Effects of egg composition and prey density on the larval growth and survival of lake whitefish (Coregonus clupeaformis Mitchill)

Eggs were collected from two stocks of lake whitefish, Coregonus clupeaformis , in Lakes Michigan and Huron to assess the effect of egg composition and prey density on larval growth and survival. Egg composition parameters including wet weight (mg egg⁻¹), dry weight (mg egg⁻¹), percent water, total caloric content (cal egg⁻¹), caloric density (cal egg⁻¹), percent lipid content, and total lipidcontent (mg egg⁻¹) were measured. Fish hatched from six parental females in each stock were fed one of four rations (0, 18, 24, 50 brine shrimp larva⁻¹ day⁻¹) after yolk sac absorption. Length at hatch, endogenous growth, exogenous growth, and survival were measured during a 42-day laboratory experiment. Length at hatch of larvae was positively related to egg caloric content ( r ²=0.780). Endogenous growth for lake whitefish larvae was positively related to percent lipid content ( r ²=0.896) and total egg lipid content ( r ²=0.876) of parental females. Exogenous growth and survival of larval lake whitefish was positively related to prey availability. Larval fish growth was accurately modelled ( r ²=0.973) as a function of prey abundance using a threshold-corrected hyperbolic equation. These results indicate that both egg composition and prey availability have the potential to influence the growth and survival dynamics of larval lake whitefish significantly.     

Reactions in individual fish to strobe light. Field and aquarium experiments performed on whitefish (Coregonus lavaretus)

This study describes how individual whitefish Coregonus lavaretusreact to strobe light. Field experiments were performed in a net enclosure on fish tagged with ultrasonic transmitters. A strobe light array was switched on near the tagged fish. The fish moved away from the light and increased their swimming speed. Aquarium experiments under controlled conditions were carried out in rearing tanks at Saimaa Fisheries and Aquaculture Station in Finland. A strobe light was directed from the side of the basin just ahead of, directly at, and behind the fish at a close range. In the first two cases fish responded by a distinct turn and a change in swimming direction away from the light. The fish did not change its swimming direction when light was aimed from behind. It is concluded that strobe light may be used to prevent fish from swimming into a specific area. Implications for development of new fishing equipment and research concerning fishes in areas with water power stations is briefly discussed.     

Human‐induced changes in the reproductive traits of Lake Constance common whitefish (Coregonus lavaretus)

Size‐selective fishery harvest leads to phenotypic changes in fish reproductive traits. When these changes represent an evolutionary response of a stock, they may have severe consequences for future stock dynamics and yields. In freshwater ecosystems, reproductive traits may also be affected by other human impacts such as changes in system productivity. The present study uses regression analysis to evaluate the impacts of changes in lake trophy and of an intensive size‐selective harvest over several decades on the reproductive traits of common whitefish in Lake Constance between 1963 and 1999. Fecundity was strongly linked to lake trophy but was also related to the calendar year, suggesting an evolutionary response to size‐selective harvest and to massive stocking of the lake with hatchery‐reared larvae. The present study is an example of how fish reproductive traits are influenced by the combined action of various human impacts: changes in system productivity, size‐selective harvest and massive stocking.     

How normal is abnormal? Discrimination between deformations and natural variation in gonad morphology of European whitefish Coregonus lavaretus

The gonad morphology of whitefish Coregonus lavaretus collected in Lake Thun, Switzerland, and two neighbouring lakes was assessed in order to differentiate between 'normal' and 'abnormal' character states of gonad morphology, which had been previously described in C. lavaretus from Lake Thun (constrictions, asymmetries, aplasia, compartmentations, fusions and hermaphroditism). In total, 4668 fish were collected and analysed using two complementary sampling schemes: (1) monthly samples of catches by the commercial fishermen and (2) samples of ripe spawners of all known 33 spawning sites of the three lakes. Considerable variation in gonad morphology in C. lavaretus populations of all lakes was found. Notably, all deviation types were observed in fish of all three lakes. Asymmetries and constrictions were frequent in all three lakes and showed systematic differences in frequency between the two sampling strategies. This indicates that asymmetries and constrictions represent to a large extent natural variation in gonad morphology of C. lavaretus and are also prone to considerable measurement error. In contrast, aplasia, fusions, compartmentations and hermaphroditism occurred predominantly in one C. lavaretus form of Lake Thun and in particular in populations spawning at great depths. This suggests that these deviation types are probably reliable indicators for gonad deformations and supports the interpretation that Lake Thun harbours a unique case of deformed gonads in C. lavaretus of yet unknown origin.     

