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1 | INTRODUCTIO N
1.1 | Coral reef light habitats
Coral reefs are one of the most spectrally diverse habitats on Earth,
and vision is critical for the survival of many of its multicoloured
inhabitants. Coral reef fishes have been shown to rely on visual
cues for habitat selection (Booth, 1992; Huijbers et al., 2012; Igulu
et al., 2011), camouflage (Marshall & Johnsen, 2011), feeding (Job
& Bellwood, 2000; Stieb et al., 2023) and communication (Mitchell
et al., 2023; Siebeck et al., 2010 ). Interestingly, the light habitat on
coral reefs is not uniform, with differences found between micro-
habitat s such as in the water column with depth, inside the reef ma-
trix and under ledges and overhangs (Marshall et al., 2003). These
varied visual demands and the specifics of the microhabitat light
environments have led to substantial differences in morphologies,
ecologies, and, more specifically, colour vision systems in reef fishes
(reviewed in Cortesi et al., 2020). In this perspective, we provide a
timely overview of the variety of coral reef fish visual systems and
their link to fish ecology. We first review the known plasticity of
Received: 3 May 2024
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Accepted: 9 September 2024
DOI : 10.1111/1365-243 5.1466 8
PERSPECTIVE
Visual Ecology in Challenging Environments
Coral reef fish visual adaptations to a changing world
Abigail Shaughnessy | Fabio Cortesi
This is an op en access ar ticle under the terms of the Creative Commons Attribution-NonCommercial License, which permits use, distribution and reproduction
in any medium, provide d the original work is properly cited an d is not used for co mmercial purposes.
© 2024 The Author(s). Functional Ecology publishe d by John Wiley & S ons Ltd on behalf of British Ecological Society.
School of the Environm ent and
Queensland Brain Institute, The University
of Queensland, St. Lucia, Queensland,
Australia
Correspondence
Abigail Shaughnessy
Email: a.shaughnessy@uq.edu.au
Fabio Cortesi
Email: f.cortesi@uq.edu.au
Funding information
Australian Research Council Discovery
Early Career Research Award, Grant/
Award Number: DE200100620; The
University of Queensland Amplify
Fellowship
Handling Editor: Sara Stieb
Abstract
1. Coral reef ecosystems show fluctuations in their prevailing light environment in
response to both regular (e.g. between seasons) and more prevalent stochastic
events (e.g. human- induced sediment runoff). In these shifting environments,
phenotypic plasticity provides an essential mechanism for coral reef fishes to
adjust their visual capability to meet changing sensory requirements.
2. Here, we evaluate the growing area of research that highlights the many genetic
and ecological mechanisms that affect the plastic responses of coral reef fish vi-
sion to environmental cues.
3. With an increasing number of disturbances in the marine environment, it is criti-
cal to understand the extent and limits of visual plasticity under natural and dis-
turbed conditions. With our current knowledge and drawing upon a large body of
work in freshwater fishes, we speculate whether coral reef fishes can adapt to the
changes to their visual environment and where the limitations could lie.
4. Whilst coral reef fishes have shown visual adaptations under different light en-
vironments, the degree of plasticity is inconsistent between species. Thus, plas-
ticity may not only be functionally significant in maintaining the performance of
visually guided behaviours for single species but, more broadly, is likely key to
sustaining ecosystem function.
KEY WORDS
colour vision, coral reef fishes, light environment, phenotypic plasticit y, visual ecology
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SHAUGHNESSY an d CORTESI
reef fish colour visual systems when exposed to more regular and
spatial environmental changes, i.e. seasons, development and depth.
Second, we will explore how coral reef fish have adapted to stochas-
tic changes to their environment and what functional significance
this has on their behavioural response. Finally, we will draw upon the
different sensory strategies that may enable reef fishes to sur vive
in a human- driven changing world. This perspec tive aims to guide
future research into this quickly developing, fascinating topic.
1.2 | Coral reef fish vision
Many key visual adaptions of coral reef fish vision are found in the
retinal tissue layer at the ba ck of the eye. The retina consists of multi-
ple layers that contain several types of neuronal cells, including gan-
glion, amacrine, horizontal, bipolar and photoreceptor cells (Baden
et al., 2020; Masland, 2012; Sanes & Masland, 2015). Depending
on the ecological niche, behaviour and diet, the visual systems of
coral reef fishes at a retinal level can differ in the functional type,
number or distribution of retinal cell t ypes (Collin & Shand, 2003;
Yoshimatsu et al., 2020). Notably, there are two types of photore-
ceptors: rods and cones. Rods facilitate scotopic (dim light) vision,
while cones mediate vision under photopic (bright light) conditions.
