Influence of lighting environment on social preferences in sticklebacks from two different photic habitats. II. Shoaling and mate preferences of lab-bred fishes.

Different environmental conditions may lead to diverse morphological, behavioral, and physiological adaptations of different populations of the same species. Lighting conditions, for example, vary vastly especially between aquatic habitats, and have been shown to elicit adaptations. The availability of short-wave ultraviolet (UV) light is especially fluctuating, as UV wavelengths are attenuated strongly depending on water properties. The island of North Uist, Scotland, comprises 2 differential habitat types, tea-stained and clear-water lakes, varying considerably in UV transmission. In previous studies, wild-caught 3-spined stickleback Gasterosteus aculeatus populations (3 populations of each habitat type) were tested with respect to their shoaling and mate preferences for fish viewed under UV-present and UV-absent conditions. The results revealed a habitat-dependent preference of UV cues during shoal choice (tea-stained populations: preference for UV-absent condition in tea-stained water; clear-water populations: no preference in clear-water) but an overall preference for UV-present conditions during mate choice. To assess genetic influences on these behavioral patterns, similar experiments were conducted with lab-bred F1-generations of the same stickleback populations that were raised in a common environment (i.e. standardized clear-water conditions). Offspring of sticklebacks from tea-stained lakes tended to prefer shoals viewed under UV-absent conditions (only in tea-stained water), while sticklebacks from clear-water lakes showed a significant preference for the shoal viewed under UV-present conditions in clear-water but not in tea-stained water. Mate-preference experiments demonstrated that females from the tea-stained lakes significantly preferred and females from the clear-water lakes preferred by trend the male viewed under UV-present conditions in the clear-water treatment. The results for both shoaling- and mate-preference tests were largely similar for wild-caught and lab-bred sticklebacks, thus hinting at a genetic basis for the preference patterns.

Influence of lighting environment on social preferences in sticklebacks from two different photic habitats. I. mate preferences of wild-caught females.

Ultraviolet (UV) A signals (320-400 nm) are important in mate choice in numerous species. The sensitivity for UV signals is not only assumed to be costly, but also expected to be a function of the prevailing ecological conditions. Generally, those signals are favored by selection that efficiently reach the receiver. A decisive factor for color signaling is the lighting environment, especially in aquatic habitats, as the visibility of signals, and thus costs and benefits, are instantaneously influenced by it. Although ecological aspects of color signal evolution are relatively well-studied, there is little data on specific effects of environmental UV-light conditions on signaling at these shorter wavelengths. We studied wild-caught gravid female 3-spined sticklebacks Gasterosteus aculeatus of 2 photic habitat types (tea-stained and clear-water lakes), possessing great variation in their UV transmission. In 2 treatments, tea-stained and clear-water, preferences for males viewed under UV-present (UV+) and UV-absent (UV-) conditions were tested. A preference for males under UV+ conditions was found for females from both habitat types, thus stressing the significance of UV signals in stickleback's mate choice decisions. However, females from both habitat types showed the most pronounced preferences for males under UV+ conditions under clear-water test conditions. Moreover, reflectance measurements revealed that the carotenoid-based orange-red breeding coloration in wild-caught males of both habitat types differed significantly in color intensity (higher in clear-water males) and hue (more red shifted in clear-water males) while no significant differences in UV coloration were found. The differential reflection patterns in longer wavelengths suggest that sticklebacks of both habitat types have adapted to the respective water conditions. Adaptations of UV signals in a sexual context to ambient light conditions in both behavior and coloration seem less evident.

Not just shades of grey: life is full of colour for the ocellate river stingray (Potamotrygon motoro).

Previous studies have shown that marine stingrays have the anatomical and physiological basis for colour vision, with cone spectral sensitivity in the blue to green range of the visible spectrum. Behavioural studies on Glaucostegus typus also showed that blue and grey can be perceived and discriminated. The present study is the first to assess visual opsin genetics in the ocellate river stingray (Potamotrygon motoro) and test whether individuals perceive colour in two alternative forced choice experiments. Retinal transcriptome profiling using RNA-Seq and quantification demonstrated the presence of lws and rh2 cone opsin genes and a highly expressed single rod (rh1) opsin gene. Spectral tuning analysis predicted these vitamin A1-based visual photopigments to exhibit spectral absorbance maxima at 461 nm (rh2), 496 nm (rh1) and 555 nm (lws); suggesting the presence of dichromacy in this species. Indeed, P. motoro demonstrates the potential to be equally sensitive to wavelengths from 380 to 600 nm of the visible spectrum. Behavioural results showed that red and green plates, as well as blue and yellow plates, were readily discriminated based on colour; however, brightness differences also played a part in the discrimination of blue and yellow. Red hues of different brightness were distinguished significantly above chance level from one another. In conclusion, the genetic and behavioural results support prior data on marine stingrays. However, this study suggests that freshwater stingrays of the family Potamotrygonidae may have a visual colour system that has ecologically adapted to a riverine habitat.

