Contrasting post-settlement selection results in many-to-one mapping of high performance phenotypes in the Hawaiian waterfall-climbing goby <Emphasis Type="Italic">Sicyopterus stimpsoni</Emphasis>

Natural selection drives adaptive evolution, but contrasting environmental pressures may lead to trade-offs between phenotypes that confer different performances. Such trade-offs may weaken the strength of selection and/or generate complex fitness surfaces with multiple local optima that correspond to different selection regimes. We evaluated how differences in patterns of phenotypic selection might promote morphological differences between subpopulations of the amphidromous Hawaiian waterfall-climbing goby, Sicyopterus stimpsoni. We conducted laboratory experiments on fish from the islands of Kaua‘i and Hawai‘i (the “Big Island”) to compare patterns of linear and nonlinear selection, and the opportunity for selection, that result from two contrasting pressures, predator evasion and waterfall climbing, which vary in intensity between islands. We found directional and nonlinear selection were strongest when individuals were exposed to their primary selective pressures (predator evasion on Kaua‘i, waterfall climbing on the Big Island). However, the opportunity for selection was greater for the non-primary pressure: climbing on Kaua‘i, predator evasion on the Big Island. Canonical rotation of the nonlinear gamma matrix demonstrated that individuals from Kaua‘i and the Big Island occupy regions near their local fitness peaks for some traits. Therefore, selection for predator evasion on Kaua‘i and climbing on the Big Island may be less effective in promoting morphological changes in this species, because variation of functionally important traits in their respective environments may have been reduced by directional or stabilizing selection. These results demonstrate that despite constraints on the opportunities for selection, population differences in phenotypic traits can arise due to differences in selective regimes. For S. stimpsoni, sufficient variation exists in other locomotor traits, allowing for necessary levels of performance in the contrasting selective regime (i.e., climbing on Kaua‘i and predator evasion on the Big Island) through many-to-one-mapping, which may be essential for the survival of local populations in an evanescent island environment.     

Proximate causes of intraspecific variation in locomotor performance in the lizard Gallotia galloti.

To understand the evolution of biological traits, information on the degree and origins of intraspecific variation is essential. Because adaptation can take place only if the trait shows heritable variation, it is important to know whether (at least) part of the trait variation is genetically based. We describe intra- and interindividual variation in three performance measures (sprint speed, climbing, and clambering speed) in juvenile Gallotia galloti lizards from three populations and examine how genetic, environmental (incubation temperature), and ontogenetic (age, size) effects interact to cause performance variation. Moreover, we test whether the three performance traits are intercorrelated phenotypically and genetically. Sprint speed is highest in juveniles incubated at the lowest temperature (26 degrees C) irrespective of population. Climbing speed differs among populations, and the differences persist at least until the lizards are 30 wk old. This suggests that the three populations experience different selective pressures. Moreover, mass, snout-vent length, and hindlimb length seem to affect climbing performance differently in the three populations. The variation in sprinting and climbing ability appears to be genetically based. Moreover, the two performance traits are intercorrelated and thus will not evolve independently from each other. Clambering speed (i.e., capacity to climb up an inclined mesh) varies among individuals, but the origin of this variation remains obscure.     

Muscle fiber type distribution in climbing Hawaiian gobioid fishes: ontogeny and correlations with locomotor performance

