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KIN AND KIND this has led to the evolution of For a century after Darwin,

an unusual way of sharing food. altruism remained a paradox.

A fight about the genetics of If a vampire bat fails to find a The first glimmers of a solution
altruism. victim during the night, it will arrived in a Bloomsbury pub in
begin licking under the wings the nineteen-fifties. According
By Jonah Lehrer
and on the lips of a chosen to legend, the biologist J. B. S.
Darwin saw the existence of colony member. The animals Haldane was several pints into
altruism as a challenge to his then lock mouths, and the the evening when he was asked
theory of natural selection, successful hunter starts vomiting how far he would go to save the
yet the biological success of warm blood. If such sharing did life of another person. Haldane
ants is directly related to not take place, scientists thought for a moment, and then
their ability to form complex estimate that more than eighty started scribbling numbers on
coperative societies. per cent of adult vampire bats the back of a napkin. I would
would die of starvation every jump into a river to save two
____________________ year. brothers, but not one, Haldane
_ said. Or to save eight cousins
but not seven. His drunken
answer summarized a powerful
Charles Darwin regarded the
scientific idea. Because
The vampire bat emerges from problem of altruismthe act of
individuals share much of their
its cave at the darkest hour of helping someone else, even if it
genome with close relatives, a
night, after the moon has set. It comes at a steep personal cost
trait will also persist if it leads to
flies low across the landscape, as a potentially fatal challenge to
the survival of their kin.
hunting by smell and sonar. his theory of natural selection.
According to Haldanes moral
Once the bat finds a victimit After all, if life were such a
arithmetic, making a sacrifice
can feed on most warm-blooded cruel struggle for existence,
for a family member is just
animals, from songbirds to cattle then how could a selfless
another way of promoting our
it starts stalking its prey. The individual ever live long enough
own DNA.
bat lands silently a few feet to reproduce? Why would
away, then runs on its wings natural selection favor a
toward the sound of a pulsing behavior that made us less likely
vein. A pair of teeth sharper than to survive? In The Descent of Haldane never expanded his
a scalpel cut into the flesh. Man, Darwin wrote, He who napkin calculations into a formal
Blood leaks from the wound; the was ready to sacrifice his life, as mathematical theory. That task
bat laps it up. Sometimes the bat many a savage has been, rather fell to William Hamilton, a
consumes its weight in blood than betray his comrades, would young graduate student at
during the night. often leave no offspring to University College London. He
inherit his noble nature. And struggled for years on the
yet, as Darwin knew, altruism is project, often working late at
everywhere, a stubborn anomaly night on a bench in Waterloo
Although the vampire bat has
of nature. Bats feed hungry Station, where the commuting
traditionally been seen as a
brethren; honeybees commit crowds eased his loneliness. In
ghoulish predator, it interests
suicide with a sting to defend 1964, he submitted a pair of
biologists for another reason: it
the hive; birds raise offspring papers to the Journal of
is deeply altruistic. The bats live
that arent their own; humans Theoretical Biology. The papers
in expansive colonies, with
leap onto subway tracks to save hinged on one simple equation:
hundreds or thousands sharing
strangers. The ubiquity of such rB > C. Genes for altruism could
the same dark cave. Bats must
behavior suggests that kindness evolve if the benefit (B) of an
feed constantlythey starve to
is not a losing life strategy. action exceeded the cost (C) to
death within sixty hoursand

the individual once relatedness understand the altruism at work Hamiltons death, in 2000, the
(r) was taken into account. The in ant colonies, and he became Oxford biologist Richard
equation confirmed the truth of convinced that Hamilton had Dawkins referred to Hamilton as
Haldanes joke: once kinship solved the problem. To further the most distinguished
was part of the calculation, the cause of inclusive fitness, Darwinian since Darwin.
