NEWS & VIEWS RESEARCH

respond to light ON and so are sensitive to light
edges, whereas T5 cells respond to light OFF
and are sensitive to dark edges. The authors’
genetic-knockout experiments not only con-
firm this optical observation but also show
that T4 and T5 are the sole pathways mediat-
ing these functions, with no other cells being
able to step in and carry the message. Thus, a
fly initially breaks down moving visual inputs
into a total of eight components: bright edges
moving up, down, forward or backward, and
dark edges moving along the same four axes.
But how do T4 and T5 actually detect the
direction of movement? Takemura and collab-
orators’ fly connectome suggests a place to look
for the answer. They show that just upstream of
T4 lies a pair of neurons, termed Mi1 and Tm3,
which report on narrowly separated points in
visual space. Because of that separation, the
pair could provide the inputs that T4 needs to
discriminate direction (Fig. 1b). Using Maisak
and colleagues’ recording technology, it may
be possible to optically record from Mi1 and
Tm3. If all goes well, this could bring the
50-year search for the mechanism of direction
selectivity to an end.
Very many structural problems could be 1. Helmstaedter, M. et al. Nature 500, 168–174 (2013).
attacked using the same original material. ■ 2. Takemura, S. et al. Nature 500, 175–181 (2013).
3. Maisak, M. S. et al. Nature 500, 212–216 (2013).
4. Masland, R. H. Neuron 76, 266–280 (2012).
Richard H. Masland is in the Departments 5. Wässle, H., Puller, C., Müller, F. & Haverkamp, S.
of Ophthalmology and Neurobiology, J. Neurosci. 29, 106–117 (2009).
Harvard Medical School, Boston, 6. Reichardt, W. in Sensory Communication
(ed. Rosenblith, W. A.) 303–317 (MIT Press, 1961).
Massachusetts 02114, USA. 7. Barlow, H. B. & Levick, W. R. J. Physiol. (Lond.) 178,
e-mail: richard_masland@meei.harvard.edu 477–504 (1965).
SO L A R SYST E M
Saturn’s tides control
Enceladus’ plume
Data obtained by the Cassini spacecraft show that the plume of ice particles at the
south pole of Saturn’s moon Enceladus is four times brighter when the moon is
farthest away from the planet than when it is closest. See Letter p.182
JOHN SPENCER relationship between the brightness of the plume
and Enceladus’ position in its orbit around

The connectomic approach has now proved
its importance for studying fly eyes and mouse
retinas, but sceptics will still doubt that we can
make the jump from these miniature neuronal
circuits to ‘real brains’; the intrinsic circuits of
the cerebral cortex are some ten times larger
than those of the retina, and this spatial scale
is dwarfed again by the distances that connect
different brain regions.
One obstacle is the need for very large tissue
sections. Another difficulty is that of image
segmentation, which is required for tracing
thin neuronal processes through the thicket
A
n analysis of images of Saturn’s moon
Enceladus taken from the Cassini
Saturn orbiter, reported by Hedman
et al.1 on page 182 of this issue, shows that
the output of the giant plume of ice particles,
which jets out of fractures in the south polar
region of the moon, is controlled by diurnal
changes in the tidal stresses from Saturn*. The
authors find a remarkably strong and simple
*This article and the paper under discussion1 were
published online on 31 July 2013.
Saturn, providing dramatic confirmation of
predictions made in 2007 from a tidal-stress
model2.
Tidal forces from Saturn are the ultimate
power source for the extravagant geological
activity on Enceladus3, a small ice world just
500 kilometres in diameter. Enceladus’ 1.37-
day orbit around Saturn is slightly eccentric,
as a result of the periodic gravitational influ-
ence of the larger moon Dione. The daily vari-
ations in the tidal stresses from Saturn due to

