dynamic phase diagram of possible behav-
iours of the filaments. Just how universal this
diagram will be — whether it describes the
behaviour of everything from flocks of birds
to catalytic colloids — can be tested only by
comparison to experiments on the many
realizations of active matter.
Jean-François Joanny is in the Physical Chemistry
Unit (UMR 168, CNRS UPMC), Institut Curie,
Paris 75248, Cedex 5, France. Sriram
Ramaswamy is at the Centre for Condensed-
Matter Theory, Department of Physics,
Institute of Science, Bangalore 560 012, India.
1. Schaller, V., Weber, C., Semmrich, C., Frey, E. & Bausch, A.
Nature 467, 73–77 (2010).
2. Joanny, J. F. & Prost, J. HFSP J. 3, 94–104 (2009).
3. Ramaswamy, S. Annu. Rev. Condens. Matter Phys. 1,
4. Szabó, B. et al. Phys. Rev. E 74, 061908 (2006).
5. Dombrowski, C. et al. Phys. Rev. Lett. 93, 098103 (2004).
6. Narayan, V., Ramaswamy, S. & Menon, N. Science 317,
7. Deseigne, J., Dauchot, O. & Chaté, H. Preprint at
http:/ /arxiv.org/abs/1004.1499 (2010).
8. Kudrolli, A., Lumay, G., Volfson, D. & Tsimring, L. S. Phys.
Rev. Lett. 100, 058001 (2008).
9. Mallouk, T. E. & Sen, A. Sci. Am. 300, 72–77 (2009).
10. Vicsek, T. et al. Phys. Rev. Lett. 75, 1226–1229 (1995).
11. Toner, J. & Tu, Y. Phys. Rev. Lett. 75, 4326–4329 (1995).
12. Chaté, H., Ginelli, F., Grégoire, G. & Raynaud, F. Phys. Rev. E
77, 046113 (2008).
13. Mishra, S., Baskaran, A. & Marchetti, M. C. Phys. Rev. E 81,
Hyun Youk and Alexander van Oudenaarden
A charitable deed by a few cells in a bacterial culture can help the rest of
that population survive in the presence of antibiotics. This finding can aid
further research into a major problem in public health.
Bacterial resistance to an antibiotic
arises when mutations in the DNA of a
few cells in a bacterial popu lation ena-
ble them to fend off the harmful effects
of the antibiotic. This gives such cells
a selective growth advantage through,
for example, being able to pump out
the antibiotic faster than it can kill
them. The mutant bacteria pass on the
advantage to their daughter cells and to
subsequent generations. Through such
amplification, they can outgrow their
non-resistant neighbours and eventu-
ally come to dominate the population,
making an antibiotic generally ineffec-
tive. This, then, is the cause of the global
challenge of widespread resistance to
On page 82 of this issue, Lee et al.2
offer a new angle from which to view
this picture, one that reveals collec-
tive action on the part of bacteria exposed to
an antibiotic threat. The work highlights the
importance of quantitatively understand-
ing microbial population dynamics in devel-
oping the correct strategies for prescribing
antibiotics for patients.
Imagine that a cell that developed resistance
can help its non-resistant neighbours by, say,
secreting some substance that assists those
neighbours in fighting off an antibiotic. This
type of behaviour might make it easier for the
whole bacterial population to avoid extinction.
From the point of view of the bacterial species
as a whole, it would avoid the need to wait for
the rare resistant mutant to come to dominate
the population. It would also ensure that popu-
lation-level genetic diversity is maintained.
Lee et al.2 describe just such a charitable
deed carried out by individual bacteria in a
population of Escherichia coli subjected to an
antibiotic. The authors show that mutations
that develop in a few cells enable them to assist
neighbouring cells that have not undergone
resistance-conferring mutations. The actions
of these few cells lead to a population-wide
resistance. But they do so in a roundabout
way that illustrates how antibiotic resistance
can emerge from seemingly harmless and
Lee et al. used the antibiotic norfloxacin,
which targets a protein that is essential for
DNA replication, and so for bacterial cell divi-
sion and population growth. They studied
how mutant cells within initially genetically
identical E. coli cells arise over time under the
selective pressure imposed by the presence of
norfloxacin in the growth media. When kept
on media containing a moderate, not fully
lethal, level of norfloxacin, the bacterial cells
initially suffered from stunted growth. But
within a couple of days, the overall popula-
tion growth rate increased because the pool of
cells had together become more resistant to the
constant level of antibiotic.
