The Price of Altruism

by Oren Harman

  He was working on a model of an optimal mutation rate: It was “far superior to Kimura’s,” he wrote to the recent Nobel laureate MIT economist Paul Samuelson, with whom he’d resumed a short correspondence.23 But then came a surprise. In the spring a letter from John Maynard Smith arrived in the mail. In it were computer printouts of John’s latest attempts to model animal combat. To George it was practically a godsend.

  “Fascinating. Congratulations,” he replied, unable to hide his excitement, “it looks as though you’ve gotten well beyond the point I’ve reached.” Combat, after all, had always been close to his heart. Even better was the offer of joint authorship, though George would only accept if John’s name was listed first. At the moment he was trying to finish a paper of his own on “Sex and Rapid Evolution,” so wouldn’t have time to work on the combat paper just now; why didn’t John write a first draft for Nature that he could look over and comment on? As for the different behavioral strategies of the model’s animals in conflict, George suggested easily understood names like Dove, Hawk, and Prober.24

  Once again the problem was how to explain why, in fighting over dominance rights, territory, or mates, animals often seem to stop short of actually hurting each other. George’s original paper had concentrated on antlers in deer, but this was a general problem. The males of many snake species, for example, were known to fight each other by wrestling without using their fangs, a kind of “benevolence” not unknown even to praying mantises. The most absurd case was that of the Arabian oryx with the rhyming name, Oryx leucoryx: Its extremely long horns, pointed so absolutely in the wrong direction, forced males of this species to kneel down with their heads between their knees in order to direct their horns forward. How could kneeling oryxes, wrestling snakes, and deer that refused to strike “foul blows” be explained? How had Nature, in her wisdom so infinitely greater than man’s, invented limited combat?

  Maynard Smith and George took their cues from John von Neumann. For once again, as in poker, nuclear proliferation, or for that matter—Vietnam—animal conflict could be modeled as a game; the trick was to see that the strategy of each “player” depended on the other. Two contestants, for example, A and B, could adopt “conventional” tactics, C, that were unlikely to lead to injury or “dangerous” tactics, D, that were sure to lead to serious harm. And of course, they could simply retreat, R. A possible conflict between them could therefore look like this:

  A’s move

  CCCCCCCCCCCDCCCCCCCD

  B’s move

  CCCCCCCCCCCCDCCCCCCCR

  with A probing on the twelfth and twentieth moves, B retaliating after the first probe, then retreating, and losing out, after the second. Each contest ended with particular “payoffs” to each contestant, measures of the contribution the contest has made to the reproductive fitness of the individual. Spurred on by Maynard Smith, George returned to the computer in the summer of 1972. Together they programmed five distinct strategies, sets of rules that ascribe probabilities to the C, D, and R plays as a function of what happened in the previous moves. There were five such strategies: “Dove,” which never plays D; “Hawk,” which always plays D; “Bully,” which plays D if making the first move, D in response to C, C in response to D, and R following an opponent’s second D; “Retaliator,” which plays C on the first move, C in response to C, D in response to D, and R if the contest has lasted a preassigned number of moves; and finally “Prober-Retaliator,” which if making the first move or in response to C plays C with high probability and D with low (but R if the contest has lasted a preassigned number of moves), following a probe reverts to C if the opponent retaliates but takes advantage by continuing to play D if the opponent plays C. There were fifteen types of two-opponent contests, and John and George simulated two thousand contests of each using pseudo-random numbers generated by an algorithm to vary the contests. With fixed payoffs and probabilities that seemed biologically sound,25 the following matrix presented itself:

  Average Payoffs in Simulated Intraspecific Contests for Five Different Strategies

  To see whether a strategy is an ESS against all others, all one had to do is examine the corresponding column. In a population full of “Hawks,” for example, does “Hawk” do better than the alternatives?

  “Dove” “Hawk” “Bully” “Retaliator” The table showed clearly that it didn’t: Both “Dove” (19.5) and “Bully” (4.9) did better against “Hawk” than “Hawk” against itself (-19.5). “Dove” too was not an ESS: “Hawk” (80.0), “Bully” (80.0), and “Prober-Retaliator” (56.7) averaged higher payoffs in a population almost entirely of “Dove.” In fact “Retaliator” alone was an ESS, since no other strategy did better (though “Dove” ties), and “Prober-Retaliator” came in a close second.

  How would such a population be expected to evolve? The answer was that “Retaliator” and “Prober-Retaliator” types would increase in frequency at the expense of “Hawk,” “Dove,” and “Bully.” These last three types wouldn’t become extinct, though, but rather remain in low numbers due to a constant flow of mutation but also because of senility, youthful inexperience, injury, and disease. The balance between “Retaliator” and “Prober-Retaliator” would depend on the frequency of “Dove,” since probing was only an advantage against the meek: If the frequency of “Dove” was greater than 7 percent, “Prober-Retaliator” would replace “Retaliator” as the predominant type. The only way to make “total war” behavior advantageous would be to significantly alter probabilities for serious injury, or to give the same payoff penalty for retreating uninjured as for serious injury. Otherwise the simulations made abundantly clear that under individual selection “limited war” was superior to unbridled aggression.

