Altruistic Wasps?

Raghavendra Gadagkar
Science 333, 833 (2011);
DOI: 10.1126/science.1210420

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 PERSPECTIVES

action limits how precisely the parameter
can be determined—the so-called standard
quantum limit.
There are tricks, however, that can be
performed so that the standard quantum
limit can be circumvented. These back-
action-evading, or quantum nondemolition
(QND), techniques (6, 7) typically involve
the measurement of an observable A with
strongly suppressed noise, most famously
perhaps a quadrature (related to the phase
of an optical signal) of a squeezed state, and
the unavoidable quantum noise being there-
fore in the conjugate variable B, the other
quadrature of the squeezed state (the conju-
gate variables A and B that characterize an
electromagnetic field are called its quadra-
experiments but, so far, have been largely
limited to electromagnetic fields. This is
because a number of exquisite techniques
have been developed to generate and char-
acterize the types of quantum optical fields,
such as squeezed states, that are required for
back-action-evading measurements.
The recent breakthrough experiments
open the way to the extension of these tech-
niques to mechanical sensors. The suc-
cessful cooling of macroscopic mechani-
cal systems to their motional ground state
is an exciting and essential step toward that
goal, but it is “beyond ground state” physics
that promises to be most exciting in bring-
ing the emerging field of quantum acous-
tics to the level of sophistication of quan-
an explosion of new results expected soon.
This bright future results from a cross-
fertilization between quantum optics, laser
cooling, nanotechnology, gravitational wave
detection, and quantum measurement sci-
ence and engineering—demonstrating the
power of interdisciplinary science at its best.
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10.1126/science.1208322

EVOLUTION

Altruistic Wasps?
Is the widely invoked theory of kin selection
necessary to explain the apparent altruism
in social wasp colonies?
Raghavendra Gadagkar

P
olistes dominulus is one of the most
common social wasps in Europe and
is an invasive species in the United
States. Its wide prevalence has made it one
of the best-studied social wasps. In most
social wasps, the female wasps live in a col-
ony and organize themselves into a behav-
ioral dominance hierarchy such that only
the dominant alpha individual (the queen)
reproduces while the rest function as appar-
ently altruistic, sterile subordinates (work-
ers), building the nest, foraging for food and
pulp, and feeding and caring for the brood.
Why should workers invest their time and
energy helping to rear the queen’s brood,
rather than found their own nests and rear
their own brood—something they are quite
capable of? On page 874 of this issue, Lead-
beater et al. (1) show that the subordinates
indeed produce their own offspring and this
raises interesting questions about the links
between altruism, direct reproduction, and
the evolution of social behavior.
Using nine microsatellite markers to
genotype pupae from 228 natural nests of
CREDIT: E. LEADBEATER
foundresses, including dominants (queens), inherit the nest and thereby produce their
subordinates (workers) and solitary nest own offspring. What does this mean for the-
foundresses (not in a multifemale nest). ories about the evolution of social behav-
They found that subordinate foundresses ior that were built upon the assumption that
produced more offspring per
capita than an average solitary
foundress who reproduced (and
reared her brood) on her own.
Thirty-two percent of the sub-
ordinates’ offspring came from
“sneaking” eggs into the joint
nest (the queen usually prevents
subordinates from laying) while
the dominant was still alive, and
68% came from inheriting the
dominant’s position in the nest
after the latter had died. Thus,
the behavior of the subordinates
is not altruistic at all. Subordi-
nates that nest with a queen in
a cooperative manner stand to
Helping themselves. Subordinate
female wasps (P. dominulus) share
a nest with a dominant queen, rear-
ing the queen’s brood. This behavior

P. dominulus from Spain, the authors mea-
sured the reproductive success of 1113
Centre for Ecological Sciences, Indian Institute of Science,
Bangalore, 560012 India. E-mail: ragh@ces.iisc.ernet.in
also benefits the subordinates, who
stand to inherit the nest, or use the
nest themselves for their own eggs,
and increase the ability to rear their
own brood.

