29. Catastrophe and Mutation 1883-1895

 From June until October 1883 large parts of the Earth were shaded beneath dark clouds of smoke. More than half of the Indonesian island of Krakatau was thrown up into the atmosphere and tsunamis killed more than 36,000 people along the coast in Java and Sumatra, 40km away. The effects on climate and weather lasted for over a year and they were global.

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The island actually increased in size with new volcanic ash up to 80m thick and in effect the whole area had been sterilised so that animal and plant life ceased. The following year someone saw a single spider and “a few blades of grass”, then more plants and a few birds and insects arrived from the island of Sebesi, 12kms away. Slowly, life started to return to something like it was before. Within fifty years the entire surface was re-colonised with forest but the changing succession, the sequence of changes to return to the stable flora and fauna, continued for several decades. Arguably it is still going on, well over a hundred years later. But a sudden event such as an explosion followed by a hundred years recovery was just one trivial catastrophe on a geological time-scale. For biological evolution it was a small turn of the screw, just one more environmental change. The eruption advanced interest in catastrophic events in nature and biologists learnt a lot. For example, when things moved onward more rapidly than was normal, it showed that there were major consequences for the environment of the region and for its biodiversity.

The immediate concerns of the scientists involved were to improve their understanding of volcanic activities so that one day they might be predictable. Others were able to use the new virgin territory to monitor the whole process of re-colonisation of the new island’s species. With his recent field experience in the Himalayas the Royal Society appointed Richard Strachey to chair a group to investigate these scientific aspects of the eruption. For evolutionary biologists the sudden event also raised interest in Frances Galton’s recently aired beliefs that sudden changes in the physical environment might cause evolutionary change. Darwin remained in favour of gradual change, not catastrophic: Galton was opening the argument again with a new theory and the volcanic event at Krakatau helped him keep the debate alive even though most of the biologists in The Royal Society remained highly sceptical.

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Richard Strachey and his family – 1890s

To help understand catastrophism, Galton had raised the metaphor of a rough stone that would “tumble over into a new position of stability” an idea that first appeared publicly as an afterthought at the back of his earlier 1869 book Hereditary Genius.1.2

Now he had good discussions with Richard Strachey about Krakatau so he brought the same metaphor into the main argument of his second important book about evolution published in1889, Natural Inheritance. Galton liked gadgets and so this time he made a wooden model of the rough stone, a few centimetres in diameter with 64 surfaces. He took it with him to demonstrate at his lectures all around the country and he called it his “polyhedron”: tipping the model to tumble onto one stable surface after another, each tumble taking it to a new position of stability until the next catastrophe.

Steve Gould called it “evolution by jerks”.

Similar thoughts were going on in the minds of a different kind of life scientists, those few new biologists beginning to measure inside reproductive cells. But cells were vulnerable to their own kind of catastrophe otherwise known as mutation, a phenomenon noticed by one of Darwin’s acquaintances, the horticulturist Hugo de Vries. This Dutchman bred varieties of evening primrose and in 1889 he observed that a larger version than normal had appeared in an instant during his breeding programme and it was stable, surviving from one generation to another. Although the Cambridge plant and animal breeder, William Bateson, suspected it to be a hybrid rather than a new species he agreed that its sudden appearance was important and should be investigated further as a possible example of a new mutant. There was also the difficulty for these breeders about what these words meant: species, variety, hybrid and mutant.

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Were such instant new forms, de Vries and Galton wondered, compatible with the more gradual changes foreseen by Darwin? They noticed a lot of continuing support for change being cause by environmental changes, especially catastrophic ones rather than the familiar gradual trends. And there was still no sign of evidence to support any other kind of selection. There seemed to be a return to the idea of straight adaptation to a new environment, more like Lamarck’s continuous changes than Darwin’s selection of one or another.

Support for natural selection was reduced even more by a split between its two most fervent backers, Galton and Lankester, respectively the quantifier and the qualifier. Lankester’s limited powers in diplomacy were not going to help even though he understood both sides and tried to keep them together. For Galton, measuring seemed to be taking over from feeling, and instead Lankester stuck with the old methods to which slow hard work describing intricate new structures would eventually answer all the important questions. There was no new evidence to bring things together. Instead there was an unhappy set-back in 1895: the death of Thomas Huxley.  Lankester was devastated: “There has been no man or woman whom I have met on my journey through life, whom I have loved or regarded as I have him, and I feel that the world has shrunk and become a poor thing, now that his splendid spirit and delightful presence are gone from it. Ever since I was a little boy, he has been my ideal and my hero.”

The conflicts became obvious just after Huxley’s death but the foundations went back at least a decade and arguably much longer. Lankester began to notice the split showing up between his friends as their personalities and experiences took them in opposite directions. One group went outwards into space and time, keeping an open mind on how things worked; the other went inwards to the cell, looking for smaller and smaller units and expecting them to hold the key to it all. Lankester knew that studying evolutionary biology was one way to get down to the brass tacks about the meaning of life.

30. Statistics against Biometrics 1895-1906

Francis Galton was 25 years older than Ray Lankester and for some years they had lived nearby on the southern side of Hyde Park where they often talked about their different attitudes to science.

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Galton believed the answer to the question of life could be measured, yet his main problem was that he didn’t have much data. He offered a challenge to Lankester about his elaborate plans for an Anthropometric Laboratory, with which he was going to collect a lot of data and analyse them. The project was being set up just down the road in South Kensington to measure physical and behavioural features from visitors to the International Health Exhibition. It was a small roving exhibition and volunteers were measured for things like head size and shape, then the results were compared to social status and other factors. Galton was also collecting data about the intelligence of whole families, though a lot of the detail seemed even then to be of dubious social, let alone scientific, acceptability. One of his expectations was to devise an index to measure the range of human intelligence.

