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.     

Development of microsatellite DNA markers of grass carp (Ctenopharyngodon idella) and their cross-species application in black carp (Mylopharyngodon piceus)

Thirty-four microsatellite DNA loci were developed for grass carp (Ctenopharyngodon idella) and used to determine the diversity of 42 individuals of grass carp and 16 individuals of black carp collected from Jingzhou fragment (the middle reach) of Yangtze River. All the 34 loci were polymorphic in grass carp. The number of alleles found in grass carp at these loci ranged from 2 to 24. The observed and expected heterozygosities varied from 0.238 to 1.000 and from 0.162 to 0.945, respectively. Thirty of the 34 loci cross-amplified the DNA of black carp (Mylopharyngodon piceus), and 21 of them revealed polymorphism (more than one allele each locus).     

Differential expression of two C‐type lectins in grass carp Ctenopharyngodon idella and their response to grass carp reovirus

The cDNAs of two C‐type lectins in grass carp Ctenopharyngodon idella, galactose‐binding lectin (galbl) and mannose‐binding lectin (mbl), were cloned and analysed in this study. Both of them exhibited the highest expression level in liver, whereas their expression pattern differed in early phase of embryonic development. Following exposure to grass carp reovirus (GCRV), the mRNA expression level of galbl and mbl was significantly up‐regulated in liver and intestine.     

Suction volume and filtering efficiency of silver carp (Hypophthalmichthys molitrix Val.) and bighead carp (Aristichthys nobilis Rich.)

Experiments were conducted to measure the suction volume of silver carp and bighead carp of age 1 + with respiratory chamber, and to calculate the suction volume and the filtering efficiency with respect to changes in concentrations of food particles. Suction volume (B. ml/mouth) and filtering efficiency (E. %) were calculated using the following formula: C ₁=C₀(1-BE/v)ⁿ where C₀ and C₁ were the concentrations of specific food particles at the beginning and at the end of experiment, respectively, V was the volume (ml) of experimental water, and n was the total number of observation of suction made during the experimental period. The relationships between suction volume (ml/mouth) of age I+ silver carp (B_h) and bighead carp (B_a) and their standard lengths (L, cm) were: B _h=0.561L-8.94, B_a= 0.627L-7.48 while those of the fingerlings were: B _h= O.l70L-0.837, B_a= 0.157L-0.418. The suction volume of the fingerlings was mainly affected by fish size, the function of temperature between 15 and 25° C being negligible. However, temperature affected filtering rate (filtered volume per unit time) through its effect on filtering frequency. The filtering efficiency of the fishes for rotifers (Brachionus caliciflorus) was 100 per cent. The relationships between filtering efficiency and sizes of food particles smaller than or equal to that of a rotifer were: E _h=25.1 ln e.s.d. -13.6, E_a=22.2 In e.s.d. -33.1 where E_h and E_a were filtering efficiency of silver carp and bighead carp, respectively, and e.s.d. was the equivalent spherical diameter (μm) of food particles.     

Biomass-dependent effects of common carp on water quality in shallow ponds

We examined the biomass-dependent effects of common carp (Cyprinus carpio) on water quality in 10 ponds at the Eagle Mountain Fish Hatchery, Fort Worth, Texas, USA. Ponds contained 0–465 kg ha⁻¹ of common carp. We measured limnological variables at weekly intervals for four weeks in early summer, after which ponds were drained and the biomass of fish and macrophytes was determined. Common carp biomass was significantly positively correlated with chlorophyll a, total phosphorus, total nitrogen, and Keratella spp. density and negatively correlated to bushy pondweed (Najas guadalupensis) biomass. In addition, we combined our data with data from comparable studies to develop more robust regression models that predict the biomass-dependent effects of common carp on water quality variables across a wide range of systems.     