altruism could be easily Wilson included the idea in a
explained in genetic terms. series of influential articles and
Hamilton referred to his model books, introducing the startling
But now, in an abrupt
as inclusive fitness theory, logic of Hamiltons equation to
intellectual shift, Wilson says
since it expanded the Darwinian biologists. I really became an
that his embrace of Hamiltons
definition of fitnesshow evangelist for the idea, Wilson
equation was a serious scientific
many offspring an individual says. And this was not an easy
mistake. Im going to be blunt:
manages to haveto include the idea to sell. Nobody wanted to
the equation doesnt work, he
offspring of surviving relatives. believe that one equation could
says. Its a phantom measure. It
The math seemed to solve the explain altruism. Eventually,
cant explain nearly as much as
biological problem, but in doing though, people saw that we were
people think it can. Back when I
so it opened up a moral right. I won that argument
first read Hamilton, inclusive
problem: altruism, it suggested, decisively.
fitness seemed to make sense of
isnt really altruistic at all;
so many different mysteries. But
rather, it is just another means of
now we know more. And Im
spreading our genes. Instead of
By the late nineteen-seventies, not afraid to admit I was
having sex, weve saving kin.
Hamiltons work was featured wrong. Wilsons apostasy,
prominently in textbooks; his which he lays out in a
original papers have become forthcoming book, The Social
At first, Hamiltons concept of some of the most cited in Conquest of the Earth, has set
inclusive fitness was entirely evolutionary biology. As Wilson off a scientific furor. The vast
ignored. Many biologists were realized, the equation allowed majority of his academic
turned off by the math, and few naturalists to make sense of colleagues are convinced that he
mathematicians were interested animal behavior using genetic was right the first time, and that
in the problems of biology. The models, giving the field a new his recantation has damaged the
following year, however, an sense of rigor. Before field. There have been
ambitious entomologist named Hamilton, there were different denunciations in the press and
E. O. Wilson read the paper explanations for every species, signed group letters in
while taking an eighteen-hour Wilson says. There was no prestigious journals; some have
train ride from Boston to Miami. overarching theory, nor was hinted that Wilson, who is
I had nothing else to do, so I there any way to connect what eighty-two, should retire. The
caught up on all my back we saw in the field to what we controversy is fuelled by a larger
journals, Wilson told me were learning about genes and debate about the evolution of
recently, when I visited him at coperation in the lab. Hamilton altruism. Can true altruism even
his office, at Harvard. When I helped solve both those exist? Is generosity a sustainable
began reading Hamiltons paper, problems. In fact, inclusive trait? Or are living things
my first response was that the fitness theory solved those inherently selfish, our kindness
equation was way too short. I problems so well that it was nothing but a mask? This is
thought, Theres no way it can soon applied to biological traits science with existential stakes.
be this easy. But then I reread completely unrelated to
the paper. And then I read it altruism, such as homosexuality,
again. And thats when I got tribal violence, and alarm calls.
The leaf-cutter ant is the best
jealous. Wilson wanted to In an obituary published after
mushroom farmer in the world.

Thriving in the tropical forests he was dissecting dozens of ant
of the New World, the dark-red bellies, searching for the source
ants live in vast, subterranean For the past sixty years, Wilson of the chemical. The work was
nests. The workers organize has been studying insects at maddening: because ant organs
themselves into seven functions, Harvard University. His office, are microscopic, Wilson was
as in an assembly line. Some in the Museum of Comparative forced to use sewing needles and
ants do nothing but cut leaves, Zoology, is down a dark hallway watchmakers forceps, carefully
harvesting up to seventeen per cluttered with metal filing plucking out each gland from
cent of the total leaf production cabinets. The room shares a wall the creatures. I wanted to steal
in a rain forest every year. with the museum, which means the ants signal and speak with it
Others haul the plant shards that the shouts of young myself, Wilson says. But
back to the nest, while others schoolkids echo through the air. nothing workedthe fire ants
tear the leaves into even smaller I dont mind the noise, Wilson were uninterested.
pieces. But the ants cannot eat says. Its like the twittering of
these leaves directly, since they birds. His hair is gray and
are laced with toxic chemicals. cropped close; like the haircut of
a little boy, it is mostly defined Wilson was running out of
Instead, they must turn the
by those strands which refuse to organs. On one of his last
mulching leaves into a fungus,
stay down. He was born in attempts, he removed the
which grows only within their
Birmingham, Alabama, and his Dufours gland, a tiny structure
colonies. One group of ants
Southern drawl has been near the ants stinger about
tends these underground
softened by decades in New which little was known. He then
mushroom gardens, weeding out
England, so that his speech is used this gland to paint a trail.
