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Sex in vertebrates is commonly determined by genotype and environmental conditions, such as temperature. A few species display intermediate systems, combining sex chromosomes with temperature effect. This phenomenon has been reported in gibel carp ( Carassius gibelio ), an invasive cyprinid fish whose invasiveness is linked to the combination of sexual and asexual reproduction. Here, we compared gonadal transcriptomes between genotypic males and temperature-induced males of C. gibelio , focusing specifically on genes related to reproduction. Many meiosis and male differentiation pathways were common to genotypic and temperature-induced males. However, the underrepresentation of reproduction- and spermatogenesis-related terms in temperature-induced males suggests reduced reproductive abilities. Our study further highlights differential regulation of key genes related to male differentiation, steroid hormone signalling, meiosis, spermatogenesis, flagellar function, and sperm-egg interaction. In particular, induced males strongly overexpressed the key sex differentiation regulator hsd17b2 and slightly overexpressed the meiotic gene mnd1 , while genotypic males overexpressed sox8a , cyp19a1a , and the crucial fertilization gene izumo . Our study highlights the importance of males in the transition from asexual to sexual reproduction in this species and contributes to understanding the molecular mechanisms underlying the reproductive plasticity and invasiveness of C. gibelio in Europe.

The genes of the major histocompatibility complex (MHC) are an essential component of the vertebrate immune system and MHC genotypes may determine individual susceptibility to parasite infection. In the wild, selection that favors MHC variability can create situations in which interspecies hybrids experience a survival advantage. In a wild system of two naturally hybridizing leuciscid fish, we assessed MHC IIB genetic variability and its potential relationships to hosts’ ectoparasite communities. High proportions of MHC alleles and parasites were species-specific. Strong positive selection at specific MHC codons was detected in both species and hybrids. MHC allele expression in hybrids was slightly biased towards the maternal species. Controlling for a strong seasonal effect on parasite communities, we found no clear associations between host-specific parasites and MHC alleles or MHC supertypes. Hybrids shared more MHC alleles with the more MHC-diverse parental species, but expressed intermediate numbers of MHC alleles and positively selected sites. Hybrids carried significantly fewer ectoparasites than either parent species, suggesting a hybrid advantage via potential heterosis.

Background Several hypotheses have been proposed to explain parasite infection in parental species and their hybrids. Hybrid heterosis is generally applied to explain the advantage for F1 generations of hybrids exhibiting a lower level of parasite infection when compared to parental species. Post-F1 generations often suffer from genetic incompatibilities potentially reflected in the higher level of parasite infection when compared to parental species. However, the presence of specific parasites in an associated host is also limited by close coevolutionary genetic host-parasite associations. This study focused on monogenean parasites closely associated with two leuciscid fish species—common bream and roach—with the aim of comparing the level of monogenean infection between parental species and hybrids representing two F1 generations with different mtDNA and two backcross generations with different cyto-nuclear compositions. Results Monogenean infection in F1 generations of hybrids was lower when compared to parental species, in line with the hybrid heterosis hypothesis. Monogenean infection in backcross generations exhibited similarities with the parental species whose genes contributed more to the backcross genotype. The distribution of monogeneans associated with one or the other parental species showed the same asymmetry with a higher proportion of roach-associated monogeneans in both F1 generations and backcross generation with roach in the paternal position. A higher proportion of common bream-associated monogeneans was found in backcross generation with common bream in the paternal position. Conclusions Our study indicated that cyto-nuclear incompatibilities in hybrids do not induce higher monogenean infection in backcross generations when compared to parental species. However, as backcross hybrids with a higher proportion of the genes of one parental taxon also exhibited high level of this parental taxon-associated parasites, host-parasite coevolutionary interactions seem to play an obvious role in determining the level of infection of host-specific monogeneans in hybrids.

Gibel carp ( Carassius gibelio ) is a cyprinid fish that originated in eastern Eurasia and is considered as invasive in European freshwater ecosystems. The populations of gibel carp in Europe are mostly composed of asexually reproducing triploid females (i.e., reproducing by gynogenesis) and sexually reproducing diploid females and males. Although some cases of coexisting sexual and asexual reproductive forms are known in vertebrates, the molecular mechanisms maintaining such coexistence are still in question. Both reproduction modes are supposed to exhibit evolutionary and ecological advantages and disadvantages. To better understand the coexistence of these two reproduction strategies, we performed transcriptome profile analysis of gonad tissues (ovaries) and studied the differentially expressed reproduction-associated genes in sexual and asexual females. We used high-throughput RNA sequencing to generate transcriptomic profiles of gonadal tissues of triploid asexual females and males, diploid sexual males and females of gibel carp, as well as diploid individuals from two closely-related species, C. auratus and Cyprinus carpio . Using SNP clustering, we showed the close similarity of C. gibelio and C. auratus with a basal position of C. carpio to both Carassius species. Using transcriptome profile analyses, we showed that many genes and pathways are involved in both gynogenetic and sexual reproduction in C. gibelio ; however, we also found that 1500 genes, including 100 genes involved in cell cycle control, meiosis, oogenesis, embryogenesis, fertilization, steroid hormone signaling, and biosynthesis were differently expressed in the ovaries of asexual and sexual females. We suggest that the overall downregulation of reproduction-associated pathways in asexual females, and their maintenance in sexual ones, allows the populations of C. gibelio to combine the evolutionary and ecological advantages of the two reproductive strategies. However, we showed that many sexual-reproduction-related genes are maintained and expressed in asexual females, suggesting that gynogenetic gibel carp retains the genetic toolkits for meiosis and sexual reproduction. These findings shed new light on the evolution of this asexual and sexual complex.

