development into a lime tree; the egg-cell as a whole is the heredity-substance" (pp. 287-8). That is, the one cell we can truly identify with the organism is the fertilized egg cell. It is, moreover, already a complete organism, and does not have to go through a gene-directed developmental process in order to become one. It's not that separate cells aggregate and "build" an organism, but rather that the unitary egg cell-organism subdivides and structures itself as it passes through developmental stages.

Russell's 1930 critique of the gene theory is truly stunning in its relevance for today. One extended passage will have to suffice for our current purposes. He writes that the presumed units of heredity, whatever their names,

are supposed to represent the parts or the characters of the developed organism, and in some way, which always remains mysterious, to give rise to them in the course of development. (The hereditary units being the purest of abstractions, it is of course natural that their relations with the characters they determine should remain obscure.)

Hereditary units and "determinants" of all kinds are pure abstractions; the process of analysis has been carried so far that it is impossible to reconstitute from these purely abstract elements the activities of the cell or the organism as a whole. All that is left then to the theorists is to smuggle back into the determinants or other "parts" the powers and functions which belong rightly to the organism as a whole, and have inevitably been dropped out during the process of analysis. The concept of the organism as a whole, which has been destroyed by unrestrained analysis, is reintroduced surreptitiously, and the qualities and powers of the organism as a whole attributed to certain abstract and subordinate parts of it.... (p. 155)

The hereditary substance, magically endowed in this way with all the powers of the organism as a whole, becomes a kind of materialized vital spirit or, in Russell's phrase, "material entelechy" (p. 154).

But the conception is an impossible one. Russell quoted in this regard marine biologist J. A. Ryder: "No creature can be supposed to have its life or germinal properties associated only with certain corpuscles within it, since we cannot suppose an organized whole dominated by a portion of it; it is not possible, for example, to conceive of individual life except from the entire organism that manifests it. There can be no 'biophors' - bearers of life - the whole organism must do that as an indivisible unit" (p. 152).

Dismissal ...

Russell thought in 1930 that biology was moving in his direction. His book testifies to the desire by many researchers of that era for a whole-organism approach to the defining questions of the day. And yet, in the short term at least, he was quite wrong. The abstract gene was at the merest beginning of its meteoric trajectory, which would light up the firmament of the scientific imagination. The decisive discovery of the double helix in 1953 seemed an irrevocable turning point, and the following years consolidated the notion of genes as physical entities mechanistically determining the traits of the organism. By the 1980s, with the neatly defined gene at the height of its influence upon scientific thinking, Russell was largely forgotten by the community of biologists. It was left to the historians of science to dispose of his legacy.

One of those historians was Nils Roll-Hansen, who in 1984 wrote a paper certifying the "failure" of the earlier opponents of mechanistic biology. "It has been claimed," he wrote, "that the approach to early twentieth-century biology changed from mechanistic to holistic, and that this change promoted the progress of biological science." He concluded, however, that while there was indeed a trend toward holistic philosophizing in biology, this did not yield any fruitful results for the actual science of biology. And one of his primary pieces of evidence for this conclusion was the fate of the work of E. S. Russell.

Russell, in Roll-Hansen's words, "rejected the theoretical speculations of mechanistic biology. Biological phenomena should be taken at face value, not explained away by underlying physico-chemical mechanisms. The idea of particulate material genes, some kind
A Critique of the Modern Gene - From 1930

When E. S. Russell published The Interpretation of Development and Heredity in 1930, the gene was still an unknown "factor" - a small segment of a chromosome of uncertain physical nature. What was known was that in sexual reproduction some of the parental chromosome segments were distributed among the offspring in general accordance with the results Mendel had obtained for certain traits of peas. There was, in these cases, a satisfying mathematical regularity seeming to govern both the occurrence of traits and the corresponding behavior of chromosomes. This made it appear obvious to many biologists that the chromosomes - that is, the unknown "factors," or genes, they contained - would fully explain the traits.