competing fungi and keeping the
now most notable for a slight The response was explosive,
chambers at an ideal
lisp and the long pauses of a Wilson says, his voice still
temperature and humidity. The
man who is used to being quickening with excitement. I
leaf-cutter ants have managed
listened to. The shelves in his got the entire colony chasing
their monoculture farms for tens
office are mostly taken up by his me!
of millions of years.
own bookshes published
twenty-four, two of which
received the Pulitzer Prizeand Wilson, along with other
E. O. Wilson has devoted his thick reference works on insects. researchers, began to identify
career to ants. As he notes, they
the specific compounds secreted
are perhaps the most successful
by the gland. Because each ant
form of multicellular life in
The discovery that made contains less than a millionth of
history, with some fourteen
Wilsons reputation occurred in a gram of the pheromones, he
thousand known species. They
1959, when he was a young had to collect tens of thousands
account for roughly the same
professor at Harvard. Wilson of fire ants, dumping entire
amount of biomass as human
was trying to understand how a colonies into a creek and then
beings. This biological success
fire-ant colony cordinated its scooping up the insects once
is especially remarkable because
behavior. He noticed that they floated to the surface. The
it depends entirely on the ability
whenever the ants encountered a field work wasnt funWilson
of ants to coperate, to form
piece of food too large to carry was stung scores of timesbut
intricate societies structured
they would drag the tip of their it allowed him to identify the
around hard work and shared
abdomen on the ground on their language of fire ants, a
sacrifice. As King Solomon
way back to the nest. Wilson vocabulary of volatile liquids
declared, in Proverbs, Go to the
assumed that the insects were consisting of up to twenty
ant, thou sluggard; consider her
laying a scent trail. Before long, communication signals.
ways and be wise.

selfless service was pure genetic
For Wilson, the exquisite logic Sociobiology sparked a bitter
of this system convinced him Ants arent the only insects to controversy. Wilson was
that biological behavior could be rely on haplodiploidy. As attacked by eminent scientists,
understood, that even something Hamilton noted, the same logic many of whom were in his
as complicated as an insect could explain the evolution of department at Harvard. There
colony could be explained in sawflies, wasps, and bees. These was a group letter in The New
terms of simple laws and insects all exhibit an extreme York Review of Books which
chemistry. I was convinced by form of altruism known as insisted that the concept of
ants that biology needed to eusociality, in which individuals sociobiology provided a
develop a theory of social live together in vast, coperative genetic justification of the status
behavior, he says. Such a societies. Although eusociality is quo and of existing privileges
theory was inevitable. It had to a relatively rare adaptation, its for certain groups according to
exist. And this is why I was so incredibly successful: only two class, race, or sex. Wilson
excited by Hamiltons papers. per cent of insect species are retained his equanimityI
eusocial, but they account for knew I was right on the
Hamilton showed that the approximately eighty per cent of science, he saysand now the
coperative nature of many all insect biomass. The ability of application of evolutionary
insect societies could be Hamiltons equation to cut theory to humans is no longer
explained by a genetic quirk across life forms suggested that controversial. Wilsons idea has
known as haplodiploidy. In it was a general principle of produced a colossal scientific-
some insect species, females social behavior, and that many research program, inspiring
emerge from a fertilized egg, of the most important examples decades of inquiry in behavioral
while males develop from of biological coperation were genetics, neuroscience, and
unfertilized eggs. (One mere by-products of genetic evolutionary psychology. Wilson
consequence of this bizarre relatedness. can barely resist gloating: That
setup is that males have half the was another argument that I
chromosomes of females. They won, he says. It took a few
also have a grandfather but no years, but, boy, did I win.
