Current Biology
Magazine
the exact same feeling as seeing
someone on a horse charging at me
with a big stick. And a former, much
braver jousting partner is my non-
scientist hero, because he (Barchan)
followed his dream, is still jousting,
and may be the oldest competitive,
full-contact jouster in the world.
Speaking of heros, do you have a
scientific hero? I have many. Darwin
(of course) but also B.F. Skinner for
being the Darwin of behavior. Most
recently, Michael Faraday has become
a personal hero for his insights into
electricity without a math background
and for predicting some of my results
in electric eels. It took me months
using high-speed cameras and kHz
data sampling rates to determine
that electric eels cause hidden prey
to twitch involuntarily, revealing
themselves to the eel. Faraday came
to a similar conclusion using his bare
hands in 1838. For living scientists it
is my friend, postdoctoral mentor, and
now colleague, Jon Kaas. Everyone
knows he’s a brilliant neuroscientist.
But he manages that while being
the most well-balanced, supportive,
positive, and fun scientist I have ever
met. It’s a delight to just sit across the
table and talk with him.
There’s a story about you in Seed
Magazine that includes practical
jokes; what’s the deal? I’m not a
big practical joker, but I know an
opportunity when I see it. My best
friend from college became a state
trooper after graduation (so many
possibilities…). I had long hair and
a beard at the time, and he looks
like the quintessential scary cop
(sorry Tom). So no one would guess
we were friends. When the Annual
Neuroscience Meeting was in D.C.
in 1992, I told my roommates that
we could get behind the scenes of
the National Zoo in exchange for
pot (actually a bag of spices from
my parent’s kitchen). Off we went
while Tom waited in his squad car.
He pulled us over, found the bag,
and said we would all go to jail.
When he ‘found out’ we were grad
students attending a neuroscience
meeting, he softened but wanted
proof. The proof required answering
obscure neuroanatomy questions that
no trooper should know. Everyone
R6 Current Biology 27, R1–R18, January 9, 2017 © 2017 Elsevier Ltd.
did their best until he broke down
laughing, revealed the joke, and
blamed me for coming up with the
idea. I was relieved that it ended like
a Candid Camera episode and no-one
seems to have PTSD.
What is the best advice you’ve been
given? Glenn Northcutt’s advice: “It’s
easy to come up with questions, but
you need to know which questions
you can answer”. These days I would
add, when you hit a wall, sometimes
you can work on something else until
new technology solves your problem.
(Can’t afford a slow motion camera?
Use your iPhone.)
What advice would you give to
students at the beginning of their
career? There are so many angles
to this question the answers could
make a book. Here are some potential
chapters. Absorb as many useful
skills and strategies as possible,
but also notice what people do
wrong. Don’t take a position with an
unsupportive mentor just to learn
something (I speak from observation,
not experience). Take some time to
study yourself. What’s the difference
between a day begun with email and
a day begun writing, reading, or doing
an experiment? Manage your digital
distractions — where would we be
if Einstein had Facebook? (PIs can
smugly manage distractions because
their ratio of party invitations to
committee assignments is small.) On
a broader note, realize that successful
scientists present and write about
their successes: this masks the heap
of failures that most science is built
upon. Keep this in mind when things
go wrong. When you start out try to
choose projects for which any result
is interesting and publication is not
too distant. Don’t be afraid to start out
describing things, you will discover
something interesting along the way.
If you see a golden opportunity, grab
it. But if you don’t yet have the tool-kit
to pursue the opportunity, start a list
of ideas for the future. Collect data as
often as possible, but don’t forget to
analyse it and write it up.
Department of Biological Sciences,
Vanderbilt University, VU Station B, Box
35-163,4 Nashville, TN 37235-1634, USA.
E-mail: kenneth.catania@vanderbilt.edu
Quick guide
Anemonefishes
William E. Feeney1,2,*
and Rohan M. Brooker3
What are anemonefishes?
The term ‘anemonefish’ refers
exclusively to fishes of the subfamily
Amphiprioninae (Amphiprion spp. and
Premnas sp.), which, with exception
of a short larval dispersal period, live
their entire lives in association with
predatory Actiniarians, known as sea
anemones (Figure 1). It is perhaps
an unintuitive pairing, as anemones
use stinging nematocysts to capture
their prey, which can include fishes.
Unlike most fishes, anemonefishes
are able to mitigate the effects of
their anemone’s nematocysts and at
least some protect their host from
potential predators. In turn, some
fish-associated anemones spend
less time contracted when hosting
fishes, increasing their capacity to
feed and photosynthesise while
also providing their fishes with extra
protection from piscivorous predators.
The degree of host specificity varies
across anemonefish species, from
extreme generalists (e.g. Clark’s
anemonefish, A. clarkii, occupies
10 anemone species) to extreme
specialists (e.g. the spine-cheeked
anemonefish, P. biaculeatus, occupies
a single species of anemone) as does
the closeness of the relationship
across anemonefish–anemone pairs.
Ultimately, it is the intimate nature of
these associations coupled with their
variability across species pairs that
has led to these relationships being
amongst the best-studied examples
of inter-species mutualisms in nature.
Are anemonefishes always
associated with anemones? Yes, and
no. There are currently 28 recognized
anemonefish species in the subfamily
Amphiprioninae (as well as a few
naturally occurring hybrids), which
are obligate associates of 10 species
of sea anemones that are exclusively
distributed throughout the tropical
Indo-Pacific. However, there are also
at least 50 species of fishes that
have been reported to facultatively
associate with anemones. These
Current Biology