Additionally, cones enable colour vision and provide high spatial and
temporal resolution (Cronin et al., 2014). Within each rod and cone
outer segment, opsin proteins are involved in light sensation when
bound to a vitamin A- derived chromophore (Bowmaker, 2008). The
opsin protein and chromophore interaction determine the wave-
length of maximum absorption sensitivity (λmax) (Kelber et al., 2003;
Yokoyama, 2000). Most coral reef fishes use a shorter- wavelength
shifted A1- based chromophore (Toyama et al., 2008). However,
longer- wavelength shifted A2- based chromophores in coral reef
fishes might be underestimated, especially in smaller cryptobenthic
species (Cortesi et al., 2020; White et al., 2004).
Within coral reef fishes, the five ancestral visual opsin types can
be found and are classed based on their phylogeny and the wave-
lengths of light they are sensitive to (Musilova et al., 2021). The rods
usually express a single RH1 opsin (rhodopsin, λmax = 447–525 nm).
The cones of coral reef fishes, especially of diurnal species, are often
arranged in a square pattern, whereby one single cone is surrounded
by four double cones (reviewed in Cortesi et al., 2020). The latter
are two morphologically connected single cones, and they may be
electrically coupled or work independently of each other to perceive
colour (Pignatelli et al., 2010). The single cones typically express
short- wavelength- sensitive opsins, SWS1 (λmax = 347–383 nm) and
SWS2 (λmax = 397–482 nm), while the double cones express middle-
wavelength- sensitive rhodopin- like 2, RH2 (λmax = 453–537 nm), and
long- wavelength- sensitive, LWS (λmax = 501–573 nm), opsins (re-
viewed in Carleton et al., 2020). Following a series of duplications,
deletions and functional mutations through evolutionary time, coral
reef fish show a vast opsin gene repertoire, with some extant spe-
cies having up to 14 opsin genes in their genome (Blackbar soldier-
fish, Myripristis jacobus) (Musilova et al., 2019). However, coral reef
fishes generally use t wo to four spectrally distinct cone photore-
ceptors for (colour) vision (Cortesi et al., 2020; Losey et al., 2003),
leaving them with what appears to be too many opsin genes for their
visual demands. Several mechanisms, including regional expres-
sion differences in the retina, the co- expression of multiple opsins
within a single photoreceptor, and phenotypic plasticity (i.e. in de-
velopment) and flexibility (i.e. reversible within- individual variation
(Piersma & Drent, 2003)) in opsin gene expression, have been sug-
gested as possible mechanisms driving this diversit y (as reviewed in
Carleton et al., 2020).
Research has shown that coral reef fish vision can indeed be plas-
tic within an individual's lifetime, such as during development, over
depth, and between seasons (Cortesi et al., 2016; Stieb et al., 2016;
Tettamanti et al., 2019). Note, that this observed plasticity is dis-
tinctive from circadian changes in opsin gene expression, which are
common in fishes (e.g. surgeonfishes (Fogg et al., 2023), cichlids
(Yourick et al., 2019), killifish (Johnson et al., 2013)).
Coral reef fishes are exposed to regular fluctuations in their pre-
vailing lig ht environment as con ditions change, suc h as diurnal cycles ,
seasons or depth. In recent years, the underwater light environment
has also been experiencing increased stochastic changes due to an-
thropogenically driven events; severe storms, sediment runoff, and
algal blooms are all increasing in frequency (Caves & Johnsen, 2021;
Dutkiewicz et al., 2019). Climate change is also causing the colour
underwater to change due to, for example, mass- bleaching events
and associated coral die- of fs (Marshall et al., 2019). As coral reef
fishes try to retain optimal visual performance across a full range
of light environments, visual plasticity and flexibility (unified under
the term plasticity from here on out) appear to have a crucial role in
sensor y adaptation (reviewed in Carleton et al., 2020).
Understanding the scope of colour vision plasticity is becoming
increasingly important in the context of anthropogenic influences
on the marine environment. As humans change the environment
(Pörtner et al., 2022) coral reef fishes will notably be presented with
new challenges to their visual habitat, which could affect their sen-
sory abilities, behaviour, and, ultimately, their survival. Therefore,
visual plasticity could provide coral reef fishes with a vital mecha-
nism to adjust their visual c apabilities to meet the changing sensor y
requirements. However, there is a significant gap in our understand-
ing of visual plasticity in these animals. We need to find out how
common visual plasticity is amongst reef fishes, where the inter- and
intraspecific differences lie (i.e. differences between species and life
stages), and the specific mechanism that drives visual plasticity.
