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Authors: Oren Harman

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John Maynard Smith (1920–2004)

 

 

William D. Hamilton (1936–2000)

 
Solutions
 

J
BS was in a London hospital bed. Rectal carcinoma, the doctors said. He was back from India to get the very best treatment available, though the “auto-obituary” he taped, sitting draped in a sari, wasn’t a very good sign. Haldane was about to leave this earth, and he knew it. Meanwhile he asked his student John Maynard Smith to go out to the bookshop to get him something to read.
1

It wasn’t easy for Maynard Smith to see his mentor in such a state. He was born in 1920 in Wimple Street, London, and childhood had been a lonely affair. He was eight when his stern and distant ex-military physician father died; an absent mother provided little comfort beyond the winter home in Berkshire and summer home in Exmoor, the means for which her wealthy Edinburgh stockbroker family could afford. Endless hours watching birds in the countryside confirmed both his love for nature and his isolation.

At Eton, by way of lore, he soon learned that there was one graduate who had attracted the particular hatred of some of his teachers by betraying his class and religion. There was a noxious blend of privilege and prejudice at the school, but to its credit

153

J. B. S. Haldane’s writings were in the library, and seeking them out, John was captivated. Haldane’s blend of atheism and reason, he thought, “never left you wallowing in a sense of misty profundity.” Almost naturally, scientific and political commitments blended: John requested
Capital
for a school prize he had won, delved into mathematics, and made peace with the absence of God in his life. He would be a “puzzle-solver,” he hoped, and a socialist. Shirking his maternal grandfather’s wish that he join the family’s stockbroker firm, he read engineering at Trinity College, Cambridge, and joined the Communist Party in 1939.
2

After spending the war years making stress calculations for Miles Aircraft near Reading, John was ready for a change. Poor eyesight meant he’d never be able to fly the planes he designed, so he could never really love them either. As for politics, it was either that or science; one’s heart could be in two places, perhaps, but the brain was more demanding. Since theoretical physics seemed too difficult and chemistry a chore, it was to biology and his old love for nature that Maynard Smith now turned. When he discovered in October 1947 that his hero Haldane was professor of genetics at UCL, he applied forthwith. “Dear Comrade,” his letter began,

My interest is mainly in evolution and genetics. My main existing qualification is that I am a competent mathematician, and in so far as I have shown any ability as an engineer, it has been in expressing physical problems in terms of mathematics. I read Huxley’s “Evolution: the New Synthesis,” and there seemed to be plenty of scope for a mathematical approach to the subject of natural selection, the origin of species, and so on.
3

 

Whether he was in a good mood that day or genuinely impressed, JBS fired back a letter of acceptance. It proved a smart decision: As an aircraft engineer John had learned to trust models and the necessary simplifications they demanded. After all, if RAF fighter planes could stay in the sky even though dry calculations on land assumed incompressible air (refuted by the pneumatic tires and inflated life jackets), simplifying assumptions could be valuable.
4
Of course, JBS had known this ever since he set foot in biology.

Maynard Smith made his way to the bookshop. Please don’t die, he thought. Not yet.

 

 

He returned from Dillon’s with a big fat book,
Animal Dispersion in Relation to Social Behaviour
, by an author with a long name. The son of the headmaster of Leeds Grammar School, who retired to become rector of the picturesque country parish of Kirkland in the Vale of York, Vero Cope Wynne-Edwards had grown up chasing rabbits in the Pennine hills. At Rugby his buddies called him “Wynne” and together, he’d remember, they “collected plants and Lepidoptera, found birds’ nests, hunted for fossils in the local cement pits, ‘fished’ in ponds for aquatic life, made drawings of ‘scratch dials’ on medieval church walls, and ‘excavated’ for pottery in a Roman camp on Watling Street.”
5
By the time he arrived as an undergraduate at New College, Oxford, in 1924, his heart was set on zoology. His teachers were giants: E. S. Goodrich for comparative anatomy, Gavin de Beer for experimental embryology, E. B. Ford for genetics, Julian Huxley for general zoology, Charles Elton for ecology. When Huxley left for King’s College, London, in 1925, it was Elton who loomed large in his education. Elton was now systematically trapping voles and wood mice in Bagley Wood near Oxford, trying to figure out their dynamics. Does the size of wild populations of animals fluctuate in a periodic cycle, he wanted to know, and if so, what was the cause?
6

