Marine and Freshwater Behaviour and Physiology

ISSN: 1023-6244 (Print) 1029-0362 (Online) Journal homepage: http://www.tandfonline.com/loi/gmfw20

Visual feeding of fish in a turbid environment:

Physical and behavioural aspects

A.C. Utne-Palm

To cite this article: A.C. Utne-Palm (2002) Visual feeding of fish in a turbid environment:
Physical and behavioural aspects, Marine and Freshwater Behaviour and Physiology, 35:1-2,
111-128, DOI: 10.1080/10236240290025644

To link to this article: http://dx.doi.org/10.1080/10236240290025644

Published online: 26 Oct 2010.

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 Mar. Fresh. Behav. Physiol., Vol. 35, No. 1–2, pp. 111–128

VISUAL FEEDING OF FISH IN A TURBID

ENVIRONMENT: PHYSICAL AND
BEHAVIOURAL ASPECTS
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A.C. UTNE-PALM*
University of Bergen, Department of Fisheries and Marine Biology,
P.O. Box 7800 N-5020 Bergen, Norway

(Received 29 June 2000; In final form 5 June 2001)

Turbidity has both positive and negative effects on prey detection, by increasing or diminishing the contrast
between prey and background due to the scattering of light. The positive effect of turbidity on prey contrast
depends on the optical properties, scattering properties of suspended particles and the visual sensitivity of
the predator.
The positive effect of turbidity is pronounced for larval fish, given that their visual field is short, leaving
fewer particles between them and their prey to scatter light and interfere with detection. This relationship,
together with a decreased risk of predation, makes turbid environments more optimal for some species and
size groups of fish (planktivores and fish larvae) and less so for others (adult piscivore fish). Thus, turbidity
might have a structuring effect on a fish community. Recently it has been demonstrated that UV light might
have positive effects on prey detection and consumption. How UV light might interact with different kinds of
particles producing turbidity is not well documented.

Keywords: Turbidity; Prey detection; Visual sensitivity; Predation; Larval fish; Planktivores; Piscivores

INTRODUCTION

Water turbidity is influenced by freshwater runoffs, with rainfall washing organic

matter, soil and sand particles into rivers, lakes and coastal sea. The increase in organic
matter raises the level of eutrophication and encourages algal blooms, which lead to
further increases in turbidity. As a result of freshwater run-off and wave action (resus-
pension of bottom sediments) coastal areas are more turbid than the open sea. Further,
human impacts have caused increases in turbidity all over the world. The use of ferti-
lisers has raised the eutrophication levels and changing weather patterns, due to global
warming, have increased the severity of rainstorms and hurricanes with the mudflows
that often follow them.
High concentrations of suspended solids can cause physiological and behavioural
stress responses in fish (Bruton, 1985) and may even be lethal (Newcombe and
MacDonald, 1991; Gregory et al., 1993). Several studies reveal that relatively small
increases in turbidity may change fish community structure (in freshwater lakes:

*E-mail: anne.utne@ifm.uib.no

ISSN 1023-6244 print: ISSN 1029-0362 online ß 2002 Taylor & Francis Ltd
DOI: 10.1080/10236240290025644
 112 A.C. UTNE-PALM

Swenson, 1978, subtropical estuaries: Blaber and Blaber, 1980; Cyrus and Blaber,
1987a,b and 1992 and temperate estuaries: Ruiz et al., 1993; Maes et al., 1998).
Newcombe and MacDonald (1991) concluded that the concentration of suspended
sediments and the duration of exposure was of importance because of the negative
effect on fish and aquatic life. The best fitting model was a concentration–duration
response model similar to those used to assess the effect of toxins.
In a study of the effects of turbidity on feeding strategies of fish in South African
estuaries, Hecht and van der Lingen (1992) found that visual piscivore predators were
affected more by an increase in turbidity than macrobenthic feeders (see also Rowe
and Dean, 1998). Although turbidity can influence several aspects of the wellbeing of
fish, in this paper, I concentrate on the effect of turbidity on the visual abilities of fish.
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Underwater light is absorbed and scattered by the water and by particles such as
sand and plankters. These particles may reduce horizontal visibility from 60 m or so
in clear water to only a few centimetres in highly turbid water. When particles scatter
light, a brighter veil is imposed between the target and the eye, reducing visibility. It is
not that the eye lacks sensitivity, but rather that the eye is sensitive to both the image-
forming light reflected from the target and the non-image-forming scattered light which
degrades the target brightness and colour contrast (Lythgoe, 1979).
Despite the generally poor quality of underwater images, most fish species depend
on vision as their main source of sensory information (Guthrie and Muntz, 1993).
The ability of both predators and prey to detect each other is frequently impaired
by turbidity in aquatic ecosystems. This paper reviews some of the studies that deals
with turbidity and predator–prey interactions in the light by theoretical knowledge
of vision in a light-scattering media.

