Political Descent. Malthus, Mutualism and the Politics of Evolution in Victorian England

Political economist Thomas Robert Malthus.
His controversial work was Darwin's inspiration.

It is well known that Darwin's theory of natural selection was inspired by Thomas Malthus's famous Essay on the Principle of Population. Darwin not only acknowledged this in Origin of Species, noting that "it is the theory of Malthus applied with manifold force to the whole animal and vegetable kingdoms", but recalled the moment when the significance of Malthus's work first struck him in his autobiography:

"In October 1838, that is, fifteen months after I had begun my systematic enquiry, I happened to read for amusement Malthus on Population, and being well prepared to appreciate the struggle for existence which everywhere goes on from long-continued observation of the habits of animals and plants, it at once struck me that under these circumstances favourable variations would tend to be preserved, and unfavourable ones to be destroyed. The result of this would be the formation of new species. Here, then, I had at last got a theory by which to work”.

It was shortly after his return from his five-year voyage on HMS Beagle in 1836 that Darwin read Malthus. He had been putting in long hours writing up his geological notes, but had been developing his theory of evolution in the few spare moments he could find. Darwin was working up an answer to that "mystery of mysteries", as it had been called, the origin of new species. Naturalists and mineralogists had long been pulling strange and wonderful beasts out of the ground in fossil form, and they needed an explanation. Darwin was already familiar with Malthus's ideas; for even though Malthus had originally developed his controversial political economy in the late eighteenth century, Malthusian economics were going through a renaissance in the 1830s. Malthus had made the case that in society, just as in nature, there would never be sufficient food for everyone to live in luxury, indeed, he had argued that there would always be those who went hungry.

The politics of Malthus's position were evident to all. He had written his essay in an attempt to undermine what he believed to be the unrealistic Enlightenment ideals of universal progress and prosperity, and had received harsh criticism from political radicals for his troubles. “An apostle of the rich,” the poet Robert Southey had called him, both amazed and affronted at the “stupid ignorance of the man”; the essay was “Adam Smith’s book in code, a confession of faith in this system; a tedious and hardhearted book.” Adam Smith's Wealth of Nations had long been the bible of liberal political economy, and Malthus was read as furthering those interests.

Indeed, half a century later, far from being decried Malthus's work was embraced by a new generation who saw the notion that individuals should have to struggle to make their way in a competitive world as speaking to their own times. In the context of the industrial revolution, and of a rising middle class of industrialists and entrepreneurs, the values of hard work, thrift and economic independence prevailed; the suggestion too that the poor were responsible for their own plight made Malthus appear all the more emblematic of the times. Historians have long noted that Darwin's theory of natural selection not only echoed, but endorsed the Whig politics of industrial England, and this, I would contend, was a large part of why evolution was accepted in England as quickly as it was. Indeed, while both historians and the public have often been led by their present preoccupations to focus upon the religious implications to Darwin's work, a full consideration of the many and diverse responses to the publication of Origin shows that evolution was read by contemporaries as having as much to say about politics as to theology.

'Darwin's Bulldog' Thomas Huxley
embraced the Malthusian politics of
evolution

Thomas Huxley, the young and talented naturalist who would later make his name as "Darwin's Bulldog", and who has long been regarded as one of the most significant scientists of the nineteenth century, welcomed Darwin's work in an early review, but did so not only as a major contrition to an important scientific question, but at the same time as "a veritable Whitworth gun in the armoury of liberalism". The analogy was telling of his enthusiasm. The Whitworth, developed by Sir Joseph Whitworth, had hexagonal rifling, and far exceeded the Enfield, which had long been the standard service rifle of the British army, in both accuracy and range. To Huxley's mind, like the Whitworth, Darwin's theory was not only the best available answer to the species question, but was certainly unanswerable by the creationist 'natural theology' offered by more conservative thinkers. Huxley was not alone in his recognition of the politics of natural selection. Although from another perspective, the noted socialist Karl Marx also thought that Darwin's book read like capitalist apologetics. Writing to his friend and long-time supporter Friedrich Engels, he made the following observation:

“It is remarkable how Darwin rediscovered, among the beasts and plants, the society of England with its division of labor, competition, opening up of new markets, ‘inventions,’ and Malthusian ‘struggle for existence.’ It is Hobbes’ bellum omniun contra omnes…” [war of all against all].

Karl Marx thought Origin endorsed capitalism

Here I should say that none of this is particularly new to historians, although as I have mentioned, - and certainly in the US - the emphasis tends to be on the religious implications of evolution. However, what it clear is that there is also an important story to tell here about the politics of evolution. From what I have said already, one might assume that this would be the familiar story of social Darwinism: that Darwin's work was read - and possibly written - as an endorsement of the ruthless economic competition of laissez-faire capitalism. This has certainly been the dominant theme of the majority of the existing work in the field. This would only be a part of the story, however, as this view of life, and of the Origin of Species, was only one among many. Notably, and what historians have allowed to go largely unrecorded, is the fact that there was a significant and vocal population who embraced both evolution and the Origin as a vindication of a very different politics, one based upon cooperation and mutual aid. I say largely unnoticed, because historians have certainly noticed the existence of Peter Kropotkin's work on mutual aid as a factor in evolution (among the best is Mark Borrello's account in his book Evolutionary Restraints, which I reviewed on this blog some time ago), which was published throughout the 1880s and 1890s, but they have not recognized that Kropotkin was one among many who embraced a mutualist reading of Origin to support and substantiate their own more socialistic politics. And this is where my new book comes in. 

Peter Kropotkin whose mutualist theory
of evolution was widely recognized 

In Political Descent. Malthus, Mutualism and the Politics of Evolution in Victorian England I show that in fact the politics of evolution were very much more contested than historians have hitherto recognized. Kropotkin was not a lone socialist voice in a capitalist wilderness, but rather was representative of an entire tradition of men and women who were interested in what the fact of our evolution had to say about the kind of creatures we are, and thus about the kind of society that it might be possible for us to live in. 