This was not at all obvious to Russell, however - nor was it obvious to the many embryologists and developmental biologists who complained that the geneticists in fact had almost nothing to say about the actual occurrence of traits in living organisms. After all, those traits arose from the embryo through elaborate processes of development, and there was already strong evidence that the complement of chromosomes remained the same in cells throughout the organism during this development. How could something that remained the same in all the different tissues and organs, from liver and skin to heart and nerves, explain the development of these organs from an original, undifferentiated zygote?

Many of these critics, including Russell, pointed out that the genetic experiments of the day showed only how differences arose when genes were mutated. Derange a particular gene, and the fruit fly's eye color changed. But to take this as evidence that the gene explains the eye color was rather like saying the ignition key not only is required for the automobile's movement, but also explains the drive train.

Russell's critique was surprisingly prescient, retaining a great deal of validity even in the post-double helix era of the molecular gene. He by no means downplayed the significance of chromosomes, saying (quite rightly) that they appeared to be crucial for maintaining the metabolism of the organism. But instead of looking to them for a causal explanation of the rest of the organism, he foresaw a time when they would, in our understanding, be fully integrated with the entire physiology of cell and organism. This would require us to "throw off all traces of the particulate conception of heredity" - a conception that sees little bits of chromosomal substance as explanations for traits.

Moreover, rebelling against the atomistic habits of thought that made genetic elements into fixed, determinate, and particulate things, transferable from one position on the chromosome to another without change in their character, he wrote that the difference between two chromosomes "may quite well be a slight though discontinuous chemical or stereochemical change affecting the chromosome as a whole." That is, we should look not merely for atomic genes, but for overall changes of significant form.

These points capture the main substance of the revolution now developing under the name of "epigenetics." The entire cell continually sculpts and re-sculpts its chromosomes without at all changing the actual DNA sequences that were so long equated with genes. This shaping activity, which we have every right to think of in artistic terms, determines what gets expressed from the DNA.     ST

of complex chemical molecules lodged along the chromosomes, was [in Russell's mind] unscientific speculation." This rejection was, for the later philosopher, definitive proof of the fruitlessness of Russell's holism. It would have been fine, says Roll-Hansen, if Russell had simply preferred some other theory of inheritance. But Russell, who believed that progress in biology was inhibited by mechanism, "argued that the chromosome theory was impossible," and therefore we now have every right to judge his views harshly on the basis of this uncompromising assertion.

The interesting thing is that Roll-Hansen finds no need whatever to cite evidence for this judgment. In 1984, it seems, the matter was settled beyond dispute. The well-defined physical gene reigned alone as First Cause and Unmoved Mover, explaining both the inheritance of traits and their actual realization in the developing organism. And so, with no need of argument, Roll-Hansen can only wonder aloud why Russell would "dogmatically reject some of the central theories of modern biology, such as the chromosome theory of inheritance" - why he would deny "a realist interpretation of the genes, maintaining that they were pure abstractions or instruments of thought and could not possibly correspond to any concrete part of the living organism."

... And Vindication

But what a difference a few years can make! Any geneticist today, hearing the view that genes are "pure abstractions or instruments of thought and could not possibly correspond to any concrete part of the living organism," would likely conclude that the words were coming from the latest issue of Nature or Science. The idea of a well-defined, causally effective gene, conceivable in isolation from the organism as a whole, has by now suffered such injury that some geneticists would prefer to jettison the word "gene" altogether (Holdrege 2005b). The situation had already reached a point by 1992 where a leading philosopher of science could wryly remark that "a gene is anything a competent biologist has chosen to call a gene" (Kitcher 1992). And at the height of the Human Genome Project, science historian Evelyn Fox Keller concluded that "the prowess of new analytical techniques in molecular biology and the sheer weight of the findings they have enabled have brought the concept of the gene to the verge of collapse" (2000, p. 54). Russell, in fact, was precisely on target in predicting a time when "the gene will cease to be regarded as a self-existent particulate unity and will be merged in the general physiological activity" of the organism (Russell 1930, p. 157).

