Pronounced heritable variation and limited phenotypic plasticity in visual pigments and opsin expression of threespine stickleback photoreceptors.

Vertebrate colour vision is mediated by the differential expression of visual pigment proteins (opsins) in retinal cone photoreceptors. Many species alter opsin expression during life, either as part of development or as a result of changes in habitat. The latter, a result of phenotypic plasticity, appears common among fishes, but its cellular origin and ecological significance are unknown. Here, we used adult threespine stickleback fish from different photic regimes to investigate heritable variability and phenotypic plasticity in opsin expression. Fish from clear waters had double cones that expressed long (LWS) and middle (RH2) wavelength opsins, one per double cone member. In contrast, fish from red light-shifted lakes had double cones that were >95% LWS/LWS pairs. All fish had single cones that predominantly expressed a short wavelength (SWS2) opsin but ultraviolet cones, expressing a SWS1 opsin, were present throughout the retina. Fish from red light-shifted lakes, when transferred to clear waters, had a ∼2% increase in RH2/LWS double cones, though double cone density remained constant. Comparison of visual pigment absorbance and light transmission in the environment indicated that the opsin complements of double cones maximized sensitivity to the background light, whereas single cones had visual pigments that were spectrally offset from the dominant background wavelengths. Our results indicate that phenotypic plasticity in opsin expression is minor in sticklebacks and of questionable functional significance.

Role of the iridescent eye in stickleback female mate choice.

Many vertebrates exhibit prominent body colours that are used in courtship and territorial communication. Some fishes also have an eye whose iris becomes iridescent during the mating season, as in the threespine stickleback. Behavioural studies in this species have focused on the redness of the throat/jaw as the primary determinant of female mate choice. Unlike the iridescent eye, however, the red throat/jaw is not present in all stickleback populations, suggesting that the colour of the eye may be equally important for female mate choice. Here, we used data on photoreceptors and environmental light to assess body conspicuousness and the colour contrast of courtship signals for stickleback populations living in a range of waters, from clear (mesotrophic) to red light shifted (dystrophic). This analysis indicated that the redness of the throat/jaw is expressed to enhance the contrast of the eye. To test the importance of eye colour as a courtship signal, we carried out mate choice experiments in which females were presented with identical videos of a courting male but for the colour of the eye and/or the throat/jaw. Females did not choose based on differences in throat/jaw redness between videos, but preferred males with the highest contrast between the eye and the throat/jaw. This result points to the blue iridescent eye as a primary courtship signal in stickleback female mate choice.

Multi-generation selective landscapes and sub-lethal injuries in stickleback.

The interaction between predation landscape and phenotypic variability within prey populations is of substantial significance in evolutionary biology. Extending from several decades of studies at a remote freshwater lake on Haida Gwaii, western Canada, we analyze the incidence of predator-induced sub-lethal injuries in 8,069 wild-captured threespine stickleback (Gasterosteus aculeatus) and using cohort analyses test whether the distribution of injuries informs the selective landscape influencing the bell-shaped frequency distribution of the traits. Our results indicate that (1) the incidence of injuries varies among phenotypes differing in the number and position of lateral plates, (2) these differences occur only among younger fish, (3) the incidence of injuries is inversely related to the estimated population frequencies of plate phenotypes, with the modal phenotype generally having the fewest injuries, (4) direct estimates of selective differentials and relative fitness based on analyses of 1,735 fish from 6 independent yearly cohorts indicates statistically informative elevated differentials in phenotypes with greater number of plates and elevated relative fitness of non-modal phenotypes, and (5) there are significant differences among yearly cohorts in strength and direction of selection, and an increased prevalence of diversifying versus stabilizing selection despite longer-term stasis (4 decades) in trait means. We conclude that the presence of multiple "optimal" phenotypes complements the renewed interests in quantifying short-term temporal or spatial variation in ecological processes in studies of fitness landscapes and intrapopulation variability.

Polymorphism and multiple correlated characters: Do flatfish asymmetry morphs also differ in swimming performance and metabolic rate?