  Of course real animal conflicts in nature were infinitely more complicated. Besides the category distinction between “Conventional” and “Dangerous” behavior, which in all probability was more subtly graded, individuals varied widely in the intensity and skill with which each kind of tactic was employed. Still, by simplifying, the model allowed for certain predictions: The table, for example, showed that the best strategy against “Hawk” is “Dove,” or in other words immediate retreat. This meant that it would be to the advantage of a given individual to simulate wild, uncontrollable rage, since those encountering him would do best to get out of his way and yield. If that was the case, a pretend “pseudo-Hawk” type would soon arise, leading in turn to selection for a second type with a special talent to “call a bluff.” To counteract this dynamic, a true sign of maniacal craziness would therefore be expected to evolve, one that could not be easily counterfeited. And this, it seemed, was precisely what sometimes happened in the wild. Elephants “on musth,” for example, were often described by local peoples as invaded by wild spirits: As they rampaged uncontrollably, often with fatal results, a dark brown tarlike fluid secreted by the temporal glands ran distinctively down their faces. Combating deviousness, this was nature’s way of signaling that here was no charade: With due respect to fakery, some things just couldn’t be bought by guile.

  These were important discoveries, George thought. Perhaps they’d prove useful to mankind.

  He was happy: not only had Maddox informed Maynard Smith that Nature was interested, but he’d have his name on a pathbreaking paper alongside a world-renowned authority. True, the insight had been his. But this was an infinitely better outcome than his first, original attempt alone.

  As exciting as it was, however, George’s heart remained elsewhere. He had no plans, he wrote to Kathy, to renew the SRC grant that was running out in July. Instead he was planning on spending the next few months working on a book about Shakespeare and the Bible.26 Most of all, he was finally giving in to fate.

  Recently he had preached a sermon on Matthew 6:25–34:

  Therefore I say unto you, Take no thought for your life, what ye shall eat, or what ye shall drink; nor yet for your body, what ye shall put on. Is not the life more than meat, and the body than raiment? Behold the fowls of
the air: for they sow not, neither do they reap, nor gather into barns; yet your heavenly Father feedeth them. Are ye not much better than they? Which of you by taking thought can add one cubit unto his stature? And why take ye thought for raiment? Consider the lilies of the field, how they grow; they toil not, neither do they spin…. Wherefore, if God so clothe the grass of the field, which to day is, and to morrow is cast into the oven, shall he not much more clothe you, O ye of little faith? Therefore take no thought, saying, What shall we eat? or, What shall we drink? or, Wherewithal shall we be clothed?…But seek ye first the kingdom of God, and his righteousness; and all these things shall be added unto you. Take therefore no thought for the morrow: for the morrow shall take thought for the things of itself.

  He couldn’t stop thinking: He held an ordinary job, lived in an ordinary flat, dressed conventionally, was paid a regular salary. He’d sort of known, then, without really knowing it, that all this would have to end. “Wishful thinking had kept me supposing that it was not going to happen to me,” he wrote to Maynard Smith in October,

  —or at least not in the extreme way that it seems always to happen in the accounts I had read by missionaries and others who had themselves lived “the life of faith.” In those accounts the saving cheque always arrives at the last possible moment when disaster is at hand.27

  “I had optimistically calculated that deliverance had to arrive around the 20th of September in order to avert disaster. However it appears that God’s standard of what constitutes ‘disaster’ are on a different scale from mine,” he continued.

  Furthermore it appears that His standards are more accurate than mine, for in fact here I am almost a month later, still with food and other necessities and with all essential accounts paid. I don’t know how much longer this is going to go on. The encouraging part is that I am now down to exactly 15p and my visitors permit for staying in the UK expires in less than a month. Thus I reassure myself by telling myself that God’s standards of disaster will shortly be satisfied. I look forward eagerly to when the 15p will be gone.28

  John was very worried. “I have less faith than you do that the Lord will provide. Please let me know at once if I can help.” Hamilton too was urging his American friend to renew his fellowship. If George got evicted he was more than welcome at Berkshire. “If there is an Almighty I think you mistake his ways!”29

  But neither Maynard Smith nor Hamilton understood the extent of the test George had put himself to, for it was far from just a matter of money. “It is now more than nine months since I last saw a doctor,” George wrote proudly to a friend back in May the previous year; when he was feeling unwell he’d go to pray at the London Healing Mission. Soon after his conversion he had stopped his periodic thyroid checkups. After all, the coincidences, the penetration of the Bible, the equation—all were signs that he’d been chosen for some task. If the Lord so wanted, the Lord would provide. By now it had been two years since he’d been to a doctor. Pushing fate, he’d begun to eat just as little as necessary and had survived for the last week on barely a pint of milk a day. But there was something worse, darkly more ominous: George had completely stopped taking his thyroid medicine.30

  It was a neighbor from Little Titchfield, Mr. Wood, who found him, collapsed, in the stairway and rushed him to the hospital teetering on the brink of unconsciousness. Luckily an alert physician at the Middlesex ICU, Dr. Webb, noticed the deficiency in his thyroxine, and administered it intravenously while George lay asleep in bed. In most probability this was what saved him: With no corrective to his lack of thyroid hormone, his energy uptake and heart rate had reached such levels as to put his life in grave danger. George had been a heartbeat away from dying.