Interested and able to get involved with this kind of analysis was Carl Pearson, a student of marine biology and statistics who Lankester had encouraged to go to Naples with Weldon. Pearson was brought up in archetypal Victorian middle class family tradition. His domineering father was a successful hard-working barrister who paid no attention to his family during term-time. Holidays were for that kind of thing and it was then that the youngsters would be taken shooting and fishing, all strictly under his control.  Carl was sent to school at Rugby and then studied mathematics at Cambridge. In 1879 he got a First and was elected to the Fellowship of King’s College Cambridge. He hated anything compulsory and argued with his colleagues about the divinity classes that he had to attend with all the other Fellows.

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The freedom of the Cambridge life-style allowed Pearson to visit Berlin and Heidelberg regularly and he took an interest in the contemporary German philosophers who were so busy then. He admired the new left wing politics and JBS Haldane argued later that it was Marx’s influence that caused him to change his name from Carl to Karl. But statistical mathematics excited him most and the new data being accumulated by people like Francis Galton presented him with very attractive applications.

Statistical analysis is an advance from what he’d been doing with Raphael Weldon at the Marine Station in Naples a few years previously. Weldon had followed Lankester’s main interest in marine life and spent time at the Naples laboratory and later at Plymouth, but was more and more aware that evolution was a statistical problem. Then, a major challenge was to provide more data for analysis and Weldon obtained measurements of the death rates of crabs and snails. In 1889 he took Lankester’s job at University College where he became well known as a great teacher. He also continued Lankester’s fight to defend the independence of University College and King’s College against their amalgamation as a new Albert University. Lankester had left London for an unhappy decade at Oxford, leaving the young scientists to take over many of his projects. It also put them under the influence of Galton and enlisted them both in the programme for a statistical solution to the problems of heredity and evolution.

Earlier, while he was an undergraduate at Cambridge, Weldon had been friendly with William Bateson who strongly favoured mutation as the cause of most change in evolution. Bateson presented the idea as his theory of discontinuous evolution, showing it as two peaks in the population of an earlier single species. He explained it in detail in what he hoped would become a student bible, Materials for the Study of Variation published in 1894. But he over-emphasising the role of mutation and gave no place for environmental influences. With Lankester out of the way in Oxford there was peace between these four quantitative scientists and they teamed up as Galton and Pearson, Bateson and Weldon. The harmony was not going to last for very long.

None of them gave much attention to Darwin’s ideas on natural selection until Alfred Wallace was asked to review Bateson Materials book. The next day he had the good chance to bump into Weldon in London and they talked about Bateson’s dismissal of many ideas that Darwin had cherished. Sharing their anger after that initial exchange Wallace and Weldon went away intent on going public with their criticisms of the biased approach and they both gave bad notices for the book. What had been a strong friendship at Cambridge between Bateson and Weldon became a bitter and nasty battle in their later lives. Galton continued to support the book because it added to the cause of his own polyhedron model and was one in the eye for his cousin’s old-fashioned insistence on gradualism.

A rare defender of mainstream Victorian values, and Charles Darwin in particular, was Sir William Thistleton-Dyer, the new Director of Kew Gardens who took over from Hooker in 1885. He shared the need for stability of the mean and agreed with Weldon about Bateson’s neglect of outside influences on evolution. Dyer made that same point in 1895 when he suggested that different coloured varieties of the ornamental flowering plant Cineraria were hybrids and not random mutants.

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His observations were that the different colours formed gradually not suddenly, though whether that meant they weren’t mutants wasn’t very clear then. This intervention attracted the wrath of Bateson in the correspondence columns of Nature and Weldon soon joined in. Bateson took the criticism personally and from then on his Cambridge group, studying mutations in what became Mendelian Genetics, were at war with the London school of statistical biometrics.

Most biologists agreed that experiments were needed to settle these questions and that some statistical analysis of the results could be very useful. The argument about mutations had divided those with a quantitative outlook on evolution, leaving hapless observers like Lankester and Wallace on the side-lines, feeling useless relics of another age. Their interests in evolution were far away from the mathematical theory that had entered the discussions and they felt isolated from the young generation. A serious divide had been made much deeper and science was moving on faster all the time.

So with this in mind, over a working lunch at his club in 1893, Herbert Spencer entertained Galton, Weldon, Wallace and Lankester to a discussion about “conducting statistical enquiry into the variability of organisms.” They agreed that the statistical method was the only one by which Darwin’s hypothesis could be experimentally checked.

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Later this group became the Evolution Committee of the Royal Society, which confirmed to Lankester that he was one of the few biologists then who still wanted to look outside at the whole organism and then towards the physical environment. Most others were on the new mission to the smallest part of the cell, expecting to join with the chemists and physicists and find mathematically defined laws. It frightened him that both directions of study were so vast, suggesting that progress in understanding the biology of life was going to be very slow, however fast one part of the science might be moving.

Despite the attempts of Wallace and Weldon to bring the sides together, the last few years of the nineteenth century saw a rapid decline in support for natural selection, encouraged by the Bateson affair and the continuing absence of evidence for adaptation. Galton worried that he might die before the elusive agent of heredity was discovered and to add to that despair his wife did die in 1898. Accidentally, he was kept going by stumbling across some very exciting new data, the pedigree of a large family of Basset hounds with white and yellow patches and he was able to trace the heritage back several generations. He used the data to establish the distribution of each parent’s contribution going back four and more generations. Overjoyed with such good data for analysis, Karl Pearson described the patterns of inheritance for the white and yellow patches with a series of new equations. To cheer up his depressed boss he incorporated them into a Happy New Year card, and through Galton’s warm reaction it gave all the London biometricians a new lease of life for the next battle with Cambridge.

That came sooner than they expected because Bateson was the referee of the manuscript setting out Pearson’s equations, and though the mathematics may have been original, the biology was bad. Bateson took great delight in rejecting the manuscript. In anger, the London group decided to begin a journal of their own for such articles on topics crossing the boundaries of traditional disciplines. They called it Biometrika, a name like Pearson’s own, spelt with a k not a c (and which is still publishing high quality work). Even with this divided opposition, Lankester was side-lined in his defence of Darwin and the holistic view he still had for all of nature.

