Some of the following people are well worth remembering for their great achievements, and the way they did them; others are not. But all were well known in their time and exercised undue influence. I have already described the parallel cases of Ernst Mayr and Huey Newton, while Bill Drury was the most important influence in my life but barely known to the outside world. All of the following people I knew personally. Let me start with the greatest.
All great minds have their unique style and Bill Hamilton was no different. While Huey Newton would blast you against the far wall with the force of his argument, you had to lean in to hear what Bill was saying, so soft was he spoken. It was almost as if he clutched his thoughts close to the chest, but the effort on your part was well worth it. His every thought on every topic was worth your close attention.
In 1969, William Hamilton came to Harvard to lecture. He was coming from a “Man and Beast” symposium at the Smithsonian in Washington where he had presented some of his latest thinking on spiteful behavior in an evolutionary context, which was the same talk he gave us. There were perhaps eighty or ninety people, almost filling the lecture hall, most of us with eager anticipation. Hamilton got up and gave one of the worst lectures I had ever heard in my life.
For one thing, he lectured for a full fifty-five minutes without yet getting to the point. It was abstruse and technical; he often had his back to us while he was writing things on the board; you had difficulty hearing his voice; you did not get any overview of where he was going or why he was going there. When he realized that he was five minutes over time and still had not gotten to the point—or indeed very near it—he looked down at Ed Wilson, his host, and asked if he could have some more time, perhaps an extra fifteen minutes. Of course Professor Wilson granted more time, but he also made a rolling, “let’s-try-to-speed-this-up” motion with his arms. Hamilton then called for slides. The room went dark, and there was a rumble and a roaring sound as about 90% of the audience took the opportunity to escape. I remember walking home with Ernst Mayr, both of us shaking our heads. It was obvious that the man was brilliant, a deep thinker, but whoa, was he bad in public.
Hamilton was not unaware of this problem. He once told a class we taught together at Harvard that after hearing him lecture many students would doubt that he understood even his own ideas.He did improve considerably in subsequent years, but he still showed the touch of a true master. He was invited to lecture to law professors in Squaw Valley, as a guest of the Institute for the Study of Law and Behavioral Sciences, and there he introduced a trick that I had never seen before. He showed a number of interesting but complex slides on parasite-host interactions. He had a hand-held microphone but no pointer, so he used the microphone as the pointer. Often all you would hear him say was, “Here, as you can see … ” and then the microphone would point to various parts of the slide, while his mouth continued to move. Then you might hear, “And then in the next slide … ” and once again you would not hear anything about the slide, though you could see Dr. Hamilton pointing animatedly to various places in it with his microphone while moving his lips.
I first met him at a party after his Harvard talk at Mary Jane West Eberhard’s home. I gave him a paper that was a very short version of the logic on reciprocal altruism, perhaps nine pages long. I had set up genes at two separate loci, so that kin effects would not be an obvious confound. He later complimented me on that but suggested my paper would be stronger without the “maths”, as gentle a way of saying that they added nothing of value (in fact, they added only a set of errors).
My first impression of Bill was that he was physically strong. I remember thinking that if the argument ever turned physical, the contest I would least like to be engaged in against him was a shoving contest. I felt that he would dig in his heels, you would be unable to move him, and he would lean forward and shove you slowly and stubbornly to wherever he wished to get you. Intellectually I imagine the interaction may have gone along similar lines.
I thought of Bill as perhaps the greatest evolutionary theorist since Darwin. Certainly, where social theory based on natural selection is concerned, he was our deepest and most original thinker. His first work in 1964—his theory of inclusive fitness—was his most important, because it is the only true advance since Darwin in our understanding of natural selection. Hamilton’s work is a natural and inevitable extension of Darwinian logic. In Darwin’s system, natural selection refers to individual differences in reproductive success in nature, where reproductive success is the number of surviving offspring produced. Hamilton enlarged the concept to include effects on other relatives—that is, not just fitness or reproductive success but inclusive fitness, defined (roughly) as an individual’s reproductive success plus effects on that of relatives, each devalued by the appropriate degree of relatedness (r).