2 | VISUAL SYSTEM PLASTICITY
2.1 | Depth
The absorption properties of water and the scattering effect of
suspended particles are two processes that govern the attenua-
tion of sunlight through seawater, dictating the spectral compo-
sition and intensity of the underwater light environment. With
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SHAUGHNESSY and CORTESI
increasing depth, the light rapidly at tenuates and firstly affects the
wavelengths at the ends of the UV–visible spectrum (i.e. <390 nm
(ultraviolet, UV) and >600 nm (red)) (Levine & MacNichol, 1982).
Thus, at a deeper depth, the light spectrum is narrower, centred
around blue- green wavelengths, along with a decrease in light
intensity.
Stieb et al. (2016) demonstrated subtle differences in opsin
gene expression of damselfishes (Pomacentridae) when the light
environment changes over depth on tropical coral reefs. Individuals
from the same species that occur in deeper depths (>10 m) shift
their opsin expression towards the blue/green wavelengths of light
(Figure 1a). However, these changes were limited to only a few of
the investigated damselfish species (Pomacentrus spp.) and not oth-
ers (Dascyllus spp.), suggesting that plasticity is species- specific and
could be further governed by a species' ecological needs or phylo-
genetic inertia. However, further attention needs to be paid to tease
apart how ecology and phylogeny shape visual function.
2.2 | Seasons
A predictable shift in the overall light environment occurs seasonally
amongst coral reefs, both in light availability and spectral content
(Jerlov, 1977; McFarland, 1986). During the winter, when the oce-
anic water is clear with little to no suspended or dissolved matter, a
broad spectrum of light from UV to red is available. In contrast, the
temperature of the water and light intensit y rises throughout sum-
mer, supporting an increase in phytoplankton growth and organic
matter suspended or dissolved in the water column. The greater the
amount of these substances, the more red- shifted, i.e., greener the
light spectrum of the water becomes (Jerlov, 1977; Lythgoe, 1979;
McFarland, 19 86). In freshwater fishes, such relatively red- shifted
habitats, e.g., in the clear and murky parts of the Amazon River
(Escobar- Camacho et al., 2019), between crater lakes in Nicaragua
(Härer et al., 2017), or between seasons in Canadian lakes and
streams (Allen et al., 1982) lead to a switch from chromophore A1 to
A2 and changes in opsin gene expression to form longer- wavelength
sensitive pigments.
To the best of our knowledge, only one study has investigated
seasonal shifts in visual opsin gene expression in coral reef fishes
(Stieb et al., 2016). The Nagasaki damsel, Pomacentrus nagasakiensis,
was shown to increase the expression of its UV- sensitive opsin, sws1,
during winter, thus correlating with more UV light availabilit y during
those months (Figure 1c). More examples can be found in freshwater
fishes, where seasonal changes in light intensity and day length have
shown different plastic changes to the visual system. For example,
the opn4 (opsin4: melanopsin)- related genes in zebrafish have been
shown to var y in response to changes in photoperiod (Matos- Cruz
et al., 2011). Also, Medaka rice fish, Oryzias latipes, are less active
and rarely feed during winter. Shimmura et al. (2017) demonstrated
the downregulation of visual genes involved in phototransduction,
which may allow Medaka to save energy during winter. In contrast ,
the summer breeding season is associated with body colouration and
ornamentation to attract mates; thus, an upregulation in their pho-
totransduction genes is seen to allow for these essential visual func-
tions to occur. Interestingly, they suggested that seasonal changes
in lws opsin gene expression in medaka were due to temperature
differences rather than shifts in the light environment.
For many coral reef fishes, especially smaller, site- attached spe-
cies, changes in opsin expression and other genes involved in visual
function will likely occur between the seasons. However, the extent
of these shifts could ultimately depend on the fish's ecology and be-
haviour. For instance, coral reef fishes with UV vision may exhibit
different opsin expression shifts to accommodate the seasonal at-
tenuation of UV wavelengths. In contrast, fishes that cannot see UV
might show no such shift. Nevertheless, it is still being determined
how widespread seasonal visual plasticity is across the more than
8000 coral reef fish species.