Three species of Hawaiian amphidromous gobioid fishes are remarkable in their ability to climb waterfalls up to several hundred meters tall. Juvenile Lentipes concolor and Awaous guamensis climb using rapid bursts of axial undulation, whereas juvenile Sicyopterus stimpsoni climb using much slower movements, alternately attaching oral and pelvic sucking disks to the substrate during prolonged bouts of several cycles. Based on these differing climbing styles, we hypothesized that propulsive musculature in juvenile L. concolor and A. guamensis would be dominated by white muscle fibers, whereas S. stimpsoni would exhibit a greater proportion of red muscle fibers than other climbing species. We further predicted that, because adults of these species shift from climbing to burst swimming as their main locomotor behavior, muscle from adult fish of all three species would be dominated by white fibers. To test these hypotheses, we used ATPase assays to evaluate muscle fiber type distribution in Hawaiian climbing gobies for three anatomical regions (midbody, anal, and tail). Axial musculature was dominated by white muscle fibers in juveniles of all three species, but juvenile S. stimpsoni had a significantly greater proportion of red fibers than the other two species. Fiber type proportions of adult fishes did not differ significantly from those of juveniles. Thus, muscle fiber type proportions in juveniles appear to help accommodate differences in locomotor demands among these species, indicating that they overcome the common challenge of waterfall climbing through both diverse behaviors and physiological specializations.     

Sexual Selection and Dynamics of Jaw Muscle in Tupinambis Lizards

Sexual dimorphism patterns provide an opportunity to increase our understanding of trait evolution. Because selective forces may vary throughout the reproductive period, measuring dimorphism seasonally may be an interesting approach. An increased male head size may be important in intersexual and intrasexual interactions. In Tupinambis lizards, a big head is attributed in part to a large adductor muscle mass. Competition for mating can differ in species with different sex ratio and different degrees of sexual size dimorphism. We examined sexual differences in mass of the pterygoideus muscle, its temporal variation throughout the reproductive period and the relationship between muscle and reproductive condition in Tupinambis merianae and T. rufescens. We characterized sexual size dimorphism and sex ratio in both species. Mature males had larger jaw muscles than mature females in both species, mainly during the reproductive season. The dimorphism in jaw muscle was due to an increase in muscle mass in sexually active males. Seasonal increases in muscle mass and variation between immature and mature individuals suggest that the jaw muscle might be a secondary sexual character. We propose that the pterygoideus muscle may act as a signal of reproductive condition of males because it is associated with testis size and sperm presence. The patterns of sexual dimorphism in jaw muscle in both species were similar; however, the comparison shows how sexual characters remain dimorphic in different competition contexts and in species with different degrees of body size dimorphism. Our results suggest that jaw muscle as sexual character could be influenced by inter- and intrasexual selective pressures.     

Hydrologic regimes as potential drivers of morphologic divergence in fish

Fishes often exhibit phenotypic divergence across gradients of abiotic and biotic selective pressures. In streams, many of the known selective pressures driving phenotypic differentiation are largely influenced by hydrologic regimes. Because flow regimes drive so many attributes of lotic systems, we hypothesized fish exhibit phenotypic divergence among streams with different flow regimes. We used a comparative field study to investigate the morphological divergence of Campostoma anomalom (central stonerollers) among streams characterized by highly variable, intermittent flow regimes and streams characterized by relatively stable, groundwater flow regimes. We also conducted a mesocosm experiment to compare the plastic effects of one component of flow regimes, water velocity, on morphology of fish from different flow regimes. We observed differences in shape between flow regimes likely driven by differences in allometric growth patterns. Although we observed differences in morphology across flow regimes in the field, C. anomalum did not exhibit morphologic plasticity in response to water velocity alone. This study contributes to the understanding of how complex environmental factors drive phenotypic divergence and may provide insight into the evolutionary consequences of disrupting natural hydrologic patterns, which are increasingly threatened by climate change and anthropogenic alterations.     

Phenotypic plasticity in gene expression contributes to divergence of locally adapted populations of Fundulus heteroclitus

We examine the interaction between phenotypic plasticity and evolutionary adaptation using muscle gene expression levels among populations of the fish Fundulus heteroclitus acclimated to three temperatures. Our analysis reveals shared patterns of phenotypic plasticity due to thermal acclimation as well as non‐neutral patterns of variation among populations adapted to different thermal environments. For the majority of significant differences in gene expression levels, phenotypic plasticity and adaptation operate on different suites of genes. The subset of genes that demonstrate both adaptive differences and phenotypic plasticity, however, exhibit countergradient variation of expression. Thus, expression differences among populations counteract environmental effects, reducing the phenotypic differentiation between populations. Finally, gene‐by‐environment interactions among genes with non‐neutral patterns of expression suggest that the penetrance of adaptive variation depends on the environmental conditions experienced by the individual.     

