Hybridization of crucian carp, Carassius carassius (Linnaeus, 1758), in Ukrainian reservoirs and genetic structure of hybrids

Hybridization of crucian carps Carassius carassius in polyspecific crucian populations of reservoirs of Ukraine and genetic structure of the hybrids were investigated using biochemical gene marking and cytometric procedure. The fact of wide hybridization between C. auratus and C. carassius was proved to be true by large number of hybrids which can form populations consisting only from hybrid individuals. Hybrids C. auratus x C. carassius were diploid, tryploid and in exceptional cases tetraploid; females and males which most likely breed by hybridogenesis. Besides, some clonal hybrids C. carassius x C. gibelio-1 appearing as tetraploid females, and one triploid female C. carassius x Tinca tinca were revealed. It is supported that hybridization of alien C. auratus with endemic C. carassius became one of mechanisms of replacement and depressions of populations of the last.     

An experimental study of body size and food size relations in crucian carp,Carassius carassius

Synopsis In dense, single-species assemblages of crucian carp, competition is intense and results in populations of stunted fish. To explore mechanisms underlying this competition, we measured handling times, return rates, and prey choice for five sizes of crucian carp feeding on six sizes of a standardized food. Handling times increased with prey size and decreased with fish size. Return rates (dry mass ingested per unit handling time) increased dramatically with fish size, and generally decreased with increasing prey size, especially for small fish. Patterns of return rates among size-classes suggested that one or more size-related shifts in feeding efficiency exist for crucian carp; combined with physiological stresses related to winter anoxia, the inability of fish to make these shifts may contribute to size structures observed in high-density populations. Comparisons of relations among fish size, prey size, and return rates for crucian carp and bluegill, Lepomis macrochirus, suggest that similar intraspecific competitive relations exist between generalist species with size-structured populations. Despite differences in return rates among prey sizes, the extent of food selectivity based on prey size exhibited by crucian carp in two types of choice trials was less than predicted. Crucian carp commonly take in several items before mechanically processing food with their pharyngeal apparatus; this multiple prey processing may contribute to disparities between observed choice patterns and those predicted based on return rates for single prey.     

Predator recognition and defence strategies in crucian carp, Carassius carassius

Crucian carp from populations that lack piscivores are extremely vulnerable to predation. However, in the presence of piscivores these fish develop an inducible morphological defence, a deep body. This switch from a vulnerable, shallow-bodied morph to a morphologically defended morph makes this species very suitable for investigations of anti-predator strategies, and trade-offs between morphological and behavioural defences. To address these questions, we performed eight different experiments. We found that crucian carp exhibited fright responses to chemical cues from unfamiliar predators (northern pike, perch) when these were fed prey that contained alarm substance (for northern pike: crucian carp, roach; for perch: crucian carp). Cues from small pike that were fed prey that lacked alarm substance (swordtails) caused no significant fright response whereas cues from larger pike with the same diet did. Perch on a chironomid diet elicited weaker but significant fright responses. Starved predators caused as strong fright reactions as recently fed ones did, whereas no response was exhibited towards nonpredatory fish (roach, crucian carp). Crucian carp were able to detect the presence of pike after cues had been diluted to an equivalent of 21 000 l, and larger predators elicited stronger fright responses. Prior experience of predators decreased fright responses. In particular, individuals from populations that coexisted with northern pike responded less to chemical cues from northern pike than individuals without prior experience did. Thus, crucian carp may use both alarm-substance related and predator-related cues to identify predators. Further, they were able to discriminate between large and small predators. Finally, individuals from populations that coexist with predators exhibit less pronounced fright responses. These fish have an induced morphological defence, a deep body, which most likely decreases the need for strong antipredator behaviour.     

Seasonal feeding activity and ontogenetic dietary shifts in crucian carp,Carassius carassius

Synopsis A pronounced seasonal cycle in the activity of the intestinal alkaline phosphatase indicated a 6 month pause in feeding by crucian carp in an anoxic pond during winter. Zero enzyme activity was not reached until ice cover of the pond was complete, although even in the preceding two months no food was found in the guts of the fish. The summer diets of four size classes of fish consisted of varying proportions of planktonic and benthic arthropods, including Odonata which were consumed exclusively by the largest size class of crucian carp. Chironomid larvae were consumed even by the smallest fish investigated, although planktonic Cladocera were preferred by this size class. Benthic Cladocera were almost totally absent in the diet of fish> 10 cm. Habitat segregation between size classes developed synchronously with the shift to benthic food. The stunting of crucian carp populations in small ponds is interpreted as resulting from limited resources.     