No-one would have guessed anything special was going to happen when William Bateson caught the train from Cambridge to London one morning in May 1900. He had been invited to give a lecture to the Royal Horticultural Society in Chelsea, about what was being called Galton’s Law. It came from the conclusions of Galton’s analysis of the white and yellow patches on Basset hounds that had just been published and stated that parents contributed equally to their offspring’s inherited matter. On the slow train journey Bateson happened to turn to some papers that he had bundled into his bag and there he found an unread reprint of an article published 34 years earlier. The author was Gregor Mendel.

Bateson realised that the characters that Mendel had counted over several generations of peas showed a pattern was carried from one generation to the other. Whatever this mechanism, Bateson thought the forty year old manuscript by Mendel had some important messages for plant breeders, a different kind of detail to Galton’s and that difference was important. Legend has it that before his train reached Liverpool Street Station in London he had decided to change his lecture and share Mendel’s work with his audience at the Chelsea Society. The audience remained silent, unaware of the importance of the monk’s breeding experiments, let alone why such dull work should be presented with such excitement. It was an uncanny repeat of the silence after the Darwin and Wallace paper was read out to the Linnean Society 42 years earlier.


Wanting to share his discovery further, Bateson wrote to alert Galton to the manuscript “in case you may miss it. Mendel’s work seems to me one of the most remarkable investigations yet made on heredity, and it is extraordinary that it should have got forgotten.” Like the audience in Chelsea, Galton did not respond and it took several years for the penny to drop in other peoples’ minds. To be fair to these observers, it was easy to miss the point from Mendel’s funny little experiments crossing different varieties of peas. For the uninitiated, and that was the vast majority, counts of which pea types had shown up in the next generation were a long way from finding Darwin’s missing units of inheritance.

Attracting hardly any interest, Bateson was left feeling that he had over-reacted to the Mendel article and that his own idea was best after all. It was six years since he had first suggested that hereditary features were transmitted by vibrations rather than by discrete particles. He became so pre-occupied with this hypothesis that he preferred to dismiss all other explanations of evolution out of hand. He summarised Darwin in one line: “Selection is a true phenomenon; but its function is to select, not to create”. He was convinced that Darwin had got it wrong and the London group around Galton and Pearson were beyond the pale.

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After all, there was still a possibility that his own theory of vibrating messages would show Mendelian ratios and that he had been right to get excited about the re-discovered manuscript. So with this always in his mind he never did accept the chromosomal account of inheritance.

Some people, like de Vries, saw mutation as hereditary change showing up as splitting one species into two, big evolutionary jumps, more dramatic than anything Darwin had anticipated in the world of gradual change. Later in the decade “mutation” became associated with smaller changes, encouraged by replacing the difficult expression “Mendelian Factor” with the simple term “gene”.

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These changes in single characters were usually happening within a species, one simple character controlled by one gene. Even Darwin had expected that mechanisms such as mutation might account for evolutionary change in addition to the more important process of natural selection. In the last sentence of the Introduction to the Origin he wrote: “I am convinced that Natural Selection has been the main but not exclusive means of modification.”

In all parts of the life sciences the quantitative approach was gaining ground. With this trend were Pearson and Weldon who used mathematics to test the validity of Mendel’s own data and whose results appeared to be too good to be true: there was nearly 100% validity, an unheard-of result that made them suspect that someone had fiddled the calculation. But who would want to do such a thing, and with what motive?  Bateson did not accept the criticism and saw Weldon’s questioning of Mendel’s conclusions as an over-reaction. Ever more determined, Weldon went on looking for evidence of adaptation in his snails and crabs, measuring their death rates, looking for signs of extinction of old species and origin of new ones. But he didn’t find any new evidence and blamed that on the complexity of the way his organisms showed their adaptation and selection. This was no way of gaining support from an already disillusioned group of Darwin supporters.

Bateson took an even stronger line against the London statisticians in Pearson’s circle and the bitter dispute about the nature of evolution and the value of the statistical method continued to rage. The battle over Mendel’s data reached its climax at the 1904 British Association meeting that happened to be in Cambridge where the audience was hostile to the London side. In response, a complacent Bateson was the proud host and felt confident that his students would give some good talks. In the event there was a series of dull displays of statistical analysis from the Londoners and the presentations of breeding experiments from Bateson’s Cambridge group were not much better. At the end, Pearson offered to bring the two sides together: the chairman looked around at the blank faces of the Cambridge audience and saw little enthusiasm for this: “But what I say is: let them fight it out!” The audience broke up into small groups, each wondering which side another was on, and many thinking that strong leadership could have led the scientists out of this hole. Instead, most of the positive work was being done miles away in New York and the English contribution was about to turn into tragedy.

Weldon was eager to accept the challenge from that meeting and his first chance for reconciliation came later in the year when the Royal Society asked him to referee a manuscript from one of Bateson’s supporters. It took data from the pedigrees of race horses and found what the author argued to be Mendelian ratios in the coat colours: bay and brown being dominant to the chestnut recessive. Weldon was unhappy with the author’s colour assignments, fearing they had been altered to give good results, and asked the author to clarify the methodology and the definition of his characters. The great row that inevitably followed ended in the work eventually being published, but with a brief footnote. It highlighted some of the changes that Weldon had demanded, but the fact that the article was published at all gave victory to Bateson’s group.

In despair at this outcome Weldon put all his energy into disproving the article’s results, and that meant him finding evidence of a chestnut mare, crossed with another chestnut, giving birth to a brown or a bay foal. He worked frantically, saying “I cannot leave this thing unsettled” and causing his good friend Pearson to measure the seriousness of the situation by observing that “he used stronger language than I have ever heard him use.” Eventually Weldon found the information he had hoped for and was persuaded to take a family holiday with the Pearsons. That was when the 45 year old Weldon caught a chest infection and he died a few weeks later. It was generally agreed this was from the exhaustion, worry and over-work.

All these incidents between London and Cambridge were based on uncertainty and the battles they created were really about their lack of belief in what Darwin or Mendel had been telling them. More particularly, Darwin’s theory was aggressively rejected by Bateson’s group and Mendel’s was not accepted by Weldon and his London friends. But neither group had anywhere else to turn, other than to the hope that somehow the numbers games would come up with something unexpected.

