Lingua Franca - Volume 10, No. 5
James Schwartz
July/August 2000
text is a cache of http://www.linguafranca.com/0007/altruist.html
IN A RUTHLESS, Darwinian world, human heartlessness is easy to
explain. After all, natural selection eliminates the weak and rewards the
strong. Unselfish behavior, on the other hand, is baffling. Compassion,
kindness, and loyalty ought to be weeded out almost as soon as they arise.
To explain the evolution of altruism, Charles Darwin suggested that natural
selection could act on groups as well as individuals -- an idea known as group
selection. Within a tribe, it could hardly be doubted that "selfish and
treacherous parents" would have the most children, he wrote in his 1871
classic, The Descent of
But
Despite his remarkable scientific achievements and the intense drama of his
personal life, George Price has remained a relatively obscure figure in the
history of science. And yet he played a key role in shaping the conceptual
basis of sociobiology and its offshoot, evolutionary psychology. He was the
first to apply the principles of game theory to the analysis of animal
conflicts, and he discovered an elegant formula to describe evolutionary change
that both simplified and improved
In his last years, even Price himself lost interest in securing recognition
for his scientific achievement. Altruism, which had begun as an intellectual
problem, became an all-consuming personal concern. In the midst of an
extraordinary burst of scientific creativity in the summer of 1970, Price
abruptly converted from militant atheist to fundamentalist Christian. After his
conversion, he combined his work in genetics with a passionate interest in
biblical exegesis. Later, he decided that the mission of a true Christian was
to help his fellow man, and he spent increasing amounts of time aiding homeless
alcoholics and the elderly in his adopted city of
In the years before his own untimely death this past spring, William
Hamilton often wrote and spoke about Price in an effort to draw attention to
his old friend's ideas. Earlier this year,
A PHYSICAL chemist and a journalist before he studied genetics,
George Price took a circuitous path to evolutionary biology. Born in 1922, he
was only four years old when his father died. His mother, a former opera singer
and actress, struggled to keep the family's lighting company afloat through the
Depression, and it was difficult for her to support George and his older
brother, Edison. After attending public school in
In 1955, the year of his divorce, Price published his first magazine piece,
a long article in Science in which he questioned the quality of the
evidence used to demonstrate ESP. It was the first of several highly visible,
and often farsighted, forays into journalism. The following year Price
published "How to Speed Up Invention" in Fortune magazine. In
the age of punched cards and Teletypes, the article described in detail a
hypothetical "design machine," which would feature a graphic display,
a cursorlike light pen, and a mouselike device to rotate, shrink, and enlarge
shapes.
In 1957, Price sent Senator Hubert Humphrey an early draft of an essay he
was writing for Life titled "Arguing the Case for Being
Panicky." The article warned that a decline in
But Price's career soon took a dramatic turn. In 1966, he was treated for
thyroid cancer. Price believed that the doctor, an old friend of his from
IN NOVEMBER 1967, Price sailed on the Queen Elizabeth, pleased
to be the sole occupant of a stateroom meant for two. In
Among the articles Price read in his library visits was William D.
Hamilton's now-classic study "The Genetical Evolution of Social
Behavior." A watershed in the history of evolutionary biology, it would be
Price's point of entry into the field.
According to traditional Darwinism, natural selection is the survival of the
fittest -- with "the fittest" defined as those organisms who leave
the most descendants. It was easy for
In March 1968, Price wrote
Left to his own devices, Price set out to find a simpler, more direct way of
achieving
AT THE HEART of Price's equation is the mathematical concept of
covariance -- a statistical measure of the relationship between any two sets of
data. Take, for example, the rainfall and temperature in a given place over
thirty days. If rainfall is independent of temperature, then the covariance of
rainfall and temperature will be close to zero. If days with more than average
rain also tend to have higher-than-average temperatures, then the covariance
will be positive. But if days with more rain than usual tend to have lower
temperatures than usual, then the covariance will be negative.
How can a covariance measure natural selection? Price's equation describes
the change in a gene's frequency from one generation to the next. Specifically,
it relates that change to the covariance between an individual's possession of
the gene and the number of children he or she has. If having the gene leads to
having more offspring, the frequency of the gene will increase in the next
generation.