Consumption of crucian carp (Carassius carassius L., 1758) by restocked pike (Esox lucius L., 1758) in a lake with frequent winter hypoxia

To assess the effects of stocked pike (Esox lucius L.) on crucian carp (Carassius carassius L.) biomasses, the annual consumption of pike was estimated and compared with removal fishery catches. The studied lake, Lake Savijärvi, is a small (40 ha), shallow and eutrophicated lake in southern Finland with frequent algal blooms during summers and fish kills during winters. Until the 1980s, the fish fauna consisted of pike, perch (Perca fluviatilis L.), roach [Rutilus rutilus (L.)], crucian carp, and tench (Tinca tinca L.). Since 2003, when crucian carp were abundant in the extreme and with only a few roach and tench individuals, the lake has been biomanipulated by removal seining. To enhance the effects of seining, piscivorous pike were restocked in the spring of 2008. During the 3‐year study, the catch of crucian carp decreased from 243 to 136 kg while the catch of pike increased from 0.1 to 5.7 kg per seining hectare due to their spawning in the lake as of 2009. The total crucian carp consumption by captured pike was 588 kg during the 2008–2010 study period. When applying the number of pike estimated with the mark‐recapture method, consumption estimate increased to 917 kg, or 22.9 kg per lake hectare in 2010. Thus, pike consumption of crucian carp was about 17% of the removal catch in 2010.     

Development and application of environmental DNA surveillance for the threatened crucian carp (Carassius carassius)

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New cell types of the pancreatic islets in the crucian carp,Carassius carassius

Summary The pancreatic islets ofCarassius carassius have been studied by electron microscopy. 1. Besides A-, B- and D-cells, two new cell types, the fourth and the fifth, have been identified. The fourth cell type is numerous; it occurs interposed among the other types of islet cells or in small clusters. The secretory granules (90–280 mg in diameter) are round or oval and usually with much lower electron density than α- and δ-granules. The secretory granules of the fifth type of cell (approximately 140–240 mμ in diameter) contain finely granular material and an electron dense core that is round or often tetra- or hexagonal. 2. The islet cells with clear cytoplasmic matrix generally contain large numbers of fine, agranular and cored vesicles 400–680 ? in diameter. They appear, in bead-like chains, or randomly scattered throughout the cytoplasm, or often clustered in aggregates close to the secretory granules and show evidence of incorporation into the secretory granules. The two types of vesicles may be formed by constriction or pinching-off of the tubular smooth endoplasmic reticulum.     

A molecular approach to detect hybridisation between crucian carp (Carassius carassius) and non-indigenous carp species (Carassius spp. and Cyprinus carpio)

1. Releases of non‐native fish into the wild is an increasing problem posing considerable ecological and genetic threats through direct competition and hybridisation. 2. We employed six microsatellite markers to identify first generation hybrids and backcrosses between native crucian carp (Carassius carassius) and introduced goldfish (C. auratus) and common carp (Cyprinus carpio) in the U.K. We also investigated the genetic characteristics of the taxonomically controversial gibel carp (Carassius spp.) from sites across Europe. 3. Natural hybridisation between goldfish and crucian carp occurs frequently, although hybrids between all other species pairs were observed. Only 62% of British crucian carp populations (n = 21) consisted exclusively of pure crucian carp. In some populations hybrids were so frequent, that no pure crucian carp were caught, indicating a high competitive ability of hybrids. 4. Most hybrids belonged to the F1 generation but backcrossing was evident at a low frequency in goldfish × crucian carp hybrids and goldfish × common carp hybrids. Furthermore, some local populations had high frequencies of backcrosses, raising the opportunity for introgression. 5. Gibel carp from Germany and Italy belonged to two triploid clonal lineages that were genetically closely related to goldfish, whereas all individuals identified from British populations proved to be crucian carp × goldfish hybrids. 6. Our study suggests that the release of closely related exotic cyprinids not only poses a threat to the genetic integrity and associated local adaptations of native species, but may also contribute to shifts in community structure through competitive interactions.     