Different growth trends of whitefish (Coregonus lavaretus) forms in the northern Baltic Sea

The sea growth of two whitefish forms, anadromous (Coregonus lavaretus lavaretus) and sea‐spawning (Coregonus lavaretus widegreni), was analysed using samples collected from the commercial sea catch in the Gulf of Bothnia (GoB) in the northern Baltic Sea during 1998–2014. In the GoB area, these two forms are possible to identify because the gill‐raker number and size at maturity vary between forms. The growth rate of the forms is linked to their feeding area. Sea‐spawning whitefish, which has a feeding migration near its home site, was shorter in the northern GoB (66°N–64°N) at the ages of 3–11 than those in the southern GoB (64°N–60°30′N). In the data, most whitefish were caught with gill nets in the GoB. The mesh sizes of gill nets capturing the anadromous form were mostly 35–45 mm, while those capturing the sea‐spawning form were <35 mm in the northern GoB. It is likely that the different growth trends for small and large whitefish were connected with differences in their recruitment for fishing. The length of anadromous females at the age of four sea years increased significantly, but the length of six‐year‐old anadromous female whitefish decreased over the catch years from 1998–2014. In contrast, the length of slow‐growing sea‐spawning whitefish of six years or older increased significantly in relation to the catch year in the gill‐net catch. The increase in the growth of young age groups in both forms was probably associated with the increasing temperature and the low fishing pressure on small fish. The decreasing age at capture for both forms and the depression of the mean size of old anadromous whitefish are signs of high fishing pressure with a high gill‐net effort that selectively removes the largest and oldest individuals of both forms.     

Genetic-biochemical analysis of hybrids between Arctic cisco Coregonus autumnalis migratorius and lake-river whitefish C. lavaretus pidschian from Lake Baikal

Among the natural hybrids of coregonid fishes in Lake Baikal, the hybrids of the first generation from crossing Arctic cisco Coregonus autumnalis migratorius and lake-river whitefish (pidschian) C. lavaretus pidschian are the most abundant. They are well-identified morphologically. The data of the analysis of allozyme variations confirm the validity of the morphological determination of different Baikal coregonid forms and their hybrids, but do not allow a strict evaluation of its accuracy.     

Habitat use and growth of age-0 whitefish,Coregonus lavaretus,and cisco,C. albula

Synopsis Local American eel stocks were studied by mark-recapture methods along 600m of tidal creeks in Great Sippewisset Marsh, Falmouth, Massachusetts, during summer 1979. The estimated stock density was 350 eels, equivalent to 875 fish ha^-1, and movement of individual American eels over the five week study was usually less than 100 m. Large American eels were found to predominate in the wide marsh creeks whereas smaller American eels predominated in narrower creeks at the landward side of the marsh. Territoriality is suggested as a mechanism for maintaining differences in distribution of size classes and a limited home range.Senior author     

Parallel and non‐parallel morphological divergence among foraging specialists in European whitefish (Coregonus lavaretus)

Parallel phenotypic evolution occurs when independent populations evolve similar traits in response to similar selective regimes. However, populations inhabiting similar environments also frequently show some phenotypic differences that result from non‐parallel evolution. In this study, we quantified the relative importance of parallel evolution to similar foraging regimes and non‐parallel lake‐specific effects on morphological variation in European whitefish (Coregonus lavaretus). We found evidence for both lake‐specific morphological characteristics and parallel morphological divergence between whitefish specializing in feeding on profundal and littoral resources in three separate lakes. Foraging specialists expressed similar phenotypes in different lakes in both overall body shape and selected measured morphological traits. The morphology of the two whitefish specialists resembled that predicted from other fish species, supporting the conclusion of an adaptive significance of the observed morphological characteristics. Our results indicate that divergent natural selection resulting from foraging specialization is driving and/or maintaining the observed parallel morphological divergence. Whitefish in this study may represent an early stage of divergence towards the evolution of specialized morphs.     