 Darwin, provided an evolutionary
account of altruistic behaviour in Descent of Man.

What I think many people will find compelling about this story is that, just as Kropotkin argued was the case in the last years of the nineteenth century, it is clear that Darwin was much more sympathetic to those who saw mutualism and cooperation as the hallmarks of what had made humans 'fit' in the struggle for existence, than either Huxley or Marx thought. Indeed, and as Kropotkin had sought to point out to his readers, Darwin had said as much in the 1871 book The Descent of Man in which he had tackled human evolution head on. By the 1870s, the most important question about the evolution of mankind had become the origin of mind, morals and conscience. These apparently uniquely human characteristics were the last bastion of a 'human exceptionalism' among those who still sought a role for the divine in origin of man. After all, if the world worked to favour only the most competitive individuals, then it seemed logical that any resources invested in helping others survive would only weaken the altruistic. If generosity and altruism were not a divinely inspired character, however, conventional accounts of the origin of human morals emphasized the benefits that might accrue to the apparent altruist by pursuing a course of self-interest. This again would fit with the politics of liberal political economy. Darwin, though, was of a different mind. By providing a naturalistic account of the evolution of these highest of human qualities, one grounded in a history of the evolution of genuinely altruistic regard for others, Darwin sought to undermine the rampant individualism that others were using his own work to support. "Thus the reproach of laying the foundation of the most noble part of our nature in the base principle of selfishness is removed", he wrote. 

 Piers J. Hale's new book Political Descent. Malthus, Mutualism and the Politics of Evolution in Victorian England in published this month by Chicago University Press. It is available from the press; from online book sellers and through all good bookstores.

Evolving the Neighborhood

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D.S. Wilson, The Neighborhood Project. Using Evolution to Improve My City, One Block at a Time, (New York: Little, Brown and Company, 2011). Hb $25.99 ISBN 9780316037679

David Sloan Wilson’s Neighborhood Project is a provocative read. It will outrage those who think that culture is too complex to be accounted for in terms of biology, and annoy biologists who think that Wilson’s adherence to group selection and pro-sociality is misguided.

The things that will offend are also exciting, however. Wilson takes the levels of selection debate into new territory, utilizing the people of his home town, Binghamton, New York, as the model super-organism in an ongoing social experiment. Whereas hitherto sociobiology and evolutionary psychology have roused opposition for their reductive portrayal of humans as driven by self-interested and often highly sex-specific instincts, here Wilson seeks to ground pro-social behaviour in our evolutionary history.

D.S. Wilson

Offering the ‘Binghamton Neighborhood Project’ as a model, he promises not only the revitalization of our neighborhoods, our cities, and of this and other nations, but most controversially, he holds out the prospect of facilitating the further evolution of human altruism. The point is not merely to understand the world, he argues, “but to change it”. We must become wise managers of evolutionary processes if we are to effect meaningful social change.

No mere recipe for the cookshops of the future, Wilson’s manifesto is a report from the barricades. The ongoing ‘Binghamton Neighborhood Project’ is based upon extensive surveys of school children, neighborhood reconnaissance, interviews with and genetic testing of respondents.

Measuring the sociality of neighborhoods by their participation in occasions such as decorating their houses for Hallowe’en and Christmas (the latter included to catch any Christians who might otherwise be highly social but object to celebrating Hallowe’en), and with the aid of GIS technology, Wilson has mapped the social (or anti-social) character of neighborhoods across the city to produce a map of hills and valleys. Wilson’s mission is to raise Binghamton’s anti-social valleys into pro-social hills and he is keen for others to follow his lead.

But can the social behaviour really be explained in terms of our biological evolution? Wilson offers four ‘parables’ from nature in support of his belief that it can. He offers the parable of the water strider (Aquarius remigis)—self interested individualism is certainly one possible outcome of natural selection, but it is not the only one.


The social wasp Polistes

In a second parable, that of the social wasp (Polistes), Wilson presents a very different behavioral strategy. It is unsurprising that Wilson should turn to the social insects, but even here he admits that the lesson they teach is not self evident. This has been exacerbated by their frequent misrepresentation as an analogy for human society. This was most famously—and most egregiously—the case in Bernard Mandeville’s Fable of the Bees (1705).

Mandeville used the hive analogy to portray mankind as a group of inherently self-interested “knaves” who were each concerned only to maximize their own outcomes (p.77). It seems likely that Adam Smith had Mandeville in mind when he wrote Wealth of Nations in 1776, his conception of the ‘invisible hand’ enlarged on the notion that the social good is best obtained by each attending only to their own self interest. “It is not from the benevolence of the butcher, the brewer or the baker that we expect our dinner, but from their regard to their own interest. We address ourselves, not to their humanity but to their self love, and never talk to them of our own necessities but of their advantages”—Smith had a way with words.

This resonates well with the nascent capitalism that Smith sought to promote, of course, but, as readers of this journal will recognize, it does so too with the predominantly individualist message not only of Darwin’s Origin of Species (1859), but of the ‘gene’s-eye-view’ account of evolution promoted first by William Hamilton in his 1963 and ‘64 papers on inclusive fitness, and which was subsequently repeated in John Maynard Smith’s interpretation of kin selection, Robert Trivers’s reciprocal altruism and most famously in Richard Dawkins’s popular metaphor of the selfish gene.

However, while the predominant theme in Origin was the unremitting competition of each against all, in chapter seven Darwin first discussed the social insects as an exception to this apparent state of nature. Darwin argued that in their case group selection prevailed - each worked not for its own good but for the benefit of the community (Origin 1859:238).