Differences in biochemical profiles among spawners of eight common carp breeds

Eight breeds of common carp (Cyprinus carpio L.) spawners reared under identical conditions and sampled in spring after over‐wintering were examined in order to compare their basic biochemical blood profiles. The breeds compared were: Amur wild carp (AS), Ropsha scaly carp (ROP), Ukraine scaly carp (US), Northern mirror carp (M72), South Bohemian mirror carp (BV), Israeli mirror carp (Dor 70), Hungarian mirror carp (M2) and Tata scaly carp (TAT). Significant differences were found among breeds in glucose concentration (GLU), total protein concentration (TP), triacylglycerols concentration (TAG), and calcium (Ca) and phosphorus (P_i) concentration. No differences were observed in aspartate transaminase activity (AST) or alanine aminotransferase activity (ALT). The highest glucose, total protein, and calcium (Ca) concentrations were found in AS (GLU 8.3 ± 1.2 mmol L^?1, TP 32 ± 3 g L^?1, Ca 2.42 ± 0.22 mmol L^?1). High values of triacylglycerol concentration (TAG) were found in ROP (1.94 ± 0.52 mmol L^?1). Phosphorus (P_i) concentration was highest in M2 (3.82 ± 1.34 mmol L^?1). Amur wild carp and breeds originating therefrom (ROP, US, and M72) had significantly higher values of TP (P < 0.05), TAG (P < 0.05), and Ca (P < 0.01) and significantly lower values of P_i (P < 0.05) than did the other breeds. Scaly breeds had higher values of glucose (P < 0.01), TP (P < 0.01), ALT (P < 0.01), and Ca (P < 0.01) and significantly lower values of P_i (P < 0.01) than did mirror carp. Significant (P < 0.01) sex‐related differences were found in GLU, TAG and Ca concentrations.     

Polymorphism of microsatellite markers in Russian common carp (<Emphasis Type="Italic">Cyprinus carpio</Emphasis> L.) breeds

Using five microsatellite loci, genotyping and genetic diversity estimates were obtained for nine samples representing seven common carp breeds most widespread in Russia. For comparison, the samples of Amur wild common carp (Cyprinus carpio haematopterus) and a sample of European Hungarian carp were used. In the samples examined (n = 148) a total of 78 alleles were revealed. The highest mean allele number per locus (7.3) was identified in Amur wild common carp, while the lowest number was found in Cherepets carps (4.0). In different breeds, the observed heterozygosities varied from 0.819 (Altai carp) to 0.651 (Cherepets scaly carp). Three out of five microsatellite loci (MFW-24, MFW-28, and MFW-19) revealed a high level of population differentiation. In the dendrogram of genetic differences, all breeds clustered into two groups. One of these groups was composed of the two strains of Ropsha carp, Stavropol carp, Amur wild common carp, and the two samples of Cherepets carp. The second cluster included Altai carp (Priobskii and Chumysh populations), two Angelinskii carp breeds (mirror and scaly), and Hungarian carp. The pairs of breeds/populations/strains, having common origin, were differentiated. Specifically, these were two populations of Altai carp, two strains of Ropsha carp, as well as the breeds of Angelinskii and Cherepets carps. The reasons for genetic differentiation of Russian common carp breeds, as well as the concordance of the evolutionary histories of these breeds, some of which originated from the European breeds, while the others contain substantial admixture of the Amur wild common carp, are discussed.     

不同捕食者暴露对鲫幼鱼行为、运动和形态特征的影响

自然界中鱼类的捕食者种类繁多,为验证猎物鱼针对不同捕食者是否会做出相异的反捕食策略选择,选取鲫（Carassius auratus）幼鱼为猎物鱼,乌鳢（Channa argus）和胡子鲶（Clarias fuscus）为捕食者,将鲫幼鱼分别暴露于空白（对照）、乌鳢、胡子鲶和双捕食者（同时存在乌鳢和胡子鲶）环境中持续2个月,随后考查不同组鲫幼鱼的生长、行为特征、形态和运动能力等指标之间的差异。结果表明：经过2个月的捕食胁迫处理,鲫幼鱼的自发运动、隐蔽场所利用率和勇敢性等行为指标在4个组之间均未表现出显著性差异。相比对照组,乌鳢、胡子鲶和双捕食者暴露组鲫幼鱼的快速启动反应时滞显著缩短（P<0.05）;但所有处理组中仅乌鳢暴露组鲫幼鱼的快速启动120ms移动距离（S_120ms）和体高（BD）显著高于对照组（P<0.05）。另外鲫幼鱼的临界游泳速度（U_crit）和活跃代谢率（M_O2active）在各组之间也未表现出显著差异。乌鳢暴露组鲫幼鱼快速启动能力的提升可能与形态上体高的提高有关;另外相比临界游泳能力,快速启动能力可能在鱼类逃避捕食者过程中更为关键;鲫幼鱼表现出对乌鳢的反应最为明显,可能与乌鳢口裂较大,给鲫幼鱼带来的威胁更大有关。整体而言,应对捕食胁迫时,鲫幼鱼仅表现出形态和快速启动能力的反捕食响应,其行为特征的保守性可能是对生长作出的妥协,这对于鱼类适应生境中长期存在的捕食胁迫十分关键。     

Linking silver carp habitat selection to flow and phytoplankton in the Mississippi River