It sounds so straightforward as to be almost tautological. But Price's
mathematics was strikingly original. Furthermore, in the process of inventing a
new algebra of natural selection, Price also devised a more sophisticated and
accurate way of understanding relatedness. In his own work,
Price's approach bore no relation to previous work. At first, even Price was
convinced that his equation was too simple to be new, so he decided to check it
with an expert at the Galton Laboratory at
WITH A TITLE, an office, and the support of certified experts, Price
set off with new confidence to write up his results. He believed that
university backing would help him get funding and make it easier to publish his
work. But outside of work, it was a difficult period for Price. His shoulder
was troubling him again, and he was anxious over money, since he had spent most
of his insurance settlement. To top it off, his mother had fallen seriously
ill.
In March 1969, his mother's health took a precipitous downturn, and he flew
to
In his 1996 book, Narrow Roads of Gene Land,
But Price would eventually cause
But it is thanks to this second term that Price's equation sheds such
powerful light on group selection. As Price hinted to
As it happened, one of the first well-documented examples of group selection
had recently been reported. In
Despite the strain in their first conversation,
Although Price's equation was strikingly original, its publication, which
would be Price's first in his new field, was by no means assured. Hamilton, who
had felt isolated and unappreciated while working out his theory of nepotistic
altruism, was anxious to help his friend avoid a similar fate. Together they
devised a clever strategy to break into Nature, one of the premier
science journals. Price would submit his paper on the mathematics of natural
selection first. One week later,
It came as no surprise when Price's paper was returned immediately. The
editors had not seen fit to send it out for review. No less surprising, the
paper by
EARLY IN THE summer of 1970, at the age of forty-seven, Price
underwent a sudden religious conversion. "On June 7th I gave in and
admitted that God existed," he explained to friends. He viewed his conversion
as a logical necessity, the result of a series of coincidences that had
befallen him. After calculating the odds of their occurrence and finding them
to be "astronomically low," he was convinced that there had been
supernatural manipulation. One week later, he attended his first service at All
Souls at Langham, a particularly evangelical branch of the Church of England,
located around the corner from his apartment.
Over the course of the next year, Price's scientific work was accompanied by
a new passion -- biblical exegesis. Adopting a highly literal approach to the
Bible, Price set out to reconcile discrepancies among the four Gospels. Nearly
a year after his conversion, he completed a fifty-page article, "The
Twelve Days of Easter," which proposed to replace the traditional
eight-day Holy Week with a new chronology. He believed he had resolved several
of the long-standing puzzles of biblical scholarship.
He sent the article to Hamilton, who was impressed by Price's reconstruction
of the events of Easter week and encouraged him to publish it. But he did not
accept Price's arguments for the existence of God, nor did he convert, as Price
had hoped he would. In a letter, Hamilton likened his resistance to
Christianity to "the Irishman who was asked whether he liked oysters and
he replied, no, he didn't like oysters and he was glad he didn't like them
because if he did he'd be eating them all the time when he hated the damned
things."
Price waited four months before replying. "The question is not whether
you like it but whether it is true," he wrote. "What difference does
it make whether you approve of it or not? Do you think that is something that I
wanted to believe in?"
MEANWHILE, Price's second major breakthrough in evolutionary biology
was at last about to see the light of day. Back in 1968, the same summer he
happened upon his covariance equation, Price had become intrigued by the fact
that male animals of the same species rarely fight to the death. In July of
that year, anxious that his windfall insurance payment was running out, Price
had worked night and day to complete an article setting forth a new idea, namely,
that game theory might help evolutionary biologists understand animal conflict.
At the time, Price hoped that a quick and dazzling academic paper would pave
the way for profitable magazine sales.
Price's key insight had been to see that the genetically optimal behavior
for an animal could depend on the behavior of other animals. In a population
made up of animals genetically programmed to make war, for example, an animal
programmed to retreat from a threat might actually be at an advantage. On the other
hand, in a population of less aggressive males, a confrontational male would
have the advantage. Price had titled his paper "Antlers, Intraspecific
Combat, and Altruism" and sent it off to Nature on the last day of
July 1968. The following February, he learned that Nature had accepted
the article, provided it was shortened. But several years passed, and Price
never bothered to undertake the revision.
