Current Biology
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7. Shadmehr, R., Smith, M., and Krakauer, J.
(2010). Error correction, sensory prediction,
and adaptation in motor control. Annu. Rev.
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(1985). Neural, mechanical, and geometric
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(2005). Applications of prism adaptation: a
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Zoology: At Last an Exit for Ctenophores
Gonzalo Giribet
Museum of Comparative Zoology and Department of Organismic and Evolutionary Biology, 26 Oxford Street, Cambridge, MA 02138, USA
Correspondence: ggiribet@g.harvard.edu
http://dx.doi.org/10.1016/j.cub.2016.08.036
Ctenophores, one of the most basal branches in the tree of life, have been found to have a through-gut,
complete with mouth and anus. Basal animals are surprisingly complex and simplification has been
rampant in animal evolution.
It has been generally accepted that life
in general and animals in particular
evolved from simple forms to more
complex ones. Early evolving animals
lacked tissues or a basement membrane
isolating their internal milieu from the
environment. Then with tissue-level
integration, guts and nervous systems
evolved, and the gut, which started as a
simple blind sac-like structure, became
more complex, regionalized, and
acquired a second opening, with a
mouth and an anus. But does this
evolutionary sequence reflect what
really happened? In an elegant new
article in this issue of Current Biology,
Presnell and collaborators [1]
demonstrate that comb jellies (members
of the animal phylum Ctenophora)
possess a through-gut, with a mouth for
ingesting their prey and two fully-
functioning anal pores — the latter is the
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12. Choi, J.T., and Bastian, A.J. (2007).
Adaptation reveals independent control
networks for human walking. Nat. Neurosci.
10, 1055–1062.
13. Reisman, D.S., Bastian, A.J., and Morton, S.M.
(2010). Neurophysiologic and rehabilitation
insights from the split-belt and other locomotor
adaptation paradigms. Phys. Ther. 90,
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14. van Ingen Schenau, G.J. (1980). Some
fundamental aspects of the biomechanics of
overground versus treadmill locomotion. Med.
Sci. Sports Exerc. 12, 257–261.
15. Vazquez, A., Statton, M.A., Busgang, S.A., and
Bastian, A.J. (2015). Split-belt walking
adaptation recalibrates sensorimotor
estimates of leg speed but not position or
force. J. Neurophysiol. 114, 3255–3267.
16. Reisman, D.S., Wityk, R., Silver, K., and
Bastian, A.J. (2009). Split-belt treadmill
adaptation transfers to overground
key discovery. These anal pores were
already reported in the nineteenth
Century by Louis Agassiz [2] but had
been thought to be mostly non-functional.
Now, Presnell and collaborators [1]
were able to demonstrate the active use
of the anal pores in two ctenophore
species, Mnemiopsis leidyi and
Pleurobrachia bachei, kept in captivity
in laboratories on the east and west
coasts of the U.S. (Figure 1). This behavior
was also observed in free-living
specimens of a multitude of ctenophore
genera.
Comb jellies are mostly macro-pelagic
(from a few centimeters to almost 1.5
meters in body length) and exclusively
marine animals of a gelatinous
consistency, nearly all of which spend
their lives in the water column. A handful
of benthic species, including a sessile
one, are known. There are between
walking in persons poststroke.
Neurorehabil. Neural Repair 23,
735–744.
17. Mazzoni, P., and Krakauer, J.W. (2006). An
implicit plan overrides an explicit strategy
during visuomotor adaptation. J. Neurosci. 26,
3642–3645.
18. Reisman, D.S., Block, H.J., and
Bastian, A.J. (2005). Interlimb coordination
during locomotion: what can be adapted
and stored? J. Neurophysiol. 94, 2403–2415.
19. Finley, J.M., Bastian, A.J., and
Gottschall, J.S. (2013). Learning to be
economical: the energy cost of walking
tracks motor adaptation. J. Physiol. 591,
1081–1095.
20. Selinger, J.C., O’Connor, S.M., Wong, J.D.,
and Donelan, J.M. (2015). Humans can
continuously optimize energetic cost
during walking. Curr. Biol. 25,
2452–2456.
150 and 250 described species, but
due to their gelatinous nature, they are
difficult to collect and study, and thus
many remain to be formally described.
Ctenophores have been reported to live
at the water’s surface, where they can
be locally abundant, to depths exceeding
7000 meters.
Ctenophores are the largest animals
to swim by ciliary action. For this, they use
their characteristic rows of
iridescent ciliary plates (hence their name
combs or ctenes) that are present at least
during some part of their life cycle. In
addition to ciliary action, ctenophores can
also move by muscular contraction of
their lobes, or use their musculature to
withdraw the retractable tentacles into
tentacular sheaths. They are carnivorous,
feeding on other small pelagic and
planktonic animals, often captured by a
pair of tentacles loaded with a special
Current Biology
Dispatches
Figure 1. Ctenophores with a full belly.