Unto Others, (1998)

In their Unto Others (1998) Wilson and the philosopher Elliott Sober made the case that we need to take Darwin's account of group selection seriously, and that post-Wynne Edwards formulations of group selection are more than just a fuzzy description of what could be more accurately described in terms of selfish genes.

Thus, and echoing Darwin, Wilson tells us that "the true fable of the bees is about a conflice between the levels of selection. Within-group selection produces individuals that are designed to benefit themsleves at the expense of their neighbors, the essense of knavery [,but] between-group selection produces individuals that behave for the good of their groups, the essense of solid-citizenry" (p.78).

In his third parable, draw from the human immune system, Wilson argues that it is wrong to presume that our genetic inheritance makes us inflexible in light of the hand that nature has dealt us—it does not. Recognition of its “innate flexibility” reveals its resemblance to “a social-insect colony inhabiting our body as its environment”, he argues (p.129). The speed with which antibodies respond to rapidly evolving disease is indicative of the sort of flexibility that can result from innate biological systems. “Alone among species we have the capacity to imagine new worlds and make them a reality”, he says. (p.139).

Wilson clearly wants to take this further, though. The parable of the crow is that we can evolve a new humanity. Despite the ‘apartheid’ that social scientists might insist upon between biology and culture the cultural adaptations of crows reveal corresponding genetic changes. The creation of “niches for niceness” in our own communities will make a difference of the same order, he argues, where cultural adaptation leads, genetic evolution will follow.

Teilhard de Chardin

The significance of evolution for our own behavioral norms was not missed by Darwin, of course, but surprisingly, it is not Descent of Man that Wilson cites to back his position, but the theologian and scientist Teilhard de Chardin. In The Phenomenon of Man (1959) Teilhard argued that we alone of all creatures have entered what he called the ‘Noosphere’, a term by which Teilhard—and now Wilson—denote not only the surface layers of the earth that reveal the paleo-anthropological evidences of our evolved capacity for culture, but also the period of mankind’s ability to reflect upon our own origins and history. If there is an ‘Omega Point’, as Teilhard believed, Wilson argues, it is us. We are “the process of evolution reflecting fully on itself” (p.113).

While Wilson notes Adam Smith’s treatment of praise and blame in his Theory of Moral Sentiments (1759) as being significant for fostering change in our normative values, he seems unaware that Darwin utilized exactly these passages in his own account of human evolution to discuss the very issues that Wilson is concerned with—the conflict between the levels of selection and the evolution of altruism.

But what are we to make of all this? Does it work, or is this all just fantasy? Wilson has already started to use the Binghamton project in such a way as to effect changes in the map, to turn anti-social valleys into social hills, and thus to clearly improve people’s lives, but even if these cartographic changes do really reflect substantive behavioral change on the ground, the question remains how much they are the result of reciprocal self-interest and how much they are the result of the pro-sociality that Wilson is clearly keen to see.

Further, we would clearly want to see evidence of the claim that behavioral change is in any way significant at the genetic level, and it seems that Wilson’s project is set up to provide exactly this—one way or another. One thing is certain, however—the Binghamton Neighborhood Project is certainly a new chapter in the debate about the levels of selection and the evolution of pro-sociality, but it will expand rather than resolve the controversy.

On inference from evidence

TweetThis is not really a question of politics and evoluion, but it interested me, so I shall pass it along... I was reading my friend John Wilkins's blog Evolving Thoughts.

Evolving Thoughts. A great blog written and edited by Dr. John Wilkins.

Today he was considering a letter sent from the nineteenth-century anatomist, "Darwin's Bulldog", Thomas Henry Huxley to the Anglican Priest, amateur naturalist and novelist Charles Kingsley. The letter he quotes I have pasted below (the authorial insertions are John's) :

Thomas Henry Huxley, as portrayed in Vanity Fair

"With respect to the sterility question [between species], I do not think there is much doubt as to the effect of breeding in and in [we would now use the neologism "inbreeding"] in destroying fertility. But the sterility which must be obtained by the selective breeder in order to convert his morphological species into physiological species – such as we have in nature – must be quite irrespective of breeding in and in.
There is no question of breeding in and in between a horse and an ass, and yet their produce is usually a sterile hybrid.
So if Carrier and Tumbler [varieties of pigeon], e.g., were physiological species equivalent to Horse and Ass, their progeny ought to be sterile or semi-sterile. So far as experience has gone, on the contrary, it is perfectly fertile – as fertile as the progeny of Carrier and Carrier or Tumbler and Tumbler.
From the first time that I wrote about Darwin’s book in the Times and the Westminster [Review] until now, it has been obvious to me that this is the weak point of Darwin’s doctrine. He has shown that selective breeding is a vera causa [Whewell's term for a true cause, taken from Newton] for physiological species.
But I entertain little doubt that a carefully devised system of experimentation would produce physiological species by selection – only the feat has not been performed yet."

Charles Kingsley in Vanity Fair

This is a really interesting letter, for many reasons - not least that Huxley is discussing the matter with the Chaplain to the Queen of England! Where John's consideration of this letter takes him to consider the evolution of sterility - an interesting quesiton in evolutioanry biology, what I find interesting abou this letter is what it reveals about the different expectations of a scientific theory that are evident between Huxley's thinking on the one hand, and Darwin's on the other. Notbaly Huxley clearly has the expectation of experimental proof of speciation, and, as he mentions in this letter, had makde this point in his early reviews of Darwin's work. 

