Harman Oren Solomon. The Price of Altruism: George Price and the Search for the Origins of Kindness

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Williams himself emerged as a pioneer late in his career.

Slowly but surely

further examples began to surface.

And while group selection remains hotly

contested, and good experimental examples are still few and far between, there’s

no arguing that it’s making a comeback, at the very least in theory.

Today even

the entomologist Edward O. Wilson, whose 1975 book Sociobiology was a central

pillar of the gene’s-eye-view approach, has come to believe that selection works

at different levels. “Superorganisms” like bee and ant communities are literally

giant vehicles, and the altruism they foster, Allee and Emerson would have been

glad to know, is in that sense genuine and real.

The possibility of selection at many levels, of course, makes the covariance

equation all the more important. And so, drawn from the closet, dusted, and

sparkled, the “Price equation,” as it has become known, is increasingly gaining

new life. Theorists, among them notably Steven Frank, Alan Grafen, and the

philosopher Samir Okasha, are finally joining in Hamilton’s enchantment,

showing just how powerful a tool George invented and building further on its

foundations.

Ben Kerr and Peter Godfrey-Smith have recently even contributed

a small improvent, adding a second covariance on the right hand side of the

equation to make the causal structure more symmetrical and therefore more

general, a development Price himself would undoubtedly very much have liked.

Still, like any tautology, it’s sometimes unclear whether the Price equation is

valuably invaluable or invaluably valuable: to really explain the natural world, it’s

clear, the pristine mathematical beauty of the equation needs to be filled with the

messiness of biology.

But if the Price equation is better for framing a problem

than for figuring out its messy biological details, it remains a crucial tool for

understanding evolution in structured populations, where interdependence is just

as important as either dependence or independence. Its generality means that it

can be applied to evolution not only of genes but also to other forms of nongenetic

evolution, based on epigenetic or behavioral inheritance. Alongside its ability to

represent selection working on different levels simultaneously, this is a major, if

still untapped, attribute. Besides altruism, the Price equation has already proved

useful in attacking problems as diverse as reproductive value, evolutionary

epidemiology, genetic programs, human cultural evolution, rogue genes, meiotic

drive, biodiversity, and ecology. And the more we think of it the more it stands a

chance to help us understand many other problems.

Hindsight is a wise master. Having thrown down the rope of game theory, it’s

now apparent, George Price helped others scale Mount Modern-Evolutionary-

Biology with a stamina and strength that have proved enduring. But even more

important, by providing the penetrating spectacles of covariance through which to

measure all other ropes, he has helped us see more clearly than ever before that we

are all climbing the very same mountain. It’s a legacy that will continue to

transform the view of what Darwin called the “entangled bank,” a dramatically

exciting window into the complexity and grandeur of life.

They could hardly have imagined it when they met behind the curtain at the

Belasco Theatre, but William Edison “Isak” Price and Alice “Clara” Avery would

have much to be proud of in their son.

What about altruism in humans: Is any of this relevant to us?

Return to the hamadryas baboons playing a prisoner’s dilemma. Imagine first that

each baboon is either altruistic or selfish by no fault of its own: Each is born the

way it is, and can no more change its nature than a vegetable become a fruit. But

imagine further that each baboon has both capacities and can begin to choose

whether it wants to be a Good Samaritan or a cheat. Suddenly we are in a

completely new game: We have entered the world of mind.

It was Trivers who saw where this was heading. If cooperation and ultimately

altruism are games of reciprocal good faith, what better way to police the system

than to be able to detect deceit? Benevolence, he reasoned, requires a strong sense

of justice, because a sense of justice is necessary in order to appreciate dishonesty.

Over evolutionary time, and beginning with animals, a genuine instinct of fairness

would have been born out of the need to distinguish the trustworthy from the

swindlers. This has been demonstrated in recent experiments with capuchin

monkeys: Experimenters rewarded monkeys with sweet grapes when the monkey

handed them a stone, but when the monkeys were divided into two groups, one of

which was handed grapes in return for the rocks and the other much less attractive

cucumbers, many of the monkeys in the second group would not accept the offer.

In fact the sense of unfairness could even be magnified: When, in return for their

rocky offering monkeys were given absolutely nothing, some began pelting the

experimenters with stones in outrage. Both our human sense of right and wrong

and our discretion over when and where to use it are, it seems, legacies of natural

selection.