This idea had been briefly advanced by R. A. Fisher and J. B. S. Haldane, but neither took it seriously and neither provided any kind of mathematical foundation. That foundation was not as obvious as it sounds. For a rare altruistic gene, it is clear that Br > C will give positive selection, where B is the benefit conferred, C the cost suffered and r the chance that a second copy of the altruistic gene is located in the recipient by direct descent from a common ancestor; but the matter is not so obvious at intermediate gene frequencies. As the altruistic gene spreads, should not the criterion for positive selection be relaxed?
Hamilton showed that the answer is ‘no’ and that his simple rule worked for all gene frequencies. He once told the story of sitting down as a doctoral student to write to Haldane, but to formulate each question more precisely he had to do additional work and after a couple of years he never sent the letter because he had by then worked out all the answers himself. A noteworthy implication of Hamilton’s work was that in almost all species the individual was no longer expected to have a unitary self-interest, because genetic elements are inherited according to different rules, a man’s Y chromosome going only to sons while the X goes only to his daughters, for example.
He soon followed this work with major advances in understanding selection acting on the sex ratio, senescence, the aggregation and dispersal of organisms, social insects, dimorphic males, and the origin of higher taxonomic units in insects. For the latter he argued that the more-or-less closed spaces created by rotting wood imposed a system of small, inbred subpopulations in insects inhabiting it, leading to a great diversity of homozygous forms, often with arbitrary, novel characters (such as a second complete metamorphosis in many male scale insects). In 1981 with Robert Axelrod he laid the mathematical foundation for the study of reciprocal altruism, when they showed that the simple rule of tit-for-tat in playing iterated games of Prisoner’s Dilemma was itself evolutionarily stable.
Thirty years ago Bill devoted himself to the theory that parasites play a key role in generating sexual reproduction in their hosts, recombination being a defense against very rapidly and antagonistically co-evolving parasites. In his memorable phrase, sexual species are “guilds of genotypes committed to free, fair exchange of biochemical technology for parasite exclusion”. He then argued that finding parasite-resistant genes must be a key function of mate choice and that in areas of high parasite abundance, species of birds evolved bright color and complex song to reveal such genes since it is hard to sing well or be brightly colored when ill.
It is hard to capture on paper the beauty of the man and the reason that so many evolutionists felt such a deep personal connection to him. He had the most subtle, multi-layered mind I have ever encountered. What he said often had double and even triple meanings so that, while the rest of us speak and think in single notes, he thought in chords. He was modest in style, with a warm sense of humor. His letters were laced with humorous asides. He once sent me a news clipping of a human father-to-son testicle transplant, along with the comment, “New vistas for parent-offspring conflict?”. The last time I saw Bill, at Oxford in December 1998, he pointed with pride to the two, and possibly three, species of moss growing on his Volvo—indeed on its front windows—and told me that this was a clear advantage of Oxford over Cambridge, the latter being too dry.
Certainly one of the most creative minds I have ever met in biology. I still remember the day a graduate student came running down the hall saying “Have you heard Hamilton thinks that bacteria use clouds for dispersal? As quick as you can say “Bill Hamilton”, I asked “Has he shown how the bacteria get the rain to fall where they want it to?” And indeed his idea humbled me because ever since I had been coming to Jamaica I had heard rural people tell me “trees draw rain” as in, don’t cut them down, and I had thought to myself you poor benighted souls, you have the correlation right but causality wrong—naturally, where it rains more, trees are more apt to grow. Now Bill suggested they Jamaicans may well have had it right all along—lower temperatures over wooded areas could itself be a useful signal.