Darwin, of course, was - as Michael Ruse has long since demonstrated, - strongly under the influence of the Cambridge philosopher of science, William Whewell, especially when it came to inductive reasoning and the question of what made for a sound explanation in science. In his 1840 work The Philosophy of the Inductive Sciences (1840), Whewell held to what he called the 'concillience of inductions' - ie: the conviction that if the hypothesis expalined enough facts then it was likley to be near the mark, but that if it explained enough facts from different fields and especially if it explained facts that the theory had not been framed to explain, then one could be really confident it was a true explanation, or, a vera causa, as he termed it. - Thus Darwin didn't need to actually see speciation to be convinced he was on the right track, - common ancestry explained so much from so many diverse fields of natural history, and speciation was a logical outcome of this view in light of the present day diversity of life.  Malthus just gave him the 


Cambridge philosopher William Whewell

mechanism, - competition and extinction. The issue of sterility was less of an immediate issue for Darwin, even though he did acknowledge the problem.

Huxley, on the other hand, was much more of the empiricist school of John Stuart Mill, who wrote in the philosophy of science as well as poltical economy. The most convincing - and indeed for Huxley - the necessary evidence had to be observed, and preferably observed under experimental conditions. This was always going to be a problem for him, especially where speciation was concerned -  which left Darwin perplexed. Darwin realised that the geological time involved and the selective nature of the fossil record made it unlikley that speciation would ever be demonstrated to Huxley's criteria.

Kingsley, of course, had written to Darwin on the eve of the publication of Origin to say that he was ready to be convinced. Following Darwin's analogy from domestic animals, he wrote to say that he could readily follow Darwin's line of reasoning from long familiarity with the breeding of dogs and horses - he also added, almost as an after thought that a God who made things make themseleves was a much more noble conception of the deity than a God who had to interfere all the time. It was Kingsley's expression of this last sentiment that Darwin included in the second edition of Origin, suggesting that on such authority there were no grounds for thinking that the views he put forward in the book should offend the religious feelings of anyone.

"Adios theory": on the importance of careful reading.

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Charles Darwin, pencil sketch, 1839.
[Courtesy of Cambridge University Library]

People care very much about what Charles Darwin wrote. They should, his theory of evolution by natural selection is a fairly big deal, after all. People care a lot about what he thought, and about how he came to think what he thought - and why not? As an historian I certainly think it's important to work out the history of ideas as well as events - and the interactions between the two.

People who care about what Darwin wrote tend to read a lot of what he wrote - those of us who care about the Origin, for instance, read the various editions that he wrote across his lifetime. If you care to do so you can read every edition of all of Darwin's published works online, beautifully scanned and searchable thanks to the work of Dr. John van Wyhe, at the Darwin Online Project.

There are even variorum editions published that track these changes, but my very favourite has to be this online research tool, courtesy of Ben Fry.

Fry's "Origin of Species: The Preservation of Favoured Traces" is wonderful - it makes research easy and fun, and I have yet to teach a class that has not contained at least one student who has instantly gone out and used it as the basis for a research essay. Nice work Mr. Fry, thank you.



Ben Fry's "Origin of the Origin"
Changes in each edition are colour-coded

We also have the Charles Darwin Correspondence Project. Big green weighty tomes in the library, on my bookshelf, and now online too. We can track changes in the various editions of Origin, for instance, and then find out what Darwin was writing to his friends, family and colleagues for added insight. My favourite example of this is the famous insertion into the Origin's last paragraph of 'the Creator' from the second edition onwards. It has often been cited as evidence of Darwin's religious belief.

He also cited the Anglican naturalist Charles Kingsley in support of this reading of his intentions. Kingsley had written to him saying that Darwin's theory gave him an even greater conception of God than he had held before (I've given you a picture here, but go and check it out... all the tools are right there for you!! )









Page 488 of the second edition of Origin. Darwin quotes Kingsley.
Insert shows an excerpt of the letter from Kingsley to Darwin,
November 18th 1859.
[Origin picture courtesy of the 'Darwin Online Project'.
Kingsley letter courtesy of the British Library.]

So, what might we make of this? Darwin clarifying things in light of all the accusations of the atheistic tendency of his theory? - I live in Oklahoma where many of my students have deep religious convictions, and this is a happy conclusion for many of them - an embrace of evolution without challenging their faith. However, let's turn to the correspondence.

 http://www.darwinproject.ac.uk/entry-4065 A few years down the line, when things were a little more settled, Darwin wrote to his friend Joseph Dalton Hooker about inserting the whole "creator" thing. "I have long regretted that I truckled to public opinion", he wrote.

Most recently, courtesy of a collaboration between Cambridge University and the American Museum of Natural History, we now have access to the notes and marginalia that Darwin made in the books he read at the Darwin Manuscripts Project.

Overkill? - I don't think so. Here we get to see not only what Darwin read, but what he thought about it, and - by cross referencing with his reading notebooks (Appendix IV of Volume IV of the correspondence) - when he read it!

How cool is that?  Very cool. - Cool enough to have excited a recent NPR blog post by Robert Krulwich.


Robert Krulwich wonders... but doesn't read as carfully as he might

The subject of Krulwich's blog is original thoughts - or, as he puts it "Thinking Thoughts No One Has Thunk" - What has attracted Krulwich's attention is some wonderful marginalia added to the second volume of Charles Lyell's Principles of Geology (1832). It was in the second volume of Lyell's work that the famous geologist had given a clear outline of the scandalous evolutionary views of the French naturalist Jean Baptist Lamarck, only to refute them. Krulwich notes that alongside an account of Lamarck's views Darwin scrawled in his makeshift Spanglish, "If this were true adios theory". Darwin even from the first, then, saw his own theory as distinct from that of Lamarck - Right? Well, er actually, no.

Darwin was certainly anxious to distance his views from those of Lamarck when he published Origin in 1859 - in part to preserve his claim to originality, in part to prevent association with the radical political views of many Lamarckians, but that was not what was going on here.

On the importance of careful reading:
I've often told my students about the importance of careful reading, and the dangers of quote mining - even when it's late and the essay is due first thing in the morning. "Put the text in context" I tell them, by which I mean that they should both look at such juicy small excerpts of text in the context of the broader argument that the author is trying to make, as well as putting the argument as a whole in the context of the intellectual and social debate of which it is a part. Mr. Krulwich would get an 'F' on the first of these, I'm afraid.