Morphological selection in an extreme flow environment: body shape and waterfall-climbing success in the Hawaiian stream fish Sicyopterus stimpsoni

Flow characteristics are a prominent factor determining body shapes in aquatic organisms, and correlations between body shape and ambient flow regimes have been established for many fish species. In this study, we investigated the potential for a brief period of extreme flow to exert selection on the body shape of juvenile climbing Hawaiian gobiid fishes. Because of an amphidromous life history, juvenile gobies that complete an oceanic larval phase return to freshwater habitats, where they become adults. Returning juveniles often must scale waterfalls (typically with the use of a ventral sucker) in order to reach the habitats they will use as adults, thereby exposing these animals to brief periods of extreme velocities of flow. Hydrodynamic theory predicts that bodies with larger suckers and with lower heights that reduce drag would have improved climbing success and, thus, be well suited to meet the demands of the flows in waterfalls. To test the potential for the flow environment of waterfalls to impose selection that could contribute to differences in body shape between islands, we subjected juvenile Sicyopterus stimpsoni to climbing trials up artificial waterfalls (～100 body lengths) and measured differences in body shape between successful and unsuccessful climbers. Waterfalls appear to represent a significant selective barrier to these fishes, as nearly 30% failed our climbing test. However, the effects of selection on morphology were not straightforward, as significant differences in shape between successful and unsuccessful climbers did not always match hydrodynamic predictions. In both selection experiments and in adult fish collected from habitats with different prevailing conditions of flow (the islands of Hawai'i versus Kaua'i), lower head heights were associated with exposure to high-flow regimes, as predicted by hydrodynamic theory. Thus, a premium appears to be placed on the reduction of drag via head morphology throughout the ontogeny of this species. The congruence of phenotypic selection patterns observed in our experiments, with morphological character divergence documented among adult fish from Hawai'i and Kaua'i, suggests that differences in morphology between subpopulations of adult climbing gobies may result, at least in part, from the selective pressures of high-velocity flows encountered by migrating juveniles.     

Phenotypic divergence and inter-specific trait correlation in a plant-pollinator/seed predator mutualism

Plant-pollinator interactions have been suggested as key drivers of morphological divergence and speciation of the involved taxa. These interactions can also promote sexual dimorphism in both the plant and pollinator, particularly if the pollinator is also a seed-eater and/or exerts different selection pressures on male and female plants. Here we tested the hypotheses that plant-pollinator interactions can be reflected in trait variation and sexual dimorphism in both organisms within and across populations. Across nine European populations, we examined intraspecific variation and sexual dimorphism in phenotypic traits potentially involved in the plant–insect interaction of the dioecious white campion Silene latifolia (Caryophyllaceae) and its specialist pollinator Hadena bicruris (Noctuidae). This interaction is expected to entail sex-specific selective pressures, as female moths lay eggs on female plants and the larvae predate on the seeds during their development. We compared divergence in phenotypic traits among populations and between sexes within populations, examined correlations between plant and pollinator traits, and between phenotypic distances and genetic distances among co-occurring populations for both plants and insects. We found key differences in phenotypic traits across populations of both the plant and moth, though only in the moth were these differences correlated with geographic distances. We also found evidence for sexual dimorphism in the plant but not in the pollinator. Evolution of floral sexual dimorphism in S. latifolia most likely results from the joint contribution of different selective forces, including biotic interactions with H. bicruris moths.     