THEORETICAL BACKGROUND

The visibility of a prey item requires the predator to detect a difference in contrast
between the prey and the background, which is dependent upon the optical properties
of the object, the background and the medium.
Because water absorbs long and short-wavelengths more than middle-wavelengths
(i.e. <430 nm and >530 nm) (Jerlov, 1968) light tends to be monochromatic at moder-
ate depths (Lythgoe, 1975). Brightness contrast is therefore most often the determining
factor for the visibility of objects underwater (Hemmings, 1965; Lythgoe, 1975), for
objects that are lighter or darker than the background (i.e. it must reflect more or
less light).
The inherent contrast (C) between an object and its background can be expressed as


Ep  Eb
C¼ ð1Þ
Eb

where Eb is background radiance and Ep is radiance of the object.

As indicated, turbidity has a veiling effect on the visual image. The sighting range of
targets underwater is determined by the exponential degradation of the inherent target
contrast along the line of sight (Duntley, 1963).

Ca ¼ C expðcxÞ ð2Þ
 VISUAL FEEDING IN TURBID WATER 113

where C is the inherent and Ca the apparent prey contrast at distance x from the
prey, and c the beam attenuation coefficient of the water (beam attenuation is
defined as c ¼ a þ b, where a is absorbtion and b is the scattering coefficient
(Zaneveld et al., 1979)).
Target detection is facilitated if the apparent contrast (Ca) exceeds a specific contrast
threshold (Ct). By knowing a fish’s contrast threshold one can estimate the visual
range (r):

Ct ¼ C expðrcÞ or r ¼ ln ðC=Ct Þ=c ð3Þ

From Eq. (3) visual range (r) decreases with increasing turbidity (c), decreasing inher-
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ent contrast (C) or increasing contrast threshold (Ct).

Several studies have shown that measured contrast thresholds vary with factors
such as target size, ambient light level and turbidity (Hester, 1968; Anthony, 1981a;
Utne-Palm, 1999). It is questionable whether this reflects a true change in the contrast
threshold of the fish or merely indicates that factors such as target size, ambient light
level and turbidity affect visibility in addition to contrast. Aksnes and Giske (1993)
and Aksnes and Utne (1997) suggest a detection criterion that combines the effects
of retinal contrast (Cr), retinal target size (Apr), and retinal background light level
(Ebr), so that the product of the three must exceed an energetic threshold value (
Sr)
in order for a target to be detected:

Cr Apr Ebr 
Sr ð4Þ

Under this assumption the contrast threshold relates to the energy threshold
value (
Sr):

Ct ¼
Sr =Apr E br ð5Þ

This result is consistent with the experimental findings of Hester (1968) and Anthony
(1981a), that contrast threshold decreases with increasing target size (Riccos law)
and ambient light level (Aksnes and Utne 1997).

Theoretical Models
Considerable attention has focused on visual feeding in relation to prey selection
and optimal foraging strategy, and several models describe foraging and prey selection
strategies of visually foraging planktivorous fish (Werner and Hall, 1974; Confer and
Blades, 1975). Eggers (1977) introduced a more quantitative model based on a fish’s
visual capability or visual range, in which prey encounter was the product of the
predator’s visual field swimming speed and prey density (according to Holling, 1966).
Eggers developed three equations that predicted the sighting distance of a predator
under different sets of combinations of prey size, inherent contrast, ambient illumi-
nation and turbidity. More recently, Aksnes and Giske (1993) presented a similar
model (later modified by Aksnes and Utne, 1997). In this model the effects of the
above-mentioned factors on visual range were expressed in a single equation. Using
this model, Giske et al. (1994) studied theoretically several aspects of mortality risk
 114 A.C. UTNE-PALM