Bill Hamilton was a naturalist of legendary knowledge, especially of insects, but he was also an acute observer of human behavior, right down to the minutiae of your own actions in his presence. Had I noticed, he asked, that lopsided facial expressions in humans are usually male? No, but I have seen it a hundred times since then. He was an evolutionist to the core, and was always heartened by news of fellow evolutionists enjoying some reproductive success. In a similar spirit I take joy in the lives of his three daughters, Helen, Ruth and Rowena, not to mention his many surviving siblings. But the loss of this ‘gentle giant’ is very great. Bill died at the age of 63 on 7 March 2000, from complications after contracting malaria during fieldwork in the Congo in January, work which was designed to locate more exactly the chimpanzee populations that donated HIV-1 to humans, as well as the mode of transmission. This was in service of a theory of HIV-1 spread into East African children via polio vaccination, one I regarded as doubtful from the outset and now firmly disproved, so in one sense, he died in service of trying to prove a falsehood, but he was strong in mind, body and spirit, with many new projects and thoughts under way, and he has been sorely missed ever since.
Bill chose to describe his preferred burial and its aftermath in biologically vivid and poetic terms. He would die in the Brazilian rain forest, his body to be scavenged by burying beetles so that he would later fly out as buzzing beetles “into the Brazilian wilderness between the stars”. But it was not to be. He died in the UK and was buried in Wytham at Oxford and it took the love of the second half of his wife, Luisa, to add her poetry, drawing on his bacterial/cloud/dispersal vision so that “eventually a drop of rain will join you to the flooded forest of the Amazon”.
I am no W.D. and my burial plan is very simple. If dead outside of Jamaica kindly cremate me—inexpensive and no place to point to. If in Jamaica dig a circular hole beneath my favorite large pimento tree, three feet wide and preferably ten feet deep and drop me head-first into the hole. Throw in some dirt and call it a day—no plaques please. I will not become a bright buzzing burrowing beetle or bacterial cloud, just a few more pimento berries when in season. I add the details on positioning my body mostly to annoy my Jamaican friends. They think I should be resting comfortably on my side in a conventional coffin, but if my way—why not standing up?—the strain on my neck upside down is too much for them to bear. I tell them all the nutritional goodness now is in my brain and upper body, hardly a thing of value is below my waist—they can trust me on that—so let’s go deepest with the best.
I first met S. J. Gould when he was a freshly minted Assistant Professor in Invertebrate Paleontology at Harvard and I a graduate student in evolutionary biology. Invertebrate Paleontology was well known then as a backwater in evolutionary biology, 80% devoted to the study of fossil foraminifera whose utility was that they predicted the presence of oil. In this environment, it was obvious that Gould would go far. New York City Jewish bright, verbiage pouring from his mouth at the slightest provocation, he would surely make a mark here.
This was not why I was visiting him. I had heard he was an expert in ‘allometry’—indeed had done his PhD thesis on the subject. Back then I wanted to know everything in biology, so I sought him out. Allometry refers to the way in which two variables are associated. It can be 1:1—the longer the fore-arm, the longer the total arm, or it can show deviations. For examples, the larger a mammal is, the more of its body consists of bone. Why? Because the strength of bone only goes up as the square of bone length whereas body weight goes up as the cube—thus larger bodies, weighing more, require relatively more bone. But what about antler size, I wanted to know, why is it that the larger the body size of the deer, the relatively larger his antlers? Why would natural selection favor that?
Gould leaned back in his chair. No, you have this all wrong, he said. This is an alternative to natural selection, not a cause of natural selection. My head spun. Natural selection was unable to change a simple allometric relationship regarding antler size that it had presumably created in the first place? Had it not already done so in adjusting bone size to body size? As I left his office, I said to myself, this fool thinks he is bigger than natural selection. Perhaps I should have said, bigger than Darwin, but I felt it as bigger than natural selection itself—surely Stephen was going for the gold!!