2.3 | Development
Ontogenetic changes in the visual system of coral reef fishes en-
able a species to adapt to shifts in their habitat or feeding be-
haviour as they progress from larvae to adults. Coral reef fishes
start in the open ocean as larvae (Helfman et al., 2009; Job &
Bellwood, 2000), where the light environment is bright and broad
spectrum (McFarland, 1986). Here, they typically feed on zooplank-
ton. After this phase, the fish transition to the reef, where there is
often a change in diet (planktivory, carnivory, herbivory, corallivor y)
and light environment (reviewed in Cortesi et al., 2020).
Developmental plasticity in opsin gene expression and A1/A2
chromophore switches have been found in some, but not all (e.g.
freshwater [e.g. stickleback (Novales Flamarique, 2019), cichlids
(Carleton et al., 2008; Nandamuri et al., 2017; Wilwert et al., 2023),
killifish (Fuller et al., 2004, 2005)], catadromous [e.g. salmonids
(Cheng & Flamarique, 2007; Cheng & Novales Flamarique, 2004;
Novales Flamarique, 2005), eels (Archer et al., 1997; Wood &
Partridge, 1993)], estuarine [e.g. black bream (Shand et al., 2008)]
and marine fishes [e.g. flounder (Savelli et al., 2018)]). Ontogenetic
changes in opsin gene expression also seem common in coral reef
fishes, often correlating with changes in ecological demands. For
example, the Dusky dottyback (Pseudochromis fusucs) changes its
expression between pelagic larvae, juvenile and adult stages on
the reef to correlate with shifts in habitat and colouration (Cortesi
et al., 2016). Pre- set tlement holocentrid larvae have high cone opsin
gene expression levels well adapted for photopic conditions. As the
adults switch to a nocturnal, reef- dwelling lifest yle, a scotopic vi-
sual system is developed with higher levels of rod opsin expression
(Fogg et al., 2022a). In addition, morphologic al and cellular changes
to the retina have also been shown to further adapt their eyes to
the dim light of adult life (Fogg et al., 2022b) (Figure 1b). However,
for surgeonfish (e.g. Naso brevirostris (Tettamanti et al., 2019) and
Acanthurus triostegus (Besson et al., 2017; Fogg et al., 2022a)), the
opsin gene expression and retinal topography mainly changed be-
tween the larval and juvenile stages, with fewer changes after that.
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SHAUGHNESSY an d CORTESI
FIGURE 1 Major environmental drivers of visual plasticity in coral reef fishes. (a) The differences in ambient light at various depths can
reflect a reef fish's opsin expression profile. At shallow depths, damselfishes (Pomacentridae) experience a full spectrum of light and express
high levels of their green- sensitive rh2a (green dot) and ultraviolet- sensitive sws1 (UV, purple dot) opsin genes. The light rapidly attenuates
with increasing depth, affecting the wavelengths at either end of the fish's visible spectrum (UV <390 nm and red >600 nm). With ample
blue wavelengths of light available, damselfishes increase the expression of their blue- sensitive opsin, rh2b (blue dot) (Stieb et al., 2016). (b)
Changes to retinal gene expression and morphology can also be tied to ontogeny. Pre- set tlement holocentrid lar vae are strongly adapted to
photopic conditions, showing high cone densities and a broad cone opsin gene repertoire (multicoloured bar). As they settle onto the reef
and adopt a nocturnal lifestyle, adults have a rod- dominated multibank retina and a reduced cone opsin gene expression (grey bar) (Fogg
et al., 2022a, 2022b). (c) Seasons and associated changes in the light environment can shift the opsin gene expression. During winter (top),
the clear oceanic water has little to no suspended or dissolved matter, creating a broad ‘blue’ light environment with ample wavelengths
ranging from UV to red. Under this light environment, the Nagasaki damsel (Pomacentrus nagasakiensis) expressed high levels of sws1 (purple
dot). In contrast, the water becomes murkier and red- shifted ‘greener’ in summer as algae and suspended sediments dominate. Thus, with
fewer UV wavelengths available, P. nagasakiensis decreases its expression of sws1 (Stieb et al., 2016). (d) Anthropogenically driven changes
can also af fect coral reef fish vision. For example, increases in ocean acidific ation can slow the speed of vision. The Spiny chromis damselfish
(Acanthochromis polyacanthus) individuals exposed to control (natural CO2 levels; left column) and high- CO2 levels (right column) can follow
a low flickering light stimulus (Hz) at 20 Hz. However, when the frequency increases, individuals exposed to elevated CO2 conditions can no
longer resolve the light stimulus above 80 Hz, seeing it as a constant illumination instead of a flickering light (flat line; Chung et al., 2014).