Pleistocene glaciations and contemporary genetic diversity in a Beringian fish,the broad whitefish,Coregonus nasus (Pallas): inferences from microsatellite DNA variation

The contemporary distribution of genetic variation within and among high latitude populations cannot be fully understood without taking into consideration how species responded to the impacts of Pleistocene glaciations. Broad whitefish, Coregonus nasus, a species endemic to northwest North America and the Arctic coast of Russia, was undoubtedly impacted by such events because its geographic distribution suggests that it survived solely within the Beringian refuge from where it dispersed post‐glacially to achieve its current range. We used microsatellite DNA to investigate the role of glaciations in promoting intraspecific genetic variation in broad whitefish (N = 14 localities, 664 fish) throughout their North American range and in one Russian sample. Broad whitefish exhibited relatively high intrapopulation variation (average of 11.7 alleles per locus, average H_E = 0.61) and moderate levels of interpopulation divergence (overall F_ST = 0.10). The main regions assayed in our study (Russia, Alaska, Mackenzie River and Travaillant Lake systems) were genetically differentiated from each other and there were declines in genetic diversity with distance from putative refugia. Additionally, Mackenzie River system populations showed less developed and more variable patterns of isolation‐by‐distance than populations occupying former Alaskan portions of Beringia. Finally, our data suggest that broad whitefish dispersed from Beringia using coastal environments and opportunistically via headwater stream connections that once existed between Yukon and Mackenzie River drainages. Our results illustrate the importance of history (e.g. glaciation) and contemporary dispersal ecology in shaping the current genetic population structure of Arctic faunas.     

A proteomics approach reveals divergent molecular responses to salinity in populations of European whitefish (Coregonus lavaretus)

Osmoregulation is a vital physiological function for fish, as it helps maintain a stable intracellular concentration of ions in environments of variable salinities. We focused on a primarily freshwater species, the European whitefish (Coregonus lavaretus), to investigate the molecular mechanisms underlying salinity tolerance and examine whether these mechanisms differ between genetically similar populations that spawn in freshwater vs. brackishwater environments. A common garden experiment involving 27 families in two populations and five salinity treatments together with a large-scale, high-resolution mass spectrometry experiment that quantified 1500 proteins was conducted to assess phenotypic and proteomic responses during early development, from fertilization until hatching, in the studied populations. The populations displayed drastically different phenotypic and proteomic responses to salinity. Freshwater-spawning whitefish showed a significantly higher mortality rate in higher salinity treatments. Calcium, an ion involved in osmotic stress sensing, had a central role in the observed proteomic responses. Brackishwater-spawning fish were capable of viable osmoregulation, which was modulated by cortisol, an important seawater-adaptation hormone in teleost fish. Several proteins were identified to play key roles in osmoregulation, most importantly a highly conserved cytokine, tumour necrosis factor, whereas calcium receptor activities were associated with salinity adaptation. These results imply that individuals from these populations are most likely adapted to their local environments, even though the baseline level of genetic divergence between them is low (F(ST)=0.049). They also provide clues for choosing candidate loci for studying the molecular basis of salinity adaptation in other species. Further, our approach provides an example of how proteomic methods can be successfully used to obtain novel insights into the molecular mechanisms behind adaptation in non-model organism.     

Comparative study of bile-acid composition in the bile of the European vendace Coregonus albula L. and whitefish Coregonus lavaretus L. under conditions of technogenic water pollution

Changes of the bile acid composition were studied in bile of the vendace Coregonus albula L. and the whitefish Coregonus lavaretus L. under effect of sewers of an ore mining and processing plant, the main pollutin agents of the sewers being heavy metals. The revealed variability of the bile acid composition indicates different species-related sensitivity to the studied pollutants as well as a lower resistance of males as compared with females to this type of pollution. Mechanisms of effects of industrial sewers on the bile formation processes and a possibility of using parameters of the steroid lipid metabolism at evaluation of the state of water ecosystems are discussed.     