Elton saw that size did fluctuate, but he never solved the mystery: Was it due to weather, the food supply, migration, predator-prey oscillations? No one seemed to know. It was just around then that he gave Wynne-Edwards a book to read,
The Population Problem
. Written in 1922 by a former student of Huxley’s who would go on to become the director of the London School of Economics and be knighted for public service, it made a revolutionary claim. Contrary to Malthus, Alexander Carr-Saunders argued, humans could do without disaster. Neither plagues nor war nor famine nor any form of “natural corrective” was necessary to provide the perfect fit between what the world can feed and the number of hungry mouths. When there was plenty, man procreated generously; when there was dearth, he procreated less. Neither a slave to the elements nor a victim of the earth, he was perfectly attuned to his environment. This much evolution had taught us: The primitive tribes that survived into modernity were exercising population control. And density was always at its optimum.
7

Forty years later, when Wynne-Edwards was writing the book Maynard Smith now brought to the dying Haldane, he suddenly understood. He was a professor at Aberdeen, an Englishman with a Welsh name who had lived half his life in Scotland. Like Kropotkin, he had made expeditions to northern lands, where the harsh elements had driven animals to cooperate.
8
If competition existed in nature, the Arctic taught him, it was directed at the environment, and animals had developed a myriad of social mechanisms to cope. Already in 1937, on the Baffin Islands’ coastline, he observed that only between one-third and two-fifths of the fulmars in the breeding colony mated while the rest were pushed into marginal territories and often died.
9
What a clever way to prevent overexploitation! Even more ingenious was the chorus of singing accompanying each breeding season: Short of using a calculator, it was the best way for the flock to assess its size and, surveying its resources, reproduce accordingly. Now, in 1962, as he sat down to explain such phenomena after years of thought, he fathomed his debt to Carr-Saunders. Birds, just like primitive man, were regulating their numbers.
10

It was an idea, some thought, that flew smack in the face of Darwinism. Man might exercise birth control, but birds? Surely their brains were no match for the inexorable natural imperative to procreate, the ultimate arbiter of the survival of the strong. In the 1930s Julian Huxley had sent another of his students to the tropics to study Darwin’s old finches. What David Lack saw there was that competition for food was rampant, but that slight differences between geographically isolated groups might reduce it.
11
Each group of finches specializing in a particular food in a particular habitat made for a wonderful mechanism to get out of the others’ way; gradually, genetically, the populations became distinct. Nature, in other words, would go to great lengths to avert conflict, even to the end of divining new species. But as powerful as the force of competition, more powerful—since more fundamental—was the instinct to procreate. Individuals were out to maximize their fitness, to sire just as much as they could. The idea of altruistic birds passing up a number for the greater good made absolutely no sense.
12

Unless, of course, natural selection was operating on the group, which was exactly what Wynne-Edwards was arguing. Drawing on Wright’s model of group selection, Wheeler’s superorganism, Emerson’s homeostasis, and Allee’s fowl hierarchies, he made a nonmathematical case for the collective. Individuality was important but subordinate: When the physiology of the singleton came up against the “viability and survival of the stock or race as a whole,” group selection was bound to be the victor and individual reproductive restraint the result. It was just like with fishing: If every fisherman set his net to catch just as many fish as he could, the village folk would quickly find themselves “entering a spiral of diminishing returns.” If, however, an agreement was reached over the maximum catch for each, depletion of the villagers’ maritime food source could be happily avoided.
13
As with fishermen and their catch, so with fulmars (and red grouse and many other birds) and their environment: To prevent exhausting limited resources, numbers could be regulated by social convention for the benefit of all.
14