Invasive silver carp (Hypothalmichthys molitrix) occurs throughout much of the Mississippi River and threatens the Laurentian Great Lakes. To quantify habitat selection relative to river flow and potential phytoplankton food, 77 adult silver carp were implanted with ultrasonic transmitters during spring 2008 through spring 2009 in adjacent upstream dammed and downstream undammed reaches (48 km total) of the Mississippi River. Sixty-seven percent of the fish were located. Selection of major river habitat features (dammed vs. undammed, backwaters, channel border, wing dikes, island side channels, and the main channel) was quantified. Flow rates and chlorophyll a concentration were compared between silver carp locations and random sites. Foregut chlorophyll a concentrations plus presence of macrozooplankton and detritus of 240 non-tagged silver carp were quantified. About 30% of silver carp moved upstream into the dammed reach, where average flow was slower and chlorophyll a concentration was higher. Silver carp selected wing dike areas of moderate flow (about 0.3 m/s) and elevated chlorophyll a (about 7 μg/L) relative to random sites. No silver carp occurred in areas where flow was absent. Wing dikes were preferred while the main channel was avoided. Chlorophyll a concentrations in guts were positively related to temperature and were unrelated to flow or river chlorophyll a concentration. Macrozooplankton and detritus were rare in guts. Silver carp seek areas of low flow and successfully forage across a range of temperatures, flows, and chlorophyll a concentrations that occur in rivers and large lakes.     

Experimental induction of Sphaerospora renicola (Myxosporea) infection in common carp (Cyprinus carpio) by transmission of SB-protozoans

The authors have experimental evidence that the protozoa causing the swimbladder inflammation (SBI) of the common carp (Cyprinus carpio) are indentical with presporogonic stages of Sphaerospora renicola Dyková et Lorn, 1982 parasitizing the renal tubules. Homogenates prepared from the thickened and inflamed swimbladder of naturally infected common carp, when injected into the abdominal cavity of fish, produced renal sphaerosporosis in the infection-free common carp if the homogenates contained the parasites described by Kovács -Gayer et al. (8). By intraperitoneal injection, the Unidentified Blood Organisms (UBOs) living in the blood of the common carp were transmissible to common carp, from the blood of which they were demonstrable for a long time. However, they were not transformed into Sphaerospora. To other cyprinids (gibel carp, silver carp, grass carp, tench, roach) neither the blood stages nor the swimbladder stages were transmissible from the common carp.     

Effect of total dissolved gas supersaturation on the survival of common carp (Cyprinus carpio) and silver carp (Hypophthalmichthys molitrix)

Flood discharge results in total dissolved gas (TDG) supersaturation downstream of a dam during the flood period. Fish suffer death from gas bubble disease (GBD) caused by TDG supersaturation. Nonetheless, current studies mainly attach importance to the survival of benthic fish affected by TDG supersaturation in the Yangtze River in China. Few studies have attempted to investigate the survival of pelagic fish influenced by TDG supersaturated water and compare the tolerance characteristics to TDG supersaturation between benthic and pelagic fish. To identify the survival of fish species that inhabit the various water layers affected by TDG supersaturation, silver carp (Hypophthalmichthys molitrix) (pelagic fish) and common carp (Cyprinus carpio) (benthic fish) were chosen to conduct an acute exposure experiment of four different TDG supersaturation levels (125%, 130%, 135% and 140%). The findings illustrated that the two fish species both exhibited evident aberrant behaviours of maladjustment in TDG supersaturated water. Obvious GBD symptoms were also found in the test fish. The survival probability of silver carp and common carp decreased with increasing levels of TDG supersaturation. The median survival time (ST₅₀) values of the silver carp exposed to four levels of TDG supersaturated water (125%, 130%, 135% and 140%) were 26.84, 7.96, 5.56 and 3.62 h, respectively, whereas the ST₅₀ values of common carp were 53.50, 26.00, 16.50 and 11.70 h, respectively. When compared with common carp, silver carp had a weaker tolerance to TDG-supersaturated water and were vulnerable to GBD. It shows that levels above 125% are not safe for common carp survival. In terms of the tolerance threshold value, silver carp merits further investigation because it showed lower tolerance to TDG than did common carp.     