Magazine

species are characterised by their

taxonomic diversity (17 families) and
small body size. They are also much
more widely distributed than their
Amphiprioniae counterparts, with
representatives throughout the tropical
Indo-Pacific and Western Atlantic,
as well as the temperate North East
Pacific and Mediterranean.

How do fishes avoid being stung

by their host anemones? Not being
vulnerable to the nematocysts of
their common hosts, while not always
innate, is a ubiquitous attribute of
obligately anemone-associated fishes.
In some species, their skin mucous
facilitates acclimation towards
specific anemone species, while
others possess a thicker layer of skin
mucous that acts as an inert barrier
between the fish and anemone.
Regardless of the mechanism that
underpins nematocyst invulnerability,
these species all exhibit characteristic
acclimation behaviours, in which
the fish progressively increases
its contact with the anemone until
it is able to swim freely among its
tentacles. These behaviours allow
anemonefishes to first ‘test’ their
anemone before commencing with
acclimation.
Far less is known about the
relationships between facultatively
anemone-associated fishes and their
host anemones. Reports suggest
Figure 1. Anemonefishes.
that about a quarter of these species Blackfin anemonefishes (A. nigripes) with their host anemones on the reefs surrounding the Mal-
are able to touch the tentacles of dive Islands (Photo: Ocean imaging, www.theocean.com.au).
their anemones without being stung,
while another quarter are not, and it being evicted from the anemone, with fishes. Perhaps most importantly,
is not known for the remaining half. experience higher mortality; and that anemonefishes have been shown
Interestingly, species that can touch both anemonefishes and facultatively to promote higher densities of
their anemones include representatives anemone-associated fishes hide in, or photosynthetic zooxanthellae within
from seven (of 17) families and are near their anemone when threatened. their anemone’s tentacles. This is
found in numerous geographically Anemone associations also appear to done by enriching nutrients such as
disparate locations. have a positive effect on fish longevity, ammonia as well as increasing the
with dominant anemonefish females rate of localized water flow. As a
What benefits do anemonefishes estimated to live up to 6 times longer consequence of this, anemones that
receive from this relationship? The than similarly-sized fishes that do not host fishes experience increased
most broadly accepted benefit to associate with anemones. Increased growth, regeneration and reproduction.
anemone-associated fishes is that survival benefits also extend to their Some species of anemonefishes have
the anemone provides protection eggs, which are protected from also been shown to vigorously defend
from predators. While this has proven predators when laid near anemones. For their anemones from predators, such
difficult to experimentally demonstrate obligate anemonefishes, the main cost as butterflyfishes (Chaetodontidae).
among the obligate anemonefishes, of this relationship appears to be their Aggressive defence against
observations suggest: that they dependence on anemones for habitat. predators has not been observed
experience low predation compared to amongst the facultatively anemone-
non-anemone-associated fishes; that And what’s in it for the anemones? associated fishes, suggesting that
socially lower-ranking anemonefishes, Anemones also obtain benefits these relationships may be less
which have a higher likelihood of through their symbiotic relationships mutualistic than are those with their