Wynne-Edwards was certain that he was walking in the path of a giant. Darwin had translated Malthus back into nature as he had translated Carr-Saunders. Darwin had used the analogy of artificial selection to explain natural selection as he had used fishing to explain population regulation. But Darwin had also written:

Whatever the cause may be of each slight difference in the off spring from their parents—and a cause for each must exist—it is the steady accumulation, through natural selection, of such differences,
when beneficial to the individual,
that gives rise to all the more modifications of structure, by which the innumerable beings on the face of this earth are enabled to struggle with each other, and the best adapted to survive.
15

 

Now Wynne-Edwards felt, after more than a century, that he’d understood what even the great master had failed to fathom: Adaptations work for the good of populations, not of individuals.

On his deathbed JBS just chuckled. “Smith, do you know what this book says?” he asked his devoted student, with his usual mischievous air:

Well, there are these blackcock, you see, and the males are all strutting around, and every so often, a female comes along, and one of them mates with her. And they’ve got this stick, and every time they mate with a female, they cut a little notch in it. And when they’ve cut twelve notches, if another female comes along, they say, “Now, ladies, enough is enough!”
16

 

When it became clear that the best medicine in the world wouldn’t save him, Haldane returned to India. Clearly he hadn’t been impressed by the biological argument for the greater good, a fact that did not stop him from dying a devoted Marxist on December 1, 1964.

 

 

Maynard Smith was less of a mule than his teacher. The Lysenko affair, the purges, and the invasion of Hungary in the fall of 1956 had been enough for him. Without giving up the sentiment, he gave up the Party, increasingly turning to model evolution and nature. It wasn’t easy: “Why do theory when Haldane is sitting in the room next door?”
17
In the beginning he stuck to fly genetics. But gradually, with Haldane’s move to India in 1957, John’s confidence had grown. He was ready to take on evolution on his own.

John hadn’t read Wynne-Edwards’s book before he brought it to Haldane, but now decided to pay attention. Reviews of the tome had been mixed. Lack, of course, hated it. So did Wynne-Edwards’s beloved teacher Elton who judged it “messianic” and “rather wooly.”
18
Many, however, found the notion of a “balance of nature” plausible, and, more importantly, deeply relevant to man. Wynne-Edwards himself egged them along. That summer he had written an article for
Scientific American
that began: “In population growth the human species is conspicuously out of line with the rest of the animal kingdom.”
19
Man was virtually alone in showing a long-term upward trend in numbers. It was a bright red warning sign: However highly he thought of himself, compared to fulmars and red grouse his social skills were retarded. Modern individual freedom, alas, had trampled tribal homeostatic wisdom.

Just like Darwin, he was trying to bring animals and man closer together; he had shown convincingly, an anonymous reviewer in the
Times Literary Supplement
wrote, that “social life in man…is no unique affair, but the culmination of a very widespread biological phenomenon.”
20
The trouble was that they were drifting apart. This much, at least, was clear to the anxious reviewer in
The Nation
:

In Wynne-Edwards’ proofs we can see reflected the breakdown of relations between parents and children, the male’s and female’s diminished attachment, the constant migration of peoples, the female’s objection to being just a breeder, the male’s resentment of being just a provider, smaller families, divorces, desertions, minorities escaping from “ghettos,” elites struggling to keep out the invaders, the increase of homosexuality and neuroticism, alcohol and drugs, and above all, the evidence that young people, the group most sensitive to social stress, desire violence, especially if it is unprofitable and senseless: in all this we see that human society is reacting just as Wynne-Edwards says a crowded society should. It is giving a warning which nobody heeds; even when they see the Sunday cars jamming the highways, as in a dance of gnats, or a swarming of locusts.
21

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