Finotypic plasticity: Predator-induced plasticity in fin size, darkness and display behaviour in a teleost fish.

Fish fins are remarkable devices of propulsion. Fin morphology is intimately linked to locomotor performance, and hence to behaviours that influence fitness, such as foraging and predator avoidance. This foreshadows a connection between fin morphology and variation in predation risk. Yet, whether prey can adjust fin morphology according to changes in perceived risk within their lifetime (a.k.a. predator-induced plasticity) remains elusive. Here, we quantify the structural size of five focal fins in crucian carp (Carassius carassius) following controlled manipulations to perceived predation risk (presence/absence of pike Esox lucius). We also assess if crucian carp respond to increased predation risk by shifts in dorsal fin colouration, and test for differences in how fish actively use their dorsal fins by quantifying the area of the fin displayed in behavioural trials. We find that crucian carp show phenotypic plasticity with regards to fin size as predator-exposed fish consistently have larger fins. Individuals exposed to perceived predation risk also increased dorsal fin darkness and actively displayed a larger area of the fin to potential predators. Our results thus provide compelling evidence for predator-induced fin enlargement, which should result in enhanced escape swimming performance. Moreover, fin-size plasticity may evolve synergistically with fin colouration and display behaviour, and we suggest that the adaptive value of this synergy is to enhance the silhouette of deep-bodied and hard-to-capture prey to deter gape-limited predators prior to attack. Together, our results provide new perspectives on the role of predation risk in development and evolution of fins.

The functional nasal anatomy of the pike, Esox lucius L.

Olfactory flow in fishes is a little-explored area of fundamental and applied importance. We investigated olfactory flow in the pike, Esox lucius, because it has an apparently simple and rigid nasal region. We characterised olfactory flow by dye visualisation and computational fluid dynamics, using models derived from X-ray micro-computed tomography scans of two preserved specimens. An external current induced a flow of water through the nasal chamber at physiologically relevant Reynolds numbers (200-300). We attribute this externally-induced flow to: the location of the incurrent nostril in a region of high static pressure; the nasal bridge deflecting external flow into the nasal chamber; an excurrent nostril normal to external flow; and viscous entrainment. A vortex in the incurrent nostril may be instrumental in viscous entrainment. Flow was dispersed over the olfactory sensory surface when it impacted on the floor of the nasal chamber. Dispersal may be assisted by: the radial array of nasal folds; a complementary interaction between a posterior nasal fold and the ventral surface of the nasal bridge; and the incurrent vortex. The boundary layer could delay considerably (up to ~ 3 s) odorant transport from the external environment to the nasal region. The drag incurred by olfactory flow was almost the same as the drag incurred by models in which the nasal region had been replaced by a smooth surface. The boundary layer does not detach from the nasal region. We conclude that the nasal bridge and the incurrent vortex are pivotal to olfaction in the pike.

Adaptations of early development to local spawning temperature in anadromous populations of pike (Esox lucius).

BACKGROUND: In the wake of climate change many environments will be exposed to increased and more variable temperatures. Knowledge about how species and populations respond to altered temperature regimes is therefore important to improve projections of how ecosystems will be affected by global warming, and to aid management. We conducted a common garden, split-brood temperature gradient (4.5 °C, 9.7 °C and 12.3 °C) experiment to study the effects of temperature in two populations (10 families from each population) of anadromous pike (Esox lucius) that normally experience different temperatures during spawning. Four offspring performance measures (hatching success, day degrees until hatching, fry survival, and fry body length) were compared between populations and among families. RESULTS: Temperature affected all performance measures in a population-specific manner. Low temperature had a positive effect on the Harfjärden population and a negative effect on the Lervik population. Further, the effects of temperature differed among families within populations. CONCLUSIONS: The population-specific responses to temperature indicate genetic differentiation in developmental plasticity between populations, and may reflect an adaptation to low temperature during early fry development in Harfjärden, where the stream leading up to the wetland dries out relatively early in the spring, forcing individuals to spawn early. The family-specific responses to temperature treatment indicate presence of genetic variation for developmental plasticity (G x E) within both populations. Protecting between- and within-population genetic variation for developmental plasticity and high temperature-related adaptive potential of early life history traits will be key to long-term viability and persistence in the face of continued climate change.