That was exactly what happened when new calculations laid another of Darwin’s ghosts to rest, the one about the age of the earth and the time that had most likely been available for animal and plant evolution to have happened. In 1904, during one of his public lectures, Rutherford announced his new radiometric dating techniques which measured the half-life of elements such as radium present in particular rocks. His first results suggested a much older age of the earth than had been advocated by Thomson.

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Afterwards, Rutherford is said to have described his shocked reaction to seeing Thomson in the audience as he began the lecture. “To my relief Thomson fell fast asleep, but as I came to the important part, I saw the old bird sit up, open an eye and cock a baleful glance at me. Then a sudden inspiration came, and I said Thomson had limited the age of the earth, provided no new source of heat was discovered. That prophetic utterance refers to what we are now considering tonight, radium!” Just as Thomson saw through Lyell’s shaky edifice of uniformity, Rutherford had seen through Thomson’s single-tracked thinking about the cooling planet earth: more than one thing was going on at once and that had clouded each process.

31. Uncertainty About Mendel 1904-1907

Useful though these biometric methods were to become, another fruitful line of enquiry was beginning in the United States. The mutations first described in plants by de Vries, and then recognised elsewhere by Bateson, were to be tackled really seriously by one of Lankester’s marine biology acquaintances, one who took delight in the switch from qualitative to quantitative outlooks. Thomas Hunt Morgan (1866-1945) was to win a Nobel Prize for his “discoveries concerning the role played by the chromosome in heredity” particularly in fruit flies. Although the penny that dropped on Bateson’s train journey in 1900 was heard clearly by Morgan, it took until 1933 for the Prize to be awarded and full public approval to be granted.

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When Morgan was a student at Woods Hole Marine Biological Laboratory in Massachusetts he had been interested in the embryology of sea spiders, to find out whether they were crustaceans or arachnids. But when he followed the footsteps of Lankester in 1894 to the marine laboratory in Naples, he became more interested in the chemical and physical changes that happened as the little creatures developed. From then on, TH Morgan saw no need for natural selection and believed that species had no reality in the flow of nature. “It appears that new species are born; they are not made by Darwinian methods, and the theory of natural selection has nothing to do with the origin of species, but with the survival of already formed ones.”  To him, a “naturalist” was the opposite of a “scientist” and biology could be explained in terms of physics and chemistry. This was despite him knowing very little of either and at the same time insisting that “genetics can be studied without any reference to evolution”. These were purist views, typical of the way an enthusiast supports a new idea if only to get it across. But they also showed the power of reduction to the smallest detail and the growing popularity of the quantitative approach.

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Morgan’s view was a restatement of the old supporters of Lamarck who thought that selection could not create but only reject. They failed to see that it was through this rejection that new forms are created and it was to be three decades later that Morgan’s students realised this. They agreed that evolution worked at many levels, whether it was with the mathematics of molecules and populations of organisms, with the physics and chemistry of genes, the chemistry of physiology and with the influence of different environments. By change at any of these levels, biodiversity came from the splitting of lineages, by speciation, and that gave discontinuity in nature. But it was mutations that mattered most to Morgan because he thought they created new species immediately, despite the environment. They also occurred in single genes and would become extinct only if the change was harmful for all individuals.

From 1904 Morgan moved to New York aiming to find the commonly suspected patterns of change that seemed to be continuously passing from one generation to another. By then the talk of Mendel’s ratios validating the role of sudden mutations gave him new hope and he started to look for evidence of what might be in control of these hereditary characters. That was when he started to work with fruit flies, which were easy to use in his laboratory, or the ‘fly room’ as he called it.

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This was more than just a room, rather a very well-organised and well-led group of enthusiasts who were in at the beginning of experimental genetics. Drosophila flies were cheap, quick and easy to breed in milk bottles, and the features they inherited showed up very noticeably. They also had just four very large chromosomes that showed changes in shape and colour in different parts, ideal for examining chromosomal events during the sexual and asexual cycles of cell division, how they related to the structural features of the adult. Mutations came easily and bred true as variously coloured eyes, striped bodies, wings of different shapes and such like. But these techniques were going to need thousands of experiments before any trends emerged and even then the data were not going to be easy to analyse and interpret. Like the processes they were monitoring the experiments needed to be fully controlled and monitored.

In 1907 Bateson visited Yale and set out his post-Mendelian statement about genetics and evolution. He fought for the slow and gradual recombinations that Mendel’s work had described and for which he had proposed the word “genetics” two years earlier. But there was still no evidence that any particles on the chromosomes or anywhere else were the agents of inheritance. Bateson kept quiet then about his thoughts on the train in 1900 when he also remembered another article by a German cytologist Theodor Boveri. That was about structures inside the cell nuclei of sea urchin embryos, structures the author had called chromosomes. Could the recombined ratios, he wondered, come from a re-sorting of particles on these chromosomes during sexual reproduction, both in peas and sea urchins?

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Bateson preferred to think that vibrations were the more likely agent: “In Mendelian analysis we have now, it is true, something comparable with the clue of chemistry, but there is little prospect of penetrating the obscurity which envelops the mechanical aspect of our phenomena.” Inheritance must be transmitted by a force from physics, vibration. “Patterns mechanically produced are of many and very diverse kinds. One of the most familiar examples, and one presenting some especially striking analogies to organic patterns, is that provided by the ripples of a mackerel sky, or those made in a flat sandy beach by the wind or the ebbing tide.”

It came as no surprise that Bateson and Morgan were very different kinds of people, and they did not get on. Morgan looked a bit like Weldon with a droopy moustache, and he had a casual and laid-back southern outlook on life. Bateson thought Morgan “rough”, “of no considerable account” and “dreadfully small” and even reported to his wife that “TH Morgan is a thickhead”. At first, in 1907, about the only thing they had in common was that chromosomes were not of much significance in genetics.