Fiber type composition of the human male trapezius muscle: enzyme-histochemical characteristics

The human trapezius muscle has an origin that is more extensive than that of any other body muscle; it has a complex macroscopic structure with fibers running in different directions. Histochemical analysis of multiple samples, obtained from different parts of the trapezius muscle from five males, showed marked differences in the distribution and the cross-sectional fiber area of the fiber types among different parts of the muscle as well as among individuals. As revealed by the mATPase activity, after different levels of alkaline and acidic preincubations, the lower third of the descending portion, the transverse, and the ascending portions of the muscle had a predominance of type I fibers (low mATPase activity at pH 9.4), whereas the most superior parts of pars descendens had a higher frequency of type II fibers (high mATPase activity at pH 9.4). The fibers of the most superior parts of the muscle were considerably smaller compared with those in all the other parts. In sections stained for NADH-TR, moth-eaten fibers were observed within parts of the descending portion. Their location and their larger fiber area, compared with that of ordinary type I fibers, may be related to frequent and/or continuous use of these fibers. In conclusion, the differences in fiber type composition between the different parts of the muscle probably reflect different functional demands on the trapezius muscle in various head, neck, and shoulder movements. We suggest that the interindividual differences in muscle fiber composition are due, at least in part, to genetic factors.     

Quantifying the temporospatial expression of postnatal porcine skeletal myosin heavy chain genes.

Postnatal skeletal muscle fiber type is commonly defined by one of four major myosin heavy chain (MyHC) gene isoforms (slow/I, 2a, 2x, and 2b) that are expressed. We report on the novel use of combined TaqMan quantitative real-time RT-PCR and image analysis of serial porcine muscle sections, subjected to in situ hybridization (ISH) and immunocytochemistry (IHC), to quantify the mRNA expression of each MyHC isoform within its corresponding fiber type, termed relative fiber type-restricted expression. This versatile approach will allow quantitative temporospatial comparisons of each MyHC isoform among muscles from the same or different individuals. Using this approach on porcine skeletal muscles, we found that the relative fiber type-restricted expression of each postnatal MyHC gene showed wide spatial and temporal variation within a given muscle and between muscles. Marked differences were also observed among pig breeds. Notably, of the four postnatal MyHC isoforms, the 2a MyHC gene showed the highest relative fiber type-restricted expression in each muscle examined, regardless of age, breed, or muscle type. This suggests that although 2a fibers are a minor fiber type, they may be disproportionately more important as a determinant of overall muscle function than was previously believed.     

Fatty acid composition in adductor muscle of juvenile scallops (Pecten maximus) from five Norwegian populations reared in the same environment

Adductor muscle fatty acid (FA) composition was compared in Pecten maximus juveniles originating from five locations along the Norwegian coast from 59°N to 65°N to detect possible population differences. Broodstock sized scallops were translocated to sea conditions by a scallop hatchery near Bergen (60°N) before spawning in February 2006. The scallop larvae and juveniles were reared in the same environment for two years and 10 individuals from each population were sampled in May 2007 and in May 2008 for analysis of the FAs in the adductor muscle. The total lipid content determined as total amount of FAs were 5.7 ± 0.3 mg per g tissue, and no significant difference was found among the five populations. The polyunsaturated FAs made up close to 60% of the total, with 20:5n3 and 22:6n3 dominating. The saturated FA content was approximately 30%, while the monounsaturated FA were less abundant (7–10%). The FA composition of the muscles of the five populations was similar within each year, with larger differences between the years. Multivariate, supervised learning method PLS, applied pairwise, showed distinct FA composition, between the scallops from the five locations, indicating population differences. The relatedness between the populations was different in the two years, but the distinct FA profiles of the adductor muscle could be used to distinguish between scallop populations on a local scale. The results indicate habitat-specific lipid metabolism which may have important implications for the scallop aquaculture industry in the context of producing well adapted individuals for the specific locations.     