Body coloration as a dynamic signal during intrasexual communication in a cichlid fish.

BACKGROUND: Intrasexual competition over access to resources can lead to aggression between individuals. Because overt aggression, i.e. fights, can be costly for contestants, the communication of aggressive motivation prior to engagement in a physical fight is often mediated by conventional signals. Animals of various taxa, including fishes, display visual signals such as body coloration that can dynamically be adjusted depending on the individual's motivation. Male individuals of the West African cichlid Pelvicachromis taeniatus express a yellow body coloration displayed during courtship but also in an intrasexual competition context. RESULTS: Within-individual variation in male yellow body coloration, as quantified with standardized digital photography and representation in a CIELab color space, was examined in a mating context by exposing males to a female and in a competitive intrasexual context, i.e. in a dyadic contest. Additionally, spectrometric reflectance measurements were taken to obtain color representations in a physiological color space based on spectral sensitivities of our model species. Exposure to females did not significantly affect male color expression. However, analysis of body coloration revealed a change in within-individual color intensity and colored area after interaction with a male competitor. In dominant males, extension of coloration was positively correlated with restrained aggression, i.e. displays, which in turn explained dominance established between the two contestants. CONCLUSION: Body coloration in male P. taeniatus is a dynamic signal that is used in concert with display behavior in communication during intrasexual competition.

Differential investment in pre- and post-mating male sexual traits in response to prolonged exposure to ambient UVB radiation in a fish.

Increasing UVB radiation (UVB) reaching earth's surface following stratospheric ozone depletion is linked to serious consequences for organisms. While studies have focused on direct cytocidal and immunomodulatory effects of UVB, indirect consequences for fitness-related life-history traits are largely unexplored, although knowledge is needed to understand organismal responses to climate change. The present study investigates the effects of developmental exposure to enhanced, but ecologically relevant, ambient UVB levels on growth (body size), parental behavior (nest-building), fitness-enhancing traits of pre-mating (sexual ornamentation) and post-mating (sperm traits) sexual selection as well as their interrelationships in male three-spined sticklebacks (Gasterosteus aculeatus). Moreover, potentially underlying direct UVB effects, testicular antioxidant capacity (TAC) and testes and skin melanization were quantified. Juvenile full-siblings were split into two groups and raised until adulthood in a semi-natural set-up under light conditions including either natural (UVBnormal) or elevated UVB levels (UVBenhanced). When becoming reproductive, males were kept individually before data collection took place at their reproductive peak. The results showed that males from the UVBenhanced-group were smaller than their brothers from the UVBnormal-treatment, whereas no treatment differences were observed for sexual ornamentation, sperm traits and TAC. Moreover, UVB-stressed males built their nests faster and the relationship between body size and nest size was negative for UVBenhanced-males while being positive for UVBnormal-males. These results demonstrate that, depending on physical state, UVB-stressed males adjusted their behavior to some extent. Additionally, a trade-off between the investment in pre- and post-mating traits was found for UVBenhanced-males, i.e. the intensity of their breeding coloration was negatively correlated with sperm number whereas this relationship was reversed for UVBnormal-males, thus showing an interaction between pre- and post-mating traits regarding exposure treatment. The interaction provides first experimental evidence that differential allocation to energetically demanding pre- and post-mating components of male fitness is triggered by a key environmental stressor of climate change.

Mate assessment behavior is correlated to learning ability in female threespine sticklebacks.