SPACE SCIENCE INSTITUTE/JPL/NASA

of neighbours in serial sections. Because digi-
tal solutions have failed, the task is currently
assigned to large teams of human observers,
but this will be impractical on larger spatial
scales. Improvements in fixation and stain-
ing might make the processes more easily
discriminable, and digital technology may yet
save the day, because in principle any task that
can be done by human observers can be done
by a computer. Tracing processes is essentially
a problem of pattern recognition, for which the
technology is evolving rapidly.
A final question concerns the cost-effective-
ness of the connectomic approach. Is it to be
the exclusive province of a few deep-pocketed
laboratories? Here, the answer is clear: the
researchers involved have stressed that the
connectomic reconstructions will be public
resources, which can be used for different pur-
poses by anyone. To be useful, an archive may
need to be accompanied by a user-oriented
interface, because computer code this com-
Figure 1 | Enceladus’ ice-particle jets.  The image shows jets of ice particles emerging from the four
plex will be hard for workers other than its ‘tiger-stripe’ fractures at Enceladus’ south pole, photographed in visible light by cameras on the Cassini
creators to use. The effort involved in creating Saturn orbiter in 2009. Hedman and colleagues’ analysis1 of hundreds of lower-resolution infrared
and curating such a public resource should be images, taken by Cassini’s VIMS instrument, of the plume that results from the combination of these
worth it, because the archives could be the big- jets, reveals that the plume responds to daily changes in the tidal stresses from Saturn on the fractures,
gest contribution of this work to neuro­science. brightening when the fractures are in tension.