in zooplankton and macroplankton vis-à-vis environmental variables (beam and diffuse

light attenuation and ambient light level) and prey characteristics (body size, inherent
contrast). Their model predicted turbidity to be a beneficial factor for macroplankton
and small pelagic fish (1–10 cm) feeding on microplankton (0.1–1 mm), as they gain
protection from the turbidity without reducing to the same extent their encounter
rate with zooplankton. The effects of turbidity on predation risk were highly dependent
on prey size due to the size-dependent impact of the diffuse attenuation coefficient (k)
(Kirk, 1980) and beam attenuation coefficient (c) (Giske et al., 1994) (Fig. 1a).
Similarly, in an ongoing modelling study Fiksen et al. (submitted) predict the search
volume of herring larva feeding on copepods and fish feeding on herring larva. Their
model predicts that if turbidity improves prey contrast only slightly, the larva will
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increase its search volume, while the predator’s search volume will be reduced by
turbidity (Fig. 1b). However, when the beam attenuation coefficient reaches higher
levels the veiling brightness effect will begin to degrade the target image even over
the short distances which a fish larvae ‘‘sees’’ plankton. It therefore seems unlikely
that the effect of turbidity on prey contrast will persist at high beam attenuation
values, as assumed in Figure 1b.

FIGURE 1a Redrawn from Giske et al. (1994). Predicted effect of beam attenuation and prey size (0.1 mm
to 10 cm) on the mortality risk of zooplankton. Light (1000 mmol m2 s1) equals a sunny summer day.
 VISUAL FEEDING IN TURBID WATER 115
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FIGURE 1b Redrawn from Fiksen et al. (submitted). Predicted effect of the beam attenuation coefficient
(turbidity), on the search volume of a herring larva eating nauplia (circles) and a fish eating herring larvae
(squares). When contrast is not effected by turbidity (open symbols), both predator and prey experience
reduced feeding abilities. If the contrast is slightly improved as turbidity increases (black symbols), the larvae
will improve its search volume, while the predator’s search volume is still reduced by the turbidity.

NEGATIVE EFFECTS OF TURBIDITY

Experiments have shown that increased turbidity has a negative effect on fish that
feed visually (e.g. Vinyard and O’Brien, 1976; Confer et al., 1978; Crowl, 1989;
Gregory and Northcote, 1993; Utne, 1997) as predicted (Duntley, 1943; Zaneveld
et al., 1979; Witherspoon et al., 1988). In most of these studies increasing turbidity
had a log-linear/hyperbolic relationship (Vinyard and O’Brien, 1976; Confer et al.,
1978; Gregory and Northcote, 1993; Benfield and Minello, 1996; Miner and Stein,
1996; Utne, 1997; Vogel and Beauchamp, 1999) (Fig. 2). A hyperbolic relation has
been predicted theoretically (Aksnes and Utne, 1997).
Suspended particles between predator and prey scatter light and interfere with
detection in the same way that fog affects long-distance vision, but has little effect on
detection of close objects. The negative effect of turbidity is dependent on the type of
prey (size, contrast, prey behaviour) that a fish is hunting. For a large piscivore that
detects its prey from a long distance, increasing turbidity will be more disruptive
than for a small planktivore which detects its planktonic prey at a short distance
(Chesney, 1989; Giske et al., 1994; Fiksen et al., submitted) (Fig. 1a and b). Turbidity
may thus have a positive (anti-predator) effect on small species and juvenile fish, but a
negative effect on larger piscivore species; this might explain observed changes in
community structure as a function of turbidity (Blaber and Blaber, 1980; Levy and
Northcote, 1982; Cyrus and Blaber, 1987b). Thus turbid areas are found to be avoided
by large predators because of their lowered foraging ability and greater physiological
stress (Ruiz et al., 1993, Chesapeake Bay; Maes et al., 1998, Zeescheld Estuary,
Belgium). In a study of species composition relative to environmental gradients in
some floodplain lakes of the Orinoco River in Venezuela, Rodrı́guez and Lewis
 116 A.C. UTNE-PALM
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FIGURE 2 Redrawn from Gregory and Northcote (1993). Effect of turbidity on the reaction distance of
juvenile chinook salmon (Oncorhynchus tshawytscha) for Artemia sp. prey. (regression based on median
values; bars include 50% of data points).