Many of us theoretical biologists who knew Stephen personally thought he was something of an intellectual fraud precisely because he had a talent for coining terms that promised more than they could deliver, while claiming exactly the opposite. One example was the notion of “punctuated equilibria”—which simply asserted that rates of (morphological) evolution were not constant, but varied over time, often with periods of long stasis interspersed with periods of rapid change. All of this was well known from the time of Darwin. The classic example were bats. They apparently evolved very quickly from small non-flying mammals (in perhaps less than 20 million years) but then stayed relatively unchanged once they reached the bat phenotype we are all familiar with today (about 50 million years ago). Nothing very surprising here, intermediate forms were apt to be neither very good classic mammals, nor good flying ones either, so natural selection pushed them rapidly through the relevant evolutionary space.
But Steve wanted to turn this into something grander, a justification for replacing natural selection (favoring individual reproductive success) with something called species selection. Since one could easily imagine that there was rapid turnover of species during periods of intense selection and morphological change, one might expect species selection to be more intense, while during the rest of the equilibrium stabilizing selection would rule throughout. But rate of species turnover has nothing to do with the traits within species—only with the relative frequency of species showing these traits. As would prove usual, Steve missed the larger interesting science by embracing a self-serving fantasy. Species selection today is a small but interesting topic in evolutionary theory, not some grand principle emerging from paleontological patterns.
Recently something brand new has emerged about Steve that is astonishing. In his own empirical work attacking others for biased data analysis in the service of political ideology—it is he who is guilty of the same bias in service of political ideology. What is worse—and more shocking—is that Steve’s errors are very extensive and the bias very serious. A careful reanalysis of one case shows that his target is unblemished while his own attack is biased in all the ways Gould attributes to his victim. His most celebrated book (The Mismeasure of Man) starts with a takedown of Samuel George Morton. Morton was a scientist in the early 19th Century who devoted himself to measuring the human cranium, especially the volume of the inside, a rough estimate of the size of the enclosed brain. He did so meticulously by pouring first seeds and then ball bearings into skulls until they were full and then pouring them out and measuring their volume in a graduated cylinder. He was a pure empiricist. He knew brain size was an important variable but very little about the details (indeed, we do not know much more today). He thought his data would bear on whether we were one species or several, but in any case he was busy creating a vast trove of true and useful facts.
I love these people—they work for the future and gather data whose logic later generations will reveal. Precisely because they have no axes to grind or hypotheses to prove, their data are apt to be more reliable than the first wave after a new theory. I have benefitted from them in my own life, most memorably when I was shown a large and accurate literature on ratios of investment in 20 ant species, gathered long before anyone appreciated why these facts might be of some considerable interest, as indeed they were.
In any case, Morton grouped his data by population according to best estimates of gross relatedness, Amerindians with Amerindians, Africans with Africans, Nordic Europeans with Nordic, and so on. It is here, Gould alleged, that all sorts of errors were made that supported preconceived notions that among the smaller cranial capacity (and therefore stupider)) peoples would be Amerindians and Africans. For example, Gould claimed that Morton made more subgroups among Nordic people than tropical ones, thus permitting more of them to be above norm, but in fact, the opposite was true. Morton reported more Amerindian subsamples than European and routinely pointed out when particular Amerindian subsamples were as high or higher than the European mean, facts that Gould claimed Morton hid.
In other cases, Gould eliminates all samples with less than four individuals in order to reduce the number of sub-samples with only one sex—a statistically meaningless goal but one that happened to be biased in his favor and permitted him to make additional errors in his favor by arbitrarily eliminating some skulls while including others. If you are comparing group means, you may not wish to use means of less than four, but if you are adding up sub-samples to produce a larger sample, there is no reason not to aggregate all data. Morton is made to look careless and incorrect when it is really Steve who is arbitrarily biasing things in his own favor.