Koi herpesvirus infection in experimentally infected common carp Cyprinus carpio (Linnaeus, 1758) and three potential carrier fish species Carassius carassius (Linnaeus, 1758); Rutilus rutilus (Linnaeus, 1758); and Tinca tinca (Linnaeus, 1758) by quantitative real‐time PCR and in‐situ hybridization

Only single cells in the carrier fish species Carassius carassius (Linnaeus, 1758) for koi herpesvirus (KHV) are infected in contrast to large numbers in the susceptible species common carp Cyprinus carpio (Linnaeus 1758). Several species of the family Cyprinidae have been described as virus carrier species, showing no clinical signs of a KHV disease but able to transmit the virus to other susceptible fish. In this study, 72 common carp Cyprinus carpio (Linnaeus, 1758), 36 tench Tinca tinca (Linnaeus, 1758), 36 crucian carp Carassius carassius (Linnaeus, 1758) and 36 common roach Rutilus rutilus (Linnaeus, 1758) were experimentally infected with KHV (isolate “Israel”) by immersion and kept at 20°C. The fish were euthanized at 12 timepoints over a period of 90 days and virus DNA was quantified in tissues by a real‐time TaqMan PCR. Whereas KHV‐DNA was found in Cyprinus carpio for up to 90 days, the virus DNA was detectable only in single individuals of Rutilus rutilus, Tinca tinca and Carassius carassius for up to 25 days after experimental virus exposure. Tissue samples of Cyprinus carpio and Carassius carassius were screened by in‐situ hybridization. Positive signals were found in various organs of the common carp tested crucian carp. In the latter species a much smaller number of virus‐positive stained cells was detected compared to the infected carp.     

Anaerobic wintering of crucian carp (Carassius carassius L.)--II. Metabolic products

In long-term experimental anoxia (up to 140 days) crucian carp excreted ethanol and acetic acid. The maximum concentration of ethanol found in fish blood was ca 0.1%. The excretion rate of ethanol at temperatures from -0.5 to +5 degrees C was ca 25 micrograms/g fish wet wt/hr, but increased rapidly with increasing temperature. At 2 and 5 degrees C the ethanol excretion rate was independent of fish size, but at 12 and 18 degrees C a higher rate was observed in smaller fish. The decrease of water pH below 5 in the experimental vial during long anoxia was assumed to be due to excretion of acid compounds by the fish.     

Intrinsic contractile properties of the crucian carp (Carassius carassius) heart during anoxic and acidotic stress

The crucian carp (Carassius carassius) seems unique among vertebrates in its ability to maintain cardiac performance during prolonged anoxia. We investigated whether this phenomenon arises in part from a myocardium tolerant to severe acidosis or because the anoxic crucian carp heart may not experience a severe extracellular acidosis due to the fish's ability to convert lactate to ethanol. Spontaneously contracting heart preparations from cold-acclimated (6-8°C) carp were exposed (at 6.5°C) to graded or ungraded levels of acidosis under normoxic or anoxic conditions and intrinsic contractile performance was assessed. Our results clearly show that the carp heart is tolerant of acidosis as long as oxygen is available. However, heart rate and contraction kinetics of anoxic hearts were severely impaired when extracellular pH was decreased below 7.4. Nevertheless, the crucian carp heart was capable of recovering intrinsic contractile performance upon reoxygenation regardless of the severity of the anoxic + acidotic insult. Finally, we show that increased adrenergic stimulation can ameliorate, to a degree, the negative effects of severe acidosis on the intrinsic contractile properties of the anoxic crucian carp heart. Combined, these findings indicate an avoidance of severe extracellular acidosis and adrenergic stimulation are two important factors protecting the intrinsic contractile properties of the crucian carp heart during prolonged anoxia, and thus likely facilitate the ability of the anoxic crucian carp to maintain cardiac pumping.     