Isolation,characterization and cross‐salmonid amplification of 31 microsatellite loci in the lake whitefish (Coregonus clupeaformis,Mitchill)

Coregonine fish represent the most successful evolutionary lineage of salmonids with Coregonus as the most speciose salmonid genus inhabiting numerous postglacial lakes across the northern hemisphere. We isolated and characterized 31 polymorphic microsatellite loci in Coregonus clupeaformis with an average number of 5.3 alleles per locus (range three to eight) and an overall expected heterozygosity of 0.74 ± 0.11. Two loci revealed significant linkage associations through analyses of mapping families. Six additional salmonid taxa assessed for cross‐species amplification revealed between 18 and 26 positive amplifications and between two and 12 polymorphic loci per species.     

Comparative study of bile acid composition of bile of the European vendace <Emphasis Type="Italic">Coregonus albula</Emphasis> L. and the European whitefish <Emphasis Type="Italic">Coregonus lavaretus</Emphasis> L. under conditions of technogenic water reservoir pollution

Changes of the bile acid composition were studied in bile of the vendace Coregonus albula L. and the sea whitefish Coregonus lavaretus L. under effect of sewers of an ore mining and processing plant, the main polluting agents of the sewers being heavy metals. The revealed variability of the bile acid composition indicates different species-related sensitivity to the studied pollutants as well as a lower resistance of males as compared with females to this type of pollution. Mechanisms of effects of industrial sewers on the bile formation processes and a possibility of using parameters of the steroid lipid metabolism at evaluation of the state of aquatic ecosystems are discussed.     

Modifying effects of a cobble substrate on thermal environments and implications for embryonic development in lake whitefish (Coregonus clupeaformis)

A laboratory flume was constructed to examine substrate effects on aquatic development. The flume was designed as a once-through system with a submerged cobble-filled corebox. Lake whitefish (Coregonus clupeaformis) embryos and temperature probes were deployed at multiple sites within the cobble and in the open water channel. Embryos were incubated in the flume for two different experimental periods: one to examine substrate impacts during natural lake cooling (37 days: 5 December 2016 to 10 January 2017) and the second to investigate substrate effects while administering a twice weekly 1 h heat shock (51 days: 11 January to 2 March 2017). During incubation, no significant difference was found in the average temperature between locations; however, temperatures were more stable within the cobble. Following both incubation periods, embryos retrieved from the cobble were significantly smaller in both dry mass and body length by up to 20%. These results demonstrate differences between embryos submerged in a cobble substrate and in the open water column, highlighting the need to consider the physical influences from the incubation environment when assessing development effects as part of any scientific study or environmental assessment.     

Predator-induced respiratory responses in juveniles of vendace Coregonus albula, whitefish C. lavaretus, perch Perca fluviatilis and roach Rutilus rutilus

Predator-induced respiratory responses of juvenile vendace, whitefish, perch and roach were studied in an intermittent-flow respirometer by creating visual contact between test fishes and predators (juvenile northern pike). Vendace and whitefish always responded to the presence of pike with increased oxygen consumption. The response of perch was the opposite: oxygen consumption decreased to near the standard respiratory level. Roach responded more variably: some individuals increased their oxygen uptake while others decreased it. Changes in the rate of respiration during exposure to predation are supposed to be caused by the differences in locomotory activity due to induced antipredator behaviour. According to their responses, these species could be grouped into escapers (coregonids), hiders (perch) or indeterminates (roach). The species-specific differences in the antipredator behaviour originate in morphological and physiological characteristics which together determine the effectiveness of a particular antipredator style (i.e. either hiding or escaping).     