Cranial muscles of the anurans leiopelma hochstetteri and ascaphus truei and the homologies of the mandibular adductors in lissamphibia and other gnathostomes

The frogs Ascaphus truei and Leiopelma hochstetteri are members of the most basal lineages of extant anurans. Their cranial muscles have not been previously described in full and are investigated here by dissection. Comparison of these taxa is used to review a controversy regarding the homologies of the jaw adductor muscles in Lissamphibia, to place these homologies in a wider gnathostome context, and to define features that may be useful for cladistic analysis of Anura. A new muscle is defined in Ascaphus and is designated m. levator anguli oris. The differences noted between Ascaphus and Leiopelma are in the penetration of the jaw adductor muscles by the mandibular nerve (V³). In the traditional view of this anatomy, the paths of the trigeminal nerve branches define homologous muscles. This scheme results in major differences among frogs, salamanders, and caecilians. The alternative view is that the topology of origins, insertions, and fiber directions are defining features, and the nerves penetrate the muscle mass in a variable way. The results given here support the latter view. A new model is proposed for Lissamphibia, whereby the adductor posterior (levator articularis) is a separate entity, and the rest of the adductor mass is configured around it as a folded sheet. This hypothesis is examined in other gnathostomes, including coelacanth and lungfish, and a possible sequence for the evolution of the jaw muscles is demonstrated. In this system, the main jaw adductor in teleost fish is not considered homologous with that of tetrapods. This hypothesis is consistent with available data on the domain of expression of the homeobox gene engrailed 2, which has previously not been considered indicative of homology. Terminology is discussed, and “adductor mandibulae” is preferred to “levator mandibulae” to align with usage in other gnathostomes. J. Morphol., 2011. © 2011 Wiley‐Liss, Inc.     

Internal morphology and function of paired fins in the epaulette shark,hemiscyllium ocellatum

Paired fins and associated internal structures of the epauletic sharkHemiscyllium ocellatum, were described on the basis of three specimens. A comparison with other genera showed the epaulette shark to be, characterized by two elongated basal cartilages articulating with a distally projecting articular condyle on the coracoid, a loosely separated radial series with an intermediate series, a levator pectoralis inferior muscle and an anterolaterally developed depressor pectoralis muscle in the pectoral fin, and an elongated anterior pelvic basal cartilage articulating with a distally projecting articular condyle and an anterolaterally developed depressor pelvicus muscle in the pelvic fin. In captivity, the sharks exhibited both upright and crawling behavior on the bottom by using the pectoral and pelvic fins and bending the body. The distinctive morphological characters are shared by otherHemiscyllium species and are suggested as important factors enabling their unique behavior associated with a complex coral reef habitat.     

Morphological correlates of river velocity and reproductive development in an ornamented stream fish

Adaptive phenotypic divergence can arise when environments vary in ways favoring alternative phenotypic optima. In aquatic habitats, the costs of locomotion are expected to increase with water velocity, generally favoring a more streamlined body and the reduction of traits that produce drag. However, because streamlining in fish may come at the cost of maneuverability, the net benefits of drag reduction can differ not only among habitats, but also among individuals (or classes of individuals) that rely on locomotion for different uses (e.g., males vs. females or adults vs. juveniles). We tested these predictions by exploring relationships among river velocity, body streamlining, ornamental fin size, and male reproductive condition in the steelcolor shiner (Cyprinella whipplei), a small-bodied North American cyprinid. Overall, males in peak reproductive condition (defined by the development of sexually dimorphic tubercles) had less streamlined bodies and larger ornamental fins than males in lower reproductive condition or individuals lacking these secondary sexual characters (females and immature males). There was a relationship between river velocity and body streamlining only for males in peak reproductive condition, but it was in the opposite direction of our predictions: these males were less streamlined in faster rivers. We found only weak support for the prediction that ornamental fin size would be negatively associated with river velocity. Overall, these results suggest either that drag is not an important selective pressure in these habitats, or that the sexual selection advantages of a deep body and large fin compensate any natural selection costs for C. whipplei males. This study highlights the often overlooked diversity of selective pressures acting on streamlining in fishes, and can offer novel insights and predictions allowing a more nuanced understanding of fish ecomorphology.     