As it happened, the reviewer of Price's paper had been John Maynard Smith,
the head of the biology department at the newly created University of Sussex.
Maynard Smith saw the potential of Price's unpublished idea, and he wanted to
use it in a paper he was writing. In 1971, he wrote a letter asking for
permission to thank Price for showing him an "unpublished
manuscript." Price wrote back that he would rather Maynard Smith referred
to a discussion between them and not a manuscript. "If one mentions an
unpublished manuscript," Price explained, somewhat mysteriously,
"then someone might wonder whether it was used with permission."
Price's concern was a delicate reference to a long-running, acrimonious
dispute between Maynard Smith and Hamilton, whose paper on nepotistic altruism
Maynard Smith had reviewed for the Journal of Theoretical Biology a few
years earlier. It was Hamilton's belief that Maynard Smith had in effect stolen
his idea. "His account of the matter," Price explained to Maynard
Smith in the fall of 1972, "is that you refereed his 1964 paper for the Journal
of Theoretical Biology, and required a major revision...that caused a
nine-month delay in publication and meanwhile you sent Nature a letter
with the term 'kin selection' that has received much of the credit for the
idea."
"I seem to have this fate of getting ideas from other people's
manuscripts when I referee them," the genial, white-haired Maynard Smith
explains today. Asked about Hamilton in particular, he is somewhat more
defensive: "I wasn't trying to steal his idea, or I don't think I was, so
it wasn't conscious." At first, Price doubted Maynard Smith's integrity
and suspected him of delaying the "Antlers" paper as he had
Hamilton's. But he changed his mind after meeting Maynard Smith. In fact,
Maynard Smith was scrupulous about crediting Price -- he offered to make Price
a co-author of the paper he was writing.
In the fall of 1972, "The Logic of Animal Conflict," co-written by
John Maynard Smith and George Price, was accepted by Nature. On
receiving this news, Price wrote to Maynard Smith that "I think this is
the happiest and best outcome of refereeing I've ever had: to become co-author
with the referee of a much better paper than I could have written by
myself." The paper was one of the first to set forth the ideas of
evolutionary game theory, ideas that in the years since have been used to
analyze everything from reciprocal grooming in African antelopes to egg
swapping among hermaphroditic fish.
IN JUNE 1972, the three-year grant that Price had obtained from the Science
Research Council came to an end, and he chose not to seek to renew it. He
preferred to give more time to his Christian work and less to mathematical
genetics. By the fall, Price had decided to live according to his literal
interpretation of the teaching of Jesus. Inspired by Jesus' advice in the
Sermon on the Mount to take no heed of the morrow, Price was pushing himself to
the brink of disaster. He was almost joyous in anticipation of the extreme
deprivation that his faith had brought upon him. "I am now down to exactly
fifteen pence," he wrote to Maynard Smith that October. "I look
forward eagerly to when that fifteen pence will be gone."
A month later, Price's diet consisted of one pint of milk a day, and he was
weak from malnutrition. He stopped taking the thyroxine pills that his thyroid
cancer had rendered necessary to his survival. Price believed that if God
wanted him to continue living, He would provide the missing hormone. In early
December, Price seems to have attempted suicide; in any case, he ended up in a
hospital. An alert doctor noticed that he was suffering from myxedema, a result
of his thyroxine deficiency, and provided the missing hormone without informing
the patient. Taking this as a sign that he was meant to go on living, Price
followed the doctor's orders and started taking his pills again.
Price made his final revisions to "The Logic of Animal Conflict"
the following February. In a cover letter, he explained to Maynard Smith that
he had made a few changes to accommodate his newfound belief in creationism.
"I think I found wordings that you won't object to, and that won't shock Nature's
readers by making them suspect what I believe," he wrote.
Later that month, Price's religious crisis deepened. In what he described as
"an encounter with Jesus," he saw that he had misunderstood the real
nature of Christianity and that his true duty was the care and love of people
rather than biblical study. He began to devote himself to the needy. He helped
out at various old people's homes and gave away all his money to homeless
alcoholics on the street, often inviting them to stay in his flat. He also set
out to make amends for the failures in his private life, apologizing to his
elder daughter, Annamarie, for deserting her and being a poor father. For a
short period, he hoped to remarry his former wife, Julia, and reunite the
family in London.