The authors then increased the amount of
norfloxacin, which slowed the overall popula-
tion growth again. But within a couple of days
the population developed resistance once
more, and the growth rate recovered. This
tug-of-war, presenting an ever-increasing
challenge to the cells, was continued for ten
days, at the end of which the population could
withstand some five times higher levels of nor-
floxacin than it could initially. But that did not
mean that every bacterial cell in the population
had become resistant through mutations.
The novelty of Lee and colleagues’ study2
lies in their temporal characterization of the
mechanisms responsible for resistance to nor-
floxacin in individual cells as they developed.
The authors found that almost all of the resist-
ant mutants were less effective at staving off
the growth defect imposed by norfloxacin than
was the whole population from which
they were isolated. In other words, when
almost all of the isolated mutants were
subjected to norfloxacin, their growth
rate was lower than that of the overall
population containing mixtures of both
resistant and non-resistant cells. But a
small minority, just one or two, of the
isolated resistant mutants grew faster
than the whole population when they
alone were exposed to norfloxacin.
Lee et al. show that this is due to these
few, highly resistant mutants producing
a small molecule called indole, which
readily diffuses into neighbouring cells
and triggers molecular defence against
norfloxacin. Notably, the mutations in
these highly resistant mutants do not
directly cause this altruistic behaviour.
Indole is a molecule that is usually pro-
duced by the bacteria when antibiotic
is not present, but is shut down when it is. The
authors demonstrate that the highly resistant
mutants developed mutations that could help
them survive in the presence of norfloxacin
in the first few days as well as allow indole
production to continue. This enabled them
to assist their much less resistant neighbours
in counteracting the antibiotic. Because the
highly resistant mutants invest energy to pro-
duce indole, which is not required for their
own resistance, their help for their kin can
be considered a form of altruism. Lee et al.
repeated their experiments with another anti-
biotic, gentamicin, and found that the bacterial
response was the same (that is, it was not drug
Research such as this — tracking how and
Escherichia coli — a test case
for antibiotic resistance.
NATURE|Vol 467|2 September 2010
NEWS & VIEWS
© 20 Macmillan Publishers Limited. All rights reserved10
which mutations develop in the presence Download full-text
of an antibiotic by isolating individual cells
from a bacterial population — is essential for
optimizing a dynamic strategy for prescribing
antibiotics. Such information can help in assess-
ing the need for changes in the dosage and dura-
tion of treatments, for example. The approach
also highlights the fact that inter action between
different communities of resistant mutants that
form in a bacterial population can enable them
to mount a more formidable defence against
antibiotics. Single-cell behaviour that is mark-
edly different from that at the population level
has been a subject of intense investigation in
systems biology. Lee et al. provide another
valuable example of such studies.
Furthermore, the collective behaviour of
single-celled organisms — as seen in the phe-
nomena of quorum sensing3 and metabolism
in a biofilm4, and now in antibiotic resistance
— shows that a pool of microbes can act in con-
cert. Apart from its implications for research
in tackling antibiotic resistance, the new work2
adds to previous studies in challenging the
conventional definition of what constitutes a
Hyun Youk and Alexander van Oudenaarden
are in the Departments of Physics and of
Biology, Massachusetts Institute of Technology,
Cambridge, Massachusetts 02139, USA.
1. Martínez, J. L., Banquero, F. & Andersson, D. I. Nature Rev.
Microbiol. 5, 958–965 (2007).
2. Lee, H. H., Molla, M. N., Cantor, C. R. & Collins, J. J. Nature
467, 82–85 (2010).
3. Bassler, B. L. & Losick, R. Cell 125, 237–246 (2006).
4. O’Toole, G., Kaplan, H. B. & Kolter, R. Annu. Rev. Microbiol.
54, 49–79 (2000).
Unexpected warm water
The detection of water vapour in a carbon star has challenged the
understanding of ageing stars. The discovery that such water can be
warm shows that our knowledge of these objects is still rudimentary.
In the short time since its launch on 14 May
2009, the European Space Agency’s Herschel
Space Observatory has delivered several
astronomical discoveries in the infrared and
submillimetre regions of the electromagnetic
spectrum. On page 64 of this issue, Decin
et al.1 report yet another of Herschel’s exciting
findings: the detection of warm water vapour
in the circumstellar envelope of the carbon star
Carbon stars were first recognized in the
1860s by William Huggins and Angelo Secchi.