Cultured Mnemiopsis leidyi: adult (left) and 9-day post hatching mid-transition between juvenile cydippid form and adult form (right). The white particles in the
center of the body constitute the gut contents. Photos: William E. Browne.
type of sticky cells called ‘colloblasts’,
and ingested alive.
Based on all these characteristics,
ctenophores have been considered
relatively complex animals — they have
discrete muscles and a diffuse but
highly integrative nervous system — at
least when compared to other basal
offshoots of the animal tree of life, such
as placozoans, sponges and cnidarians
(jelly fishes, anemones, corals, etc.).
In fact, some authors considered them
to be more closely related to the
‘higher animals’ or bilaterians due to
their complex anatomy [3]. However,
phylogenetic evidence based on
molecular data has unambiguously
shown that ctenophores ought to be more
basally placed than originally thought,
either as the sister group to all other
extant animals [4–8] or the first offshoot
after the divergence of sponges from
other animals [9–11]. Irrespective of which
of these positions is correct, all modern
studies suggest that placozoans, which
are the anatomically simplest animals,
and cnidarians, which lack a through-
gut, are more derived phylogenetically
than the more complex ctenophores.
However, due to their phylogenetic
position — ctenophores were once
regarded to form a common clade called
‘Coelenterata’ together with cnidarians —
it was assumed that their digestive
system, despite having two aborally-
positioned (at the opposite end to the
mouth) anal pores, functioned like those
of cnidarians, i.e., the food was ingested
by the mouth and the digested remains
were also expelled by the same oral
opening.
The new work by Presnell and
colleagues [1] elegantly shows that
the ingested food passes through the
different sections of the ctenophore’s
complex digestive tract, and that after
being digested, its remains are actively
evacuated through the anal pores.
Food remains are only expelled through
the mouth under specific stressful
conditions similar to those leading to
vomiting in many other animals, including
humans. The normal path followed by
the ingested prey of these voracious
animals is thus unidirectional, as in
most bilaterians.
The implications of this discovery are
numerous. If ctenophores are the sister
group to all other animals, as many
phylogenetic studies still indicate, the
extant representatives of this lineage thus
constitute fairly complex animals, with a
Current Biology 26, R913–R936, October 24, 2016 R919
unique nervous system that probably
evolved in parallel to that of other
animals, muscles and a through-gut [6,8].
This would provide further evidence for
secondary simplification in sponges
and placozoans, and a convergence in
body plan with cnidarians. Some of
these implications would still stand even
if sponges were more basally placed than
ctenophores. Nonetheless, a question
remains with respect to the origins of the
anus in ctenophores and that of a
clade of bilaterian animals called
‘Nephrozoa’ — all bilaterally symmetrical
animals, except a small clade called
Xenacoelomorpha [12]. Nephrozoans
were thought to have incorporated two
anatomical innovations to their body
plan: discrete excretory organs for
eliminating nitrogenous waste more
efficiently (e.g., different types of
nephridial organs, including the kidney),
and the through-gut with mouth and
anus. Whether the anus of ctenophores
is homologous to that of nephrozoans
remains unanswered.
The specific series of events by which
principal features of animal bodies
evolved is still missing key pieces, but it
now seems clear that Darwin’s ‘‘from so
simple a beginning’’ may not necessarily

apply to the base of the animal tree of
life. Instead, placozoans and cnidarians
are probably simpler than some of their
ancestors (i.e., the common ancestor they
shared with ctenophores). This should
actually not come as a surprise following
the description of many independent
cases of extreme simplification in
parasitic metazoans, as exemplified by
orthonectids [13] and myxozoans [14].
Ctenophores and their troubled
phylogenetic position may hold the key to
resolving some of these recalcitrant
mysteries.
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Episodic Memory: Rats Master Multiple Memories
William A. Roberts
Department of Psychology, Western University, London, Ontario N6A 5C2, Canada
Correspondence: roberts@uwo.ca
http://dx.doi.org/10.1016/j.cub.2016.08.042
A new study in which rats had to discriminate odors according to whether they were novel for a
particular environmental context has found that they can accurately discriminate a large number of odors
and multiple context transitions, suggesting that they are able to form and remember multiple episodic
memories.
People show phenomenal memory for
pictures: when shown a list of over 2000
pictures, each briefly exposed, people
accurately recognize seen pictures from
unseen pictures with over 90% accuracy
on a subsequent test [1]. Such an ability
is undoubtedly underlain by a primate
visual system highly developed for the
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If a person is exposed to a sequence of
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images will seem familiar because they
were first seen earlier in the list, and more
recently seen faces will seem more
familiar. The presentation of an historical
figure, say Abraham Lincoln, will retrieve
semantic or reference memories of
general information about slavery and the
American civil war. Still another picture in
the sequence might be that of a close
friend: this image retrieves multiple
episodic memories of shared experiences
Current Biology
Dispatches
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that occurred at different times and in
different contexts. In this issue of Current
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that rats too can remember multiple
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contexts.
It is widely acknowledged that
animals form semantic memories, as
they learn to make Pavlovian responses
to conditioned stimuli or to make
instrumental responses of lever pressing