Both men had nothing to say about natural selection or Darwin or the link between his ideas and Mendel’s ratios of inheritance. Instead, based as much on envy as on reason, Bateson gave Morgan, and the audience at Yale, his own eccentric idea, but with no new evidence. Instead, because he couldn’t leave his vibration theory alone, he could only offer a sad and weak attempt to get the idea across. “I think we are entitled to the inference that in the formation of patterns in animals and plants mechanical forces are operating which ought to be, and will prove to be, capable of mathematical analysis.”

Bateson left America without changing anyone’s mind about the cause of evolution, let alone his own. Many of the different ways of explaining evolution were still possibilities though they all had different levels of support and very few people could appreciate how each idea might fit into the whole picture of a living system. There was some support for vibrations, and evidence for degeneration, a case for eugenics and mutation as well as for Darwin’s more gradual theory of natural selection. Without evidence from genetics, geological dating, biogeography, migration or ecology, no single theory or investigator stood out as the most acceptable. There was still all to play for in the game of trying to understand how life worked. Although new disciplines had begun in the four decades since the Origin was published, none had given any major new advance. What little seemed acceptable was split up, all in bits.

One thing was for sure, the way to understand Mendel’s results was going to be through some quantitative assessment, and Lankester organised a series of monthly dinners at his club to help find a way through. On such occasions there was a lot of talk about the biometric work coming from Galton’s laboratory and how it might inform those hoping for selective human breeding. Two mathematicians in the group, Whitehead and Russell, were preparing to work on their monumental Principia Mathematica, and they wanted to link Lankester’s descriptive biology to their own quantitative methods, but it was going to be difficult at first. th

Bertrand Russell (1872-1970) was best placed to link these topical issues because he was one of the few people who could understand both languages. As a philosopher with mathematical skills, he was in the right place at the right time. Together with Whitehead he advocated three requirements to explain the history of life: a concept of infinity, the flexibility of choice, and the desire to reduce explanations to the smallest component. This rigorous and optimistic programme was raising the stakes of biologists, pushing qualitative description to one side and claiming biometricians as heroes.

Even Lankester appreciated that this was the way things were going and was pleased to explain that the new methods relied on descriptive data. And all around in science there was a shortage of data; numbers were highly sought after and soon there were plenty of non-scientists interested in mathematics solely as a mental exercise. This was summed up by one presentation in Cambridge that attracted a large audience when Bertrand Russell teamed up with TS Eliot reciting the value of pi for ten minutes. There was then a period of meditation before they continued with another hundred or more decimal places of the constant’s value. It was going to be important for biologists to keep mathematics under control.

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What the old brigade of Wallace and Lankester saw as being even more difficult than describing life by equations was the link these mathematicians, outsiders to their classical biology, had with their philosopher friends in Vienna. It was the group that Rudolf Carnap was to lead for the next two decades and which at this early stage became known as the First Vienna Circle. These men met at the Cafe Central and had a very clear vision of where philosophy was going. They saw all knowledge coming together as a single language of science, becoming more and more precise and leading to a single truth, one Law of Life being verifiable by experiments and taken further by mathematical modelling.

Russell was never an inside member of this group and later went out of his way to distance himself, despite the excellent credentials that his three explanation of life gave him for membership. Instead, he experienced an incident that his Principia Mathematica could not account for and which would have been rejected as irrelevant by the First Vienna Circle. It was from a simple incident of seeing the pain felt by a lonely woman as she was growing old, all alone. “Having for years cared only for exactness and analysis, I found myself filled with semi-mystical feelings about beauty …. to find some philosophy which should make human life endurable.”

By then, measuring was seen to be an essential part of the scientific routine and few young biologists were sympathetic to Russell’s feelings about the old lady’s values of life: instead they searched for other explanations. One such person was in the same city of Vienna at the same time as the philosophers, Paul Kammerer (1880-1926), an experimental zoologist. He wanted to prove Lamarkian theories of evolution by breeding reptiles such as toads and salamanders. Kammerer’s first experiments involved breeding midwife toads in warm water and after several new generations he noticed the growth of black nuptial pads on the males’ feet to keep grip on the female.

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Other toad species also had these pads, and Kammerer explained them all as adaptations to the slippery conditions. This didn’t prove or disprove Lamarck, but it did help understand something about toad life-styles. Usually the males could grip their mate easily and didn’t slip on dry land, but in the moist conditions of Kammerer’s experiments they kept sliding around. The question people were asking was whether the pads came from the expression of an existing trait, inherited from a line of ancestral species that also mated in water, or whether the pads were from a new mutation. Kammerer himself was unsure of the answer when he began this work in 1906 and didn’t come to favour the explanation that they were from rapid mutations until after the war.

With the measured scientist and philosophers at the Cafe Central and experimentalists like Kammerer, Vienna was hardening into a place of scientific rigour for the twentieth century. Not least of importance was an ear-nose-and-throat doctor, Wilhelm Fliess, who also reckoned to have helped turn biology into a science describable by mathematics. For more than ten years he had gathered data from cyclical body patterns such as 28 day menstruation and an associated periodicity of 23 days. He compared these to times of nasal bleeding, and even dates of birth and death and concocted an arithmetic formula to account for many other bodily functions as well. Fliess’s best friend was Sigmund Freud who entrusted his own nose to Fliess’s surgery, despite another patient nearly dying from similar treatment. The surgery was thought to be a cure for Freud’s arrhythmic heart beat and was regarded as a great success. It was also at the time of Freud’s entrance into life science, when he began to link human psychology to evolutionary biology.

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(Rudolph Carnap)               (Central Cafe, Viena)                                           (Freud and Fliess 1895)

Francis Galton had already been thinking about this other ghost of Darwin, how study of the mammalian mind might fit in with evolutionary mechanisms. To open up his enquiries he began to subject some of the results from his Anthropometric Laboratory to his first statistical methods of analysis. He had the height measurements of thousands of parents and their children and plotted them out in different ways. Not surprisingly, taller parents tended to have taller children, but the children were rarely taller than the parents. Equally, shorter parents had only slighter shorter children. There was a tendency to revert back towards the average.