This may help to explain why we are so good at detecting cheats. A number of

neat experimental designs suggest that this betrays a human cognitive bias: We

are better at solving logical riddles that have to do with exposing shysters in

real-life situations than we are solving the very same puzzles presented in the

abstract. Whether detecting cheats also requires a system of punishment is a

question that is argued. While some find punishment requisite for cooperation,

others claim that it’s effective, but only for establishing dominance. True

egalitarian cooperation, on the other hand, is best served by withholding

recrimination: At least when it comes to peers (as opposed to the state), taking the

law into your own hands is unprofitable.

Whether punishment is necessary or not, it’s clear that people often cooperate

altruistically even when they have absolutely nothing to gain: The classical

economic assumptions of self-interest cannot explain the richness of human

behavior, Adam Smith’s long shadow notwithstanding.

This may not be all that

surprising if one considers how valuable regard, and its subtler cousin,

self-regard, is to human beings. But why should this be so?

Following in Trivers’s footsteps, the economist Robert Frank proposed that it all

comes down to trust. To do well in life, our simian and hunter-gatherer ancestors

needed to forsake the short-term temptation of self-interest, since they lived in

small, mutually dependent groups. Emotions, or what used to be called “moral

sentiments,” helped to ease the retreat from egoism by establishing commitment

(“I love you and won’t leave you”) and eliciting reciprocity (“I love you, love me

back!”). Since in evolutionary terms life is a game in which one seeks trusted

partners to play with, emotions that trigger loyalty and altruism (even irrational

ones) are adaptations for regulating behavior. According to this view we have a

genuine conscience underlying our emotional repertoire in order to help us join

hands with other conscientious sorts to everyone’s mutual advantage.

But the logic is tricky. The possibility of deceit raises the probability of ever more

subtle mechanisms for spotting deceit, which raises the probability for

mechanisms of self-deceit. Trick yourself to trick another; what better way to

conceal true intentions? This in fact seems rampant in unsuspecting man: Ask a

hundred people whether they are above average in intelligence, or beauty or

kindness or virtue, and considerably more than fifty, hand on heart, will tell you

that they are. Still, if an arms race was put in place in evolution between cheaters

and detectors, perhaps the result has ultimately been felicitous: After all, the best

way to seem trustworthy and giving is to actually be trustworthy and giving.

Which is it, then: pure genuine selflessness or an egoism so cunning that even the

trickster is tricked? Some argue that the genuineness of goodness can be

demonstrated scientifically, though their arguments seem really quite feeble. The

strongest case that can be made for pure unsullied selflessness is that it is just as

plausible as unadulterated egoism. Proponents call this “pluralism,” but in truth it

is more of an admonition: There are questions that, however much we’d like it to,

science simply cannot solve.

Whether goodness is genuine or not, it was the father of the theory of evolution by

natural selection who instructed that the moral sense has natural foundations. “Mr

J.S. Mill speaks, in his celebrated work, ‘Utilitarianism,’” Darwin wrote in The

Descent of Man in 1871,

of the social feelings as a “powerful natural sentiment,” and as “the natural basis

of sentiment for utilitarian morality.”…But…he also remarks, “…the moral

feelings are not innate, but acquired.” It is with hesitation that I venture to differ

from so profound a thinker, but it can hardly be disputed that the social feelings

are instinctive or innate in the lower animals; and why should they not be so in

man?…[Several thinkers] believe that the moral sense is acquired by each

individual during its lifetime. On the general theory of evolution this is at least

extremely improbable. The ignoring of all mental qualities will, as it seems to me,

be hereafter judged as a most serious blemish in the works of Mr. Mill.

In private, rhetorical deference could duly be abandoned: “He who understands

baboon,” Darwin scribbled in his personal notebook years before, “would do

more towards metaphysics than Locke.” The mind had been crafted by millions of

years of evolution. If man wanted to comprehend his own morality, he’d need to

take a careful look back at the animals.

Everyone says humans are special, and, undoubtedly, this is so. Even if each and

every species is idiosyncratic, if uniqueness is really not unique, however we like

to spin it, our culture does set us apart.