There is an additional contrast between Morton and Gould worth noting. To conjure up Morton’s mistakes, Gould lovingly describes the action of unconscious bias at work: “Morton, measuring by seed, picks up a threateningly large black skull, fills it lightly and gives a few desultory shakes. Next, he takes a distressingly small Caucasian skull, shakes hard, and pushes mightily at the foramen magnum with his thumb. It is easily done, without conscious motivation; expectation is a powerful guide to action.” Indeed it is, but careful re-measures show that Morton never made this particular mistake—only three skulls were mis-measured as being larger than they were and these were all either Amerindian or African.
The same can’t be said of Gould. He came across distressingly objective data of Morton, and by introducing biased procedures (no sample size below four) he was able to get appropriately biased results. And by misrepresenting the frequency of Nordic vs Amerindian subpopulations, he was able to create an illusion of bias where none existed, by mere emphatic assertion (no one bothered to check).
Where are the unconscious processes at work here? Is Steve flying upside-down on auto-pilot, unconsciously making the choices (substitute Nordic for Tropical, delete all samples smaller than four) that will invite the results he wants while hiding his bias? Is the conscious organism really completely in the dark while all of this is going on? Hard to imagine—but at the end the organism appears to be in full self-deception mode—a blow-hard fraudulently imputing fraud, with righteous indignation, coupled with magnanimous forgiveness for the frailties of self-deception in others.
In response to the criticism of Lewis et al, the keeper of Gould’s Tomb—his longtime editor at Natural History, Richard Milner—had some choice comments in defense of Stephen. Gould acted with “complete conviction and integrity” (that is, with full self-deception). “He was a tireless crusader against racism in any form.” (In what way is misrepresenting the true facts about population differences—and then hiding this misrepresentation—a contribution to anti-racism?) And then, fully in flight, he says that any bias was “on the side of the angels”. Who of us is in any position to say what is on the side of the angels? We barely know what is in our own self-interest.
A general point is that it is often very hard to draw the line between conscious and unconscious deception—or to define the precise mixture of the two. Linguistic analysis in 2010 suggested that the architects of the U.S. 2003 war on Iraq were speaking deceptively when they warned that Saddam Hussein caused 9/11 and Iraq possessed WMDs. I naively thought that this analysis showed conscious deception (Trivers 2011) but I no longer agree with myself—unconscious deception could cause the same symptoms—reduced use of the word “I”, less qualifiers, and so on.
Professor Lewontin was quite the dominant figure in the 1970’s. He strode back and forth in front of audiences, hands on his suspenders, belly pushed forward, and expounded on the importance of “dynamical equations” in evolutionary thinking. None of us knew what a dynamical equation was, but we knew we sure better find out quickly. Although apparently expert in population genetics theory, his true move to fame was to ally himself with Dr Hubby so as to give—for the first time—what appeared to be a more or less unbiased measure of the frequency of heterozygosity in nature, that is, the degree to which the two sides of an organism’s genome (maternal and paternal) are different from each other. This is an important variable that had never been measured before. Inbreeding produces similarity, outbreeding heterozygosity (vide President Obama). Hubby and Lewontin’s conclusion was that there was a lot of heterozygosity in natural populations. This was an important finding.
I first heard him talk when he visited Harvard in 1969 to lecture on the new work. He gave a masterful talk, both in content and in style, and polished off the work with a series of slides showing evidence of selection along gradients in nature, acting similarly in several Drosophila species. But within five years he turned his back on natural selection and decided to emphasize the importance of random factors, which of course produced no patterns of particular interest, nor any insight into the function of genes and traits. This I believe he did for political grounds, emasculating his own discipline in order to render it sterile regarding human behavior and genetics.
In later years, doing less and less science, he spent more of his time on politics and philosophical writing whose meaning was difficult to locate, in part because there was often no meaning there. Consider the following obscurantist thought:
“Throughout the history of modern biology there has been a confusion between two basic questions about organisms: the problem of the origin of differences and the problem of the origin of state. At first sight these seem to be the same question, and taken in the right direction, they are. After all, if we could explain why each particular organism has its particular form, then we would have explained, pari passu, the differences between them. But the reverse is not true. A sufficient explanation of why two things are different may leave out everything needed to explain their nature.”