This way of measuring association between any two like-with-like variables, such as heights of parents with heights of children, helped Galton find a standard that he applied to other closely comparable data. Two variables are often closely associated for correlations as well as regressions, similarities as well as differences. They formed the basis of much more sophisticated biometric applications, weather forecasts, economic indicators and even public opinion polls. But in all cases the quality of the prediction was no better than the quality of the data used in the first place.

32. Early Eugenics 1903-1911

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In 1903 the Daily Chronicle carried a brief article under the headlines: “Our National Physique – Prospects of the British Race – Are We Degenerating?” It was written by Galton, just after The Royal Society awarded him the Darwin Medal, and urged that “a material improvement in our British breed is not so Utopian an object as it may seem”. The next year he wrote that “the aim of eugenics is to represent each class or sect by its best specimens; that done, to leave them to work out their own civilisation in their own way.”

If it had been written specially to niggle HG Wells (1866-1946) and his new friend Lankester then the trick had certainly worked. Their fury was just about balanced by the view that Galton had always so many balls in the air at once that none of them ever really hit their target with much force. In this case, for example, he had mixed up two different meanings of ‘degeneration’ and compared them to even more confused meanings of the word ‘race’, much more difficult to define socially and genetically. Wells responded immediately and publicly: ‘I believe … it is in the sterilisation of failure, and not in the selection of successes for breeding, that the possibility of an improvement of the human stock lies.” He had hinted at these ideas a decade earlier in The Time Machine with its degenerate man-creatures.

Francis Galton was not to be silenced and when he addressed the new Sociological Society he made clear which characteristics he favoured: energy, ability, manliness, health and courteous disposition. He concluded that “what nature does blindly, slowly and ruthlessly, man may do providently, quickly and kindly. Galton was still analysing the data he had collected over the years about health and family backgrounds.

Karl Pearson was appointed as the first Galton Professor of Eugenics at University College. Statistical in 1911, some saying it was proof that racial degeneration was a threat to the Empire: improved health allowed genetically inferior people to survive; it stopped natural selection by “propagating unfitness. The right to live does not connote the right of each man to reproduce his kind. As we lessen the stringency of natural selection, and more and more of the weaklings and unfit survive, we must increase the standard, mental and physical, of parentage.” Pearson expected life to be a struggle and welcomed was as part of that struggle to survive by fitness. “National progress depends on racial fitness, and the supreme test of this fitness was war. When wars cease mankind will no longer progress for there will be nothing to check the fertility of inferior stock.”

Support came from a respectable source when Bertrand Russell first published on eugenics, applying Pearson’s statistical reasoning and the principles of Galton’s law of ancestral heredity. In conclusion, Russell advocated direct payments from the state to “desirable” parents, a plan he mooted again in the form of scholarships paid to qualifying parents while “undesirable” parents were to be discouraged from procreating and to receive no financial aid for their children from the State. He accepted the “dangers” of the so-called “differential birth rate” – the Edwardian concern that the poor sections of society reproduced much faster than the wealthy – that conditioned Russell’s understanding of parental desirability.

Only a few senior biologists rose up in protest at the even more threatening programme of legislation that was underway to restrict the breeding of the feebleminded. One who did protest was Ray Lankester who demanded that more should be known about the origin of the characteristics being controlled, whether they were inherited or learnt. He challenged what some of the terms meant, words such as “racial quality” and “improvement” meant different things to different people, and he called for a serious study of human races and their origin, before parliament intervened. Lankester made several appeals for these further considerations in his weekly Daily Telegraph column and they seemed to have a considerable effect on the public perception of the “intellectual eugenicists”. Like the spiritualists and magicians, they came across as people who could not be entirely trusted.

Galton also started to write a novel that he provisionally titled Kantsaywhere, about a eugenic utopia and its Eugenics College founded by Mr Neverwas. The college awarded diplomas for heritable gifts, physical and mental and encouraged graduates to marry one another by offering “appropriate social and material rewards” to “relieve the cost of nurturing the children.”. The professor of Vital Statistics arrived in the colony and met Miss Augusta Allfancy. Failures went to labour colonies “under conditions that were not onerous” but they had to work hard and remain celibate. Others received a certificate and could mate only “with reservations”. The plot told of what happened when the fragmentary pedigrees of immigrants caused trouble with the loves and desires of some of the new graduates of the college.

Galton was seen by his contemporaries as the founder of their “science” of eugenics in contrast to the German philosopher Friedrich Nietzsche who wrote about the “religion of eugenics. He wrote that: “Society should in many cases actually prevent the act of procreating and may without any regard for rank and under certain circumstances have recourse to castration.”

It fell to another young new Cambridge biologist to bring the London biometricians and Cambridge Mendelians together and to examine eugenic methods more seriously. Even before he graduated Ronald Fisher (1890-1962) got involved establishing a Cambridge University Eugenics Society with the advice of Maynard Keynes and Horace Darwin. It was intended that the group should be addressed by the leaders of mainstream eugenics organisations such as the Eugenics Education Society, but the eager young Fisher beat them to it and talked himself.

From the start Ronald Fisher was set for an unusual life at an extraordinary time. His family were partners in a firm of London art auctioneers and owned a big house in Hampstead. This intellectual background introduced him to the fashionable side of London society, but his happiness was rudely broken by his mother’s death and his father’s financial ruin. Fisher was still at Harrow School and buried himself in work. He won a scholarship for mathematics at Cambridge and from school, a prize of Darwin’s complete works.

In his short address to the small group at the Eugenics Education Society the very young Fisher had something good to say for both sides of the biometrics-Mendelian struggle, as well as something cautionary. Mendel had published unusually accurate results, conforming almost too much to the expected ratios, more like demonstrations than real results. Biometricians could be ambiguous, too, and they “could squeeze the truth out of the most inferior data”. As we shall see, Fisher did succeed in uniting these two factions of evolutionary biology but the specialists who recognised and described new species were still left out and the new specialists who looked at the environment were not even considered.

Later, Fisher reflected on his role as arbiter: “Each generation, perhaps, found in Mendel’s paper only what it expected to find; in the first period a repetition of the hybridization results commonly reported, in the second a discovery in inheritance supposedly difficult to reconcile with continuous evolution. Each generation, therefore, ignored what did not confirm its own expectations.” His generation of young men were more hard-minded than most, expecting that mathematics would be involved in any scientific problem. Physics and chemistry were going to be heavily involved in the forthcoming war and there was no time for soft bits of evolutionary biology.