(1997) found that fish with good sensory adaptation to low light (rod-cells dominant in
retina) were dominant in turbid lakes, while visually oriented fish predominated in clear
lakes (cone-cells dominant in retina). Furthermore, seasonal changes produced a
decline in the proportion of visually oriented fish concomitant with a fall in transpar-
ency. Similarly, in a comparative study of the community structure of fjords on the
west coast of Norway, Eiane et al. (1999) found jellyfish to be dominant and fish
predators to be absent in a fjord with permanently high light absorbency. This jellyfish
fjord was more influenced by coastal water (a shallow sill maintained a low exchange
rate with oceanic water), resulting in much higher light absorption than in the other
fjords.

POSITIVE EFFECTS OF TURBIDITY

Although increasing turbidity tends to decrease the visual range of fish, a certain degree
of turbidity, albeit relatively low, may increase the reaction distance (distance between
predator and prey at time of detection), the growth and the feeding rates of both fish
(Gregory and Northcote, 1993; Utne, 1997; Rowe and Dean, 1998) and fish larvae
(Boehlert and Morgan, 1985; Miner and Stein, 1993; Bristow and Summerfelt, 1994;
Bristow et al., 1996) when compared to clear waters (Fig. 3).
The two most common explanations for enhancement of feeding at low levels of
turbidity are (1) the physical effect hypothesis, where turbidity increases the contrast
between the prey and its background (Hinshaw, 1985) and (2) the motivation hypoth-
esis, that increased turbidity leads to increased feeding motivation caused by a lowered
risk of predation (Gregory and Northcote, 1993).
 VISUAL FEEDING IN TURBID WATER 117
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FIGURE 3 Redrawn from Boelhert and Morgan (1985). Effect of periodic resuspensions of estuarine
sediment upon the feeding success (rotifers consumed per feeding larvae) in larval Pacific herring (Clupea
harengus pallasi). Error bars indicate  1 standard error. Numbers of feeding larvae range from 54–259.

In agreement with these findings an increase in reaction distance occurred at inter-

mediate turbidity in a study of the two-spotted goby (Gobiusculus flavescens) (Utne-
Palm, 1999) and rainbow trout (Oncorhynchus mykiss) (Barrett et al., 1992).
In both the goby and rainbow trout the initial increase in reaction distance occurred
at a lower turbidity level than in larval bluegills (Lepomis macrochirus) (Miner & Stein,
1993) and larval pacific herring (Clupea harengus palasi) (Boehlert & Morgan, 1985),
supporting the physical effect hypothesis. Accordingly, juveniles of many marine and
anadromous species seem to prefer rivers and estuaries with high concentrations of sus-
pended sediments that could provide protection from larger piscivorous fish (Blaber
and Blaber, 1980; Levy and Northcote, 1982; Cyrus and Blaber, 1987a; Gregory and
Levings, 1998). Although experimental and field observations support the physical
effect hypotheses, they do not refute the motivation hypothesis.
Results show that turbidity has a positive effect on juvenile fish by enabling them to
engage in activities that would be risky under clearer conditions, including increased
feeding rates (Gregory and Northcote, 1993; Gregory, 1994), migratory activity (Ginetz
and Larkin, 1976), reduced use of shelter (low-risk area) (Gradall and Swenson, 1982;
Gregory, 1993) and increased use of open water (high-risk area) (Miner and
Stein, 1996).
It was concluded that young fish and crustaceans used the highly turbid Zeeschelde
estuary in Belgium as a refuge from predators (Maes et al., 1998). Experiments have
demonstrated that animals reduce or eliminate anti-predator behaviour under turbid
conditions. Since this behaviour is costly, a reduction in anti-predator behaviour
should also result in a compensatory increase in feeding rates (Abrahams and
Kattenfeld, 1997).
 118 A.C. UTNE-PALM

Analysis of visual predation in relation to light and turbidity by Giske et al. (1994)
and Fiksen et al. (submitted), showed a notable decrease in predation risk of fish
from 1–10 cm, versus a smaller decrease in predation risk of individual 1–10 mm
copepods. This analysis supports the motivation hypothesis and combines it with the
physical effect hypothesis where the size difference between juvenile or larval fish
and their prey is a possible explanation for the larva fish’s preference for a turbid
environment (Fig. 1a and b).