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This suggests that visual plasticity is reserved for the development
of some species, a phenomenon also seen for other traits such as
colour change (Cheney et al., 2017).
2.4 | Stochastic events
Stochastic turbidity events can occur on shallow coral reefs, as par-
ticulate matter can be resuspended by energetic processes such as
wind- driven waves, strong tidal currents, river discharge plumes or
severe weather events (as reviewed in Zweifler et al., 2021). Other
environmental drivers such as water temperature, riverine outputs
or terrestrial runof f can further impact the suspended particulate
load on the reefs. Studies investigating the consequences of these
turbidity events have highlighted noticeable direct effects on coral
reef fishes, such as loss of habitat and reduction of food sources
(Erftemeijer et al., 2012; Fabricius et al., 2005). However, in turbid
environments, visual information available to coral reef fishes de-
clines as suspended par ticles absorb and scatter light, reducing the
overall light availability, obscuring visibility and changing the spec-
tral composition of the water. For instance, chromophoric (coloured)
dissolved organic matter and non- algae particles tend to absorb UV
and shor ter violet and blue wavelengths (Babin et al., 2003; Binding
et al., 2008; Bricaud et al., 1981; Kirk, 1985). Algal turbidity, formed
by larger biotic particles, shifts the spectrum of light towards me-
dium (green/yellow) wavelengths of light (Cronin et al., 2014;
Jerlov, 1977 ). Therefore, since turbidity significantly alters the visual
environment, it is likely that fish vision is af fected in many ways, in-
cluding detecting brightness (luminance), colour (colour discrimina-
tion) and motion (directional selectivity).
As for the other factors that may cause visual plasticity, most
of our knowledge on how visual systems adapt to turbid environ-
ments derives from freshwater fish. Frequently, the visual pigments
in freshwater fish are longer- wavelength- shifted to adjust the visual
system to the red- shifted light environment caused by high sedi-
ment or chlorophyll concentrations. For instance, CYP27C1 is the
enzyme that catalyses the conversion of the A1 chromophore to A2
(Enright et al., 2015). Lake Malawi (Terai et al., 2002) and Lake Xiloa
(Härer et al., 2017) are clear lakes, and the resident cichlids exhibit
a low expression of the cyp27c1 gene. However, high expression
levels are found in cichlids living in the turbid Lake Managua and
Lake Nicaragua. Chromophore and opsin gene expression plasticity
correlating with changes in water clarit y have also been found in
fishes living in the Panama Canal (Escobar- Camacho et al., 2020).
Furthermore, adult killifish (Lucania goodei) altered their opsin gene
expression when inhabiting clear or murky waters to match the avail-
able light spectrum (Fuller et al., 2004). Remarkably, sudden light
habitat changes resulted in rapid shifts within a few days (Fuller &
Claricoates, 2011). Evidence suggests that long- wavelength sensitiv-
ity is associated with motion detection (Krauss & Neumeyer, 2003;
Mizoguchi et al., 2023; Schaerer & Neumeyer, 1996). Thus, increased
lws expression in killifish exposed to tea- stained waters could main-
tain or enhance the ability to detect motion in a turbid environment.
Whether coral reef fishes can rapidly adjust their visual systems
when experiencing turbidity remains to be investigated. Some in-
sights can be gained from laborator y experiments that have placed
various reef fish species (damselfish and cardinalfish (Luehrmann
et al., 2018), surgeonfish (Fogg et al., 2023)) under different light en-
vironments for several weeks (blue, green, red, broad light, dim light,
etc.), which suggests that short- term, reversible visual plasticity is
possible, at least in some species. However, we know more about the
functional impact of turbidity on coral reef fish visual behaviour. For
example, planktivorous coral reef fishes under sediment- induced
turbid conditions showed a decrease in total foraging rates, implying
the performance of visual predators could be compromised by in-
creases in turbidity (Johansen & Jones, 2013). High levels of turbid-
ity can also inhibit coral reef fish lar vae from visually distinguishing
suitable coral habitats from dead coral during settlement (O'Connor
et al., 2016). Similar studies in freshwater fishes have shown that
high turbidity levels can also reduce the activit y levels of prey, in
this case, guppies (Poecilia reticulata), causing them to exhibit more
solitary behaviours, thus making them more vulnerable to predation
(Borner et al., 2015).