Effects of freshwater to seawater transfer on osmoregulation,acid-base balance and respiration in river migrating whitefish (Coregonus lavaretus)

Osmoregulation, acid-base balance and respiratory parameters were investigated in whitefish following transfer from freshwater to salt water. Whitefish acclimated successfully to 25 ppt brackish water but died after direct transfer to 32 ppt sea water. Transfer to brackish water induced rapid (<6 h) and permanent increases in plasma [Na⁺], [Cl^–], total [Ca] and [Mg]. The extracellular hyperosmolality effected a transient (<3 days) muscle tissue dehydration and red blood cell shrinkage. Exposure to brackish water decreased both the arterial O₂ tension and whole body O₂ uptake. The extracellular acid-base status changed from an initial respiratory acidosis at 1 h towards a pronounced metabolic acidosis at 48 h of brackish water exposure. Red cell pH_i decreased in parallel with extracellular pH_e, but the in vivo pH_i/pH_e was only 0.26, suggesting some selective protection of red cell pH_i. Plasma cortisol concentration and gill Na⁺, K⁺-ATPase activity increased after exposure to high ambient salinity, reflecting the induction of hypo-osmoregulatory mechanisms. The physiological changes in whitefish are discussed in relation to salinity-induced effects in other salmonid fishes.Abbreviations CO₂ solubility in plasma - water O₂ capacitance coefficient - BW brackish water - C _T total CO₂ content in plasma - FW fresh water - Hb hemoglobin - Hct hematocrit - M _b body mass of fish - MCHC mean cellular hemoglobin concentration - PCO₂ carbon dioxide tension - pH _e extracellular pH - pH _i intracellular pH - PO_{2 in} oxygen tension in water flowing in - PO_{2 out} oxygen tension in water flowing out - ppt parts per thousand - RBC red blood cell(s) - SW sea water - V _m water flows through chamber - OV ₂ ml O₂ consumed per kg per hour     

Development of the embryonic heat shock response and the impact of repeated thermal stress in early stage lake whitefish (Coregonus clupeaformis) embryos

Lake whitefish (Coregonus clupeaformis) embryos were exposed to thermal stress (TS) at different developmental stages to determine when the heat shock response (HSR) can be initiated and if it is altered by exposure to repeated TS. First, embryos were subject to one of three different TS temperatures (6, 9, or 12 °C above control) at 4 points in development (21, 38, 60 and 70 days post-fertilisation (dpf)) for 2 h followed by a 2 h recovery to understand the ontogeny of the HSR. A second experiment explored the effects of repeated TS on the HSR in embryos from 15 to 75 dpf. Embryos were subjected to one of two TS regimes; +6 °C TS for 1 h every 6 days or +9 °C TS for 1 h every 6 days. Following a 2 h recovery, a subset of embryos was sampled. Our results show that embryos could initiate a HSR via upregulation of heat shock protein 70 (hsp70) mRNA at all developmental ages studied, but that this response varied with age and was only observed with a TS of +9 or +12 °C. In comparison, when embryos received multiple TS treatments, hsp70 was not induced in response to the 1 h TS and 2 h recovery, and a downregulation was observed at 39 dpf. Downregulation of hsp47 and hsp90α mRNA was also observed in early age embryos. Collectively, these data suggest that embryos are capable of initiating a HSR at early age and throughout embryogenesis, but that repeated TS can alter the HSR, and may result in either reduced responsiveness or a downregulation of inducible hsps. Our findings warrant further investigation into both the short- and long-term effects of repeated TS on lake whitefish development.     

Morphological divergence and origin of sympatric populations of European whitefish (Coregonus lavaretus L.) in Lake Femund, Norway

Combining morphological and genetic analysis, we compared patterns of diversification within and between morphs among sympatric European whitefish (Coregonus lavaretus L.) populations in Lake Femund, Norway. Seven external populations, from potential colonization routes into Lake Femund were included. We found that deep-, shallow-, river- and bay spawning populations are distinct morphs in Lake Femund. Within morphs, populations range from being similar genetically (Fst=0-0.005) among deep-spawning populations to being highly differentiated (Fst=0.153) between bay-spawning populations. Between morphs, genetic differences ranged from a low (Fst=0.008-0.022) between deep- and shallow-spawning populations to high difference (Fst=0.125-0.143) between shallow- and bay-spawning populations. A higher proportion of molecular variance was seen among (3.9%) than within morphs (2.8%). The adaptive gene combinations behind the four morphs seem to have originated within the lake, although the lake could have been colonized from more than one source population.