In many species, males signal quality with elaborate traits, but females often show inter-individual variation in preference for these traits. Choosing a mate requires multiple cognitive steps; therefore, cognitive style (how an individual processes information) likely influences the perception of sexual signals and ability to choose a high-quality mate. An important component of cognitive style is flexibility; cognitively flexible individuals are more perceptive to shifts in cues. We hypothesized that cognitively flexible individuals would acquire more information about potential mates, better discern between two quality-signaling traits, and thus be more discriminatory. Here, we show that mate assessment is correlated to other cognitive traits. Although we did not detect an effect of cognitive style on mate preference or discrimination, we found that female threespine sticklebacks Gasterosteus aculeatus that spent more time assessing potential mates (more responsive) in a dichotomous mate choice task reached both the initial and reversal learning criterion in a spatial learning task with fewer errors. However, these highly responsive females made more consecutive mistakes immediately at the beginning of the reversal phase, suggesting that they did not quickly adapt to the environmental change but instead rapidly formed strict routines during the learning task that were eventually reversible after repeated errors. Furthermore, we found evidence for condition-dependent mate preference, with larger females preferring the high-quality male. These are among the first results that illustrate how cognitive traits might influence mate choice, which has implications for the strength and direction of sexual selection.

Enhanced ambient UVB radiation affects post-mating, but not pre-mating sexual traits in a fish.

Organisms inhabiting shallow aquatic habitats currently experience increasing levels of solar ultraviolet B radiation (UVB). UVB causes damage on cellular and molecular levels and can affect associated life-history traits either through direct exposure or indirectly through oxidative stress generation. We examined UVB effects on pre- and post-mating sexual traits in three-spined stickleback fish (Gasterosteus aculeatus). Adult, reproductively non-active males were assigned to two exposure treatments under semi-natural conditions in an outdoor experiment; one group received natural radiation (UVBnormal) whilst the other group received additional UVB (UVBenhanced). After two months, colour metrics were used to quantify male breeding colouration as pre-mating trait. At the post-mating stage, sperm morphology, number and movement as well as testes mass were determined. Males did not significantly differ in sexual ornamentation between treatments, but UVBenhanced fish had smaller testes as well as fewer and shorter sperm than UVBnormal fish. Sperm movement was not significantly different between treatments. However, in UVBenhanced males, linear and progressive movement of sperm was positively correlated with sperm morphology (head-to-tail length ratio), whereas in UVBnormal males this relationship was negative (but not significant). Additionally, there was a significant treatment by body condition interaction concerning head-to-tail length ratio, i.e. head-to-tail length ratio increased with condition in UVBnormal males whereas there was no relationship in UVBenhanced fish. Our findings reveal that increased UVB levels influence post-mating fitness-relevant traits in males whilst having no significant impact on pre-mating sexual traits, suggesting selective UVB-effects at the gamete level with consequences for reproductive performance.

Shoal sex composition and predation risk influence sub-adult threespine stickleback shoaling decisions.

Group living reduces individual predation risk most effectively when group members are behaviorally and phenotypically similar. Group preferences are influenced by the individual, the members of the shoal, and the environmental conditions. While shoaling behavior has been studied extensively in the threespine stickleback (Gasterosteus aculeatus), it is unclear whether the sex of shoal mates influences the shoal preference of non-reproductive males and females and how this changes under increasing predation risk. Although non-reproductively active sticklebacks are sexually monochromatic in appearance, sex-related differences may result in sexual segregation when shoaling. Here we show that male and female sub-adult threespine sticklebacks had contrasting preferences for shoal mate sex, and that this preference was dependent on the level of predation risk during standardized experimental choice tests. In detail, test fish shoal with the opposite sex within low predation risk trials and with same-sex shoals within high predation risk trials. This difference might be linked to activity patterns; test males were more active than females. Our results demonstrate that differences between the sexes in a species with a sexually monochromatic non-reproductive stage can result in sex-related shoaling preferences. Most studies examining sexual segregation focus on sexually dimorphic species, but these results highlight the potentially widespread occurrence of sexual segregation beyond the sexually dimorphic reproductive stage.

Long-term UVB exposure promotes predator-inspection behaviour in a fish.

Ultraviolet-B radiation (UVB) reaching the earth's surface has increased due to human-caused stratospheric ozone depletion. Whereas the harmful effects of UVB on aquatic organisms are well studied at the molecular and cellular level, recent studies have also begun to address behavioural changes caused by sublethal amounts of UVB. However, the behavioural consequences of long-term exposure to ecologically relevant UVB levels over several life stages are virtually unknown, particularly with regard to predator-prey behaviour. We found increased predator-inspection behaviour together with a smaller body length in three-spined sticklebacks (Gasterosteus aculeatus) after fish were exposed for about seven months to natural sunlight conditions with enhanced UVB, compared with full siblings exposed to natural sunlight only. The observed change in antipredator behaviour may reflect a direct behavioural response mediated through UVB-induced oxidative stress during development. Alternatively, the smaller body size in UVB-exposed fish may result in an increased inspection effort allowing them to spend more time foraging. Our findings suggest that, within the scope of environmental change, UVB radiation constitutes an important stress factor by eliciting behavioural responses that influence crucial ecological processes, such as predator-prey interactions.