Whilst co ral reefs can nat urally be relati vely turbid enviro nments,
notably those inshore (reviewed in Zweifler et al., 2021), the increas-
ing frequency and severity of elevated turbidity and algal blooms are
recognised as a significant local stressor (Griffith & Gobler, 2020;
Syvitski et al., 2005). Increases in sedimentary, nutrient, and other
organic particle inputs are primarily caused by changes in land use
(e.g. agriculture, dredging, coastal development), human activities
(e.g. shipping traffic and fishing trawlers) or climate- change- driven
events (e.g. rainfall and cyclones) (Syvitski et al., 20 05). Therefore,
it highlights the impor tance of understanding the scope of coral
reef fish vision under these exacerbated turbid environments and
to what extent this limits their ability to perform critical behavioural
tasks (Figure 2).
3 | ANTHROPOGENIC STRESSORS
3.1 | Light pollution
Artificial light at night (ALAN) is a widely distributed form of anthro-
pogenic pollution (Falchi et al., 2016). In the oceans, ALAN can be
caused directly by shipping, oil platforms, and coastal developments
(e.g. towns, cities, and harbours). The scattering and reflection of
upward emitting light, termed skyglow, can further extend the spa-
tial influence of ALAN on marine environments (Kyba et al., 2011;
Tidau et al., 2021). The effects of ALAN are predicted to be further
amplified due to a societal shift towards ‘white’ or broad- spectrum
lights, such as energy- efficient light- emitting diodes (LEDs; Gaston
et al., 2012; Tamir et al., 2017). Thus, the increasing number and
variety of spectrally distinct lighting types amplify nocturnal irra-
diance and the available light spectrum. For example, white broad-
band LEDs have a short- shif ted spectrum with an abundance of blue
wavelengths of light compared to moonlight (Elvidge et al., 2010).
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Moreover, shorter wavelengths of light penetrate deeper into the
water column (Jerlov, 1977), which can further exacerbate the
problem.
Thus far, research on the impacts of light pollution on coral reef
fishes has focused on the effects on ecology and behaviour. Reef
fish rely on lunar cycles to regulate their biological rhythm, which is
intrinsically tied to reproduction (spawning synchrony), settlement
and trophic interactions. For instance, ALAN has been shown to
disrupt sp awning cycles, clutch size, egg quality and hatching success
in anemonefishes (Fobert et al., 2019, 2021). Many larval reef fishes
also have phototactic behaviours to support their foraging eff iciency
during high lunar illumination (Shima & Swearer, 2019), or they time
their metamorphosis to allow them to set tle on dark, moonless or
overcast nights (Besson et al., 2017; Shima et al., 2018). Thus, ALAN
can disrupt natural light cues that are important for growth and set-
tlement (O'Connor et al., 2019). Also, as ALAN increases, smaller
coral reef fishes are at increased risk of predation due to changes in
their effectiveness to camouflage and reduced shelter use at night
(Georgiou et al., 2024). Other predator–prey interactions might
also be disrupted, such as the familiar sight of invasive lionfishes in
the Caribbean and giant trevally and groupers on the Great Barrier
Reef hunting sleeping prey lit up by the torchlights of divers. Finally,
ALAN might also disorient fish accustomed to navigating in the dark.
For instance, the vertical migration patterns of larval coral reef fish
through the water column (Hawes et al., 2020) could be affected by
ALAN, as seen similarly in arctic fishes (Berge et al., 2020).
As far as we know, only one study has studied the influence of
ALAN on the visual system of coral reef fishes. Recent work in the
convict surgeonfish, A . triostegus, revealed a shift in opsin gene ex-
pression and retinal ganglion cell density in developing fish under
ALAN (Fogg et al., 2023). Unsurprisingly, this work shows that
changes in light exposure can affect coral reef fish vision, with the
mechanism and functional consequences remaining to be investi-
gated for most species.
3.2 | Ocean acidification
Rising atmospheric carbon dioxide (CO2) levels from anthropogenic
fossil fuel, land- use change (deforestation and agriculture), and in-
dustrial emissions are quickly making our oceans more acidic (Sabine
et al., 2004). Several studies have indicated that sensory systems
such as vision can be affected, in which a change in response to
visual cues and altered retinal function has been observed under
elevated CO2 levels. For example, coral reef fishes have shown be-
havioural changes such as increased boldness (Jutfelt et al., 2013),
homing ability and set tlement behaviour (Munday et al., 2012).