Darwin's copy of the second volume of
Charles Lyell's Principles of Geology 

Thanks to the wonderful people at Chicago University Press I have on my shelf a facsimile copy of the first edition of Lyell's Principles of Geology. I saw one once in a second hand book shop, but didn't have a house to mortgage at the time, and so I have been ever grateful for these affordable facsimile copies.  

(In the intersts of full disclosure I should confess that I have a contract with Chicago for one book I am writing and meet with one of their editors about a contract for my Political Descent book in a couple of days... as I say, they really are nice people!)

Well, thanks to CUP I turned to this passage only to find that Lyell is not in mid rant against Lamarck at this point, that was his theme for the previous two chapters, certainly. Here, however, Lyell is laying out his own views on the species question. He makes six points, the section that Darwin annotated is the fourth. Lyell writes:

"4thly. The entire variation from the original type, which any given kind of change can produce, may usually be effected in a brief peiod of time, after which no farther variation can be obtained by continuing to alter the circumstances, though ever so gradually,-indefinite divergence, either in the way of improvement or deterioration, being prevented, and the least possible excess beyond the defined limits being fatal to the existence of the individual." (Lyell, Principles, Vol. II, Chicago UP, p.65).

Lyell concludes (in point six) that "From the above considerations, it appears that species have a real existence in nature, and that each was endowed, at the time of creation, with the attributes and organization by which it is now distinguished." (Lyell, Principles, Vol.II, p.65).

So, as historians have long known, Lyell was determined to undermine Lamarck, and in the process gave a careful outline of exactly what Lamarck had said. He then - in this six point summary, gave his own conclusions as to why Lamarck was so wrongheaded. Darwin, however was more convinced by the Frenchman's argument than by Lyell's rebuttal.
When he read Lyell's account, he noted in the margin:

"If this were true, adios theory."


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Please note: although I think marginal notes in books are way cool, this is only the case if you own the book. If it's a library book, DON'T.

Evolutionary Restraints

Mark E. Borrello, Evolutionary Restraints.  The Contentious History of Group Selection.  Chicago: Chicago University Press, 2010.

One of the best books I've read in a while that has given me real food for thought is Mark Borrello's Evolutionary Restraints (2010). In fact, the more I think about it the book I am working on, Political Descent, seems almost to work as a 'prequel' to Borello's study. My focus is on debate about the politics of evolution (and selection) from Darwin up to the mid 1920s when the marxist geneticist J.B.S. Haldane mooted the possibility of a genetic model for the evolution of a genuine "other-regarding" altruism, Borello's focus is on the later debate about group selection involving Vero Copner Wynne Edwards.

Borello's work is particularly relevent now given the present and heated debate about kin selection that resulted from the recent publication by Harvard mathematician Martin Nowak in Nature (This links to a great blog discussion of of the paper and the response).  Anyhow, here's the essence of Borrello's book - I highly recommend it:

Evolutionary Restraints is a detailed genealogy of the ‘contentious history of group selection’ in evolutionary biology, which Borrello explores, in part, through the biography of Vero Copner Wynne Edwards, the man whose name has become synonymous with the concept following the 1962 publication of Animal Dispersal in relation to Social Behaviour.  As Borrello shows, though, the history of group selection is deeper and broader than this.  Not only did the Russian geographer and zoologist Peter Kropotkin clash with Thomas Huxley over this issue in the 1880s and ‘90s, but Darwin also explicitly invoked group selection both in relation to the evolution of both the social insects and the social instincts -  both of the higher animals and of man.  Thus, in contrast to the partisan opinions of Michael Ruse (1980) and more recently of Helena Cronin (1991), Darwin and Wallace were not the fountainhead of individualist orthodoxy, and each was quite comfortable appealing to group selection when it suited them.  Further, so too was almost every other nineteenth and early-twentieth century biologist of significance, August Weismann among them.  As Borrello points out, this much was hardly surprising given debate about the compound individuality of primitive “communal organisms” such as the marine invertebrate Portuguese Man-O’-War and coral polyps.

V.C. Wynne Edwards

It is chapter three before we meet Wynne Edwards, and Borrello allows the story he tells of the development of Wynne Edwards’ ideas to echo those of Kropotkin quite explicitly.  Wynne Edwards’ own history took him from Oxford to McGill in Canada before later returning to Aberdeen.  Like Kropotkin, who had surveyed the expansive Siberian Steppes, the environment in which Wynne Edwards pursued his field studies were similarly harsh and sparsely populated; to both men it was evident that the ‘struggle for existence’ went on between organisms and their environment much more so than between competing organisms.  By 1939, in his paper on the intermittent breeding of Wynne Edwards was already framing his observations of bird populations in the context of the group selectionist theory with which his name was to become inextricably linked.  Wynne Edwards estimated that only 1/3^rd – 2/5^ths of the breeding colony appeared to be engaged in rearing offspring and hypothesized that this non-breeding behavior was an evolved strategic behavior to regulate their numbers in order to avoid the threat of over-population. 

This explanation of the social behavior of breeding populations brought Wynne Edwards into conflict with fellow ornithologist David Lack.  Lack was the first to make the case that the variable beak morphology of the finches were adaptations resulting from selection operating not – as Wynne Edwards had argued regarding his Fulmars, to prevent the population running up against limited resources, - but precisely because population ran up against limited resources.  As Rosemary and Peter Grant would later illustrate, what we now know as ‘Darwin’s finches’ are a perfect illustration of Darwin’s theory of divergence.  The battle lines between Lack and Wynne Edwards were publically drawn at a session of the British Ornithological Union on population ecology, the proceedings of which were later published in the journal Ibis in 1959.  As Borrello tells it, the battle of words was carried on by proxy, with Wynne Edwards stuck in the United States at the time, George Dunnett, one of his students, presented in his stead.  It was in “Control of Population Density through Social Behaviour, A Hypothesis”, the paper that Dunnett read, that Wynne Edwards suggested that populations self -regulated through social conventions informed by what he called ‘epideictic’ behavior – behavior through which populations actively assessed their own population density (p.69). With Lack on home ground, he sealed the advantage, as Borrello’s research shows, however, Lack had no better empirical grounding for his own theory.  