The dynamics of color signals in male threespine sticklebacks Gasterosteus aculeatus.

Body coloration and color patterns are ubiquitous throughout the animal kingdom and vary between and within species. Recent studies have dealt with individual dynamics of various aspects of coloration, as it is in many cases a flexible trait and changes in color expression may be context-dependent. During the reproductive phase, temporal changes of coloration in the visible spectral range (400-700 nm) have been shown for many animals but corresponding changes in the ultraviolet (UV) waveband (300-400 nm) have rarely been studied. Threespine stickleback Gasterosteus aculeatus males develop conspicuous orange-red breeding coloration combined with UV reflectance in the cheek region. We investigated dynamics of color patterns including UV throughout a male breeding cycle, as well as short-term changes in coloration in response to a computer-animated rival using reflectance spectrophotometry and visual modeling, to estimate how colors would be perceived by conspecifics. We found the orange-red component of coloration to vary during the breeding cycle with respect to hue (theta/R50) and intensity (achieved chroma/red chroma). Furthermore, color intensity in the orange-red spectral part (achieved chroma) tended to be increased after the presentation of an artificial rival. Dynamic changes in specific measures of hue and intensity in the UV waveband were not found. In general, the orange-red component of the signal seems to be dynamic with respect to color intensity and hue. This accounts in particular for color changes during the breeding cycle, presumably to signal reproductive status, and with limitations as well in the intrasexual context, most likely to signal dominance or inferiority.

Context-dependent dynamic UV signaling in female threespine sticklebacks.

Color signals, including ultraviolet (UV) signals, are widespread throughout the animal kingdom and color changes can be influenced by reproductive and motivational state. However, studies on dynamic changes of UV signals are scarce. Three spine sticklebacks (Gasterosteus aculeatus) that show intraspecific UV communication were used to study dynamic UV signaling in females. Reflectance measurements were taken from the distended abdomen, which serves as signal of female fecundity and readiness to spawn for courting males, and the melanized dorsal region. Scans were taken during egg maturation as well as before and after stimulation with a male to investigate context-dependent color changes. We used a physiological model of vision to determine how females might be perceived by conspecifics and quantified chromatic contrasts among both body regions and between body regions and the background for all stages. Females showed a significant increase in abdominal UV intensity during egg maturation and in response to a courting male. Measures of chromatic contrast among body regions (abdomen vs. dorsal region) and against the background (abdomen vs. background) were also increased during egg maturation and in response to the male stimulus (abdomen vs. background). Our results provide evidence for dynamic UV signaling in females in a reproductive context.

Dynamic resource allocation between pre- and postcopulatory episodes of sexual selection determines competitive fertilization success.

In polyandrous mating systems, male reproductive success depends on both mate-acquisition traits (precopulatory) and sperm competitive abilities (postcopulatory). Empirical data on the interaction between these traits are inconsistent; revealing positive, negative or no relationships. It is generally expected that the investment in pre- and postcopulatory traits is mediated by environmental conditions. To test how dietary resource availability affects sexual ornamentation, sperm quality and their interrelationship in three-spined sticklebacks (Gasterosteus aculeatus), full-sibling groups were raised under three conditions differing in food quantity and/or quality (i.e. carotenoid content): (i) high-quantity/high-quality, (ii) high-quantity/low-quality or (iii) low-quantity/low-quality. After 1 year of feeding, food-restricted males developed a more intense breeding coloration and faster sperm compared with their well-fed brothers, indicating that they allocated relatively more in pre- and postcopulatory traits. Moreover, they outcompeted their well-fed, carotenoid-supplemented brothers in sperm competition trials with equal numbers of competing sperm, suggesting that food-restricted males maximize their present reproductive success. This may result in reduced future reproductive opportunities as food-restricted males suffered from a higher mortality, had an overall reduced body size, and sperm number available for fertilization. In accordance with theory, a trade-off between the investment in pre- and postcopulatory traits was observed in food-restricted males, whereas well-fed males were able to allocate to both traits resulting in a significantly positive relationship.