Although some of the behavioural changes might have been
due to experimental ar tefacts (Clark et al., 2020), an increase in
acidity can lead to changes in retinal function and, by extension,
visual behaviour. Fast flicker fusion is the rapid response of a vi-
sual system tracking a flickering light. When the flickering light
appears continuous, this is the critical flicker fusion (CFF), and the
visual system can no longer follow the flashing light. The CFF can
vary between fishes as it correlates with lifest yle, environmental
illumination level and physiological processes in the eye, which
are sensitive to temperature and acidity (Fritsches et al., 2005;
Horodysky et al., 2008). Under elevated CO2, the CFF of the
Spiny chromis, Acanthochromis polyacanthus, slowed down, thus
potentially reducing reaction time to fast events such as an in-
coming predator or a fast- moving prey item (Chung et al., 2014)
FIGURE 2 Coral reef fish vision is affected by changes in the
intensity and wavelengths of light available underwater. Turbidit y
events such as algal blooms or sediment suspension due to floods
or storms shift the clear blue light environment (top) towards
medium (green/yellow) wavelengths (bot tom). These shifting
light environments can have behavioural consequences such as
reduced feeding success, predator avoidance, and recognition
of conspecifics (bottom). Phenotypic plasticity is one way to
maintain visual performance in rapidly changing light environments.
However, it may only be implemented by some species on the
reef, with significant implications where ecosystem function may
change.
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(Figure 1d). The underlying mechanism is suggested to be associ-
ated with altered GABA A receptor function, as these receptors are
sensitive to changes in pH and control the flicker fusion response
in fishes (Mora- Ferrer & Neumeyer, 2009).
3.3 | Rise in ocean temperature and associated
coral bleaching
As mentioned previously, a change in the temperature of the water
might have profound impact s on vision, especially in ectotherm ani-
mals such as coral reef fishes. Moreover, declining coral cover and
the associated loss of structural complexity due to ocean warming
have been widely reported across the world's coral reefs (Hughes
et al., 2018). Many contemporary reefs are shifting towards a non-
coral substratum, where turf algae and rubble are widespread
(Bellwood et al., 2019 ; Graham et al., 2015). When considering the
visual environment, these shifts can have profound implications for
reef- associated fishes that rely on the colour and structural com-
plexity that coral- dominated habitats provide.
Many coral reef fishes have evolved colouration and patterns
that allow them to blend into a reef background (reviewed in
Marshall et al., 2019). As such, blue and yellow are commonly seen
colours in coral reef fishes (Marshall et al., 2015). For nearby observ-
ers, this colouration is conspicuous. Yet, from a distance, due to the
poor resolving power of reef fish eyes (Collin & Pettigrew, 1989), the
colours begin to merge into the background (Marshall et al., 2003).
This interaction distance and the structural complexit y of the reef
are also critical when viewing high spatial frequenc y patterns used
for camouflage, such as in the Banded humbug, Dascyllus aruanus
(Phillips et al., 2017). Predator–prey interactions might also be
impacted by the change in background colour associated with a
bleaching and dying reef. For example, yellow Dusky dottyback can
be spotted more readily by the predator y Coral trout (Plectropomus
leopardus) when seen against a rubble background compared to a
live coral background (Cortesi et al., 2015). As the world's coral reefs
are becoming less colourful, so are the fish communities inhabiting
them (Hemingson et al., 2022). The visual systems that evolved to
perceive these colours are expected to change along the way.
There is some evidence to sug gest that coral reef fishes can
adapt their thermal tolerance fast enough to survive the warming
surface waters caused by climate change (Donelson et al., 2012).
However, it needs to be clarified whether all reef fish species can
acclimate to global warming over a short timescale. Thus, for spe-
cies that may not have adaptive potential in warmer waters, range
expansions and contractions of coral reef fish communities are
expected to occur as their environment changes regularly. More
specifically, some coral reef fishes have exhibited a poleward shift
in their distribution to escape to colder waters (Fear y et al., 2014).
However, the success of settling into temperate habitats may be
limited to reef fish species with a large set tlement body size, high
swimming ability or pelagic spawning behaviour (as reviewed in
Feary et al., 2014). Alternatively, to find cooler waters, coral reef
fishes may move deeper in the water column, leading to a bathy-
metric shift in distribution (Caves & Johnsen, 2021). Once shallow-
water adapted species start moving deeper, there are several ways
in which they may plastically respond to changes in the overall light
environment (Caves & Johnsen, 2021). At greater depths, they may
depend more on their other senses, such as olfaction or mechanore-
ception. They may also depend more on aspects of visual signals less
affected by the changing light environment, such as the brightness
contrast between stripes or spot s, rather than chromatic colour in-
formation. For example, UV- reflecting body patterns are commonly
seen in shallow- living smaller coral reef fish species such as dam-
selfishes (Siebeck et al., 2010; Stieb et al., 2017). UV sensitivity has
provided these fishes with a close- range communication channel,
aiding in species discrimination, social hierarchy and mate selec-
tion (Mitchell et al., 2024; Siebeck et al., 2010; Stieb et al., 2024).