Molecular identification of hybrids of the invasive gibel carp Carassius auratus gibelio and crucian carp Carassius carassius in Swedish waters

Both mitochondrial DNA sequence and two nuclear microsatellite markers were used to confirm the identity of the first record of Carassius auratus gibelio in the western (Swedish) Baltic Sea region. A total of 49 fishes were analysed, where 22 were from three Swedish sites connected to the Baltic Sea. The D-loop mitochondrial DNA sequences showed that 16 of 22 Swedish fishes were related to C. a. gibelio. The phylogenetic analysis of these sequences showed that these fish are probably not native, but represent different lineages of C. a. gibelio from China, Japan and Russia. All except three of these 16 fishes had microsatellite alleles suggesting hybridization with Carassius carassius. These findings suggest that a cryptic invasion of C. a. gibelio might be in progress.     

Distribution of aldehyde dehydrogenase and alcohol dehydrogenase in summer-acclimatized crucian carp, Carassius carassius L.

The tissue distribution of aldehyde dehydrogenase (ALDH) and alcohol dehydrogenase (ADH) in summer-acclimatized crucian carp showed almost the same exceptional pattern as previously found in winter-acclimatized specimens. There was a nearly complete spatial separation of ALDH and ADH; in other vertebrates these enzymes occur together. This exceptional enzyme distribution is probably an adaptation to the extraordinary ability of Carassius to produce ethanol as the major metabolic end product during anoxia. Since the crucian carp is less likely to encounter anoxia during the summer, the present results suggest that the crucian carp is unable to switch over to a 'normal' ALDH and ADH distribution in the summer. However, it is also possible that there is an advantage for the summer-acclimatized crucian carp in keeping ALDH and ADH separate, because of occasional anoxic periods.     

Genetic polymorphism and isoenzyme patterns of lactate dehydrogenase in tench (Tinea tinea), crucian carp (Carassius carassius) and carp (Cyprinus carpio)

Isoenzyme patterns and the polymorphism of lactate dehydrogenase (LDH) were investigated in 3 fish species of family Cyprinidae, i.e. tench ( Tinea tinea ), crucian carp ( Carassius carassius ) and carp ( Cyprinus carpio). The isoenzyme patterns were tissue and species specific. In crucian carp subunits with different electro-phoretic mobility are present, which are genetically controlled from the B¹, B², A¹, A²and C loci, while the set of loci in carp is B¹, B², A, C¹and C²and in tench B, A, C. The locus B of LDH in tench, the locus B²in crucian carp, and the loci B¹, C¹and C²in carp are polymorphic and have two different alleles in each case. The polymorphism did not affect the total LDH activity in the tissues. All the populations investigated were in Hardy-Weinberg equilibrium. The genetic control of the polymorphism in B¹and C¹loci in carp was proved by test matings. The polymorphism in B loci tested in erythrocytes may be utilized as genetic markers in the fish breeding.     

Genetic structure of crucian carp (Cypriniformes, Cyprinidae, Carassius L. 1758) populations of middle-Dnieper basin

Comparative analysis of Middle Dnieper crucian carp's population structure was made by means of biochemical gene marking, cytometry and biological analysis. As a result the structure was found, which may by expounded like association of the genetically related species and forms. This association consists of diploid C. auratus (which predominates in population), C. carassius (which was found in the some of north-eastern reservoirs only) and triploid C. gibelio. In Ukraine the homogeneous populations of C. gibelio can be found on the north-east of Eastern Ukraine only. Though some of individuals maybe present with diploid C. auratus. Besides were found hybrid form: diploid C. auratus-C. carassius and presumably tetraploid C. auratus-C. gibelio and C. gibelio-C. carassius. However in the tetraploid's spectra were absent a number of specific alleles, that marking C. species-one of the parental species of triploid C. gibelio. Therefore presumptive tetraploids have not difference from C. auratus and diploid C. auratus-C. carassius on the gene markers level. Nevertheless they are neatly diagnose by the erythrocytes which have good difference and are bigger then diploid species erythrocytes in 40% (C. auratus-C. gibelio) and 100% (C. gibelio-C. carassius).     

