range of substrates if they have time-sharing of

carboxylates in common with these substrate-

specific antiporters?
The answer could lie in the differences in
accessibility and local environment of cata-
lytic carboxylates in the different systems.
By analogy with crystallized multidrug-
binding transcriptional regulators
14
, catalytic
carboxy lates in secondary-active multidrug
transporters are most probably found in a
large, flexible substrate-binding surface, where
they cooperate with different sets of polar and
non-polar amino-acid residues in order to
bind structurally dissimilar substrates
8
. To
create such a binding surface in the inward-
facing conformation of a transporter, a large
part of the protein must change conforma-
tion. This causes the turnover number of the
transporter the number of molecules of
substrate that can be transported per minute
to be relatively low (14 to 1,500 per minute,
depending on the transporter and substrate), a
characteristic that facilitates the binding of large
organic molecules that have low diffusion rates
and low rates of association and dissociation
at binding sites.
By contrast, the catalytic carboxylates in
the NhaA antiporter are located between two
funnels in the protein that provide access
from the cytoplasm and the cells exterior to the
centre of the cell membrane, at a selective site
that can accommodate only Na
+
or Li
+
(ref. 13).
The selectivity of NhaA is probably supported
by fast, local conformational changes near the
binding site and by a high turnover number
(100,000 ions per minute) that is compatible
with the high diffusion rates of small inorganic
ions. Although the first examples of multi drug-
binding surfaces bound to substrates are avail-
able from crystal structures of the bacterial
RND transporter AcrB (ref. 15) and the mouse
ABC transporter ABCB1a (refs 8, 16), the anal-
ogous structural information from crystallized
MATE and MFS members is still missing. A
high-resolution structure of inward-facing
NorM bound to a substrate will therefore be
extremely valuable. A related question follow-
ing on from this is how the Na
+
selectivity of
outward-facing NorM is maintained.
The crystal structure
1
of NorM points to a
conservation of general mechanisms of trans-
port among structurally unrelated multidrug
transporters, and raises several questions. How
does Na
+
binding to NorM switch the proteins
conformation from outward-facing to inward-
facing, and what is the sequence of events that
leads to re-establishment of the outward-facing
state and release of substrate on the external
side of the membrane? How many Na
+
ions are
transported per substrate, and is this stoichi-
ometry fixed, or variable and dependent on the
chemical structure of the substrate
1012
? And
do translocation pathways in NorM allow the
transport of lipid-soluble substrates from the
cell membrane, as suggested by its crystal struc-
ture? He and colleagues work
1
will certainly
encourage further structural, biochemical,
biophysical and modelling studies on MATE
proteins in the quest for a molecular descrip-
tion of the dynamic processes under lying
substrate transport. Such a description would
not only satisfy scientists academic interest.
It would also be crucial for developing modu-
lators that bind to and inhibit the activity of
MATE proteins and other multi drug transport-
ers, thus providing a solution to the increasing
problem of multidrug resistance.
Hendrik W. van Veen is in the Department
of Pharmacology, University of Cambridge,
Cambridge CB2 1PD, UK.
e-mail: hwv20@cam.uc.uk
1. He, X. et al. Nature 467, 991994 (2010).
2. Omote, H., Hiasa, M., Matsumoto, T., Otsuka, M. &
Moriyama, Y. Trends Pharmacol. Sci. 27, 587593
(2006).
3. Brown, M. H., Paulsen, I. T. & Skurray, R. A. Mol.
Microbiol. 31, 394395 (1999).
4. Saier, M. H. Jr & Paulsen, I. T. Semin. Cell
Dev. Biol. 12, 205213 (2001).
5. Yamashita, A., Singh, S. K., Kawate, T., Jin, Y. &
Gouaux, E. Nature 437, 215223 (2005).
6. Shimamura, T. et al. Science 328, 470473
(2010).
7. Khare, D., Oldham, M. L., Orelle, C., Davidson, A. L.
& Chen, J. Mol. Cell 33, 528536 (2009).
8. Gutmann, D. A. P., Ward, A., Urbatsch, I. L., Chang, G.
& van Veen, H. W. Trends Biochem. Sci. 35, 3642
(2010).
9. Morita, Y., Kataoka, A., Shiota, S., Mizushima, T.
& Tsuchiya, T. J. Bacteriol. 182, 66946697
(2000).
10. Yerushalmi, H. & Schuldiner, S. Biochemistry 39,
1471114719 (2000).