Most of these Edwardians were intrigued by all the talk of eugenics, seeing that it joined three strands of late-Victorian science: a hereditary theory of population, the study of population statistics, and a theory of population regulation derived from population genetics. The popular belief behind the foundation of eugenics was the conviction that it was possible to scientifically intervene in the reproduction of human populations with the specific goal of biologically improving future generations.

The new century had got off to an eventful start. The new king, Edward VII, not only removed from sight Queen Victoria’s most symbolic belongings but happily strolled around Buckingham Palace with a huge cigar in his mouth. In Vienna, Sigmund Freud shocked middle class society with suggestions of unconscious sexual feelings during childhood. And in his new job as Director of the Natural History Museum, Ray Lankester felt attacked from all sides, academic and administrative, as though he could do nothing right. On one occasion he ordered a room to be repainted without consulting the Trustees. At their next meeting they severely censured him for going outside his powers: “We don’t like the new paint”, said the chairman, “and we must ask you to arrange to have it taken off.” “I thought you’d say that,” growled Lankester, “but it won’t come off.”

In his academic life there was no resolution to the struggles between nature and nurture, the endless debates between quality and quantity and the continuing confusion about whether units of inheritance had been found or not. In their different world the politicians were also unaware of similar forces acting at their more diplomatic level of the art of what was possible. But then, so little was under the direct control of both life scientists and politicians that they were all blindly walking headlong into the same great catastrophic event. 

38. Eugenics Gets Serious 1925-1936

After that Great War, Europe was left stunned more than ever before. It was a major social catastrophe and it caused many people to change their lives dramatically. One who reacted with deep insight was the ecologist Arthur Tansley, apparently secure in academia after his recent election to The Royal Society. But still troubled by the war and the haunting dream, he decided to resign his lectureship at Cambridge and move to Vienna, with his family, for intensive psychoanalysis with Sigmund Freud.

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That experience itself also had a profound impact on Tansley and he returned with new ideas for both psychology and ecology: “We must never conceal from ourselves that our concepts are creations of the human mind which we impose on the facts of nature.” Tansley was concerned at the high numbers of “feeble-minded” incarcerated in asylums to stop them contributing to national degeneracy by breeding. He had been particularly horrified by one young boy being detained for stealing a postal order and committed to a mental institution. Juvenile courts regularly institutionalised their charges for long periods and called them “morally defective”. He had a strong memory of Bloomsbury’s working class life and his rescue by the Great Ormond Street Working Men’s College.


Meanwhile, Tansley was working away quietly on gathering data from a few selected dynamic environments, listing the local movements of plant communities and checking changes in weather and species frequency. This went on to his idea of a complete “ecosystem” in the early 1920s though he didn’t coin the word until 1935. He was joining bits of his Vienna experiences of human community with his knowledge of plant communities, how separate individuals with independent powers of existence, lived together through different stages of growth and reproduction. Like Smuts’ holism, this eventually meant taking the basic laws of physics and chemistry into the realms of basic biology and then to the complexity of the human mind. They were adventures into the unknown that involved all of science and more beyond.

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One of Tansley’s acquaintances was Ronald Fisher, desperate to become his own master after the war. He rejected the suggestion of his hero Pearson that they worked together at University College and quickly found the right opportunity at Rothamsted Experimental Station. Population genetics, experimental genetics and the systematic description of species were topics that needed to be brought together, and Fisher attempted to join Darwin’s natural selection with Mendel’s genetic ratios, helped by the pre-war realisation that gene particles lined up on the chromosomes. Fisher considered all the mathematical and statistical support for that model and showed for the first time that selection was the only way in which the process could be explained. Evolution couldn’t just be driven from inside the organisms: it needed a push from outside in the environment, one that led to the kind of small-scale continuous variability that Mendel’s recombinations explained so well. That explanation also meant that selection happened on a huge scale, between every organism in all places at all times, persistently testing the way forward and each time choosing the best option

Fisher’s mathematical insight drew comparisons between this style of evolution and the second law of thermodynamics. “Both are properties of populations, or aggregates, true irrespective of the units which compose them; both are statistical laws; each requires the constant increase of a measurable quantity, in the one case the order of a physical system, and in the other the fitness …. of a biological population.” Malthus’s warnings a hundred years before were always at the back of his mind, and his mission after the war was to do something about that.

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This challenge led him to go too far and he spoiled his reputation in a book called The Genetical Theory of Natural Selection. In the spirit of the time, and doing himself no long-lasting favours, Fisher used the last part of the book for a long justification of the eugenics practices that he favoured so strongly. “The deductions respecting Man are strictly inseparable from the more general chapters.” He believed that industrial society was in decline due to “social promotion of the relatively infertile” and that the higher reproductive rate of less worthy classes will swamp the superior classes. His chapters carried statistical evidence for that view and it continued to attract support throughout the decade.

Two years after Fisher’s work was published, JBS Haldane came along with The Causes of Evolution, an easier book to read in which all the mathematics was relegated to the appendix. Like Fisher’s story, Haldane’s showed the power of natural selection and he began in characteristic mood: “I can write of natural selection with authority because I am one of the three people who know most about its mathematical theory.” Presumably the other two were Ronald Fisher and Sewall Wright. So the study of population genetics began, the mathematics of patterns of change coming from large numbers of individuals living together in the same community. Many biologists didn’t understand the mathematics, but a main reason for the intellectual movement they had accidentally created was to bridge these differences.

Although Haldane strongly rejected Lamarckian trends he did leave open the space for some evolution to happen by means other than natural selection. Hybridisation and some large mutations could make new species and Haldane urged that other processes may explain even more. He had plenty of evidence that “degeneration is a far commoner phenomenon than progress” and is usually hard to spot because it leads to extinction. From the fossil record he noticed “at any given evolutionary level we generally find one or two lines leading up to it, and dozens leading down”.