“If we could explain how each organism has evolved its particular form, by the selection of differences, then we would have explained, per stirpes, why it has its particular state. But the reverse is not true. A sufficient explanation of how an organism develops may tell us nothing about why it has its particular form.”
As for his political writing, nothing could beat a piece he wrote with Richard Levins stating that there was nothing in Marxist/Leninism that could be contradicted by objective reality. Wow, I thought, it is rare for people to fess up so quickly that there is no content to their enterprise, since if in principle it can’t be contradicted, it says nothing.
Lewontin’s story is that of a man with great talents who often wasted them on foolishness, on preening and showing off, on shallow political thinking and on useless philosophical rumination while limiting his genetic work by assumptions congenial to his politics. He ran a successful lab for many years, and easily raised large sums of research funds, so many U.S. geneticists remember him fondly for their time with him at Harvard, as a grad student or post-doc, but as an evolutionary thinker, never mind geneticist (beyond his early work on linkage disequilibrium), he has turned up mostly empty and the best of his ex-students concede he had done little of note for more than 20 years.
By the way, Lewontin would lie openly and admit to doing so. Lewontin would sometimes admit, in private at least, that some of his assertions were indeed fabrications, but he said the fight was ideological and political—they lied and so would he. On other matters, such as committee work, Lewontin could be rational and useful. Much less so, it was said was Stephen Gould, who was into self-promotion, self-inflation and self-deception full time. Not only was his science hopeless but so was much of his behavior in other contexts as well. I can remember him arguing against offering a full professorship to a truly excellent Colombian biologist because we would be discriminating against a third world country by depriving it of him. A heavy silence enveloped the room. Surely there was no brain/drain problem in evolutionary biology (Colombia to the United States) comparable to the one for nursing. Why not imagine the benefit he could confer on his own country sitting on a fat sum of Harvard money, linked to one of the best Museum collections in the world, with multiple opportunities to polish off his top scientists.
Professor Darlington was a revered and much feared personage along the hallways of the Museum of Comparative Zoology. He was the Curator of Coleoptera (beetles) the most speciose group in all animals and he was revered because he was the master of zoogeography, the distribution of all animal species through space and even time. And there were very interesting patterns. Life overwhelming and repeatedly differentiates in the tropics, spreads to the temperature zone, and a section onto the arctic but rarely the other way around. Salamanders are a temperate group that has colonized the tropics but they are the exception. The rule even holds for humans, repeated evolution out of the tropics, Africa, with minor reverse movements. And on more intimate scales, very few Canadians sneaking across the U.S. border but hordes of Mexicans and those from countries further toward the tropics.
In Darlington’s world of the 1950s the continents were stationary, but this soon gave way in the 1960s to moving tectonic plates and ‘continental drift’. For a moment, it looked like a lifetimes’ work, built on one assumption, would prove irrelevant on another, but it was not to be. Most of Darlington’s findings held whether the continents moved or not. Africa was the center of Gondwanaland, itself the center of Pangaea (when all the continents were combined) and the deeper inside the land mass, in general the more tropical. A corollary of Darlington’s discovery is that almost all groups are more speciose in the tropics, especially the smaller the individuals are, insects and then microbes.
We also feared him because he was a tall, lanky, dour, elderly character who did not invite easy banter. But there was one reason we all loved him. He had a pronounced limp on one side and he gained this, we were told, is the service of evolutionary biology. As the story went, he was walking along a rope ladder above a river in Indonesia when a crocodile leapt up and grabbed his leg and hauled him into the river. As is their style, a croc likes to pull you under water, whip you around and drown you. On his way down Darlington was alleged to have said to himself in righteous anger, “Wait a second, you don’t collect us as specimens, we collect you!” In any case, he managed to free himself and reach safety, although for the rest of his life he walked with a pronounced limp.