1H-NMR study of the metabolome of an exceptionally anoxia tolerant vertebrate, the crucian carp (Carassius carassius)

The crucian carp (Carassius carassius) can tolerate anoxia for days to months, depending on the temperature. In this study, we applied ¹H-NMR-based metabolomics to polar extracts of crucian carp brain, heart, muscle and liver samples obtained from fish exposed to either control normoxic conditions, acute anoxia (24 h), chronic anoxia (1 week) or reoxygenation (for 1 week following chronic anoxia) at 5 °C. Spectra of the examined tissues revealed changes in several energy-related compounds. In particular, anoxic stress resulted in decreased concentrations of phosphocreatine (muscle, liver) and glycogen (liver) and ATP/ADP (liver, heart and muscle) and increased concentrations of lactate (brain, heart, muscle) and beta-hydroxybutyric acid (all tissues). Likewise, increased concentrations of inhibitory compounds (glycine, gamma-amino butyric acid or GABA) and decreased concentrations of excitatory metabolites (glutamate, glutamine) were confirmed in the anoxic brain extracts. Additionally, a decrease of N-acetylaspartate (NAA), an important neuronal marker, was also observed in anoxic brains. The branched-chain amino acids (BCAA) valine/isoleucine/leucine increased in all anoxic tissues. Possibly, this general tissue increase can be due to an inhibited mitochondrial function or due to protein degradation/protein synthesis inhibition. In this study, the potential and strength of the ¹H-NMR is highlighted by the detection of previously unrecognized changes in metabolites. Specifically, myo-inositol substantially decreased in the heart of anoxic crucian carp and anoxic muscle tissue displayed a decreased concentration of taurine, providing novel insights into the anoxia responses of the crucian carp.     

Genetic polymorphism and isoenzyme patterns of lactate dehydrogenase in tench (Tinca tinca), crucian carp (Carassius carassius) and carp (Cyprinus carpio)

Isoenzyme patterns and the polymorphism of lactate dehydrogenase (LDH) were investigated in 3 fish species of family Cyprinidae, i.e. tench (Tinca tinca), crucian carp (Carassius carassius) and carp (Cyprinus carpio). The isoenzyme patterns were tissue and species specific. In crucian carp subunits with different electrophoretic mobility are present, which are genetically controlled from the B1, B2, A1, A2 and C loci, while the set of loci in carp is B1, B2, A, C1 and C2 and in tench B, A, C. The locus B of LDH in tench, the locus B2 in crucian carp, and the loci B1, C1 and C2 in carp are polymorphic and have two different alleles in each case. The polymorphism did not affect the total LDH activity in the tissues. All the populations investigated were in Hardy-Weinberg equilibrium. The genetic control of the polymorphism in B1 and C1 loci in carp was proved by test matings. The polymorphism in B loci tested in erythrocytes may be utilized as genetic markers in the fish breeding.     

Differential regulation of AMP-activated kinase and AKT kinase in response to oxygen availability in crucian carp (Carassius carassius)

We investigated whether two kinases critical for survival during periods of energy deficiency in anoxia-intolerant mammalian species, AMP-activated kinase (AMPK), and protein kinase B (AKT), are equally important for hypoxic/anoxic survival in the extremely anoxia-tolerant crucian carp (Carassius carassius). We report that phosphorylation of AMPK and AKT in heart and brain showed small changes after 10 days of severe hypoxia (0.3 mg O2/l at 9 degrees C). In contrast, anoxia exposure (0.01 mg O2/l at 8 degrees C) substantially increased AMPK phosphorylation but decreased AKT phosphorylation in carp heart and brain, indicating activation of AMPK and deactivation of AKT. In agreement, blocking the activity of AMPK in anoxic fish in vivo with 20 mg/kg Compound C resulted in an elevated metabolic rate (as indicated by increased ethanol production) and tended to reduce energy charge. This is the first in vivo experiment with Compound C in a nonmammalian vertebrate, and it appears that AMPK plays a role in mediating anoxic metabolic depression in crucian carp. Real-time RT-PCR analysis of the investigated AMPK subunit revealed that the most likely composition of subunits in the carp heart is alpha2, beta1B, gamma2a, whereas a more even expression of subunits was found in the brain. In the heart, expression of the regulatory gamma2-subunit increased in the heart during anoxia. In the brain, expression of the alpha1-, alpha2-, and gamma1-subunits decreased with anoxia exposure, but expression of the gamma2-subunit remained constant. Combined, our findings suggest that AMPK and AKT may play important, but opposing roles for hypoxic/anoxic survival in the anoxia-tolerant crucian carp.