father.) Once haplodiploidy was Wilson was entranced by the
taken into account, the extreme haplodiploidy hypothesis and
solidarity among sisters in, say, made inclusive fitness an
colonies of leaf-cutter ants important part of his book Still, even as Wilson
ceased to be a mystery. Sociobiology: The New campaigned for sociobiology, he
Normally, siblings share fifty Synthesis (1975), which began to grow dismayed with
per cent of their genes, but explored the role of evolution in the scientific framework that
female worker ants share three- shaping social behavior. In the made it possible. I noticed that
quarters of theirsall of their final chapter, Wilson set out to the foundations of inclusive
fathers genes and half of their apply biological principles to fitness were crumbling, Wilson
mothers. Crucially, sister ants human beings, attempting to says. The reasoning that had
are more closely related to one consider man in the free spirit convinced me it was correct no
another than to their own of natural history, as though we longer held. For instance, after
offspring. For Hamilton, female were zoologists from another pursuing Hamiltons
workers were willing to look planet. After all, why should haplodiploidy hypothesis,
after the queen because she was Homo sapiens be exempt from scientists discovered that many
essentially a sister-producing the selfish logic of genes and of the most coperative insect
machine. Their seemingly kin? The equation was a species, such as termites and
universal truth. ambrosia beetles, werent

actually haplodiploid. Instead of acting like randy of life through its underlying
Furthermore, tens of thousands juveniles, seeking out mates and mathematics. Nowak is now
of species that did manifest getting into territorial fights, forty-six years old, with a shiny
haplodiploidy never evolved they are content to remain at bald head and thick black
eusocialityalthough these home, in the nest of their eyebrows. He speaks with an
insects were closely related, they parents. Stranger still, they Austrian accent so perfect it
didnt share food or serve the spend much of their time feels like a put-on, his dense
queen. By the late nineteen- helping to raise their younger sentences lightened by the lilt of
nineties, the relationship siblings, incubating the eggs and his voice. Nowak is widely
between haplodiploidy and gathering food for the extended regarded as one of the most
eusociality was no longer family. This behavior, known as important mathematical
statistically significant. coperative breeding, makes biologists in the world, with
little Darwinian sense. Why are more than forty publications in
the males squandering their Nature and fifteen in Science. In
most fertile years at home rather 2003, he became the director of
What happened is that, very
than competing to reproduce? It the Program for Evolutionary
quietly, the inclusive fitness
wasnt until the rise of inclusive Dynamics, at Harvard, a think
theorists stopped talking about
fitness theory that biologists tank of theoreticians working on
haplodiploidy, even though it
could explain this altruistic biological problems.
was their best piece of
behavior. In 1976, in one of the
evidence, Wilson says. At first,
first experimental tests of
Wilson kept his skepticism to
Hamiltons hypothesis, the
himself. He had little interest in Nowaks interest in inclusive
researchers Jerry and Esther
dismantling a theory that hed fitness theory began in the early
Brown began manipulating the
persuaded so many others to nineteen-nineties, when, as a
number of helper birds in the
accept. But, after investigating graduate student at Oxford, he
babbler nests. When they
the literature in detail while caught an offhand remark by his
removed the male helpers, the
working on a textbook about mentor, the theoretical biologist
survival rate of their young
ants, he concluded that inclusive Robert May. He referred to
siblings plummeted. In fact, the
fitness was no longer a tenable inclusive fitness as a cult, not a
Browns found that each helper
concept. The flaws swept in on science, Nowak remembers. I
generated 1.6 additional
my consciousness, he says. I thought that was interesting, but
offspring, a fitness benefit that
was, quite frankly, surprised by I didnt really know what he
helped compensate for its own
how little progress had been meant. It wasnt until 2006,
reproductive loss.
made. I could no longer deny the more than a decade later, that
possibility that the field had Nowak started looking closely at
made a wrong turn. It was at Hamiltons equation. He soon
this point that Wilson heard It was stories like thisvivid grew frustrated with its
from someone outside his field, anecdotes of animal behavior, fuzziness. Everyone talked
a mathematician named Martin encapsulated by simple about this rule, but in very
Nowak, who had reached a equationsthat first got Martin imprecise ways, he said. It
similar conclusion. Nowak interested in biology as seemed like a kind of pretend
an undergraduate at the math. At the time, he was too
University of Vienna, in the immersed in other projects to
nineteen-eighties. But he didnt pursue his skepticism. But then,
The young males of the
want to be a naturalist; in October of 2007, he received
Australian gray-crowned
observation struck him as too an e-mail from a young Harvard
babbler, a small woodland bird
inefficient. Instead, Nowak mathematician named Corina
with a curved black beak, have
wanted to understand the details Tarnita, asking for a meeting.
long perplexed biologists.