Mark Borrello

It is important to recognize, and this is one of the key contributions that Borrello makes, that Wynne Edwards was not the lone group-selectionist voice in the wilderness that received wisdom would suggest.  Indeed, he was encouraged not only by the population thinking of the synthesis to see his own work as a part of the intellectual vanguard of modern biology, but by the fact that almost all the main players in the synthesis entertained some level of group selectionist thought. An analysis of key first-edition synthesis literature shows that  E.B. Ford, Theodosius Dobzhansky, Sewewll Wright, Ernst Mayr, G.G. Simpson and Julian Huxley, among others, all acknowledged that selection occurred on a number of levels.

Thus despite the loudly proclaimed unity in biology that accompanied the 1859 centenary of Origin, the reaction to the publication of Wynne Edwards’ Animal Dispersal in 1962 reveals that there was still much that remained unsettled on what might reasonably seem to be some fairly basic issues.   Animal Dispersal was a monumental work of some 653 pages, and although it provided little in the way of statistical analysis or sophisticated population modeling, it was packed with decades of detailed field observations in support of Wynne Edwards’ hypothesis.  As Borello notes, the book was consciously modeled on the concilient structure that Darwin had employed in Origin, it was ‘one long argument’ linking behavior and population size.  Selection at the level of the group, Wynne Edwards concluded, “was much more important that selection at the individual level” (80).

Public concern over broader environmental issues about sustainability and population growth that would later fuel enthusiasm for Paul Ehrlich’s 1968 Population Bomb, ensured Wynne Edward’s book a ready audience. Non-specialist commentators read his book as corroborating their own beliefs that any number of modern social ills could be explained in terms of a neurotic population stressed beyond its limits. However, public acclaim quickly gave way to professional ambivalence. Even former mentors like Charles Elton were little more than lukewarm, confessing he found Wynne Edwards’ account not only dogmatic, but its author guilty of “woolly thinking” (86). Lack was not far behind, and even less kind, affronted, perhaps, that Wynne Edwards had appropriated the term ‘dispersal’ from his own Natural Regulation of Animal Numbers (1954) only to give it an entirely different meaning than Lack had employed.  To Lack it had signified no more than the non-random distribution of species, to Wynne Edwards, it was the result of a communally policed strategy. Borrello notes again, though, that although Lack’s criticisms were perhaps the loudest, his argument relied primarily on parsimony and a sympathetic ear from the individualist inclinations of the post-synthesis community.

G.C. Williams made Lack’s rejection of Wynne Edwards orthodox in his now canonical Adaptation and Natural Selection (1966). In it Williams clearly stated that identification of adaptations “should be attributed to no higher level of organisation than is demanded by the evidence” (107-8). Further, selection would always act to maximize the mean reproductive level, over and above any role that multiple environments played in determining selection coefficients.  Most damning, though, was his rejection outright of group selection which he presented as fundamentally opposed to genic selection.  Despite attempts to appear reasonable, in private correspondence with Lack, Williams confessed he found Wynne Edwards’ thesis laughable and had found it difficult “to avoid the appearance of sarcasm or ridicule” (111).

Wynne Edwards faired little better at the hands of the ethologists Niko Tinbergen and Konrad Lorenz, who, despite the superficial similarities in their research also found reason to differ.  Tinbergen’s focus was on fitness that particular behaviours bestowed upon individuals, Lorenz stressed the invariability of instinctive behaviours.  Wynne Edwards perspective kept him always an outsider in the developing field.  The fact that Tinbergen and Lorenz shared the 1973 Nobel Prize for Physiology and Medicine with Karl von Frisch, while Wynne Edwards was only further marginalized only set the tone for future developments.

The increasing emphasis in biology upon mathematical modeling at the molecular level cannot be under-estimated in keeping Wynne Edwards work under fire.  G.C. Williams, W.D. Hamilton and John Maynard Smith proffered gene-level explanations for all the observed phenomena that Wynne Edwards described. Even though Maynard Smith’s ‘Haystack’ model showed that group selection was a theoretical possibility, the parameters of the model showed it not only to be extremely unlikely, but also indicated that any altruistic community that came about by such mechanisms would be fragile and vulnerable to invasion by selfish individuals. Indeed, it quickly became the go-to weapon in the arsenal of critics of group selection. E.O. Wilson’s Sociobiology and Richard Dawkins’ Selfish Gene took popular biology to a new level and in doing so sealed the fate of group selection in what Borrello calls “the dark ages of group selection” (136). Dawkins made explicit the contempt for group selection theory that Williams had kept private.

Wynne Edwards was not to be silenced, however, finally publishing his second magnum opus, Evolution through Group Selection in 1986, in which he attempted to explicitly align his own work with Sewell Wright’s much earlier simple quantitative model showing that individually disadvantageous, but socially advantageous traits could evolve through group selection.  “My task” Wynne Edwards wrote, “is obviously to show that the group selection I advocate is not essentially different from Wright’s in raising average individual fitness, and is not fragile either.” His intention, Borrello notes, was clearly to “salvage his theory from oblivion” (145).