TURBIDITY AND ITS INTERACTION WITH OTHER

PHYSICAL PROPERTIES OF WATER AND PREY
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Light and Turbidity

Increased light intensity has a positive effect on the visual range of fish up to a certain
saturation level (Vinyard and O’Brien, 1976; Confer et al., 1978; Utne, 1997).
Investigations of contrast thresholds in goldfish (Carassius auratus) (Hester, 1968)
and cod (Gadus morhua) (Anthony, 1981a), have shown that contrast threshold
decreases log-linearly as background radiance increases.
For the interaction between light and turbidity, the latter has a positive effect on
visual feeding when the light level is medium to high. At low light levels, however,
the detection of prey may be limited by absolute light levels rather than contrast.
Miner and Stein (1993) found reduced success of ingestion in bluegill larvae at low
light levels.
If intensity is above the light saturation level of a fish, light might have a negative
effect on prey detection, because light increases backscattering and thereby would
reduce the contrast between the prey and its background (Cerri, 1983; Loew and
McFarland, 1990; Guthrie and Muntz, 1993). Thus, above light-saturation the negative
effect from backscattering should become more pronounced with increasing turbidity.
This relation has been demonstrated experimentally in planktivores and bluegills
(Lepomis macrochirus) feeding on Daphnia spp. (Vinyard and O’Brien, 1976) (Fig. 4)
as well as piscivores, e.g. trout (Salvelinus namaycush) feeding on salmonid prey
(Vogel and Beauchamp, 1999).
However, in the goby, G. flavescens a positive relation existed between turbidity
and contrast threshold (Fig. 5) (Utne-Palm, 1999). Turbidity seemed to have the
same effect as increasing background radiance had on the contrast threshold of
cod (Gadus morhua) (Anthony, 1981a). It is questionable whether this reflects a
true change in the contrast threshold of this fish. In current visual models turbidity
is represented only by increased beam attenuation (c), which is related to reaction
distance (r) (Eq. (3)), but turbidity also affects the inherent contrast.

Turbidity and Prey Contrast

As concluded above, turbidity increases the contrast between prey and background,
concurrently with the scattering of light and subsequent loss of detection. Thus for a
fish predator to identify a target in a turbid or dim light medium, enough light must
be reflected from a sufficiently large target to activate the visual response as well as
to provide sufficient visual contrast to distinguish it from the background. The walleye
 VISUAL FEEDING IN TURBID WATER 119
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FIGURE 4 Redrawn from Vinyard and O’Brien (1976). Three-dimensional plot of juvenile bluegill’s
(Lepomis macrochirus) reactive distance vs. illuminance and turbidity. Prey being 2.5 mm Daphnia pulex.

FIGURE 5 Redrawn from Utne-Palm (1999). Contrast threshold (Ct) of the three different prey types
related to increasing turbidity or beam attenuation (c). Vertical bars indicate standard deviation. NS, No
significant difference in contrast threshold between transparent immobile and transparent mobile copepods;
NS¤, no significant difference in contrast threshold between mobile red and mobile transparent copepods.
Data points are offset slightly, so that standard deviation could be presented.
 120 A.C. UTNE-PALM

(Stizostedion vitreum) is an example of a species that has eyes that are adapted to
turbid and scotopic environment. Adult and subadult walleyes are primarily crepuscu-
lar or nocturnal foragers in clear water (Ali et al., 1977), but they may be diurnal
predators in turbid environments (Ryder, 1977). Their retina is specialised in two
ways to enhance scotopic vision or vision in a turbid environment, as both tapetum
lucidum and macroreceptors are present. The tapetum lucidum increases retinal light
sensitivity while the macroreceptors increase the ability to recognise slight contrast
differences (Locket, 1977). Both the tapetum and the macroreceptors develop during
the walleye’s first year of life, and the development of scotopic vision correlates with
the change from planktivore to piscivore (Vandenbyllaardt et al., 1991). A similar
ontogenetic change is also found in the visual system of yellow perch (Perca flavescens)
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which shifts from pelagic planktivore to demersal piscivore (Whal et al., 1993).
Ontogenetic change in the developmental timing of scotopic vision makes sense
in light of the predictions of Giske et al. (1994) and Fiksen et al. (submitted) (Fig. 1a
and b), where turbidity has a positive effect on planktivores (juvenile walleyes) and a
negative effect on piscivores (adult walleyes).