how it was actually used by mathematical tests, inclusive
biologists, Tarnita says. What fitness can be applied only under
Tarnita, who had grown up on a I discovered is that no one very specific biological
farm in rural Romania, was a actually uses the equation to circumstances, which almost
math prodigyshed won make calculations, because then never exist.
nearly every major award from it isnt very helpful.
the Harvard math department as
an undergraduate. But shed
become disenchanted with her While Tarnita was grinding out
graduate research, in an esoteric As Tarnita describes it, the the mathematical models,
branch of algebraic geometry. problem with Hamiltons Nowak was searching for allies
There were maybe five other equationand with the concept within biology. The search soon
people in the world who cared of inclusive fitness more led him to Wilson, who had
about my work, she says. And generallyis that it attempts to started publicly criticizing
so she told her adviser she analyze each action in isolation, Hamilton and inclusive fitness,
needed a break and began as a discrete deed with benefits pointing out the limitations of
browsing the shelves of the math and costs. Now, thats a very the insect data and suggesting
library, looking for another admirable goal, Tarnita says. alternative hypotheses. During
subject. Thats when she But how do you actually our first conversation, Wilson
discovered a textbook that calculate those benefits and told me that he always assumed
Nowak had written on the costs? To illustrate the the math of inclusive fitness
mathematics of evolution. I challenge, Tarnita imagines a must be very strong, because the
opened the book up and I scenario straight out of a soap biology was quite weak,
realized that this math wasnt so opera. Lets say your cousin is Nowak remembers. And I told
abstract, she told me. This was drowning, and so you risk your him the opposite. I said I always
math about life. At the time she life to save him. Thats good, assumed the biology was solid,
e-mailed Nowak, Tarnita had a right? Youve improved your because the math was very
dilemma. Shed recently inclusive fitness. But what you obscure.
received a job offer from a large dont realize is that your cousin
hedge fund, for a lucrative is actually competing with your
position as a quantitative brother for a wife. They both
The three scientists began
analyst. She was tempted by the love the same woman, but she
meeting every week, exchanging
money. I like fancy clothes and chooses your cousin. So now
stories about insects and game
fast cars, she says. I told your brother doesnt get
theory. They quickly zeroed in
myself that if Martin didnt e- married, and he doesnt have
on the paradox they wanted to
mail me back then maybe I three kids. Was it still a good
explain: if coperation is such a
would leave Harvard. idea to save your cousin from
successful strategyeusocial
species dominate their selfish
cousinsthen why is it so rare?
Fortunately, Nowak responded Why havent more creatures
and soon invited Tarnita to join Tarnitas point is that the imitated the altruistic life styles
his working group. One of her elegance of Hamiltons equation of honeybees and ants?
early assignments was to learn is a faade. When applied to the
about inclusive fitness. She real world, the math quickly
spent a year reading papers and becomes exceedingly
Wilson convinced the
poring over hundreds of studies convoluted, as it attempts to
mathematicians that they needed
on the application of Hamiltons encompass the extended
to delve into the details of living
equation. I wanted to figure out consequences of every decision.