Phenotypic polymorphisms often differ in multiple correlated traits including morphology, behavior, and physiology, all of which can affect performance. How selection acts on these suites of traits can be complex and difficult to discern. Starry flounder (Platichthys stellatus) is a pleuronectid flatfish that exhibits rare polymorphism for the direction of eye migration and resulting whole-body asymmetry. P. stellatus asymmetry morphs differ subtly in several anatomical traits, foraging behavior, and stable isotope signatures, suggesting they may be ecologically segregated, yet performance and metabolic differences are unknown.Here we tested the hypothesis that sinistral and dextral P. stellatus asymmetry morphs diverge in performance and routine metabolic rate (RMR) by comparing prolonged swimming endurance (time to exhaustion at a constant swimming speed), fast-start swimming velocity and acceleration, and rate of oxygen consumption. Based on subtle morphological differences in caudal tail size, we expected sinistral P. stellatus to have superior prolonged swimming endurance relative to dextral fish, but inferior fast-start performance.Sinistral P. stellatus exhibited both significantly greater prolonged swimming performance and fast-start swimming performance. However, sinistral P. stellatus also exhibited greater RMR, suggesting that their general swimming performance could be enhanced by an elevated metabolic rate.Divergence between P. stellatus asymmetry morphs in swimming performance and metabolic rates contributes to growing evidence of ecological segregation between them, as well as our understanding of possible ecological consequences of asymmetry direction in flatfishes. These data provide an example of the complexity of polymorphisms associated with multiple correlated traits in a rare case of asymmetry polymorphism in a marine flatfish species.

Particle size distributions, size concentration relationships, and adherence to hands of selected geologic media derived from mining, smelting, and quarrying activities.

Hand-to-mouth activity, especially in children, is a potentially significant pathway of exposure to soil contaminants. Hand-mouthing behavior is of particular concern in areas impacted by mining, smelting, and quarrying activities as these activities may lead to elevated levels of heavy metals in soil. In order to estimate potential exposures to contaminated geologic media attributable to hand-to-mouth contact, it is useful to characterize adherence of those media to skin, as contaminant concentrations in adhered media may differ greatly from unfractionated, whole media concentrations. Such an investigation has been undertaken to aid estimation of exposures to arsenic, cadmium, lead, and zinc in nine different geologic media collected in the Pacific Northwest region of the United States. After establishing the particle size distribution of each medium (fractions <63 µm, 63-150 µm, 150-250 µm, and 250 µm-2mm were determined) and target elemental concentrations within each particle size fraction, an active handling protocol involving six volunteers was conducted. Wet media always adhered to a greater extent than dry media and adhered media generally had higher elemental concentrations than bulk media. Regression analyses suggest smaller particle fractions may have higher elemental concentrations. Results of application of a maximum likelihood estimation technique generally indicate that handling of dry media leads to preferential adherence of smaller particle sizes, while handling of wet media does not. Because adhered material can differ greatly in particle size distribution from that found in bulk material, use of bulk concentrations in exposure calculations may lead to poor estimation of actual exposures. Since lead has historically been a metal of particular concern, EPA's Integrated Exposure Uptake Biokinetic (IEUBK) Model was used to examine the potential consequences of evaluating ingestion of the selected media assuming concentrations in adhering versus bulk media.

The ecology of asymmetry in stickleback defense structures.

Slight departures from bilateral symmetry are usually associated with reduced fitness. Here we show that an insular freshwater population of Gasterosteus aculeatus exhibits spatial and temporal segregation according to the incidence and direction of asymmetry in the number of bony lateral plates, which are important predator-defense structures. We analyzed 11,263 fish collected from 19 full lake transects over three years, and found that signed asymmetries for plate number were slightly left-side biased. Asymmetries occurred in 55% of the stickleback with more asymmetrics found during cold weather, and more left-biased individuals found during windy conditions, possibly due to environmentally driven activity levels that differ among asymmetric forms. Absolute plate asymmetries were randomly distributed in the lake, but there was a strong shift in signed plate asymmetry from a mean of zero in littoral zones to left-side bias in limnetic zones, probably due to microhabitat choices among asymmetric forms. Video data of avian piscivores on the lake show laterality in prey handling, providing a potential mechanism for asymmetric prey morphology. Our results imply a complex relationship between slight asymmetries and niche space that is relevant to the broader literature on the evolutionary implications of developmental instability and intrapopulation variability.