This kind of division within the biological community, between the rigid fixers and the more artistic flexers, was to continue for many years and was going to get much wider before anyone spotted the difficulties. But both sides needed to gain more knowledge about themselves before they could share their similarities and find out that way just how evolution worked. The two World Wars also kept scientists on these separate tracks, rigid armaments relied on a strong force of hard-nosed mathematicians, physicists and chemists. The more vague environmental sciences such as the new ecology and palaeontology didn’t stand much of a chance. Tansley and Huxley would have to wait.

Eugenics gave a very different outlook for all life on the planet and Fisher was hell-bent on exploiting biology in that single direction. He likened natural selection to the working of a casino where the odds were set for its own success. In his game the losers were equivalent to an extinct species, and the far right were the winners, supporting the casino owner who was only too pleased to let science make up the rules. Never one for philosophy or history, Fisher believed there was little new “that would not stand if the world had been created in 4004BC”.

Already a government committee had been set up to recommend how to sterilise the ‘feeble-minded’ in England and Wales where a quarter of a million people were classified as ‘mental defectives’ and suggestions had been made that they were suitable candidates for voluntary sterilisation. Some members of the committee thought that “Broadly speaking stupid people will produce stupid children” though Haldane and Hogben explained there was no certainty about that and that environmental factors were also likely to be a cause. The committee rejected compulsory sterilisation though even Haldane agreed with their conclusions: “Biologists may legitimately demand that a proportion of mental defectives should be prevented from breeding”.

Another popular debate between the two world wars was stimulated by Arthur Keith’s proposition that humans would become extinct without competition, and that meant war was necessary. Keith was an anatomist and anthropologist and wrote articles in the popular press advocating physical struggle and conflict as a vital biological characteristic.

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In 1927 he told the British Association that cultural differences provide a mental barrier to social groups, suggesting that territorial behavior is a strong force in human evolution. He saw scientific and political merit in his idea of an ‘in-group’ and the less favoured ‘out-group’.

That view had been expressed earlier by a Prussian General, Friedrich von Bernhardi, in 1912, and several biologists agreed that it might be a possibility. Herbert Spencer had promoted the idea with his phrase “the survival of the fittest” and many in Europe during the 1920s and 30s believed that war played a role in this kind of biological process. They believed that the individuals and communities that survived were fitter in an evolutionary sense than those who died. This argument got tied up with some of the views of the eugenicists, inevitable consequences of those frightened times.

A forceful opponent of Keith was Joseph Needham, an influential member of the Cambridge left. He deplored the way that phrases such as “the struggle for existence” and “survival of the fittest” had become “the stock-in-trade of the man-in-the-street”, arguing that they were from old Victorian attitudes, then taken in by the public as part of their justification for joining-up to the armed forces. Keith continued with the idea that it was simply nature’s way of controlling populations, but the left-wing scientists would have none of that and instead sprang on to the offensive with their positive eugenics programme. Julian Huxley aimed for “the virtual elimination of the few lowest and most degenerate types” and he argued that biology should be the chief tool for rendering social politics.

The Huxley brothers called for some different thinking as part of a strategy towards the best interests of the planet earth and the human race. It involved a universal rhythm within earthly life, something they had been talking about for the last decade. As premier intellectuals they felt a responsibility to see to it that anything like the tragedy of the First World War did not happen again and that knowledge of the common features for all biodiversity might stop humanity being dragged down into the same mud. And from their family tradition, their grandfather’s smile beaming down on them constantly, Aldous and Julian had plenty more expected of them. Even those sceptical of Darwin’s ideas, broad minded men like Lawrence and Shaw, had their influence on Julian Huxley. More than most other scientists he was the cautious centrist, enjoying the foundations of Smuts’ holism very much within the fairness and functionalism of the English tradition. So prepared, Huxley went on to give another lecture that year in which once again he considered some of these wider aspects of his world view for our own species.

He called the lecture Eugenics and Society and repeated his call for a new Social Science to find ways to improve the living standards in the poorer parts of human communities, promote policies to favour more middle class babies and to discourage large families for the poor. But despite these hopes and concerns for a joined-up future it was hard in the 1930s to see any exciting new trends developing in biology and the evolution of life. Things were on hold, digesting the full impact of genetics, natural selection, cellular biochemistry and ecology as they became mixed together, waiting for their union to be approved and their significance understood. Krebs advances in biochemistry set an example against this pessimism that appealed to the Huxleys.

Meanwhile, even the attention of evolutionary biologists was directed to ways of trying to avoid war with the racist Nazis.  The League of Nations asked if “biology can end war?”, naïve in the view that war served some biological purpose. They thought that biologists still argued for some sort of “struggle for existence”, especially those who read Sir Arthur Keith’s regular articles in The Daily Mail, ostensibly about his subject, anthropology. He delighted his readers with his belief that Piltdown Man was a valid record of British supremacy and that it was important for humans to struggle to survive. Julian Huxley and many others tried to explain that there was no such scientific explanation, either of Piltdown Man or Nazi supremacy, but Keith held his ground against these scientists who he dismissed as left-wing extremists. It was a point well-made when Communist Party members such as the biologist Joseph Needham spoke publicly against Keith’s interpretations. “These men have deserted their heroes like Herbert Spencer and even Thomas Huxley” Keith argued, reminding his readers that in biology, both groups as well as individuals compete to be part of a healthy community: surely that validated war as part of the process of social evolution? Even Keith’s bitter enemy Haldane had urged in a lecture “that a proportion of mental defectives should be prevented from breeding”, while Sigmund Freud commented that violence was a character of the whole animal kingdom “from which men have no right to exclude themselves.”

These were still sensitive topics in those years when the Soviet experiment was just beginning and many biologists hoped that population might be controlled by eugenics. Some eugenicists even saw method in the madness of the First War that the weak were overtaken by the strong, but the large number of officers who died weakened their argument. By 1936 the support for voluntary sterilization of the less valuable members of society was on the wane and Pip Blacker, the Secretary of the Royal Zoological Society, thought it was all “a biological crisis unprecedented in the history of life”.