things. What Id learned from
This is why, according to her

the failure of inclusive fitness is natural history. The reason leaving the nest. They ran
that you cant build a theory out eusociality is so rare, he computer simulations in which
of thin air, Wilson says. Our believes, is simply that it eusocial queens competed
theories need to begin with requires a long list of against solitary ones, and found
boots on the ground, with a preadaptationstraits that must that eusociality increases a
really close look at the species be in place before a further trait queens birth rate eightfold and
in question. can evolve. The most important reduces the probability of her
of these is the formation of a death tenfold. A competitive
cohesive group, which most advantage of this magnitude
often comes about because the would explain why, once
This emphasis on the empirical
daughters dont leave the nest. If eusociality emerges, it leads to
marked a shift for Wilson. Ive
the group persists for an such striking success. And yet
always been an ambitious
extended period, the female the model also documents the
synthesizer, he told me. But
insects might then construct a barriers to the evolution of
Im now wise enough to know
defensible nest. Its only at this eusociality, since it typically
the limitations of that approach.
point that the species can begin requires a set of unusual
These days, he regards the
developing the genetic mutations and very particular
books that made him famous
adaptations that enable ecological conditions.
Sociobiology and On
eusociality, such as the feeding
Human Nature (1979)as
of larvae and the division of
flawed accounts of evolution,
labor. Once that happens, the
marred by their uncritical In 2010, Tarnita, Nowak, and
logic of natural selection takes
embrace of inclusive fitness. Wilson published these ideas in
over, as the intensely altruistic
Hes prouder of an eight- Nature, in a seven-page paper
life style of the insects allows
hundred-page textbook that he titled The Evolution of
them to reproduce at an
wrote on Pheidole, the most Eusociality. (The math was
accelerated pace. The key point
abundant genus of ants. At one contained in a thirty-nine-page
is that the relatedness of the ant
point during my conversation online supplement, full of
colonyall those kin working
with him, he walked over to his abstruse equations.) The
togetheris a consequence of
bookshelf, gingerly heaved scientists knew that their paper
eusociality, not the cause. The
down the enormous volume, and might be controversial, but
sisters dont get along because
began flipping through the pages assumed that the debate would
theyre sisters. Rather, clumps of
and admiring the art work. center on the technical details of
females just happen to be the
There are six hundred and their model. Instead, the article
most likely to evolve the
twenty-four species in this book, unleashed a tempest of criticism,
necessary preadaptations. They
and I drew every one by hand, much of it aired in public. (Such
work together because they
he said. It took me twenty arguments are a regular
cant leave; they have become
years. I know that sounds occurrence in evolutionary
slaves to the queen.
obsessive, but thats what it biology, a field that seems to
takes. If you want to explain lapse into discord every decade
ants, then you have to know or so.) Andy Gardner, an
ants. This idea was attractive, but it evolutionary biologist at Oxford
lacked evidence. To test University, told the New York
Wilsons theory, Nowak and Times that it was a really
Tarnita developed a terrible article. Jerry Coyne, a
Wilsons deep knowledge of
mathematical model for the biologist at the University of
insects led him to propose a new
effect of a eusocial mutation Chicago, wrote on his blog that
model for the evolution of
one that would prevent some the only reason this paper was
altruism, which he decided was
daughters of a queen from published was because it has
rooted in the contingencies of

two big-name authors, Nowak inclusive fitness is inclusive fitness has been tested
and Wilson, hailing from Mother overwhelming, Dawkins says. in a number of ways. Its made
Harvard. . . . The lesson: if To insist otherwise is simply predictions, and those
youre a famous biologist you wrong. predictions are correct.
can get away with publishing

Tarnita and Nowak replied in For their part, the

Nature, and then again on their mathematicians seem to regret
The critiques were even harsher Web site, making it clear that some of the language used in the
in private. Robert Trivers, an they were not trying to discredit Nature paper. If I could write it
eminent biologist at Rutgers and Hamilton or disregard the again, I would make it more
a former collaborator with importance of relatedness. clear that Hamiltons equation
Wilson, wrote a strongly worded Nowak, in a forthcoming edition inspired lots of good research,
personal e-mail to Nowak. of his book Supercooperators, Tarnita says. He got people
Heres how the letter ends: expresses an eagerness to learn thinking about relatedness, and
from his detractors, but that was very important.