Current Biology 27, R1–R18, January 9, 2017 R7


obligately-associated counterparts.
For anemones, the cost of hosting
fishes appears minimal but can include
occasional biting of tentacles and
‘theft’ of anemone prey.
How did these relationships
evolve? The circumstances that
select for anemone-associations
in fishes is largely unknown. It has
been suggested that an association
between fishes and corals, close
relatives of anemones that typically
have less potent stinging cells, could
act as a precursor for fish–anemone
associations. This hypothesis
remains to be tested. While the
anemonefishes evolved from a single
common ancestor, the taxonomic
diversity of non-obligate anemone-
associated fishes suggests that these
relationships have evolved several
times. These facultative anemone-
associated fishes are less-studied
than their anemonefish counterparts,
but their taxonomic diversity offers
an exciting opportunity to investigate
the ecological circumstances that
may lead to the evolution of obligate
anemone-associations. That being
said, the evolutionary history of
the anemonefishes is relatively well
understood. This group appears
to have originated between the
late oligocene and early miocene
(about 10–22 million years ago) in
an area between the Philippines and
the Great Barrier Reef and between
Sumatra and Melanesia. The ancestral
anemonefish was likely a crevice-
dwelling species that occupied a
limited number of host anemones.
Following anemone colonization, the
proliferation of anemonefish species
took place rapidly compared to
non-anemone-associated fishes and
likely occurred through a process of
range expansion, allopatric speciation
and hybridization. In general, there
is no strong link between host
specialization and anemonefish
speciation.
What next? The relationships
between anemones and their
associated fishes provide tractable
models to study the ecology and
evolution of interspecies symbioses.
While considerable research has
been conducted into these systems,
many questions remain. For example,
R8 Current Biology 27, R1–R18, January 9, 2017 © 2017 Elsevier Ltd.
substantial baseline natural history
information is lacking regarding the
facultatively anemone-associated
fishes, such as the phylogenetic
diversity of nematocyst tolerance
and how nematocyst tolerance is
achieved: do they use the same
mechanisms as the anemonefishes,
or are they doing something
different? The dynamics of these
relationships are also unclear: are
they true mutualisms, one-sided
commensalisms, or does it vary
according to species pair? Finally,
understanding how organisms and
ecosystems will respond to rapidly
changing environmental conditions is
becoming an increasingly important
area of research. The interactions
between anemones and fishes provide
a potentially informative model for
understanding how relationships
between species are affected by
changing conditions, as well as
whether these relationships will
confer resilience or vulnerability to the
species involved.
Where can I find out more?
Collingwood, C. (1868). Note on the existence
of gigantic sea anemones in the China Sea,
containing within them quasi parasitic fish. An.
Mag. Nat. Hist. 1, 31–33.
Elliott, J.K., and Mariscal, R.N. (1997). Ontogenetic
and interspecific variation in the protection of
anemonesfishes from sea anemones. J. Exp.
Mar. Biol. Ecol. 208, 57–72.
Fautin, D. (1991). The anemonefish symbiosis: what
is known and what is not. Symbiosis 10, 23–46.
Frédérich, B., Sorenson, L., Santini, F., Slater,
G.J., and Alfaro, M.E. (2013). Iterative
ecological radiation and conversgence during
the evolutionary history of damselfishes
(Pomacentridae). Am. Nat. 181, 94–113.
Litsois, G., Sims, C.A., Wüest, R.O., Pearman, P.B.,
Zimmermann, N.E., and Salamin, N. (2012).
Mutualism with sea anemone triggered the
adaptive radiation of clownfishes. BMC Evol.
Biol. 12, 212.
Randall, J., and Fautin, D. (2002). Fishes other
than anemonefishes that associate with sea
anemones. Coral Reefs 21, 188–190.
Santini, S., and Polacco, G. (2006). Finding Nemo:
Molecular phylogeny and evolution of the
unusual life style of anemonefish. Gene 385,
19–27.
Schmitt, R.J., and Holbrook, S.J. (2003). Mutualism
can mediate competition and promote
coexistence. Ecol. Lett. 6, 898–902.
Scott, A., and Dixson, D.L. (2016). Reef fish can
recognize bleached habitat during settlement:
sea anemone bleaching alters anemonefish
host selection. Proc. R. Soc. B Biol. Sci. 283,
20152694.
1
School of Biological Sciences, University of
Queensland, Australia. 2School of Integrative
Biology, University of California, Berkeley,
USA. 3School of Marine Science and Policy,
University of Delaware, USA.
*E-mail: william.e.feeney@gmail.com
Current Biology
Magazine
Primer
Tropomyosins
Peter W. Gunning
and Edna C. Hardeman
The actin cytoskeleton provides not only
the underpinning for cell architecture
but also mechanical force and the
ability to drive movement of cells and
their organelles. It is tempting to think
of it simply as a set of stable structural
elements, but nothing could be further
from the truth. The cells of our bodies
are continually remodelling their
architecture by responding to a range
of imposed biomechanical forces and
intracellular functional demands. Studies
of the dynamic and functional properties
of the actin cytoskeleton have been
dominated by a focus on actin and the
view that actin filaments are essentially
‘generic’. However, the ‘other’
component of most actin filaments in
animals — tropomyosin — is coming
into prominence. With this discovery
is the realisation that far from being
generic, actin filaments have their own
functional individuality provided to them
by their associated tropomyosin. This is
changing the way we understand and
study the actin cytoskeleton and has
delivered a new therapeutic opportunity
in what had come to be considered a
‘no-go zone’.
The discovery of tropomyosin in the
cytoskeleton was born in complexity
and this explains why it was not readily
incorporated into molecular studies of
the actin cytoskeleton. Tropomyosin
genes were recognised as exemplars
of alternative splicing that produce a
range of isoforms. While they generated
much interest in terms of mechanisms
of splicing regulation, their gene
products were seen as too complicated
to understand with regard to their
functional impact on the cytoskeleton.
It was only when sufficient tools
were made to discriminate between
the isoforms that their functional
significance came to be appreciated.
This Primer will cover the emerging
view that the tropomyosins specify
the functional properties of individual
cytoskeletal actin filaments in an
isoform-specific manner. From
the perspective of evolution,
tropomyosins provide the mechanism