Toward the end of June 1973, Price gave up his comfortable flat and lived as
an itinerant. Once the model IBM employee, short-haired and suited, Price now
let his hair grow out and dressed in sneakers and colorful shirts with an
aluminum cross around his neck. He gave away the last of his possessions,
including his watch and coat, and lived hand to mouth. In a letter to Hamilton,
he explained that he was living according to Luke 6:30: "Give to every man
that asketh of thee; and of him that taketh away thy goods ask not again."
Elizabeth Mansell, who worked as a manager of an old people's home where
Price volunteered, recalls his arriving on Christmas Eve 1973 "like an
angel coming in." She and Price stayed up past midnight wrapping Christmas
presents for the old people, and she overslept the next morning. Rushing down
in a panic, she found Price had fed and dressed all twenty-one residents.
In the little time that remained after doing his Christian work, Price continued
his genetics research at the Galton Labs, where Cedric Smith, his first
sponsor, had procured him a one-year Medical Research Council stipend. But by
the end of the year, he was forced to avoid the lab because his charity cases
were causing disturbances there. In one incident, a belligerent alcoholic
(whose abused wife Price had been protecting) pissed publicly on the front
steps of the genetics building, smashed a bike lamp, and scattered the contents
of a student's satchel while shouting obscenities. Fortunately, Price's recent
paper with Maynard Smith in Nature had earned him credit with the lab
authorities. As he wrote to his daughter Annamarie, "I expect that one
cover-illustrated lead article in Nature compensates for one urination
at the front entrance to the building."
In March 1974, Price took up temporary residence in the home of an elderly
woman whom he'd helped. By keeping his whereabouts secret from his charity
cases, he hoped to get some work done on a joint project on sexual selection with
Hamilton. In June, he took a job as a night office cleaner. In a letter to
Hamilton, he wrote, "I thought it was about the first honest work I'd done
in my life -- working for others rather than for my own amusement or
advance." In August, Price gave up his night job and moved into a commune
located in six deserted buildings a mile north of his first flat.
This move would mark the last major transition in his life. He wrote to his
daughter Kathleen that after the summer's low in money and social prestige, he
was "heading back up" and had started acquiring possessions again,
which he was now slower to give away. He wrote to Hamilton that he believed
"Jesus wants me to do less about helping others and give more attention to
sorting out my own problems." To another friend he wrote that honesty
perhaps meant confessing to "one's deepest selfish desires." He had
fallen in love with a woman in his commune, and he hoped to move back to the
States. In November, he confided to his brother, Edison, that he had given up
his social work, returned to a more conventional Christianity, and was
considering marriage.
Meanwhile, Price was beginning to receive some long-overdue recognition.
Five years after their New York meeting, Richard Lewontin wrote Price: "It
has taken me a long time to come around to understanding the work you have been
doing, which I was too stupid to appreciate when you first showed it to
me." In October, Price also received a long and detailed letter from the
eminent American population geneticist James Crow. Like Lewontin, Crow
expressed chagrin at having been so slow to appreciate the significance of
Price's work.
Just before Christmas, Price visited the
THE SCIENTIFIC community has been slow to appreciate Price's
contributions, perhaps because he never pursued academic recognition strenuously.
The application of Price's covariance formula to group selection, for example,
has far-reaching implications, but Price never drew attention to them. Before
his own death this spring,
That may be changing. In 1995, twenty years after Price's death, the
theoretical biologist Steven Frank of the
For his groundbreaking insight into the evolution of altruism, Price merits
a special place in the history of evolutionary biology. The painful irony is
that his struggle to extinguish all selfish motives in his own life nearly
prevented him from achieving it. In Narrow Roads of Gene Land,
"Then why aren't you working on it yourself, George?"
"Oh, yes. Cedric wants me to also.... But I have so many other things
to do," Price replied. "Population genetics is not my main work, as
you know. But perhaps I should pray, see if I am mistaken."