A digital dissection of two teleost fishes: comparative functional anatomy of the cranial musculoskeletal system in pike (Esox lucius) and eel (Anguilla anguilla).

Advances in X-ray computed tomography (CT) have led to a rise in the use of non-destructive imaging methods in comparative anatomy. Among these is contrast-enhanced CT scanning, which employs chemical stains to visualize soft tissues. Specimens may then be 'digitally dissected', producing detailed, three-dimensional digital reconstructions of the soft- and hard-tissue anatomy, allowing examination of anatomical structures in situ and making accurate measurements (lengths, volumes, etc.). Here, we apply this technique to two species of teleost fish, providing one of the first comprehensive three-dimensional (3D) descriptions of teleost cranial soft tissue and quantifying differences in muscle anatomy that may be related to differences in feeding ecology. Two species with different feeding ecologies were stained, scanned and imaged to create digital 3D musculoskeletal reconstructions: Esox lucius (Northern Pike), predominantly a suction feeder; and Anguilla anguilla (European eel), which captures prey predominantly by biting. Muscle cross-sectional areas were calculated and compared between taxa, focusing on muscles that serve important roles in feeding. The adductor mandibulae complex - used in biting - was larger in Esox than Anguilla relative to head size. However, the overall architecture of the adductor mandibulae was also very different between the two species, with that of Anguilla better optimized for delivering forceful bites. Levator arcus palatini and sternohyoideus - which are used in suction feeding - are larger in Esox, whereas the levator operculi is larger in Anguilla. Therefore, differences in the size of functionally important muscles do not necessarily correlate neatly with presumed differences in feeding mode.

Photoreceptors and eyes of pikeperch Sander lucioperca, pike Esox lucius, perch Perca fluviatilis and roach Rutilus rutilus from a clear and a brown lake.

The photoreceptors and eyes of four fish species commonly cohabiting Fennoscandian lakes with different light transmission properties were compared: pikeperch Sander lucioperca, pike Esox lucius, perch Perca fluviatilis and roach Rutilus rutilus. Each species was represented by individuals from a clear (greenish) and a humic (dark brown) lake in southern Finland: Lake Vesijärvi (LV; peak transmission around 570 nm) and Lake Tuusulanjärvi (LT; peak transmission around 630 nm). In the autumn, all species had almost purely A2-based visual pigments. Rod absorption spectra peaked at c.526 nm (S. lucioperca), c. 533 nm (E. lucius) and c. 540 nm (P. fluviatilis and R. rutilus), with no differences between the lakes. Esox lucius rods had remarkably long outer segments, 1.5-2.8-fold longer than those of the other species. All species possessed middle-wavelength-sensitive (MWS) and long-wavelength-sensitive (LWS) cone pigments in single, twin or double cones. Rutilus rutilus also had two types of short-wavelength sensitive (SWS) cones: UV-sensitive [SWS1] and blue-sensitive (SWS2) cones, although in the samples from LT no UV cones were found. No other within-species differences in photoreceptor cell complements, absorption spectra or morphologies were found between the lakes. However, E. lucius eyes had a significantly lower focal ratio in LT compared with LV, enhancing sensitivity at the expense of acuity in the dark-brown lake. Comparing species, S. lucioperca was estimated to have the highest visual sensitivity, at least two times higher than similar-sized E. lucius, thanks to the large relative size of the eye (pupil) and the presence of a reflecting tapetum behind the retina. High absolute sensitivity will give a competitive edge also in terms of short reaction times and long visual range.

Changes of sperm morphology, volume, density, and motility parameters in northern pike during the spawning period.