11. Sigal, N., Fluman, N., Siemion, S. & Bibi, E. J. Biol.
Chem. 284, 69666971 (2009).
12. Schaedler, T. A. & van Veen, H. W. FASEB J. 24,
36533661 (2010).
13. Padan, E. Trends Biochem. Sci. 33, 435443
(2008).
14. Schumacher, M. A. et al. Science 294, 21582163
(2001).
15. Murakami, S., Nakashima, R., Yamashita, E.,
Matsumoto, T. & Yamaguchi, A. Nature 443,
173179 (2006).
16. Aller, S. G. et al. Science 323, 17181722 (2009).
ATMOSPHERI C PHYSI CS
Chorus keeps the
diffuse aurora humming
The origin of the diffuse aurora, whose beauty and intensity pale beside those
of the famous aurora borealis, has remained controversial. A convincing
explanation for this auroral display is now at hand. S L .943
PAT RI CK T. NE WE L L
T
he diffuse aurora, which is hard to see
and has long been neglected despite its
great importance to the energy budget
of the upper atmosphere, has been attract-
ing greater interest. On page 943 of this issue,
Thorne and colleagues
1
have finally solved
the decades-old mystery of how the diffuse
aurora manages to keep going: a type of very
low-frequency plasma wave called a chorus
wave does the job.
The diffuse aurora is created when charged
particles (mostly electrons) that surround
Earth as part of its magnetosphere collide
with the ionosphere, the ionized part of the
upper atmosphere. Although most such elec-
trons do not reach the atmosphere, some do.
Those electrons that can reach the ionosphere
are rapidly lost to it, along with their energy.
Because the electrons in question have tem-
peratures of between about 1 million and
100 million degrees centigrade, they zip along
at speeds that should empty the loss cone that
allows them to reach the ionosphere in just a
few seconds (electrons in the loss cone have
nearly all their velocity parallel to Earths mag-
netic field). Space physicists have long assumed
that some type of strong scattering process
was at work, constantly shifting new electrons
from outside the loss cone into it. In fact, this
hypothesis of strong scattering of pitch angle
(the angle between the electrons trajectory
and the direction of the magnetic field) has
been accepted for decades. Now Thorne and
colleagues
1
have convincingly demonstrated
what causes the strong scattering.
Figure 1 (overleaf) shows an auroral display
that includes bright, multicoloured curtains of
light to the left and centre. This is the discrete
aurora, or the famous aurora borealis, which
produces displays that are impressive even to
a casual observer on the ground. At the bottom
right of the picture is a dull green cloud. This
is the diffuse aurora, and although it can be
seen from the ground (with much more dif-
ficulty than the discrete aurora), it lacks the
dynamic shifts in form and intensity that are
characteristic of its more photogenic sibling.
Yet it turns out that the diffuse aurora, which
is always present, and is always widely distrib-
uted in rings around Earths magnetic poles,
collectively accounts for about three-quarters
of the auroral energy precipitating into the
iono sphere
2
. This drab aurora has most of
the energy flux simply through its persistence
2 1 O C T O B E R 2 0 1 0 | V O L 4 6 7 | N A T U R E | 9 2 7
NEWS & VIEWS RESEARCH
20 Macmillan Publishers Limited. All rights reserved 10
50 Years Ago
The discovery that some errors
of sexual development and
other congenital defects in man
were associated with karyotypic
abnormalities has aroused
considerable interest, and research
in this field is growing rapidly
Various systems of numbering of
the chromosomes have been used by
different research groups, making the
comparison of their results difficult.
A meeting with the object of trying
to devise an agreed nomenclature
was therefore arranged by
Dr. T. T. Puck, of the University of
Colorado, and held in Denver during
April 911 A system was agreed
on and a statement embodying it
and other information relating to
the identification of human mitotic
chromosomes has been published
(Lancet, i, 1063; 1960) It is hoped
that the proposed standard system
will be generally adopted, and that
any who may prefer to use another
scheme will refer to the proposed
system and show clearly in what way
their own system differs.
From Nature 22 October 1960
100 Years Ago
Fellows of the Zoological and other
scientific societies, museum officials
and others, are warned against an
individual representing himself
as a consumptive and asking for a
recommendation to a hospital and
temporary help. The modus operandi
is to call upon you with a bogus
introduction from another fellow
of your society or someone known
to you, and to mention a few other
well-known persons as interested in
his case. The individual is rather tall,
thin, of wan appearance, and has a
dark moustache. His manner shows
some refinement and education,
and is also persuasive, as proved by
the number of those known to have,
unfortunately, been victimised by
his false representations.