Hamilton saw that this meant that group

selection might play an even more central role in humans: The kind of

cohesiveness borne by living in small hunter-gatherer groups and developing

language might help solve the “free-rider” problem and would only have been

strengthened by religion and lore. This left him cold: Group selection bumping

whatever the opposite of altruism is to the next level was nothing to gain any

solace from. For in order for true goodness to be born, a ruthless rival to the group

was mandatory. This would both foster xenophobia between groups and

necessitate a closing of the ranks within, a realization that led Trivers to dub

Hamilton’s 1975 study his “Fascist paper.”

It’s a sobering thought, but it needn’t always be true. A recent study in rats

showed that the more a rat benefits from the altruism of a stranger rat, the more he

will later act benevolently towards stranger rats himself.

This happened in a

laboratory, in rats that were trained by humans to pull a lever that released food

for a stranger rat in another cage—perhaps not the best approximation of what

happens in nature. But we humans constantly find ourselves in situations where

complete strangers need our help. In fact the notion that as egalitarian morality

spreads through education and culture to an ever-greater number of people, the

costs of altruism are reduced may mean that we have developed a unique capacity

for altruism. It’s true: History continues to serve up a litany of factious wars over

religious and political influence.

But this doesn’t mean things can’t change.

Cheap airfare, integrated markets, the Worldwide Web, and the extinction rate of

languages are just a few of the harbingers of our increasing interdependence. And

even if globalism fails or infinitely worse—the promise of universal tranquillity

crashes—something else may end up uniting us. A classic Hardin “tragedy of the

commons,” the threat of a degrading environment, could graduate mankind to a

higher level. And if Earth can serve as a genuine “vehicle,” then the environment

will be a proxy for a ruthless enemy. Short of recognizing the inherent goodness

in universality and peace, coming together to protect our planet may prove a

saving grace for a still-fractured humanity. In the end it may even lead to an

expansion of our circle of morality to other creatures besides ourselves.

These are grand affairs. What direction history takes is known only to prophets

and fools, and from the dawn of time distinguishing between the two has been

impossible. Increasingly, however, science is training its gaze inward, to the

brain, in search of clues to the origin of altruism. The neuroscientists Jorge Moll

and Jordan Grafman placed volunteers in a functional MRI unit and asked them to

imagine either donating a sum of money to charity or keeping it to themselves.

They found that thoughts of generosity activated a primitive part of the brain that

lights up with reference to food and to sex, a result interpreted by them to mean

that altruism is as hard wired as our most basic functions.

The psychologist

Marc Hauser agrees. His research shows that all over the world people process

ethical dilemmas in very similar ways, suggesting that the capacity for moral

judgment, like language, should be thought of as intrinsic to our nature rather than

a product of our culture. It is a well-documented fact that injuries to particular

parts of the brain result in patients who seem to have lost their moral sense:

Perfectly intelligent otherwise, they remain unable to tell right from wrong.

But science is going further. In the most reductionist attempts to date to locate a

“moral sense,” researchers have begun searching for genes for goodness: A team

led by the psychologist Ariel Knafo, the geneticist Richard Ebstein, and their

graduate student Salomon Israel gave fifty shekels to two hundred students and

asked each of them to play what is called a dictator game: The person with the

money, the “dictator,” must make a one-shot, nonreciprocated decision on how

much of his pot to share with an anonymous “recipient.” Not surprisingly there is

quite a bit of variation: some split their loot judiciously down the middle, others

selfishly keep it to themselves, still others give most of the money away. What the

researchers found when they genotyped the students was that there is a significant

correlation between the degree of giving and the length of a tiny DNA

microsatellite in and around a gene encoding for a receptor of the neurotransmitter

oxytocin in the brain. The same was true for a related neurotransmitter called

vasopressin. According to these studies, and however incredible, altruism is

literally in (and around) our genes.

This kind of data is exciting to some, but leaves others scratching their heads.

What do students divvying up fifty shekels in a controlled experiment on a college

campus have to do with real-life altruism? If variation in the receptor gene also

affects marital status, the chances for divorce, and autism, how specific is it

really? Autistic people may find it difficult to marry, but aren’t egoism and

selflessness in a relationship relative and fluctuating? Finally, and perhaps most

important, if the different DNA profiles account for just a few percentage points

of the variation in altruistic behavior at best, in what sense do they account for the

behavior at all? Causality is a tricky beast, especially when it comes to humans.