Wavelength and Turbidity

Contrast will be detected in the form of brightness or colour. The relative importance
of colour and brightness contrast is determined by the fish’s visual pigments, the reflec-
tance characteristics of the prey, the radiance level and the spectral distribution of
the ambient light, as well as by the fish’s visual sensitivity to these properties
(Lythgoe, 1968; Munz and McFarland, 1977; Douglas and Hawryshyn, 1990). The
spectral composition of a fish’s cone pigments is often related to the water colour
and the visual tasks of the species (Lythgoe, 1980, 1984; Anthony, 1981b; Bargarinao
and Hunter, 1983). Physiologically to minimise contrast reduction a fish is faced
with two opposing visual tasks – detection of non-reflecting (dark) and reflecting
(bright) targets. One way, that many fish solve this dilemma and maximise the contrast
of both dark and bright targets is to possess photoreceptors that have different visual
pigments – one matched and the other offset from the spectrum of the ambient light
under water (see McFarland and Munz, 1975 for details). Pure seawater is blue due
to the inherent properties of the water itself, chiefly Rayliegh scattering and absorption
of short wavelengths and absorption of long wavelengths (Rayleigh, 1889; Smith
and Tyler, 1967). In contrast, in more turbid water the high amount of particles and
dissolved substances attenuates the shorter wavelengths and the spectral composition
is shifted toward longer wavelengths. Thus, coastal water with its high content of
dissolved organic matter is greener. Freshwater often contains large amounts of
suspended material (humus) which rapidly attenuates light and shifts the spectral
range of available light to wavelengths in the red or even infra-red (Bowmaker,
1995). Freshwater fish and inshore fish in turbid environments with freshwater run-
off are therefore often more red-sensitive than pelagic or deep-sea species (Munz,
1958, 1964; Hester, 1968; Munz and McFarland, 1977; Shand et al., 1988).
In a study of male mating colour display in three-spined sticklebacks (Gasterosteus
aculeatus) McDonald et al. (1995) found that sticklebacks from more red shifted
turbid lakes had a melanistic mating colour instead of the common red colour associ-
ated with lakes of relatively clear water (blue-shifted lakes). They suggested that skin
pigmentation preference, which has a sexual attraction function, is probably a function
 VISUAL FEEDING IN TURBID WATER 121

of its efficiency in generating visual contrast rather than its intrinsic spectral character-
istic. They tested female’s preference for male breeding coloration with video imaging
techniques where red and black-throated males were superimposed on either blue or
red backgrounds. Even females belonging to a red-throated population were found
to respond preferentially to the black-throated male image, when this was viewed
against a red background, indicating that preference was determined by brightness
contrast rather than spectral difference (McDonald et al., 1995). A further study by
McDonald and Hawryshyn (1995) of the same stickleback populations showed that
sticklebacks from the red-shifted lakes had monochromatic red-sensitive vision (on
and off response), while sticklebacks from the blue-shifted lakes were most sensitive
to blue (on and off response) and intermediately sensitive to red wavelengths (on
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response). Sticklebacks from a moderately red-shifted lake were most sensitive to red
wavelengths (on and off response) and had an intermediate sensitivity to blue
wavelengths (off response). These studies by McDonald and his co-authors are an
example of how colour vision in fishes has evolved to maximise contrast between
target and the ambient light in their habitat.
Ultraviolet light has been regarded of little significance in the visual behaviour of
fish, due to its rapid attenuation in water. However, recent studies have shown that
UV visual cone pigments are present in many fish (reviewed by Douglas et al., 1995).
According to Losey et al. (1999) there is sufficient UV light in clear seawater to sustain
vision to depths beyond 100 m. Even in coastal waters and many freshwater habitats,
where particulates and dissolved organics attenuate UV light further, the number of
UV photons at moderate depths remains well above the visual thresholds of most
fishes (Losey et al., 1999). Due to Rayleegh scattering, the short UV wavelengths is
highly scattered, increasing the UV background-light and causing nearby objects to
stand out in silhouette while more distant ones disappear behind a veil. Loew et al.
(1993) showed that planktivorous juvenile yellow perch (Perca flavescens) detect and
recognise prey using only near-UV visual cues, and that sensitivity in the near UV
was greater than in other regions of the spectrum (see also Loew and Wahl, 1991).
Furthermore, Loew et al. (1996) showed that larval silversides (Atherinops affinis)
not only possess a single cone with peak absorbance at about 355 nm, but importantly
that the larvae actually engulf prey when exposed to the UV light alone. This is the first
study to actually prove feeding, not just prey detection. Browman et al. (1994) found
that UV light had a positive effect on the detection of plankton in juvenile rainbow-
trout (Oncorhynchus mykiss) and pumpkinseed (Lepomis gibbosus). Interestingly,
adult P. flavescens lack near-UV receptors (Cameron, 1982). The loss of UV receptor
with growth are shown in rainbow-trout (Browman and Hawryshyn, 1994) and
Atlantic salmon (Salmon salar) (Kunz et al., 1994). This could be a result of them
becoming demersal as adults or a response to the fact that their prey size and reaction
field have increased making UV light less useful in visual prey detection.
Zooplankton are also known to both scatter and absorb ultraviolet light (Giguere
and Dunbrack, 1990). This property may make them more or less susceptible to an
ultraviolet-sensitive predator.
Studies have shown that UV light is necessary for the perception of polarised light by
fish, and that some fish with the ability to detect UV light are also capable of detecting
polarised light (Hawryshyn and McFarland, 1987; Hawryshyn, 1992; Losey et al., 1999).
Detection of polarised light enables a fish to see further under water, and enhances the
contrast between target and background (Lythgoe and Hemmings, 1967). In a recent
 122 A.C. UTNE-PALM