Natural radioactivity can explain clinal variation in the expression of melanin-based traits

A recent study shows that the expression of pheomelanin-based coloration in barn owls follows a continuous gradient across Europe as a result of local adaptation. The selective pressures that promote local adaptation remain, however, unknown. Here we hypothesize and test that natural radioactivity levels follow a similar spatial gradient to that of pheomelanin-based color in Europe and thus represents a potential selective pressure. The rationale is that the production of pheomelanin consumes glutathione (GSH), a key intracellular antioxidant, and that GSH is particularly susceptible to ionizing radiation, which depletes antioxidants. As predicted, the intensity of pheomelanin-based coloration in 18 populations of barn owls was negatively associated with terrestrial γ-dose rates across Europe. Therefore, we propose that natural selection acts against barn owls that present the molecular basis to produce large amounts of pheomelanin in those populations that are exposed to high levels of natural radioactivity, as in these populations individuals would require higher antioxidant resources to combat oxidative stress. This is the first time that natural radioactivity levels are related to the expression of a phenotypic trait.     

Phenotypic diversity,population structure and stress protein-based capacitoring in populations of Xeropicta derbentina,a heat-tolerant land snail species

The shell colour of many pulmonate land snail species is highly diverse. Besides a genetic basis, environmentally triggered epigenetic mechanisms including stress proteins as evolutionary capacitors are thought to influence such phenotypic diversity. In this study, we investigated the relationship of stress protein (Hsp70) levels with temperature stress tolerance, population structure and phenotypic diversity within and among different populations of a xerophilic Mediterranean snail species (Xeropicta derbentina). Hsp70 levels varied considerably among populations, and were significantly associated with shell colour diversity: individuals in populations exhibiting low diversity expressed higher Hsp70 levels both constitutively and under heat stress than those of phenotypically diverse populations. In contrast, population structure (cytochrome c oxidase subunit I gene) did not correlate with phenotypic diversity. However, genetic parameters (both within and among population differences) were able to explain variation in Hsp70 induction at elevated but non-pathologic temperatures. Our observation that (1) population structure had a high explanatory potential for Hsp70 induction and that (2) Hsp70 levels, in turn, correlated with phenotypic diversity while (3) population structure and phenotypic diversity failed to correlate provides empirical evidence for Hsp70 to act as a mediator between genotypic variation and phenotype and thus for chaperone-driven evolutionary capacitance in natural populations.     

Muscular and intraocular pressure responses among ocular-hypertensive subjects: Is there a rationale for biofeedback?

Several animal and human investigations have indicated that intraocular pressure (IOP) levels may be associated with extreme drug-induced changes in the extraocular muscles. Further, recent data suggest that, among individuals with normal IOP level, moderate increases in facial muscle (EMG) activity around the eye while the eye is open are associated with increases in IOP. To investigate further the relationship between facial EMG activity and IOP levels and to examine a group of individuals with elevated IOP levels, subjects were recruited from outpatients at an optometry clinic. Three groups of subjects were selected: a group of ocular hypertensive subjects who showed elevated pressures at the optometry clinic and upon the day of testing, a group of labile ocular hypertensive subjects who evinced elevated pressures during their visit to the optometry clinic but lower pressures on the day of testing, and a group of normal IOP subjects who showed normal pressures both during their optometry clinic visit and on the day of testing. To investigate anxiety differences, subjects were administered the State-Trait Anxiety Inventory, but subsequent analysis revealed no group differences. To evaluate the role of stress upon muscle (EMG) functioning around the eye, subjects were subjected to imagery and standardized mental arithmetic stressors; analyses of these results also revealed no significant group differences. Finally, subjects were given EMG biofeedback for muscle activity around the eye while IOP was assessed during five alternating periods in which they made decreases and increases in EMG activity. Results revealed significant group, period, and group by period interaction effects. The pattern of results is interpreted as implicating EMG activity in IOP fluctuations; the implications of these data for potential biofeedback and stress management treatments are discussed.This research was supported by a Faculty Research Grant of the University of Alabama at Birmingham.