  It was Christmas 1972, and he was recovering in the ward. His eyes were sore from the atropine, and when a patch was removed from one of them, it took awhile before he could stop seeing double. But most troubling at the moment was his imminently expiring visa. “If you say that I entered Middlesex Hospital in a state of ‘starvation,’” he subsequently wrote to Dr. Webb, whose explanations would be necessary, “they are likely to say, ‘Hah, he ran out of money, one of those American beatniks.’” George preferred that Dr. Webb write that he arrived in “a state of extreme bodily weakness” instead of “malnutrition.” Also, he thought that it would be better not to mention “myxedema”: Some immigration officer might know what that meant and wonder why he’d stopped taking his thyroid medication. Then he turned to the bigger picture: The reason he hadn’t renewed his grant, he’d explain, was because he was in the process of switching his career “from science to Christianity.”31

  Finally, in January, release papers in hand, he was helped into a cab by Mr. Wood and was back in Little Titchfield. Weakened and shaky, he was doing his best to nurse himself back to health. “What is it that one does wrong in cooking rice that makes it stick together?” he wrote to Annamarie in sunny California.

  Sometimes my rice sticks together and other times it doesn’t. I don’t cook it very often, so that it is difficult to remember how I did it when it worked well. The last time I cooked it it didn’t come out at all well. The time before it was very good. Now maybe the difference was that it was a different brand last time, since I was opening a new package. Or maybe the trouble was that I didn’t follow directions correctly. It said on the package to bring it to a boil, stir once, and turn it down immediately to a simmer. But mine boiled for a little while before I turned it down. Could that have caused the trouble? It stuck not only to itself, but also really stuck to the pan. So please tell me what are the critical points that make the difference between sticky and non-sticky rice?32

  He was fifty years old. Miraculously he’d survived the Lord’s test. But more than ever now he grasped what seemed to have escaped him for quite some time. He was pale and thin, his fingernails brittle and blackening. His brand of strict Christianity had obviously gotten him nowhere. In this world at least, obedience to God notwithstanding, George was utterly alone.

  Hamilton’s rule, rB>C, was good for the family or at most, after George’s correction, to those who shared similar genes. But what about altruism between nongenetically related organisms? This, after all, had been the moralist’s concern throughout the ages, sacrifice among relations being a lesser riddle to spin. Recognizing the problem, the Judeo-Christian tradition exhorted its followers to “Love thy neighbor as thyself,” but Aristotle was more of a cynic. “The friendly feelings that we bear for another,” he wrote in his Nichomachean Ethics, “have arisen from the friendly feelings that we bear for ourselves.”33

  Pagan thought was not the only hardened philosophy on the moralism block: Saint Thomas Aquinas took a surprisingly utilitarian position, arguing in the Summa Theologia that we should love ourselves more than our neighbors. His interpretation of the Pauline phrase was that we should seek the common good more than the private good since the common good is a more desirable good for the individual. David Hume, in A Treatise of Human Nature, may have spelled out at the age of twenty-six what the venerable medieval theologian was thinking: “I learn to do service to another, without bearing him any real kindness: because I foresee, that he will return my service, in expectation of another of the same kind, and in order to maintain the same correspondence of good offices with me or others.”34

  In 1776, the year of Hume’s death, his Edinburgh neighbor the economist Adam Smith put it even more directly in The Wealth of Nations: “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. Nobody but the beggar chooses to depend chiefly on the benevolence of his fellow citizens.” Most succinct of all, however, was the Dutch-born English satirist Bernard de Mandeville, who in a couplet from 1714 wrote: “Thus every Part was full of Vice, / Yet the whole Mass a Paradise.”35

  And so when a precocious so
n of Lithuanian Jewish immigrants started his education at Harvard in the early 1960s, he had quite a philosophical tradition to build on. But Robert Trivers was not interested in biology—he wanted to be a lawyer—and it would take a breakdown (his mania took the form of staying up all night, night after night, reading Wittgenstein and finally collapsing) to bring him closer to the animal world. Diagnosed with schizophrenia, Trivers took an illustrating course while recovering, and was hired to draw animals for a biology textbook. His mentor was Bill Drury, an Audubon ornithologist whom he learned to love and revere. “Bill and I were walking in the woods one day,” Trivers once recounted to a reporter, “and I told him that my first breakdown had been so painful that I had resolved that if I ever felt another one coming on, I would kill myself. Lately, however, I had changed my mind, and drawn up a list of 10 people I would kill first in that event. I wanted to know if this was going forwards or backwards. He thought for a while, then he said ‘Can I add three names to that list?’ That was his only comment.”36