Characterizations of four toll‐like receptor 4s in grass carp Ctenopharyngodon idellus and their response to grass carp reovirus infection and lipopolysaccharide stimulation

In this study, the subcellular localization, tissue distribution and response to grass carp reovirus (GCRV) infection and lipopolysaccharide (LPS) stimulation of four grass carp Ctenopharyngodon idellus toll‐like receptor 4 (tlr4) genes were investigated. All four genes were constitutively expressed in all tissues studied, but the subcellular localization and tissue exhibiting the highest expression differed for each protein. Following GCRV infection, all the four tlr4s were upregulated in all tissues examined, and stimulation of C. idellus kidney (CIK) cells with LPS resulted in downregulation of all four tlr4s. These results provide a foundation for further investigation of tlr4 genes in bony fishes.     

Polymorphism of the cytochrome oxidase b gene (<Emphasis Type="Italic">cyt b</Emphasis>) in Russian populations of common carp and wild common carp (<Emphasis Type="Italic">Cyprinus carpio</Emphasis> L.)

Polymorphism of the mitochondrial cyt b gene was examined in 35 individuals of common carp and wild common carp (Cyprinus carpio L.). The fish examined represented two natural populations from Khabarovsk krai (Ac and Am), Volga wild common carp, Don wild common carp, and two common carp breeds, Ropsha (strains BB and MM) common carp and Hungarian common carp. The highest level of nucleotide (π) and haplotype (h) diversity was detected in two strains of Ropsha common carp (MM, π = 0.67%, h = 0.7; and BB, π = 0.21%, h = 0.9) and in one population (Am) of Amur wild common carp (π = 0.26%; h = 0.6). The second population of Amur wild common carp (Ac) and Hungarian common carp were characterized by lower variation estimates (π = 0.035%, h = 0.4; and π = 0.09%, h = 0.7, respectively). Genetic homogeneity was demonstrated for the populations of Volga and Don wild common carp (π = 0, h = 0). In the sample of the cyt b sequences examined, three lineages were identified. Lineages I and II united all haplotypes of the Am Amur wild common carp along with two haplotypes of Ropsha common carp, strain MM. The third lineage (III) was formed by the haplotypes of three individuals of Ropsha common carp strain MM, all representatives of Ropsha common carp strain BB, Hungarian common carp, Ac Amur wild common carp, and Don and Volga wild common carps. Statistically significant amino acid differences were observed only for the sequences, corresponding to haplotypes of lineage III, and the sum of sequences of lineages I and II. Effectiveness of different types of markers to differentiate the two subspecies of European and Amur wild common carp (C. c. carpio and C. c. haematopterus) is discussed, as well as the issues of the origin and dispersal of Russian common carp and wild common carp breeds.     

Comparative analysis of intermuscular bones in three strains of common carp

The objective was to document the variations in the intermuscular bone counts among three carp strains: mirror carp, hybrid carp [Boshi carp (Cyprinus pellegrini) × Heilongjiang carp (Cyprinus carpio)], and a cold‐resistant strain of Hebao carp. The intermuscular bone counts, lengths, and weights were obtained from 146 fish; the bone count ranged from 55 to 110 (mean 92.85) among the three strains. Mirror carp had the lowest number of intermuscular bones and a higher coefficient of variation in counts relative to the other two species. There was no relationship between the intermuscular bone counts to standard length or body length. Similarly, there was no difference in the intermuscular bone count or shape between the left and right sides of the body. However, the count was significantly higher (P < 0.01) in the anterior region (snout to cloaca: 54% of the total count) than in the posterior region (cloaca to base of caudal fin: 46% of the total count). There was a significant difference between strains (P < 0.0001), but not between age classes; a significant difference was also observed in the posterior region counts among strains, but not in the anterior region (P < 0.0001). Similarly, the difference in counts in the posterior region was primarily related to differences in the counts of three of the six types: non‐forked (I), tree‐branch and two‐end‐bi‐fork type. Our results suggest that selective breeding protocols should target a reduction in the number of intermuscular bones in the posterior region, particularly in mirror carp.     

Genetic divergence between Cyprinus carpio carpio and Cyprinus carpio haematopterus as assessed by mitochondrial DNA analysis,with emphasis on origin of European domestic carp