On the basis of his purely visual inspection of
spectroscopic observations, Secchi defined this
new group of objects as class IV in his classifica-
tion system of stellar spectra. These extremely
red stars, he noted2, were remarkably different
from the orange stars of class III: “we cannot
identify precisely the sources of the lines and
the bands. We can say however that there is a
marked analogy with the reversed spectrum
of carbon.” Further observations by Hermann
Carl Vogel and Nils Dunér later strengthened
The distinctive spectral features that char-
acterize carbon stars — notably the dominant
spectral bands of carbon compounds, such as
the C2 Swan bands in the green part of the spec-
trum, and the lack of bands from oxides such as
TiO and H2O, which are characteristic of other
types of cool star — is due to their atmospheres
being richer in carbon than in oxygen, as was
suggested by Charles Donald Shane3 and dem-
onstrated by Henry Norris Russell4. If there is
more carbon than oxygen, oxygen is mostly
bound to carbon in the form of carbon monox-
ide (CO) because the molecule has a high bind-
ing energy (11 electronvolts). As a result, little
oxygen is left free to form other oxides in such
stellar atmospheres, whereas carbon atoms are
available to form other carbon compounds. By
contrast, in normal stars such as the Sun, the
atmosphere contains more oxygen than carbon
and the opposite occurs: carbon-containing
molecules other than CO become rare.
During the 1950s, investigators showed that
a peculiar class of ageing red giant stars known
as asymptotic giant branch (AGB) stars — to
which carbon stars belong — have important
roles in nucleosynthesis processes. For example,
the heavy ‘s-elements’ found in the Milky Way
(so called because they are created by relatively
slow, hence the ‘s’, neutron capture by heavy
atomic nuclei), as well as nitrogen and carbon5,
are believed to be produced in AGB stars and
later expelled into the inter stellar medium. But
the details of these processes have remained
unclear: we still lack a complete understanding
of key mechanisms, not least of those that drive
the intensive winds from such stars.
There is therefore good reason to study car-
bon stars — not least, as we shall see, the Milky
Way’s pulsating star IRC +10216. This visually
faint, extended object is, as seen by an observer
on Earth, the brightest source outside the Solar
System in the 5-micrometre waveband6. Radio
observations have demonstrated that the opti-
cally thick, dusty shell that surrounds the star
is a rich source of complex molecules. More
than 70 molecular species, which are charac-
teristic of a carbon-rich chemistry, had already
been detected there before the Herschel era7.
In fact, about 50% of all molecules observed in
astronomy have been detected in this object8.
Data from the IRAS infrared satellite have
shown9,10 that about 4% of all Galactic car-
bon stars have signs of silicate grains in their
expanding circumstellar envelopes. This sug-
gested — contrary to what one would expect
under thermal-equilibrium conditions and
for well-mixed gas mixtures — that the gas
envelopes of these carbon stars contain both
oxygen-rich and carbon-rich material, thereby
posing a challenge to the conventional under-
standing of the chemistry and evolution of
ageing stars. One possible explanation for this
astonishing observation is that the stars’ evo-
lutionary transition from an oxygen-rich to a
carbon-rich phase occurred quite recently, so
that the remains of previous epochs can still be
traced in the stars’ outer envelopes. The sub-
sequent detection11, with the SWAS submilli-
metre satellite, of circumstellar water vapour in
IRC +10216 — a characteristic of oxygen-rich
stars — has caused further astonishment.
This observation11 was based on the iden-
tification of a single water-vapour spectral
line of low excitation, which corresponds to a
transition between two energy levels that are
populated even at low temperatures. The fact
that the line is a low-excitation one may suggest
that the water vapour originates from the outer,
cold regions of the stellar gas envelope. Another
possibility is that the vapour arises from the
vaporization of icy bodies, such as comets
or minor planets, in orbit around the star11,12.
In their study, Decin et al.1 identify not just
one but numerous water-vapour lines in the
spectra of IRC +10216 (see Fig. 1 on page 65).
However, many of these are high-excitation
lines, which — if the water molecules are ther-
mally excited — means that the temperature of
the gas in which the lines are formed is of the
order of 1,000 kelvin. These results point to the
existence of warm water vapour in the inner
regions of the stellar envelope, and seem to rule
out models — including the vaporization-of-
icy-bodies hypothesis — that posit that water
vapour originates only from the stellar enve-
lope’s cooler intermediate and outer regions.
The strength of the newly discovered lines
also goes against another hypothesis: that the
existence of water vapour in the envelope’s
inner regions is due to shock waves that are
induced by the star’s pulsation and generate the
non-thermal-equilibrium chemistry needed to
form water in a carbon-rich gas. The authors1
suggest, instead, that the non-thermal-equilib-
rium chemistry is the result of the penetration
of ultraviolet photons into the inner regions of
the envelope, possibly from the star but more
likely from interstellar space. But for these
hypotheses to work, a highly clumpy circum-
stellar envelope is required, so that enough
of the ultraviolet radiation penetrates into its
NATURE|Vol 467|2 September 2010
NEWS & VIEWS
© 20 Macmillan Publishers Limited. All rights reserved10