8 AUG U S T 2 0 1 3 | VO L 5 0 0 | NAT U R E | 1 5 5
© 2013 Macmillan Publishers Limited. All rights reserved
 RESEARCH NEWS & VIEWS
50 Years Ago
A hazard to the animal that has
always accompanied war is when
persons unaccustomed to animals
are drafted to animal units; in the
Second World War this was a very
common occurrence, because by
then, so few persons had experience
with horses. Many loved the job
and did it well, but others were
probably disappointed in a non-
mechanical role and not very
efficient. Mules reported to be
vicious were often underworked
and overfed: in any event for a tyro
to be placed in charge of an army
mule could be described as an
experience.
From Nature 10 August 1963
100 Years Ago
A very remarkable red-water
phenomenon is at present
observable in a small pond in
Broad Oak Park, Worsley, near
Manchester, just in front of the
seventh tee on the golf course. The
surface of the pond — at any rate
at times — is covered in places
with an almost blood-red scum,
which seems to float on the surface
film like fine dust. The scum
sometimes assumes a greenish hue.
Microscopical examination shows
that it is due to the presence of
immense numbers of a large species
of Euglena, the green chlorophyll
of which … is more or less replaced
by red haematochrome … Since
writing the above I have been able
to observe how the Euglenae reach
the surface of the water. They
evidently secrete some sort of slime
in which they become entangled.
Bubbles of oxygen gas, given off
by the Euglenae in the presence
of sunlight, are also caught in this
slime, and when these reach a
certain size they rise to the surface,
trailing strings of slime, with
numerous entangled Euglenae,
after them.
From Nature 7 August 1913
1 5 6 | NAT U R E | VO L 5 0 0 | 8 AUG U S T 2 0 1 3
this eccentricity distort Enceladus and dump
gigawatts of frictional heat into its interior. This
heat powers the ejection of the plume of water
vapour and ice particles, discovered by Cassini
in 2005, from four parallel warm fractures
called ‘tiger stripes’, which surround the moon’s
south pole (Fig. 1).
Enceladus is thus one of the few places
beyond Earth where we can watch geology
happen in real time, giving us a primer for
understanding other, less active, icy worlds.
The ice particles in the plume are salty, hint-
ing at a subsurface source of liquid water and
a probable subsurface ocean4, and the plume
gases include complex hydrocarbons and other
organic compounds5. The likely presence of
liquid water and complex organic chemistry
makes Enceladus especially intriguing as a
potential habitat for extraterrestrial life, pro-
viding additional motivation for investigating
its interior.
The 2007 modelling study2 noted that the
rhythmic tidal stresses that power Encela-
dus’ activity would also put the tiger-stripe
fractures into alternating states of tension
and compression during each orbit of Saturn.
The study predicted that the majority of the
tiger stripes would be in compression when
Enceladus is closest to Saturn (periapse), and
in tension when it is farthest away (apoapse).
Tensional stresses would plausibly open
pathways for the venting of plume gases and
particles, and thus increase plume activity at
apoapse.
Cassini has so far made 20 close fly-bys of
Enceladus to investigate its surface and inter­ior
and to sample its plume. But the plume is large
enough and bright enough to be seen by remote
sensing from longer range, permitting more
frequent study both with Cassini’s visible-wave-
length cameras and with its Visual and Infrared
Mapping Spectrometer (VIMS). Hints of the
predicted variability in individual plume jets
were seen in an early study6, using a relatively
small number of images from Cassini’s cam-
eras. Hedman and colleagues’ work, which is
based on VIMS infrared images, was able to
reach more definitive conclusions, thanks to
systematic analysis of the spectro­meter’s much
larger data set of 252 plume images.
The large data set allowed the team to tease
apart orbit-related temporal variations in
plume brightness from possible brightness
variations caused by long-term changes and by
illumination geometry (the micrometre-sized
ice particles in the plume brighten greatly when
backlit). The temporal variations revealed by
this analysis are simple and dramatic — the
plume is consistently about four times brighter
when Enceladus is at apoapse than when it is at
periapse, just as predicted by the 2007 model2.
This provides strong evidence that the tiger-
stripe fractures do indeed open and close
each day in response to diurnal tidal stresses,
controlling the plume in the process.
Geology, dealing as it does with the complex
© 2013 Macmillan Publishers Limited. All rights reserved
behaviours and long memories of materials in
the solid state, tends to be a messy business.
So it is always startling and instructive when
simple patterns like this emerge. The strength
of the plume–orbit relationship is a strong con-
straint on how Enceladus works, and a valu-
able guide to future research. For instance, the
2007 tidal-stress model treats Enceladus’ crust
as a thin elastic shell that is detached from the
interior by a fluid layer, such as a global ocean.
Global-ocean models have fallen out of favour
for Enceladus, because it is difficult to keep a
global ocean from freezing7, and a regional
south polar ocean8 is now considered more
likely. Such refined models can now be tested
and constrained by their ability to match the
observed plume behaviour.
Also, Cassini’s optical cameras, with
50 times higher spatial resolution than that of
VIMS, can provide more precise constraints
on models of internal structure by tracking
the variability of the individual jets that make
up the plume9. Jet variability can thus provide
knowledge of local stresses on the fractures
from which the jets emanate, giving another
means of investigating local subsurface condi-
tions. The VIMS study itself hints at changes
in plume ejection speed at different positions
in the orbit, offering another handle on how
the plume reaches the surface. No other world
beyond Earth allows such detailed analysis of
active geophysical processes.
Enceladus displays other strikingly clear-cut
patterns. For example, its several geological
provinces, defined by age and style of surface
deformation, are arranged with almost perfect
symmetry around its spin axis and the direc-
tion of Saturn10. Equally strange is the geo-
metric simplicity of the four active tiger-stripe
fractures, all roughly the same length (about
130 km), and evenly spaced about 35 km apart.
As with the plume behaviour, such simple pat-
terns must point to important truths, but these
other puzzles remain mysterious, and defini-
tive explanations await future research. ■
John Spencer is at Southwest Research
Institute, Boulder, Colorado 80302, USA.
e-mail: spencer@boulder.swri.edu
1. Hedman, M. M. et al. Nature 500, 182–184 (2013).
2. Hurford, T. A., Helfenstein, P., Hoppa, G. V.,
Greenberg, R. & Bills, B. G. Nature 447, 292–294
(2007).
3. Spencer, J. R. & Nimmo, F. Annu. Rev. Earth Planet.
Sci. 41, 693–717 (2013).
4. Postberg, F., Schmidt, J., Hillier, J., Kempf, S. &
Srama, R. Nature 474, 620–622 (2011).
5. Waite, J. H. et al. Nature 460, 487–490 (2009).
6. Spitale, J. N. & Porco, C. C. Nature 449, 695–697
(2007).
7. Roberts, J. H. & Nimmo, F. Icarus 194, 675–689
(2008).
8. Collins, G. C. & Goodman, J. C. Icarus 189, 72–82
(2007).
9. Porco, C., DiNino, D. & Nimmo, F. 44th Lunar Planet.
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