study Shashar et al. (1998) showed that polarised light helps squid detect their trans-
parent zooplankton prey. Shashar et al. (2000) recently showed how cuttlefish uses
polarisation vision to detect silvery prey fish. They suggest that polarisation vision
can be used to brake the countershading camouflage of light-reflecting silvery fish.
This leads to the prediction that polarisation based predation may also be common
in polarisation-sensitive planktivorous and piscivorous fish. In a resent study of sockeye
salmon (Oncorhynchus nerka) Flamarique (2000) suggests that the presence of ultravio-
let cones in salmonid retina coincide with times of prominent feeding on zooplankton
and/or small fish.
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Turbidity and Prey Colour

Three-spine sticklebacks (Gasterosteus aculeatus L.) attacked transparent Daphnia
magna in preference to red coloured ones when presented against a red background,
whereas first choice on a pale background was a red D. magna (Ohguchi, 1981).
Likewise Browman and Marcotte (1987) showed that combinations of background
colour and prey colour affected feeding preferences of Atlantic salmon (Salmo salar
L.). Fish probably evolved colour vision to enhance contrast and, thus, improve the
visibility of objects against the background (McFarland and Munz, 1975).
In the two-spotted goby (G. flavescens) an increase in reaction distance at intermedi-
ate levels of turbidity was more pronounced for transparent than for red prey (cope-
pods), and the contrast threshold was similarly lower for transparent than for red
prey at low and intermediate turbidity levels (Fig. 5) (Utne-Palm, 1999). In turbid
water, light scattering illuminates transparent copepods from all directions, possibly
decreasing their apparent transparency and increasing their contrast against the
backlighting (Boehlert and Morgan, 1985). Further, if gobies can detect UV light, it
is possible that the inherent contrast of the transparent copepod was increased due
to the transparent copepod either scattering or absorbing UV light. If gobies are UV
sensitive, a prey item that differentially absorbs ultraviolet light will appear as a dark
object against a background rich in ultraviolet photons. In open water this effect
would be most pronounced when the target is viewed from below, since downwelling
light is relatively rich in ultraviolet radiation. Alternatively, a prey that differentially
scatters ultraviolet light will appear brighter against a background poorer in ultraviolet
photons.
This is an example of how the optical properties of both prey and suspended particles
might act together; making the total picture quite complicated.