Habitat-dependent olfactory discrimination in three-spined sticklebacks (Gasterosteus aculeatus).

The ability to recognize conspecifics is indispensible for differential treatment of particular individuals in social contexts like grouping behavior. The advantages of grouping are multifarious, and there exist numerous additional benefits of joining aggregations of conspecifics. Recognition is based on different signals and transmitted via multiple channels, among others the olfactory channel. The sensory system or the combination of sensory modalities used in recognition processes is highly dependent on the availability and effectiveness of modalities, which are a function of the environmental conditions. Using F1-generations of six three-spined stickleback (Gasterosteus aculeatus) populations from two habitat types (tea-stained and clear-water lakes) from the Outer Hebrides, Scotland, we investigated whether individuals are able to recognize members of their own population solely based on olfactory cues and whether the habitat type an individual originated from had an influence on its recognition abilities. When given the choice (own vs. foreign population) sticklebacks from tea-stained lakes significantly preferred the odor of their own population, whereas fish from clear-water habitats did not show any preference. Moreover, fish from the two habitat types differed significantly in their recognition abilities, indicating that olfactory communication is better developed when visual signaling is disturbed. Thus, the observed odor preferences appear to be the consequence of different selective constraints and adaptations as a result of the differences in environmental conditions that have acted on the parental generations. These adaptations are likely genetically based as the differences are present in the F1-generation that had been reared under identical laboratory conditions.

The influence of ambient ultraviolet light on sperm quality and sexual ornamentation in three-spined sticklebacks (Gasterosteus aculeatus).

Exposure to enhanced levels of ambient ultraviolet (UV) radiation (UVR) can have adverse effects on aquatic organisms including damage at the cellular and molecular level and impairment of development, fecundity and survival. Much research has been conducted on the role of the harmful UVB radiation. However, due to its greater penetration in water the more abundant UVA radiation can also act as an environmental stressor. Little is known about UVR effects on sperm characteristics although sperm cells should be especially prone to UV-induced oxidative stress. Moreover, UV-related changes in oxidative status may affect the phenotypic expression of energetically costly sexual ornaments. We investigated the effects of long-term exposure to ecologically relevant levels of simulated UVA radiation on sperm quality and sexual ornamentation in three-spined sticklebacks (Gasterosteus aculeatus). Males were assigned to three spectral exposure treatments differing in the UV spectral part so that they received either enhanced, moderate or no UVA radiation. The results reveal that exposure to enhanced ambient UVA levels had detrimental effects on both male breeding coloration and sperm velocity providing evidence that UVR affects traits targeted by pre- and post-copulatory sexual selection. By highlighting the role of UVA as a factor influencing fitness-relevant traits, our findings may contribute to a better understanding of the consequences of current and future levels of solar UVR for mating systems and life history.

Visual prey detection by near-infrared cues in a fish.

Many animal species are able to perceive light wavelengths beyond those visible to humans. While numerous species are additionally sensitive to short wavelengths (UV), long wavelengths such as the near-infrared spectrum (NIR) are supposed to be unsuitable for visual perception. Here, we experimentally show that under exclusive NIR illumination, the cichlid fish Pelvicachromis taeniatus displays a clear foraging response towards NIR reflecting prey. Additional control experiments without prey indicate that the observed behavior is not a mere response to the NIR environment. These results give first evidence for NIR visual sensitivity in a functional context and thus challenge the current view about NIR perception.

Male red ornamentation is associated with female red sensitivity in sticklebacks.

Sexual selection theory proposes correlated evolutionary changes in mating preferences and secondary sexual characters based on a positive genetic correlation between preference and the preferred trait. Empirical work has provided support for a genetic covariation between female preference and male attractiveness in several taxa. Here, we study parent and offspring visual traits in threespine sticklebacks, Gasterosteus aculeatus. While focusing on the proximate basis of mating preferences, we compare the red breeding coloration of males, which strongly contributes to female choice, with their daughters' red sensitivity measured by optomotor response thresholds. We show that the red color expression of fathers correlates well with their daughters' red sensitivity. Given that a within-population genetic correlation between signal and preference was experimentally confirmed for the red coloration in sticklebacks, our results indicate a proximate mechanism in terms of perceptual sensitivity being involved in the co-evolution of female preferences and male mating signals.

Color signaling in conspicuous red sticklebacks: do ultraviolet signals surpass others?