Sexually mature males (BW = 1600 ± 150 g and TL = 235 ± 30 mm) of northern pike (Esox lucius L.) were randomly selected from a pond to record changes in their sperm quality parameters (spermatozoa morphology, sperm volume, density, and motility parameters) during the spawning season. The morphological and motility parameters changed significantly during the reproductive season with following trends. Only, head width was not changed during the spawning season. The longest spermatozoa and its flagellar length were found at the middle of spawning period (TL = 38.24 ± 0.37 µm and 35.14 ± 0.26 µm) and shortest at the beginning of spawning period (TL = 34.81 ± 0.29 µm and 32.53 ± 0.18 µm). Other morphological characters were always the lowest at the beginning of spawning period. Sperm volume was changed from 0.33 ± 0.3 ml in February, 0.43 ± 0.2 ml in March to 0.24 ± 0.1 ml in April, and density from 16.2 ± 0.2 × 109 spermatozoa ml-1 in February, 19.4 ± 0.2 × 109 spermatozoa ml-1 in March to 4.8 ± 0.2 × 109 spermatozoa ml-1 in April. Same sperm velocity was observed in all spawning terms at 10 and 20 s after activation. Higher velocity was found at 30 and 40 s after activation in sperm collected at the middle and the end of spawning period. Significantly, higher percentage of motile sperm was observed at 20, 30, and 40 s after activation in sperm sampled at the end of spawning period. This study supports the hypothesis that longer spermatozoa swim faster.

Species richness in the African pike genus Hepsetus: a perfect match between genetics and morphology.

In this study, morphological differences were found that corroborate earlier results that showed the existence of six species within the previously monospecific African pike genus Hepsetus. Additional genetic data (coI, mtDNA and rag1, nDNA) confirm the morphology-based species delineations. Deep genetic divergences imply a relatively old age for diversification within the genus. An identification key for the six species is provided in the present study.

Evolutionary divergence of adult body size and juvenile growth in sympatric subpopulations of a top predator in aquatic ecosystems.

Evolutionary theory predicts that different selective regimes may contribute to divergent evolution of body size and growth rate among populations, but most studies have focused on allopatric populations. Here, we studied five sympatric subpopulations of anadromous northern pike (Esox lucius) in the Baltic Sea subjected to allopatric habitats for a short period of their life cycle due to homing behavior. We report differences in adult body size among subpopulations that were in part due to variation in growth rate. Body size of emigrating juveniles also differed among subpopulations, and differences remained when individuals were reared in a common environment, thus indicating evolutionary divergence among subpopulations. Furthermore, a QST-FST comparison indicated that differences had evolved due to divergent selection rather than genetic drift, possibly in response to differences in selective mortality among spawning habitats during the allopatric life stage. Adult and juvenile size were negatively correlated across subpopulations, and reconstruction of growth trajectories of adult fishes suggested that body size differences developed gradually and became accentuated throughout the first years of life. These results represent rare evidence that sympatric subpopulations can evolve differences in key life-history traits despite being subjected to allopatric habitats during only a very short fraction of their life.

Reciprocal behavioral plasticity and behavioral types during predator-prey interactions.

How predators and prey interact has important consequences for population dynamics and community stability. Here we explored how predator-prey interactions are simultaneously affected by reciprocal behavioral plasticity (i.e., plasticity in prey defenses countered by plasticity in predator offenses and vice versa) and consistent individual behavioral variation (i.e., behavioral types) within both predator and prey populations. We assessed the behavior of a predator species (northern pike) and a prey species (three-spined stickleback) during one-on-one encounters. We also measured additional behavioral and morphological traits in each species. Using structural equation modeling, we found that reciprocal behavioral plasticity as well as predator and prey behavioral types influenced how individuals behaved during an interaction. Thus, the progression and ultimate outcome of predator-prey interactions depend on both the dynamic behavioral feedback occurring during the encounter and the underlying behavioral type of each participant. We also examined whether predator behavioral type is underlain by differences in metabolism and organ size. We provide some of the first evidence that behavioral type is related to resting metabolic rate and size of a sensory organ (the eyes). Understanding the extent to which reciprocal behavioral plasticity and intraspecific behavioral variation influence the outcome of species interactions could provide insight into the maintenance of behavioral variation as well as community dynamics.

Inferring occurrence of growth checks in pike (Esox lucius) scales by using sequential isotopic analysis of otoliths.

RATIONALE: Sequential analysis of otolith oxygen isotope (δ(18) O(oto) ) values was performed by classical mechanical microdrilling and dual inlet mass spectrometry to infer the timing of growth checks in pike (Esox lucius) scales from Lake Annecy (France). It indicated that the first check on scales occurs before the first winter of life of the fish. METHODS: Eleven fish caught in February and June 2008 were studied. Intra-individual isotopic profiles exhibit cyclic variations that are consistent with that of modeled δ(18) O(oto) values using variations in epilimnion temperature and otolith-specific isotope fractionation equations. The positions of growth checks on scales were compared with the periodic variations of δ(18) O(oto) values on otolith radii using an allometric relationship. RESULTS: All individuals presented at least one check deposited before the first winter. Deposition of some checks might be linked to a shift in fish diet or habitat or to catch-and-release induced stress. CONCLUSIONS: Although coupling between sequential analysis of δ(18) O(oto) values and observation of scale growth features has been shown to be useful, there are limitations in the methodology, especially for the growth-attenuated region of otoliths associated with older ages.