maintains that inclusive fitness Nevertheless, Tarnita and
is a useless gyration, Nowak continue to insist that
Martin, do you really think a
characterized by a tendency to Hamiltons equation is not a
quick and cheap paper like this
theorize without precision. precise expression of a
is going to displace W. D.
biological phenomenon. Rather,
Hamilton and kinship theory?
its little more than a rule of
Do you think anybody actually
thumb, a truism masquerading
expert in these matters is going Still, the debate has not reached
as a truth.
to swallow this tripe? Do you a satisfying conclusion. The
think this is a worthy use of your mathematicians insist that their
time? I hope you do not. critics dont understand the
math, and the biologists insist The scientific disagreement is
that the mathematicians dont particularly hard to resolve
understand the biology. Wilson, because its rooted in the distinct
Before long, a group of
meanwhile, admits that he cant perspectives of mathematicians
biologists began drafting a
follow much of his colleagues and biologists. Tarnita and
response to send to Nature. The
math, but says that he knows Nowak want the equations of
letter was signed by a hundred
insects. One of the few scientists altruism to be literal; when they
and thirty-seven scientists, and
able to speak the language of talk about inclusive fitness, they
insisted that Wilson and the
both sides is the man who are talking about the
mathematicians misunderstood
helped to cordinate the letter to mathematical details of rB > C.
evolutionary theory and
Nature, David Queller, of Such computations, of course,
misrepresented the scientific
Washington University in St. are extremely difficulthow
literature. In particular, they
Louis. He argues that, while the does one measure the fitness of
were upset by the claim that
pages of equations employed by an action?which is why they
inclusive fitness theory has
Nowak and Tarnita might be find Hamiltons work so
produced meagre results. The
technically correct, they dont unsatisfying. I get that his
scientists cite a long list of
support their grand conclusions. equation is simple, and thats
insights that stem directly from
Martin is always going on nice, Tarnita says. Everyone
Hamiltons equation, such as
about how inclusive fitness likes simple equations. But its
why animals act spitefully and
doesnt work in the real world, too simple. Our model is
why some species have such
Queller told me. But his model messier than Hamiltons, but
uneven sex ratios. The case for
is certainly no better. The fact is, you get a lot more out of it.

Nowak is even blunter. If the theory is wrong, why wouldnt spends up to three years hacking
theory works, then it has to work one author suffice? I feel the away at the healthy wood of a
at a mathematical level, he same way. When you read their pine tree. This labor is not as
says. responses, they never say what thankless as it appears, though.
weve got wrong. And thats It turns out that drilling into a
because we didnt get anything live pine has ancillary benefits.
wrong. I dont want to sound too When the woodpecker bores its
The biologists, on the other
boastful, but I think this is a big holes, resin seeps out, coating
hand, dont mind that inclusive
paper. Its a game changer. the tree trunk in a sticky glue,
fitness often cant be calculated.
which prevents the
Instead, they see it as a
woodpeckers chief predators,
framework for making sense of
such as rat snakes, from gaining
the world, an important principle And so the argument continues,
access to its nest. The bird has
that helps us understand the with both sides promising new
built a trap.
varied behavior of bats, ants, papers that will prove the other
and other species. The equation, side wrong. The problem is, of
in other words, isnt really an course, that its hard to imagine
equation. Its just a short what such proof might look like. Although its not uncommon for
summary of a big idea, much Despite the impressive tool kit Wilson to delve into the natural
like Darwins description of of modern biology, this is still a history of a particular species,
natural selection. debate about distant history, shot his interest in this bird has been
through with ambiguous facts unusually intense. I thought it
and contested first principles. was incredible that these little
Meanwhile, nobody seems to guys drill into living trees, he
Wilson is the only one who
have noticed the irony of the says. And I began wondering if
seems to be enjoying the
situation: they are fighting over that might play a role in why the
controversy. His appetite for
the origins of kindness. young males stick around.