Even the most sterling cancer research studies claim barely a 4 percent stake for

their pet agents with regard to the occurrence of disease. And figuring out the

relationship between altruism and genetics is infinitely more complex than

figuring out the connection between lung cancer and tobacco.

What does all this mean?

Like many of us, Sally Adams of Hampstead, London, isn’t sure what it all

means, but she knows that she wants a baby. Having spent more than £15,000 on

fertility treatments, and even traveled to Crete for her cause, Sally has not given

up. Over fifty years old, a cat lover, and single, she has the requisite sperm in

hand; only a tiny egg remains missing. Recently Sally advertised the fact that she

is looking for an egg donor from among Oxford and Cambridge graduates. Her

reason: Oxbridge graduates are not only intelligent but often also “very

altruistic.”

Whether through brain imaging, animal behavior experiments, dictator games,

genetic typing, gene-culture interaction models, or advertising in the Daily

Telegraph, the timeless search for altruism continues, with “admiration and awe.”

George Price, London, fall of 1974

Last Days

After World War II the old Victorian church in the middle of Tolmers Square was

converted into a cinema. When it was being swept up after a showing, back to

front, rats would come out the front door, run around the building, and go in the

back door, every night exactly the same. Still the locals loved it. With stall tickets

going for fifteen pence and circle ones for twenty-five pence, it was the cheapest

theater in London, and queues of two hundred would gather for the

Sunday-afternoon performance. When Joe Levy and Stock Conversion bought it

out in 1972, a veil of sadness descended over the neighborhood. Appropriately

The Looters and Die Slowly, You’ll Enjoy It More were the last films ever

screened. One pensioner wrote in rage to the local paper:

Dear Sir, Again a place of enjoyment…is to be closed and pulled down, although

it had 1,000 regular patrons. Is this what we call progress? Who decides about

this property? Do the 1,000 patrons have a say? Many age-old [sic] pensioners

will miss it. We were told in 1918 and 1945 this was a land fit for heroes to live in;

and take it from me, you need to be a hero to live in it.

George moved into a squat in the square on July 15, 1974; the cinema had long

been demolished, and a barbed-wire fence erected around the site had given it the

look of a concentration camp. The entrance to the squat was through Tolmers

number 19, but the walls between apartments 19–25 had been destroyed to make

room for a large silk-screen-printing studio on the first floor. Together with two

other squatters, Vince and Dave, George could share the entire second floor.

Everything was extremely primitive: the toilet, the shower, the kitchen. But after

what he’d been through, George thought it grand. The barbed-wire-surrounded

outdoor courtyard notwithstanding, it was the nicest home he had lived in, he

thought, since his babyhood home with his mommy and daddy in Scarsdale.

The next day he sat on a bench to talk to Sylvia, the twenty-five-year-old

American printmaking artist who worked together with Petal (a man) on the first

floor. Sylvia was the daughter of a U.S. Air Force fighter pilot wing commander

and had moved to Europe with an old boyfriend; now, together with a few friends,

she had opened a gallery called Artists for Democracy on Whitfield Street. A

group of them became the Poster Collective and had set up a squat studio at 19

Tolmers. She was living at number 68 Warren Street, just between Tolmers and

the gallery. She had dark hair, a beautiful face and body, and said she came from

Colorado.

Even among the politicos, the artists, the eccentrics and the druggies, George cut

an unusual figure. He was scraggly and often unclean. His teeth were rotting, and

he suffered from a terrible raspy cough. And yet this stranger, his thin legs

crossed, had the most amazing stories! He told her about his work at the

Manhattan Project and at IBM, about Skinner, and Shannon, and Senator

Humphrey. He spoke about Bell Labs and his Design Machine, about his attack

on the Soviets in Life magazine. He told of his brother, Edison, back home, who

had invented track lighting and worked with great architects like I. M Pei, Louis

Kahn, and Buckminster Fuller.

He told her about his own journeys into genetics

and evolution, about animal conflict and the equation that solved the problem of

altruism. But mostly he spoke about the people he was trying to help, about their

addictions, their sorrows, and hopes. George was a gentleman, she thought, a

quiet, thinking man who talked in a high, squeaky voice, almost like a baby. There

was something about him, she felt, that “just wanted to ‘be.’”