BACKGROUND: The use of ultraviolet (UV) signals for communication tasks is widespread in vertebrates. For instance, there is a UV component to mate choice in several species. Nevertheless, it remains unclear how the signal value of the UV wave band compares to that of other regions of the animal's visible spectrum. We investigated the relative importance of UV signals compared with signals of longer wavelengths in the threespine stickleback (Gasterosteus aculeatus), a species using UV wavelengths in female and male mate choice as well as in shoaling behavior. In a choice experiment, female sticklebacks were simultaneously presented with four male visual appearances manipulated by optical filters. Each male lacked one wavelength range of the stickleback's visible spectrum corresponding to the spectral sensitivities of the four cone types. The resulting male appearances thus had no UV (UV-), no short-wave (SW-), no medium-wave (MW-) or no long-wave (LW-) body reflectance. RESULTS: Males without UV wavelengths and long wavelengths ("red") were least preferred. In contrast, the removal of medium and most notably short wavelengths left male attractiveness to females rather unaffected. Using color metrics, the effects of the four optical filters on stickleback perception of three male body regions were illustrated as quantal catches calculated for the four single cones. CONCLUSION: The removal of UV light (UV-) considerably reduced visual attractiveness of courting males to female three-spined sticklebacks particularly in comparison to the removal of short-wave light (SW-). We thus report first experimental evidence that the UV wave band clearly outranks at least one other part of an animal's visible spectrum (SW-) in the context of communication. In addition, females were also less attracted to males presented without long wavelengths (LW-) which supports the traditionally considered strong influence of the red color component on stickleback mate choice. Overall, the removal of medium wavelengths (MW-) and especially short (SW-) left male attractiveness for females rather unaffected. Our work suggests that, in addition to long wavelengths ("red"), the UV wave band contains important information for visual mate choice in sticklebacks. Hence, the previously suggested exclusive role of the characteristic red nuptial coloration in visual interactions between reproductively active stickleback conspecifics may be overestimated with UV wavelengths playing a more important role than previously suggested.

Males do not see only red: UV wavelengths and male territorial aggression in the three-spined stickleback (Gasterosteus aculeatus).

Animal colour signals serve important functions in intraspecific interactions, including species recognition, mate choice and agonistic behaviour. An increasing interest concerns ultraviolet (UV) wavelengths, for instance studies on the effect of UV in mating decisions. More recently, some studies also established that UV signals affect intrasexual interactions. We studied the role of UV during aggressive encounters between male three-spined sticklebacks (Gasterosteus aculeatus), a species in which UV has an effect on female and male mate choice and shoaling behaviour. To that aim, we compared the aggressive response of a territorial male to male intruders, either seen in UV-including (UV+) or UV-lacking (UV-) conditions. Our prediction was that, if UV wavelengths are used in male-male competition, a territorial male should show less competitive behaviour towards an intruder representing a lower threat, i.e. the one presented without UV light. Male sticklebacks showed significantly lower levels of aggression towards male opponents lacking an UV component to their coloration than male opponents possessing this colour component. Discrimination was not influenced by a difference in brightness between the UV+ and UV- stimuli. Finally, we present some reflectance-spectrophotometrical data of two skin regions (cheek and abdomen) of the experimental males and analysed relationships between colorimetric variables, body variables and behaviour. Our study emphasises that UV visual cues are of importance in different communicational tasks in the three-spined stickleback.

UV matters in shoaling decisions.

Shoaling behaviour in fish is influenced by numerous factors, such as familiarity, kinship, group size and shoal composition. Grouping decisions are based on both olfactory and visual cues. The visual system of many vertebrates is extended into the ultraviolet (UV) wave range as in three-spined sticklebacks (Gasterosteus aculeatus, L.). We investigated whether the presence or absence of UV wavelengths has an influence on shoaling behaviour in this species. Reproductively non-active three-spined sticklebacks were given the choice between two shoals, equal in numbers of individuals, which could be seen either through a UV-transmitting [UV(+)] or a UV-blocking [UV(-)] filter. Test fish preferred to join the shoal seen under UV(+) conditions. Due to differences in quantal flux between the UV(+) and UV(-) filters used, control experiments with neutral-density optical filters were performed in order to clarify the role of luminance. Here, test fish spent significantly more time near shoals that were seen in a darker environment, suggesting a potential trade-off between UV radiation and lower brightness during shoal choice. To our knowledge, these results demonstrate for the first time that shoaling decisions are influenced by UV wavelengths.