Although common carp is the major fish species in Asian and European aquaculture and many domestic varieties have occurred, there is a controversy about the origination of European domestic common carp. Some scientists affirmed that the ancestor of European domestic common carp was Danube River wild common carp, but others considered it might be Asian common carp. For elucidating origination of European domestic common carp, we chose two representative European domestic common carp strains (German mirror carp and Russian scattered scaled mirror carp) and one wild common carp strain of Cyprinus carpio carpio subspecies (Volga River wild common carp) and two Asian common carp strains, the Yangtze River wild common carp (Cyprinus carpio haematopterus) and traditionally domestic Xingguo red common carp, as experimental materials. ND5–ND6 and D-loop segments of mitochondrial DNA were amplified by polymerase chain reaction and analyzed through restriction fragment length polymorphism (RFLP) and sequencing respectively. The results revealed that HaeIII and DdeI digestion patterns of ND5–ND6 segment and sequences of control region were different between European subspecies C. carpio carpio and Asian subspecies C. carpio haematopterus. Phylogenetic analysis showed that German mirror carp and Russian scattered scaled mirror carp belonged to two subspecies, C. carpio carpio and C. carpio haematopterus, respectively. Therefore, there were different ancestors for domestic carp in Europe: German mirror carp was domesticated from European subspecies C. carpio carpio and Russian scattered scaled mirror carp originated from Asian subspecies C. carpio haematopterus.     

Diet spectrum of great cormorants (<Emphasis Type="Italic">Phalacrocorax carbo sinensis</Emphasis> L.) at the Donji Miholjac carp fishponds in eastern Croatia

This paper reports the results of a diet analysis from the digestive tracts of 203 great cormorants shot at the Donji Miholjac fishponds in eastern Croatia, in the period 2000–2002. Eight fish species were determined. The dominant species was common carp (Cyprinus carpio) with a relative frequency of 73.4%, followed by grass carp (Ctenopharingodon idella) (11.6%), bighead carp (Aristichthys nobilis) (7.2%), pikeperch (Sander lucioperca) (2.0%), wels catfish (Silurus glanis) (1.7%), pike (Esox lucius) (1.5%), topmouth gudgeon (Pseudorasbora parva) (1.5%) and Prussian carp (Carassius auratus gibelio) (1.1%). The diet spectrum of the great cormorants from the Donji Miholjac fishponds was in accordance with the structure of the fish population in the fishponds. The average stomach weight per bird was 244 g. When birds shot with an empty stomach were excluded, the average stomach weight increased to 286 g. The length of consumed fish ranged from 40–335 mm, with 47% of the fish belonging to the length category 100–149 mm. The study revealed no significant relationship between the weight of the consumed fish and the body weight of male and female cormorants.     

Size‐specific effects of bighead carp predation across the zooplankton size spectra

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Phenotypic plasticity in the spawning traits of bigheaded carp (Hypophthalmichthys spp.) in novel ecosystems

1. Bigheaded carp, including both silver (Hypophthalmichthys molitrix) and bighead (H. nobilis) carp, are successful invasive fishes that threaten global freshwater biodiversity. High phenotypic plasticity probably contributes to their success in novel ecosystems, although evidence of plasticity in several spawning traits has hitherto been largely anecdotal or speculative. 2. We collected drifting eggs from a Midwestern U.S.A. river from June to September 2011 and from April to June 2012 to investigate the spawning traits of bigheaded carp in novel ecosystems. 3. Unlike reports from the native range, the presence of drifting bigheaded carp eggs was not related to changes in hydrological regime or mean daily water temperature. Bigheaded carp also exhibited protracted spawning, since we found drifting eggs throughout the summer and as late as 1 September 2011. Finally, we detected bigheaded carp eggs in a river reach where the channel is c. 30 m wide with a catchment area of 4579 km², the smallest stream in which spawning has yet been documented. 4. Taken with previous observations of spawning traits that depart from those observed within the native ranges of both bighead and silver carp, our findings provide direct evidence that bigheaded carp exhibit plastic spawning traits in novel ecosystems that may facilitate invasion and establishment in a wider range of river conditions than previously envisaged.     

Comparison of plankton intake by Indian carp fingerlings under different feeding conditions

The influence of different culture systems on the amount of plankton consumed by fingerlings of three carp species (Labeo rohita, Catla catla and Cirrhinus mrigala) was examined in outdoor culture tanks during a 90 day growing period. The fish were raised under two basically different feeding conditions: (a) fed with allochthonous live plankton; (b) fed with plankton grown autochthonously in the fish growing tanks fertilized with manures. Both feeding regimes were compared with control groups. It was found that the plankton intake for carp held in the live food system was significantly higher than with the manured and control systems. Related to maximum intake, maximum abundance of plankton in the live food system was a consequence of improved water quality expressed in terms of lower values of BOD and COD and higher values of DO and pH; this was conducive to fast reproduction of some of the zooplankton which constituted the major food items for the test carp, as well as to the regular exogenous introduction of live plankton. The ingestion of plankton was found to be related to carp body weight raised to the power b. The exponent b ranged from 0.29 to 0.93 during the 90-day growing period.