At the end of a day’s work at the studio, she’d walk out into the square and set her

sights back home. Increasingly, there was George, waiting to walk her across the

Euston Road. He was lonely and she liked him; she was glad of his company. On

rare occasions he’d come up to the collective kitchen for a cup of tea. But soon

Sylvia began to see that for George this was more than just a friendship. Over in

France, his old friend Henry Noel reproached him: Of course George might be an

exception, but “when an ex-married man, over fifty, falls in love with a girl who

could be his daughter, he is not on a Godward path.”

To Sylvia it was clear that his infatuation had nothing to do with her per se. The

real reasons were mysterious, tucked deep inside his imagination. He was

obsessive and intense but at the same time somehow not connected to the “real”

world. When he asked her to marry him the first time, sitting together on the

bench, she declined with a smile. When he asked again, she tried to quiet him with

a chuckle. Really, George, she’d say. I like you. But marriage and romance are out

of the question.

To Hamilton he divulged the true scope of his transformation: He had just

recently left his job as a night office cleaner (“my supervisor was sorry to have me

leave”). He was giving up his work on genetic polymorphism (“Harry, for reasons

that I am not certain about, is not interested in having me finish”). He was giving

up on their project on sexual selection (“I think maybe you’ll be better off without

me in your beetle theorizing”). He believed that a new phase had started in his life,

“where Jesus wants me to do less about helping others and give more attention to

sorting out my own problems,” at least for a certain number of years. He was

going to marry “someone named Sylvia (from Colorado),” and move to America

to raise a family and maybe do some economics. Sylvia didn’t quite share his

feelings yet, he knew, but George was optimistic. “Let’s wait and see what

happens,” he wrote, before reminding Bill that he had some books in the squat

that he could pick up the next time he was in London.

The following week George typed up a letter to the CPL Recruitment Division at

14 Old Park Lane. “Your advertisement in Computer Weekly for a programmer-

analyst with IBM FORTRAN experience to work on ‘a complex financial

modeling system’ at City Bank,” he opened, “fits my background and present

interests just about exactly.” After describing the optimization work at IBM over

which he had corresponded with Paul Samuelson back in the sixties, as well as his

genetics work at the Galton, George explained that his current plan was to get in

touch again with Samuelson and inquire about a research appointment at MIT.

“That finishes describing the relevant parts of my rather odd occupational career,”

he wrote, explaining that his plan should not dissuade them from hiring him:

…even if an offer of an appointment should be immediately forthcoming, there is

a matter much more important than economics research or jobs or anything like

that that will keep me staying in London for an unknown length of time. To put

matters briefly, I want to marry someone, and by “someone” I don’t mean

anyone, but one particular someone, who at the present time does not particularly

seem to be close to agreeing with me in this matter. Hence the need for a

temporary job.

Of course this would prevent him from being put in charge of any long-term

project, not to mention a lower salary rung than advertised. But George wouldn’t

mind. He was a quick learner, and in view of his past experience, could be

expected to start producing useful output during the first week. Then he ended: “I

don’t think money’s all that important (tactless thing to say when seeking

employment from a bank!), and so I would expect to receive substantially less

than the 3375 pounds p.a. I was getting when I left University College, though

substantially more than the 4.90 pounds per night I was earning in my last job as

an office cleaner.”

He was feeling good about himself again, and no one could stop him. Recently

two foot “fuzz” (policemen) had detained him for questioning on the street. He

was “hairy, tired and decrepit” soon two more cops appeared in a car, one of them

six foot three or four, cold and tough and of high rank. The frisk job didn’t take

long to follow: pockets, shoes, socks, carrier bag—they even read his letters from

Smoky in jail. The big cop radioed to the station to see if he had a record or was

wanted. But George thought he had gotten the last word, he wrote to Kathleen, as

is promised in Luke 21:15: “Because I will give you the ability to speak, along

with wisdom, that none of your opponents will be able to resist or refute.” He was

following a “total honesty policy” now. What this meant Kathleen, back in

Hawaii, wasn’t certain, but George at least seemed to know where he was going.

He had shaved his mustache and trimmed his beard. He had his hair in a neat

ponytail instead of “floating in the wind.” He had started acquiring more

possessions instead of giving them away. “Am now apparently heading back

upward,” he promised.