Notably, at deeper depths, UV is progressively filtered out. In their
research, Stieb et al., 2016, found no difference in sws1 opsin gene
expression for six out of seven damselfish species investigated at a
depth of 10–15 m. This is likely because the high sensitivity of UV
single cones in fishes may be sufficient to perceive UV patterns even
when only a few UV photons are present (Yoshimat su et al., 2020).
However, UV vision will not be possible at deeper depths where UV
light disappears entirely. Thus, whether the perception and function
of colourful patterns in a dim, blue wavelength- dominated deep en-
vironment continue to be the same as those of a shallow, well- lit,
broad- spectrum coral reef remains to be investigated.
4 | CONCLUSIONS
In a habitat where vision serves as a primary source of information
for coral reef fishes, the ability to quickly alter their visual system
could make them more resilient to escalating environmental change.
However, many unknowns exist regarding how the visual system and
the corre sponding behaviours will resp ond to global and loc al threats
to the reef. The few studies conducted to date have revealed a mixed
picture whereby some species might be able to rapidly shift their
visual system in response to environmental changes, while others do
not. Importantly, synergistic ef fects have yet to be considered when
faced with multiple stressors. For example, temperature increases
vision speed while elevated CO2 levels decrease it. It is also essential
to consider whether visual plasticity responding to regular and
spatial environmental changes will remain stable as conditions vary
due to anthropogenic fac tors. For instance, will seasonal changes
in opsin expression stay consistent as average sea temperatures
rise with climate change? Thus, it's important to understand how
widespread visual plasticity is amongst coral reef fish and it s
functional significance in sensory adaptation. Also, follow- on effects
might be expected, such as when habitat complexity decreases,
coral reef fish navigational ability will be reduced, which is likely to
affect fish cognition, potentially lowering their survivability. Hence,
future research should identify which species are most affected by
sensor y pollution and the roles that ecology and phylogeny can play
8
|
SHAUGHNESSY an d CORTESI
in shaping a plastic sensory system. Additionally, attention should
be paid to whether the timing at which environmental change is
experienced alters the capacity for plasticity, what factors make
a species plastic, and the extent and limits of plasticity. Finally,
behavioural studies should be emphasised to assess the functional
implications of plasticity for species reproduction and survival.
AUTHOR CONTRIBUTIONS
Abigail Shaughnessy and Fabio Cortesi conceptualised the perspec-
tive piece. Abigail Shaughnessy led the writing and visualisation of
the manuscript. Fabio Cortesi reviewed and edited the manuscript.
All authors contributed critically to the drafts and gave final ap-
proval for publication.
ACKNOWLEDGEMENTS
We want to acknowledge the many luminaries in visual ecology
who have inspired this perspective piece and the continuing sup-
port from the Marine Sensory Ecology Group at the University of
Queensland. We would also like to recognise the brilliant studies
on visual plasticity that we will likely have missed. We would also
like to acknowledge the Traditional Owners and custodians of the
lands on which some of the reported research has been carried out.
Additionally, we would like to thank the two anonymous reviewers
and the editor for valuable feedback. Open access publishing facili-
tated by The University of Queensland, as part of the Wiley - The
University of Queensland agreement via the Council of Australian
University Librarians.
FUNDING INFORMATION
F.C. was supported by an Australian Research Council DECR A
Research Fellowship (ARC grant DE200100620) and a University of
Queensland Amplify Fellowship.
CONFLICT OF INTEREST STATEMENT
The authors declare no conflict of interest.
DATA AVAIL ABILI TY STATEMENT
This manuscript did not use data.
STATEMENT ON INCLUSION
Our perspective piece included a wide range of peer- reviewed stud-
ies. As such, no primary data were collected for this written piece.
However, efforts were made to consider relevant works across
a wide geographical distribution to represent the growing area of
research.
ORCID
Abigail Shaughnessy https://orcid.org/0000-0001-9330-0093
Fabio Cortesi https://orcid.org/0000-0002-7518-6159
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How to cite this article: Shaughnessy, A., & Cortesi, F. (2024).
Coral reef fish visual adaptations to a changing world.
Functional Ecology, 00, 1–12. https://doi .or g/10.1111/1365 -
2435.14668