It was not to be.  Wynne Edwards was either unable or unwilling to adapt his position on group selection to adequately take account of the many advances that had been made in the field since 1962.  Being seen to be talking past rather than to his colleagues undermined his professional reputation and as a result, Borrello argues, “he essentially wrote himself out of the debate” (147).  In the intervening years, philosophers and biologists had taken the debate to new and theoretically more sophisticated territory.  David Sloan Wilson and Michael Wade had embraced kin selection as one form of group selection, negating the ‘either-or’ scenario that Williams had painted. Also, D.S. Wilson and Elliott Sober (1998) challenged the limited presumptions of Maynard Smith’s Haystack model of group selection.  Selection at the level of the group, they demonstrated, was not only mathematically plausible, but a highly likely, and a highly stable possibility.

As Borello concludes, Wynne Edwards’ attempts to remain a significant player in the debate over group selection as it became reframed around the levels of selection were frustrated in part by the mathematical turn in theoretical biology, but, perhaps more seriously, were dogged by his continued refusal to engage with developments in the field, - despite the pleas of editors and referees. He was not helped by the fact that few journal editors were willing to entertain what was by now deemed out-dated heterodoxy.

But what should we make of this story?  What was the significance of Wynne Edwards’ career in biology? Certainly Wynne Edwards challenged others to think deeply about how one might account for the observed phenomena as a result of either genic or organismic selection, however, in light of more recent, if grudging, acceptance of hierarchical selection, Wynne Edwards was also an acknowledged influence for present day advocates of an albeit more sophisticated group selection. 

Further, though, this story is also about the nature of science and about the peculiar relationship between the empirical world and our best perceptions of it. Borello asks, when Kropotkin and Wynne Edwards look at the world and see what they see, and when Huxley and, say, Williams look at the world and see what they see, what is it that leads them to such different conclusions? In this instance, both Kropotkin and Wynne Edwards approached nature with a full appreciation of Darwinian natural selection. With Kropotkin and Huxley we might appeal to their deeply held political commitments. As an anarchist, Kropotkin just could not see the competition in nature that to Huxley’s liberal mind was only too evident.  To what extent is biology still deeply imbued with politics?  As Borrello points out, there remains a lot at stake about the nature of man in the question over whether altruism can ever be truly genuine, as well as about whether the group dynamic is the faciltator or the oppressor of personal liberties.  It would be strange if the politics of biology were not at work in the post World War Two and cold war years just as they were in the industrial revolution.  It is here that historians not only have an insight into what occurred in the past in this debate, but have an opportunity to contribute to ongoing discussions of the hierarchical nature of evolutionary theory.  In addition, of course, the history of a science is relevant to its practitioners too, not only in terms of thinking about the processes and practices of science, but in the sense that, as Borrello points out, many of the common misunderstandings of evolution to which students in biology frequently fall victim, are a direct result of the exclusive application of natural selection at the level of the individual.

In sum, Evolutionary Restraints is a compelling book and a challenging one.  It will be required reading for historians and philosophers of biology, but also clearly warrants the attention of behavioural ecologists. More sophisticated than Dugatkin’s Altruism Equation (2006), Borrello’s work provides us with a new perspective on the history of group selection that really does make us have to rethink its most vociferous condemnation in preference to selection at the level of the gene. The prevalence of group selectionist ideas throughout the synthesis years cannot be ignored in the same way that a solitary figure, however prominent, can be marginalized and cast as eccentric and imprecise.  Sewell Wright, and Dobzhnsky, as well as Mayr were only three notable figures who argued that selection occurred at multiple levels prior to what has been referred to variously as the ‘hardening’ or ‘constriction’ of the synthesis. Mayr was not alone in insisting that the individual organism, as the entity that interacted with the environment, must be the entity exposed to selection, and not merely the genes it carried.  Of course, counting genes is important, but this much is really just so much book-keeping without an awareness of the interactions that lead to their survival.  This, of course, demands attention not only to the gene, but to the individual in the context of the population and the environment.

The Politics of Evolution in the classroom

This week in my class on "The Darwinian Revolution", a sophomore class at the University of Oklahoma, I have been talking with my students about the controversial history of teaching evolution in American public schools. This has always been an interesting topic for me, not least because as a British citizen the whole concept of the constitutional separation of church and state is an alien idea - one I like, I might add.

We have looked at the notorious Scopes Trial from 1925, in which a young teacher named John Scopes agreed to purposefully teach a class on human evolution in direct contravention of House Bill No. 185 which had been enacted by the State of Tennessee as the Butler Act in the March of that year. The Butler Act was "An Act prohibiting the teaching of the Evolution theory" in all public schools, that specifically mandated "That it shall be unlawful for any teacher... to teach any theory that denies the story of the Divine Creation of man as taught in the Bible, and to teach instead that man has descended from a lower order of animals." {section one}.

Clarence Darrrow in Action in Dayton

The upshot of this case is not really my concern here - for the record, though, Scopes was found guilty (he did teach the class, he did break the law), but the verdict was later overturned - (not the victory for the evolutionists that it is sometimes made out to be - after all, they - and the ACLU who had sought this out as a test case - wanted the conviction to stand so they could appeal the law). What I am concerned with is the broader concern of why Butler was moved to propose House Bill No.185 in the first place, and why William Jennings Bryan - the twice presidential candidate and veteran populist, was moved to take the stand for the prosecution.

The late Stephen J. Gould, paleontologist and essayist on all things evolutionary, wrote about this case and the subsequent legal history of evolution in the classroom, including the various attempts to guarantee 'equal time' for the teaching of "Creationism", and later of "Creation Science" alongside evolutionary accounts of human origins in his book Rocks of Ages. (Although it post-dates the publication of Gould's book, we can add "Intelligent Design" to this list of attempts to get a religious message into schools by way of skirting the establishment clause - a great PBS video on the 2005 Kitzmiller court case is available here.)