scientific brawls seems, if
anything, to be increasing with
age. Wilson likes to quote
Schopenhauer on how all new A few years ago, Wilson became Before long, Wilson was
ideas go through three phases. obsessed with the red-cockaded travelling down to the Gulf
First, the truth is ridiculed, he woodpecker, an endangered Coast and looking at the nest
says. Then it meets outrage. species that inhabits the pine sites for himself. He consulted
Then it is said to have been forests of the southeastern woodpecker experts and spent
obvious all along. Were United States. Like the time wandering around the
currently in the outrage stage, Australian gray-crowned vanishing pine forests of
but well be obvious before babbler, these birds engage in Florida, searching for small
long. coperative breeding, as most boreholes and slicks of resin and
young males spend several years trying to understand what led to
raising their relatives. But coperative breeding. And
Wilson wasnt just entranced by thats when I realized one could
He seems similarly unconcerned
their altruism. He also marvelled come up with a much better
by the reaction of his colleagues.
at another peculiar habit of the explanation than inclusive
After Einstein published his
red-cockaded: it drills into living fitness, he says. According to
theory of relativity, a hundred
trees. While most woodpeckers Wilson, the driving force is the
physicists wrote a paper
construct nests in dead and scarcity of suitable nesting sites.
condemning it, Wilson says.
dying tree trunksthe rotting Species that engage in
Einsteins response was
wood makes it easier to excavate coperative breeding, he notes,
marvellous. He said, If the
the red-cockaded woodpecker all have very limited territories

and are very picky about where Darwin in The Descent of Altruistic groups beat selfish
they live. This suggests that the Manthat human generosity groups. Everything else is
young males hang around might have evolved as an commentary.
because they cant find a emergent property not of the
suitable tree for themselves, and individual but of the group.
are hoping to inherit the nest of There can be no doubt that a
Wilsons larger point is that, to
their parents. Babysitting the tribe including many members
the extent that altruism exists, it
siblings, then, is just a way of who . . . were always ready to
isnt an illusion. Instead,
paying rent, a chore performed give aid to each other and to
goodness might actually be an
in exchange for shelter. My sacrifice themselves for the
adaptive trait, allowing more
point is that I can explain these common good, would be
coperative groups to
helpers without talking about victorious over most other
outcompete their conniving
relatedness or inclusive fitness, tribes, Darwin wrote. While
cousins. In a field defined by the
Wilson says. These birds are acts of altruism can be costly for
cruel logic of natural selection,
just dealing with the challenges the individual, Darwin argued
group selection appears to be the
of real estate. that they helped sustain the
rare hint of virtue, the one
colony, which made individuals
biological force pushing back
within the colony more likely to
against the obvious advantages
In his more expansive moods, of greed and deceit. I see
Wilson goes beyond the habits human nature as hung in the
of woodpeckers to speculate on balance between these two
the larger forces at work in the This idea is known as group extremes, Wilson says. If our
evolution of altruism. He isnt selection, and its an explanation behavior was driven entirely by
content to tear inclusive fitness that most evolutionary biologists group selection, then wed be
downhe wants to replace it now dismiss, because the robotic coperators, like ants.
with something better. As usual, advantages of generosity are But, if individual-level selection
Wilsons latest proposal is much less tangible than the was the only thing that mattered,
driven by his faith in the power benefits of selfishness. (A tribe then wed be entirely selfish.
of close observation, as well as full of nice guys would be easy What makes us human is that
by a kind of intuitive empathy prey for a cheater, who would our history has been shaped by
with fauna, which seems to be quickly spread his genes through both forces. Were stuck in
the product of a life spent the population.) But Wilson between.
observing the natural world. believes that it may hold the key
While he clearly enjoys having to understanding altruism. To
the math back him up, one gets make his case, he cites recent
the sense that he wouldnt studies of coperating
change his mind if it didnt. At microbes, plants, and even
this late stage of his career, female lions. In all these studies,
Wilson is less interested in many of which have been
equations than in narratives. conducted in the controlled
Indeed, a few years ago he conditions of the lab, clumps of
published a novel, albeit one coperators thrive and replicate,
about ants. while selfish groups wither and
die. In a 2007 paper that he co-
authored, he summarizes his
new view in three terse
Wilsons current explanation for
sentences: Selfishness beats
altruism has returned to a
altruism within groups.
hypothesis first proposed by