From Nature 20 October 1910
and geographical extent. It reduces the accu-
racy of navigation using the Global Positioning
System, affects upper-atmospheric, high-
latitude chemistry (for example, it produces
copious quantities of nitric oxide), and is in fact
largely responsible for creating the ionosphere
during darkness, when solar ultraviolet light
cant do the job of separating electrons from
neutral atoms.
In recent years, and largely on the basis
of theoretical studies, two distinct types of
plasma wave electrostatic electron cyclo-
tron harmonic (ECH) waves and whistler-
mode chorus waves have been the leading
candidates for the source that keeps the diffuse
aurora going. But uncertainty has persisted
because of the intricacies of the problem. Most
researchers assumed that both types of wave
contributed substantially. Rather surprisingly,
Thorne et al.
1
have been able to provide a fairly
definitive answer to the contrary.
To do so, they combined advances that
included observations, theory and modelling.
The satellite CRRES has gathered detailed
data on the power of plasma waves. But
more work was needed, including detailed
electron-scattering calculations. Doing those
correctly, in turn, requires knowledge of
how the waves propagate (ray tracing). Thorne
and colleagues have managed to put all of
these pieces together and to produce detailed
simulation results clearly showing that chorus
waves can scatter enough electrons to maintain
the diffuse aurora, and that ECH waves cannot.
But how do we know that these calculations
and simulations can be trusted?
The proof is in the details. First, the results
correctly predict the gross morphology of
the diffuse aurora, including its peak in the
midnight-to-dawn sector. Perhaps even more
convincingly, the calculations actually account
for the observed fine details of the electron
pitch-angle distributions. In fact, the new
work can account for differences in lower- and
higher-energy particle behaviour over a wide
range of pitch angles. Thorne and colleagues
seem to have provided a compelling answer to
the question of why the bulk of auroral energy
precipitating into the ionosphere occurs when
and where it does, and why electrons in the
magnetosphere are distributed in pitch angle
in the way that they are.
In the 1950s, early rocket shots into the
auroral ionosphere recorded intense emissions
from plasma waves that had frequencies of
between a few hundred and several thousand
hertz, with complex changes in both inten-
sity and frequency occurring over a fraction
of a second. This reminded their discoverer,
Storey
3
, of a chorus of birds at dawn. Now,
more than half a century later, Thorne and col-
leagues have shown that it is this chorus that
keeps the diffuse aurora humming.
Patrick T. Newell is at the Applied Physics
Laboratory, Johns Hopkins University, Laurel,
Maryland 20723, USA.
e-mail: patrick.newell@jhuapl.edu
1. Thorne, R. M., Ni, B., Tao, X., Horne, R. B. & Meredith,
N. P. Nature 467, 943946 (2010).
2. Newell, P. T., Sotirelis, T. & Wing, S. J. Geophys.
Res. 114, A09207, doi:10.1029/2009JA014326
(2009).
3. Storey, L. R. O. Phil. Trans. R. Soc. Lond. A 246,
113141 (1953).
Figure 1. The discrete aurora includes the spectacular curtains and rays seen left and center. To the bottom right is the difuse aurora,
a faint green cloud lacking the dynamic changes in intensity and morphology of the discrete aurora. However the difuse aurora
actually contributes more to the global energy budget, because of its persistence.
Photograph courtesy of Dr. M. Yamaguchi of the Swedish Institute of Space Physics.
Figure 1 | Spot the diffuse aurora. The discrete aurora includes the spectacular curtains and rays
seen on the left and centre of this image. At the bottom right is the diffuse aurora, a faint green
cloud that lacks the dynamic changes in intensity and morphology of the discrete aurora. However,
the diffuse aurora actually contributes more to the global energy budget of the upper atmosphere
than does the discrete aurora because of its persistence and geographical extent. Thorne and
colleagues
1
demonstrate that a type of plasma wave called a chorus wave keeps the diffuse aurora going.
(Photo courtesy of M. Yamaguchi, Swedish Institute of Space Physics.)
9 2 8 | N A T U R E | V O L 4 6 7 | 2 1 O C T O B E R 2 0 1 0
NEWS & VIEWS RESEARCH
20 Macmillan Publishers Limited. All rights reserved 10
Reproducedwith permission of thecopyright owner. Further reproductionprohibited without permission.