What I particularly like about Gould's account is that after telling the story of the various trials, he goes on to ask why someone like Bryan would be so moved to oppose the teachin of science and evolutionary biology. The answer, Gould suggests, is not simply grounded in the fact that he was religious - (it was not until 1968 and Epperson v Arkansas, after all, that the teaching of a religious account of human origins was ruled an unconstitutional breach of the Establishment clause of the First Amendment. Rather his concerns lay in the politics with which evolution had come to be associated by the 1920s. These were not a politics associated with the advance of liberal values that we might associate with scientific progress, but rather the politics of the eugenics movement and the evolutionary justifications that had been given (by both sides) for the carnage of the First World War.

 Veron Kellogg had disclosed the prevalence of these views among the German High Command in his Head-Quarters Nights, - (just as George Bernard Shaw (who surely needs no wiki-link!) did in England in his long preface to Back to Methuselah). The history of evolution, we need to recognise, is not just a history of scientific discovery about the origins and ecology of the wonderful diversity of life that surrounds us and of which we are a part, but also has a history that is deeply social, deeply political, and not always pretty.

This is one reason why I think that we need to be open to and educated about the political history of evolution, - the history of the various ways in which people have taken the fact of our evolution to speak directly to questions about the sort of creatures we are, and in consequence the sort of society that we might live in. - This is especially important in light of the fact that many who oppose the teaching of evolutionary biology in public schools today argue that evolution is inherently tied to a politics that is competitive, individualistic, racist and xenophobic. The book I am writing, entitled Political Descent, will show that this has often been far from the case.  Of course, there is no guarantee that the opponents of evolution will like the idea that evolution has also been used, for instance, by Charles Darwin to endorse an inclusive liberal politics that argued that there is an evolutionary grounding for a morality that was inclusive of all people of all races, and ultimately of all sentient beings; or by the likes of Peter Kropotkin to endorse the ethics of anarchist socialism. But these are both subjects for another time...

The Politics of Descent

T

he Enlightenment was a period of previously unheard of enquiry and speculation—a quest to discover the nature of man and the nature of things through the powers of Reason. Rationality was lauded as the sole arbiter of truth; as the only sound basis for knowledge, taste, ethics and government. Having its origins in the seventeenth century, the Enlightenment encompassed the scientific revolution as well as the French and American Revolutions, it was dubbed “The Age of Reason,” by Thomas Paine, who spoke for many when he declared that “the most formidable weapon against errors of every kind is Reason. I have never used any other, and I trust I never shall.”[1] The Age of Reason spoke most directly, though, to the nature of man. Thus, although the likes of Copernicus, Galileo, Kepler and Newton looked to the stars, what they saw there altered their view of mankind—for the briefest of moments they unsettled the smug certainty that we humans were the centre of the Creator’s universe just as we are of our own. Our nature is not such that our self-obsession was for long disturbed however, and Enlightenment scholars consoled themselves with the full knowledge that even while we might not adequately speculate on the nature of God and his intentions through reasoned inquiry, it was the nature of man, after all, that was—and remains—the most interesting of questions.

Indeed, there is no greater question that this: What is the nature of man? This was true for Enlightenment scholars no less than for ourselves, troubled as we are by the sublime promises of genetic engineering, psychotropic drugs, and the deep ethical problems inherent to our reflections upon the extent and limits of autonomy.[2] But how might one even begin to answer such a question as that posed by the nature of man? The French philosopher René Descartes had drilled this down to the one presumption that he believed that man was entitled to make about himself—the fact of his existence. He was sensible of himself by virtue of his Reason—“I think, therefore I am,” he had declared in consequence; in this great world of enquiry there was no other point of certainty.[3] “Give me a lever long enough and a place to stand and I will move the Earth” the Greek mathematician Archimedes is supposed to have said—here was somewhere to stand. 

            With reason as a starting point the world could indeed move, Copernicus had proven this much, at least. Turning to man, though, what more might be said—if one could divine the nature of man one might also determine how he should live—reason had made man a moral animal, or so it seemed legitimate to presume. But how had it done so? This was a compelling question indeed! The question of the nature and the origin of the moral sentiments taxed some of the greatest minds of the enlightenment—and we shall have cause to consider the views of a number of them in this book: the associationist philosophers Adam Smith, David Hume and Erasmus Darwin, of course, as well as Nicholas de Condorcet and the English utilitarians William Godwin and Thomas Robert Malthus. These men were followed in this enquiry and in this tradition by Jeremy Bentham, James Mill and his son John Stuart Mill, the last of whom became a foundational figure in philosophical radicalism. The moral nature of man was no mere topic of idle curiosity, but rather became a fundamental question in consideration of what constituted a right and just polity. “What is the nature of man, and in consequence, how might he live?” This was the question they addressed themselves to, and the question that is my focus in this book. Further, was the nature of man fixed and unchangeable? And what did this say to the way in which he was governed? Indeed, did man make his environment, or was he made by it? All these questions and more came rushing in like a flood—rising to the task, these and other men provided no end of answers.

It is in this context that I will consider the work of the eminent nineteenth-century naturalist Charles Darwin. His 1859 book On the Origin of Species once again shook the foundations of the world. English Victorians wrestled with it, uncertain as to which was the most disturbing of the books many implications—the common ancestry with apes, or the fact that our very existence was the outcome of contingency and chance?

>>>>Are you interested in reading more?

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[1] Thomas Paine, The Age of Reason, Being an Investigation of True and Fabulous Theology, (Philadelphia, 1794), p.1.

[2] These moral questions and the challenge posed by our increasing focus, at least in the West, upon autonomy have been most recently posed by philosophers from both the right and the left of the political spectrum—Leon Kass, Life Liberty and the Defense of Human Dignity (2002)and Frances Fukuyama, Our Post-Human Future (2002) on the one hand, and Bill McKibbin Enough and Troy Duster, The Backdoor to Eugenics to name only a few from the other.

[3] René Descartes, Discourse on Method, (1637).