This merger is what the dramatic, ongoing, and accelerating discoveries in the field of epigenetics are all about. What this field has been revealing in the most striking ways is that the cell and organism as a whole determine, not only how genes will be expressed, but even what is to count as a gene at any given time. There's widespread agreement among molecular biologists today that we have lost all meaningful possibility of defining a gene as a particular stretch of DNA with a fixed identity, independent of what is going on in the larger cell and organism as a whole. We can use the word "gene" only as a convenient way of referring to an almost unfathomably complex constellation of cellular events.

Whenever and wherever we look in detail at the molecular level, we see no causal particles from which everything else can be explained. Rather, we see exactly the same sort of interwoven complexity, the same unity, the same striving toward self-realization, that is so evident in the organism as a whole. This is why, only a few years after the exaggerated hype of the Human Genome Project, the author of a review in a major biological journal could write that genome mappings and genomic comparisons of species "shed little light onto the Holy Grail of genome biology, namely the question of how genomes actually work" in living organisms (Misteli 2007). And it is also why a pair of geneticists could write that trying to define the structure and functioning of the chromosome "is like trying to define life itself" (Grewal and Elgin 2007).

We can only hope that now, benefiting from the vastly greater amount of detail available to them than was available to Russell in the 1920s and 1930s, biologists will refuse the search for isolated parts that can somehow explain the whole, and will instead heed Russell's call to discover the whole organism at work in each of its parts. Then a truly organic understanding of heredity may become possible (Holdrege 1996).

It remains a sobering fact, however, that up through the late twentieth century, biologists with all the advantage of several decades of further scientific discovery could still look back at the perfectly sound arguments of E. S. Russell and wonder how the man could have gotten it so wrong. This tells us something about how ideas, when they become ideological and rigid, can direct our thinking and research in such a way as to blind us for generations to a truth glimpsed long before by more perceptive thinkers.

This article was written as an addendum to an ongoing series of papers entitled On Making the Genome Whole, dealing with the current epigenetic revolution in biology. For fuller discussion of the bearing of epigenetics on the issues presented here, and what it means to look for the whole in every part, please see those papers.


Grewal, Shiv I. S. and Sarah C. R. Elgin (2007). "Transcription and RNA Interference in the Formation of Heterochromatin", Nature vol. 447 (May 24), pp. 399-406. doi:10.1038/nature05914

Holdrege, Craig (2005a). The Giraffe's Long Neck: From Evolutionary Fable to Whole Organism, Nature Institute Perspectives #4. Ghent NY: The Nature Institute. Available online:

Holdrege, Craig (2005b). "The Gene: A Needed Revolution," In Context #14 (fall), pp. 14-7. Available online:

Holdrege, Craig (2006). Genetics and the Manipulation of Life: The Forgotten Factor of Context. Hudson NY: Lindisfarne Press.

Keller, Evelyn Fox (2000). The Century of the Gene. Cambridge MA: Harvard University Press, p. 54.

Kitcher, Philip (1992). "Gene: Current Usages," in Keywords in Evolutionary Biology, edited by Evelyn Fox Keller and Elisabeth A. Lloyd. Cambridge MA: Harvard University Press, pp. 128-131.

Misteli, Tom (2007). "Beyond the Sequence: Cellular Organization of Genome Function", Cell vol. 128 (Feb. 23), pp. 787-800. doi:10.1016/j.cell.2007.01.028

Richards, Robert J. (2008). The Tragic Sense of Life: Ernst Haeckel and the Struggle over Evolutionary Thought. Chicago: University of Chicago Press.

Roll-Hansen, Nils (1984). "E. S. Russell and J. H. Woodger: The Failure of Two Twentieth-Century Opponents of Mechanistic Biology," Journal of the History of Biology vol. 17, no. 3 (Fall), pp. 399-428.

Russell, E. S. (1945). The Directiveness of Organic Activities. Cambridge: Cambridge University Press.

Russell, E. S. (1982). Form and Function: A Contribution to the History of Animal Morphology. Chicago: University of Chicago Press.

Russell, E. S. (1930). The Interpretation of Development and Heredity: A Study in Biological Method. Freeport NY: Books for Libraries Press. Reprinted in 1972.

Talbott, Stephen L. (2009). "On Making the Genome Whole." Available online here.

Steve Talbott :: When Holism Was the Future

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