The inner ears of Northern Canadian freshwater fishes following exposure to seismic air gun sounds.

An earlier study examined the effects of exposure to seismic air guns on the hearing of three species of fish from the Mackenzie River Delta in Northern Canada [Popper et al. (2005). "Effects of exposure to seismic airgun use on hearing of three fish species," J. Acoust. Soc. Am. 117, 3958-3971]. The sound pressure levels to which the fishes were exposed were a mean received level of 205-209 dB re 1 microPa (peak) per shot and an approximate received mean SEL of 176-180 dB re 1 microPa(2) s per shot. In this report, the same animals were examined to determine whether there were effects on the sensory cells of the inner ear as a result of the seismic exposure. No damage was found to the ears of the fishes exposed to seismic sounds despite the fact that two of the species, adult northern pike and lake chub, had shown a temporary threshold shift in hearing studies.

Causes and consequences of intra-specific variation in vertebral number.

Intraspecific variation in vertebral number is taxonomically widespread. Much scientific attention has been directed towards understanding patterns of variation in vertebral number among individuals and between populations, particularly across large spatial scales and in structured environments. However, the relative role of genes, plasticity, selection, and drift as drivers of individual variation and population differentiation remains unknown for most systems. Here, we report on patterns, causes and consequences of variation in vertebral number among and within sympatric subpopulations of pike (Esox lucius). Vertebral number differed among subpopulations, and common garden experiments indicated that this reflected genetic differences. A QST-FST comparison suggested that population differences represented local adaptations driven by divergent selection. Associations with fitness traits further indicated that vertebral counts were influenced both by stabilizing and directional selection within populations. Overall, our study enhances the understanding of adaptive variation, which is critical for the maintenance of intraspecific diversity and species conservation.

Morphological study of the northern pike (Esox lucius) tongue.

The northern pike (Esox lucius) is a fresh water species belonging to the Esocidae family. It is a carnivorous fish feeding mostly on invertebrates and fishes. Due to the scantiness of relevant literature regarding the morphology of the tongue in fish we carried out this study with the aim of providing information on the dorsal surface morphology and histological structures of the tongue in E. lucius. The tongues of five E. lucius were examined using light- and scanning electron- microscopy (SEM) techniques. The SEM studies revealed the presence of numerous teeth, longitudinal mucosal strands and scattered taste buds spread on the tongue surface. Histological studies using hematoxylin and eosin and Masson's trichrome staining showed that the musculature was not visible in the tongue of E. lucius. The tongue is composed of mucosa, and submucosa supported by osteocartilagionous skeleton. The mucosa consists of several layers of unicellular mucous cells interrupted by numerous teeth. The derivation of teeth from the underlying bronchial skeleton was visible in longitudinal section. The scattered taste buds with a typical onion shape were also present. Overall, the morphological features of the E. lucius tongue together suggested its mechanical and sensory roles. The findings of this study together with morphological and physiological data from other fishes contribute to the knowledge of the nutrition and feeding behavior in aquaculture species.

Morphological and molecular evidence of three species of pikes Esox spp. (Actinopterygii, Esocidae) in France, including the description of a new species.

This integrative taxonomy study of French pikes compares morphological characters and molecular sequence data (mitochondrial COI and nuclear Plagl2 genes). In addition to the expected E. lucius, DNA sequences and morphology both support a new species in France, E. aquitanicus sp. nov. from the Charente to the Adour drainages. It is characterized by a color pattern of sides with narrow 1-1.5-scale-wide oblique vertical bands, conferring it a marbled coat, a snout only 0.9 times larger than the postorbital length, an anal fin basis 1.1-1.2 times larger than the caudal peduncle length, 101 to 121 lateral scales, 53 to 57 vertebrae, as well as 24 diagnostic sites in the COI gene and 3 in the Plagl2 gene. Partial COI sequences (131 bp) from modern and historical specimens indicate also the presence of E. cisalpinus and E. lucius during the 19th century in Lake Geneva. Morphological and molecular data points to a possible hybridization between E. lucius with both other local pike species, representing a risk for them. Their endangerment status should be evaluated rapidly in order to take conservation measures.