www.sciencemag.org SCIENCE VOL 333 12 AUGUST 2011 833

Published by AAAS
PERSPECTIVES
subordinate wasps do not lay any eggs of
their own?
In 1964, the British evolutionary biolo-
gist William D. Hamilton proposed a new
theory that is often referred to as kin selec-
tion or inclusive fitness theory (2). Ham-
ilton argued that to deal with the apparent
paradox of altruism—whereby a behav-
ior that benefits the recipient but harms
the individual performing it—the scope
of Darwinian fitness must be expanded.
Thus, the existing direct component of fit-
ness, gained through personal reproduction,
must be summed with an indirect compo-
nent, gained through aiding close genetic
relatives. He dubbed the sum of the direct
and indirect components “inclusive fitness.”
Hamilton went on to derive a rule according
to which altruism will be favored by natu-
ral selection if the benefit to the recipient,
devalued by the genetic relatedness between
the altruist and the recipient, is greater than
the cost incurred by the altruist.
Unfortunately Hamilton’s rule has not
been subjected to adequate rigorous empir-
ical testing—costs and benefits are hard to
measure (unlike relatedness, which is read-
ily measurable). But it can be argued that if
the benefit is sufficiently greater than the
cost, then any nonzero value of relatedness
can drive the evolution of altruism. This has
BIOCHEMISTRY
Catching a Moving Target
Taia T. Wang and Peter Palese
S
easonal influenza viruses persist as
important human pathogens because
of their ability to constantly change
the structure of a key protein, the viral hem-
agglutinin, which is needed to invade cells
and is a major target of immune defenses.
This shiftiness dictates a hefty vaccine reg-
imen: Ideally, patients are vaccinated each
year with updated formulations that pro-
tect them against new virus strains. Recent
research, however, has raised the possibil-
ity of developing a universal flu vaccine
that would not have to be updated. In mice
and humans, researchers have identified
“broadly neutralizing antibodies” (bnAbs)
(1–8) that have the potential to make influ-
enza less of a moving target. Two papers in
Department of Microbiology, Mount Sinai School of Medi-
cine, New York, NY 10029, USA. E-mail: taia.wang@mssm.
edu; peter.palese@mssm.edu
834 12 AUGUST 2011 VOL 333 SCIENCE www.sciencemag.org
led to the corollary that altruism can evolve
if it is nepotistic—that is, directed to indi-
viduals with high genetic relatedness (3).
Because relatedness among individuals in
a social group or colony is usually nonzero,
kin selection always appears to be correct
and it has therefore become very difficult to
criticize this theory (4–10).
Only because zero relatedness has been
found in P. dominulus [of the workers in
their colonies, 35% have been found to have
zero relatedness to their queens (11)], Lead-
beater et al. looked for possible reasons for
the workers’ apparent altruism, outside the
framework of kin selection, and found that
there was no altruism in the first place. An
average worker (which includes full sis-
ters of the dominant egg layer) gets enough
direct fitness to justify her nest sharing with
a queen and her subordinate role in caring
for the queen’s brood, making kin selection
(or indirect fitness) entirely unnecessary for
explaining sociality in P. dominulus. The
authors argue that direct fitness benefits are
not always the main driver of subordinate
behavior, because their study also shows
that for those subordinates that are relatives
of the dominant wasp, indirect benefits usu-
ally outweigh direct benefits. But the new
result is that kin selection is quite unnec-
essary to explain the formation of social
this issue, by Ekiert et al. (9) on page 843
and Corti et al. (10) on page 850, describe
novel bnAbs that protect animals against
influenza and show how they bind to the
viral hemagglutinin protein.
Hemagglutinin has two basic structural
regions (see the figure): a stem domain that
changes relatively little, and a globular head
that often undergoes structural changes.
Current seasonal vaccines prompt the body
to create strain-specific antibodies that bind
to the globular head, preventing the virus
from attaching to host cells. In contrast, a
second type of neutralizing antibody acts
on the more stable, conserved regions of
the hemagglutinin stem, preventing a fusion
step involved in cell entry. These “stem
reactive” bnAbs are less potent, but can
act against a broader range of virus strains
and subtypes than antibodies that target the
globular head. Indeed, animal studies have
Published by AAAS
groups in P. dominulus.
These findings make the point that we
need to pay attention to the possibility that
individual selection might be adequate to
explain many instances of social behavior.
No claim is being made here that Hamil-
ton’s inclusive theory is wrong or always
unnecessary; it is likely to be very useful
when there is altruism. But we need not
invoke it when there is no altruism in the
first place. In short, all social behavior need
not involve altruism and therefore need not
be kin selected.
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1. E. Leadbeater, J. M. Carruthers, J. P. Green, N. S. Rosser,
J. P. Field, Science 333, 874 (2011).
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10.1126/science.1210420
Broadly neutralizing antibodies could help
create an influenza vaccine that would not
have to be updated.
shown that many stem-reactive bnAbs can
prevent severe disease caused by distinct
influenza viruses.
There are 16 subtypes of hemagglu-
tinin proteins that can be divided into two
phylogenetic groups. The 10 subtypes in
group 1 include the hemagglutinins found
in currently circulating H1 viruses, highly
pathogenic H5 avian influenza viruses, and
H2 subtype viruses that have previously
caused pandemic disease in humans. The
remaining six group 2 subtypes include
H3 viruses, which are also currently circu-
lating in humans, and H7 viruses that have
caused sporadic human disease. Previ-
ously described bnAbs have activity against
viruses within group 1 or within the group 2
H3 subtype only (1, 2, 5–7).
Ekiert et al. describe an antibody,
CR8020, which is the first human mono-
clonal bnAb shown to neutralize an array of