The last time I spoke with George Williams was in 2002 when I called about something and he told me he had pre-Alzheimer’s. There were simple memory tests now that were diagnostic, he said. In the background I could hear his wife Doris saying something and George said, “Doris always tells me not to tell people” and continued by saying that what he first noticed is that all words starting with capital letters were disappearing from his mind—arbitrary words for cities, buildings, people and so on.
A few months later, I sent my Selected Papers book but I never heard from him. He was gone. The person I felt for was Doris, a beautiful woman about half his size, and a very welcome complement to him. It is those closest to someone with Alzheimer’s who often suffer the most but George had a sweet disposition that, I hear, greatly reduced the cost to those closest to him.
We last saw each other when we were at the William Hamilton memorial session at Amherst in 2000 during the meetings of the Human Behavior and Evolution Society, at which both of us spoke. He was sitting behind me while Richard Dawkins was talking and I could hear Doris saying, “Now, George, don’t do what you are thinking of. Just tell the stories you have about Bill, don’t do it.” So I was full of anticipation when George got up because I knew he was surely going to do exactly what his wife thought was a bad idea. George gets up and says “I wish Bill were here today, because I have a bone to pick with him”.
And then he went and picked that bone for the entire talk. It had to do with the evolution of sex and patterns of evidence that George had pointed out years ago that contradicted (so George said) aspects of Bill’s parasite approach. I thought it was wonderful. There were those that said it was inappropriate and why didn’t he tell stories, but I thought it was perfect for the occasion, both vintage George Williams—no wasted motion with that organism!—and a tribute to the enduring importance of Bill’s ideas.
My first contact with George was when as a graduate student. I sent him my chapter then in press on “Parental investment and sexual selection”. When I wrote the paper I had completely forgotten that a key portion of the argumentation came right out of George’s 1966 book, Adaptation and Natural Selection. I had only relearned this when I reread his book in preparation for teaching my first course on social evolution. There were “sex role reversed” species (as well as female choice for genes and investment) and the relevant pages were full of underlining and marginal comments by me. None of this was acknowledged in the chapter I was sending him, so I pointed this out and said I would try to put some in before the book was printed. I was therefore feeling a little nervous when a letter came from George Williams. I braced myself for an unpleasant experience.
Instead, I found one of the warmest and most generous letters I have ever received. Among other things, he said my paper had rendered obsolete a chapter in his own forthcoming book on “Sex and Evolution”, namely the one on differential mortality by sex, which chapter he enclosed. He said nothing about not being properly cited but dealt only with scientific content. His chapter had my essential insight regarding male mortality—that higher variance in male reproductive success would often select for traits more costly in survival. The larger book was the first to systematically explore the consequences of seeing that sex usually has an immediate 50% cost in every generation (compared to asexuality) which cost has to be overcome in any successful model.
I invited him to Harvard in 1974 and he lectured on his ideas on sex. I do not say he was shy so much as reserved, but with a warm smile and sense of humor. My favorite joke of his occurred when George was telling me about the joys of grand-fatherhood. “If I could have figured out how to have grandchildren without having children first, I would have done so.” I knew just what he meant—high relatedness, no work, Or as Melvin Newton (Huey’s brother) once put it, “You can serve them ice cream for breakfast, what do you care?”
Having started with the evolution of senescence in 1957, in later life he tackled Darwinian Medicine, memorably saying that he did not think there was any compound—arsenic included—that was not beneficial if given in sufficiently small doses. This was almost surely an overstatement but a bracing and useful one. His knowledge of biology was so deep that he is the only person I know of to have predicted in advance the existence of an entire category of selfish genetic elements (genes that spread within an individual because they are advantageous to themselves, not the individual). Called ‘androgenesis’ it occurs when paternal genes eject maternal ones and take over the genome of an organism, a system now known from three very different groups of organisms.
He was a beautiful man, with a very simple and clear style of thinking, in a warm and humble personality. He was especially good at seeing through gibberish—group selection or psychoanalysis—and advancing carefully and slowly on major issues.