Prey Size and Mobility

Reaction distance is a function of prey size, given that a larger apparent size will
stimulate a larger area of the retina (Lazzaro, 1987), which in turn will lower the
contrast threshold (Ware, 1973; Eggers, 1977). Most studies support the idea of reac-
tion distance increasing with prey size (Vinyard and O’Brien, 1976; Confer et al.,
1978; Schmidt and O’Brien, 1982; Wright and O’Brien, 1982; Howick and O’Brien,
1983; Li et al., 1985; Walton et al., 1997). This relationship has been interpreted as
linear (i.e. Confer et al., 1978; Wright and O’Brien, 1982; Howick and O’Brien,
1983), which is predicted from theoretical approaches (Eggers, 1977; Aksnes and
Giske, 1993). Furthermore, the theoretical approach of Eggers (1977) and Aksnes
 VISUAL FEEDING IN TURBID WATER 123

and Giske (1993) predicts that size-dependent visual range will be valid regardless of
illumination and turbidity.
Vinyard and O’Brien (1976) observed that the negative effect of turbidity on reactive
distance of bluegills was greater for large prey (Daphnia pulex) than small prey (Fig. 6).
In bluegills, Miner and Stein (1993) found a shift towards smaller prey under conditions
of increasing turbidity in intense light situations. The shift towards smaller prey in
an increasingly turbid environment predicts a higher predation risk for smaller
zooplankton (Giske et al., 1994; Fig. 1a). In clear water larger zooplankton suffer a
higher predation risk than smaller zooplankton because they can be detected over
long distances. However, this relation will change when turbidity increases, as light
absorption and scattering will decrease the visual range, and the search volume for
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small and large prey will becomes more alike.

Motion is known to be detected by special motion-detection receptive fields (e.g.
Sekuler et al., 1978), but like prey size, prey mobility can also be expected to increase

FIGURE 6 Redrawn from Vinyard and O’Brien (1976). Relationship between juvenile bluegill’s (Lepomis
macrochirus) reaction distance and prey size (Daphnia pulex) for different turbidities (JTU) at a constant
illuminance.
 124 A.C. UTNE-PALM

the area stimulated on the predator’s retina, decreasing the contrast threshold and
increasing the reaction distance (Eggers, 1977).
Prey mobility increased the reaction distance of fish in clear water (Wright and
O’Brien, 1982; Howick and O’Brien, 1983; Holmes and Gibson, 1986; Crowl, 1989),
but for turbid water Crowl (1989) found no positive effect of prey movement on the
reaction distance of largemouth bass (Micropterus salmoides). A positive effect was
found for gobies (G. flavescens) (Utne-Palm, 1999). Crowl (1989) explained his result
as a consequence of largemouth bass using different foraging tactics as prey visibility
changes. The goby also showed the expected decrease in contrast threshold when prey
was mobile rather than stationary (Fig. 5), an effect that was enhanced by increasing
turbidity.
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Turbidity has most commonly been viewed as ‘‘pollution’’ resulting from unnatural
high levels of suspended sediment, most often caused by human activity. Studies on the
effects of extreme levels of suspended sediments do therefore pervade the literature.
However, ‘‘pollution-like’’ effects are observed for the most part only at turbidity
levels well beyond those to which the species, or the life stage in question, would
be exposed to as a normal feature of their environment. In the visual world of fish
the effect of natural turbidity levels is a double edged sword. Experimental and theor-
etical studies indicate that turbidity may enhance the visual ability of young stages of
fish (planktivore) – through enhanced background contrast – while at the same
levels, may negatively influence the visual feeding ability of adult fish (piscivore) –
through increased veiling brightness. Turbidity has therefore a contradicting effect at
different trophic levels or at different life stages, which implicates turbidity as an im-
portant ecological structuring factor in the aquatic environment.

Acknowledgements
The Norwegian Research Council (NFR) supported this work, through a strategic
program granted to the Department of Fisheries and Marine Biology, University of
Bergen. The author thanks IMC (International Marine Centre, Oristrano, Sardinia)
and P. Domenici for organising this symposium issue. The author thanks the following
journals for their kind permission to use figures earlier published in their journals:
Canadian Journal of Fisheries and Aquatic Science and the Journal of Fisheries
Research Board Canada, National Research Council Canada Research Press;
Hydrobiologia, Kluwer Academic Publishers; Journal of Fish Biology, Academic
Press and the Fisheries Society of the British Isles; Vie et Milieu – Life and
Environment, Laboratoire Arago France. I also want to thank W.N. McFarland and
R.S. Gregory for their thorough and constructive comments, which greatly improved
this paper. Further I like to thank D.L. Aksnes and Ø. Fiksen for their support and
valuable suggestions concerning the manuscript.

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