Behavioural syndrome in a solitary predator is independent of body size and growth rate.

Models explaining behavioural syndromes often focus on state-dependency, linking behavioural variation to individual differences in other phenotypic features. Empirical studies are, however, rare. Here, we tested for a size and growth-dependent stable behavioural syndrome in the juvenile-stages of a solitary apex predator (pike, Esox lucius), shown as repeatable foraging behaviour across risk. Pike swimming activity, latency to prey attack, number of successful and unsuccessful prey attacks was measured during the presence/absence of visual contact with a competitor or predator. Foraging behaviour across risks was considered an appropriate indicator of boldness in this solitary predator where a trade-off between foraging behaviour and threat avoidance has been reported. Support was found for a behavioural syndrome, where the rank order differences in the foraging behaviour between individuals were maintained across time and risk situation. However, individual behaviour was independent of body size and growth in conditions of high food availability, showing no evidence to support the state-dependent personality hypothesis. The importance of a combination of spatial and temporal environmental variation for generating growth differences is highlighted.

Effects of osmolality on sperm morphology, motility and flagellar wave parameters in Northern pike (Esox lucius L.).

Northern pike (Esox lucius L.) spermatozoa are uniflagellated cells differentiated into a head without acrosome, a midpiece and a flagellar tail region flanked by a fin structure. Total, flagellar, head and midpiece lengths of spermatozoa were measured and show mean values of 34.5, 32.0, 1.32, 1.17 microm, respectively, with anterior and posterior widths of the midpiece measuring 0.8 and 0.6 microm, respectively. The osmolality of seminal plasma ranged from 228 to 350 mOsmol kg(-1) (average: 283.88+/-33.05). After triggering of sperm motility in very low osmolality medium (distilled water), blebs appeared along the flagellum. At later periods in the motility phase, the tip of the flagellum became curled into a loop shape which resulted in a shortening of the flagellum and a restriction of wave development to the proximal part (close to head). Spermatozoa velocity and percentage of motile spermatozoa decreased rapidly as a function of time postactivation and depended on the osmolality of activation media (P<0.05). In general, the greatest percentage of motile spermatozoa and highest spermatozoa velocity were observed between 125 and 235 mOsmol kg(-1). Osmolality above 375 mOsmol kg(-1) inhibited the motility of spermatozoa. After triggering of sperm motility in activation media, beating waves propagated along the full length of flagella, while waves appeared dampened during later periods in the motility phase, and were absent at the end of the motility phase. By increasing osmolality, the velocity of spermatozoa reached the highest value while wave length, amplitude, number of waves and curvatures also were at their highest values. This study showed that sperm morphology can be used for fish classification. Sperm morphology, in particular, the flagellar part showed several changes during activation in distilled water. Sperm motility of pike is inhibited due to high osmolality in the seminal plasma. Osmolality of activation medium affects the percentage of motile sperm and spermatozoa velocity due to changes in flagellar wave parameters.

Mixed mesodermal and mesenchymal origin of myotomal muscles in pike (Esox lucius: Teleostei).

During the myotomal myogenesis in pike (Esox lucius) two phases of muscle differentiation can be distinguished. In the first phase, the somite cells-derived stock, the primary myoblasts (of mesodermal origin), fuse to form multinucleate myotubes. Participation of myotomal cells of mesodermal origin is insufficient for further muscle development. In the second stage mesenchymal cells migrate, via myosepts, into the myotome between myotubes. Immunocytochemical detection of proliferating cell nuclear antigen (marker of S phase of cell cycle) showed their mitotic activity. Transmission electron microscope analysis revealed that the differentiation of these cells depends on their position. Cells remaining in the myosepts develop into fibroblasts and produce collagen fibres, while those that have migrated into the myotomes transform into secondary myoblasts. Mesenchymal cells in the studied species are believed to participate in hypertrophy and hyperplasy of muscle fibres. Thus the muscle fibres in pike (E. lucius) are of mesodermal-mesenchymal origin.