Background Host specificity is one of the outputs of the coevolution between parasites and their associated hosts. Several scenarios have been proposed to explain the pattern of parasite distribution in parental and hybrid genotypes ranging from hybrid resistance to hybrid susceptibility. We hypothesized that host-parasite co-adaptation limits the infection of host-specific parasites in hybrid genotypes even under the condition of the high frequency of hybrids. The experimental monogenean infection in pure breeds of Blicca bjoerkna and Abramis brama and cross-breeds (the F1 generation of hybrids) under the condition of similar frequencies of pure and hybrid genotypes was investigated. We also examined the potential effect of the maternal origin of hybrids (potential co-adaptation at the level of mitochondrial genes) on monogenean abundance. Methods Pure breeds of two cyprinids and two cross-breeds (one with B. bjoerkna , the next with A. brama in the maternal positions) were exposed to infection by monogeneans naturally occurring in B. bjoerkna and A. brama . The experiment was run under similar frequencies of the four breed lines. Results We showed similar levels of monogenean infection in B. bjoerkna and A. brama . However, each species harboured specific monogenean fauna. Hybrids harboured all monogenean species specifically infecting one or the other species. Monogenean infection levels, especially those of Dactylogyrus specific to A. brama , were lower in hybrids. For the majority of host-specific parasites, there was no effect of the maternal origin of hybrids on monogenean abundance. Asymmetry was found in the distribution of specific parasites in favour of specialists of B. bjoerkna in the monogenean communities of hybrids. Conclusions Our results indicate that the maternal mtDNA of hybrids is not an important predictor of host-specific monogenean infection, which may suggest that mitochondrial genes are not strongly involved in the coadaptation between monogeneans and their associated hosts. The asymmetry of species-specific parasites suggests similarity between the molecular components of the immune mechanisms in hybrids and B. bjoerkna . Our results revealed a difference between the degree of host-parasite coadaptation in specific parasites of A. brama and the degree of host-parasite coadaptation in specific parasites of B. bjoerkna and their associated hosts.

Information about ploidy is important in both commercial and conservation aquaculture and fish research. Unfortunately, methods for its determination, such as karyology, determination of the amount of DNA in a cell using microdensitometry or flow cytometry and/or measuring erythrocytes in a blood smear can be stressful or even destructive. Some of these methods are also limited by the relatively large minimum size of the individual being measured. The aim of this study was to test a new low-stress method of determining ploidy by measuring the size of erythrocytes in the capillaries of a fish, including small individuals. First, we examined diploid and triploid loach ( Cobitis sp.) and gibel carp, Carassius gibelio (Bloch, 1782), using flow cytometry and blood smears, with these results being used as a control. Subsequently, we measured the size of erythrocytes in the caudal fin capillaries of anesthetized fishes of known ploidy under a light microscope. For both the loaches and gibel carp, direct observation of the mean erythrocyte size in epithelial fin capillaries provided a consistent and reliable determination of ploidy when compared with the controls based on flow cytometry and blood smears. This new method allows for rapid determination of ploidy in living small fish, where collection of tissue using other methods may cause excessive stress or damage. The method outlined here simply requires the measurement of erythrocytes directly in the bloodstream of a live fish, thereby making it possible to determine ploidy without the need for blood sampling. The method described is sufficiently efficient, less demanding on equipment than many other procedures, can be used by relatively inexperienced personnel and has benefits as regards animal welfare, which is especially important for fish production facilities or when dealing with rare or endangered species.

Monogenean parasites are often co-introduced with their fish hosts into novel areas. This study confirmed co-introduction of two dactylogyrids, Dactylogyrus squameus Gusev, 1955 and Bivaginogyrus obscurus (Gusev, 1955), and a newly described gyrodactylid species, Gyrodactylus pseudorasborae n. sp. into Europe along with their fish host, the invasive topmouth gudgeon Pseudorasbora parva (Temminck & Schlegel) from East Asia. All three species were observed in the lower Dnieper and middle Danube basin regions and had slightly larger haptoral hard parts than the same parasites in their native range. While dactylogyrids occurred sporadically, we recorded regular infection by G. pseudorasborae n. sp. at relatively high prevalence and abundance. This latter species was observed in both the native and non-native range of topmouth gudgeon, and resembles Gyrodactylus parvae You et al. , 2008 recently described from P. parva in China. Both species were distinguished based on genetic analysis of their ITS rDNA sequence (6.6% difference), and morphometric differences in the marginal hooks and male copulatory organ. Phylogenetic analysis of dactylogyrid monogeneans showed that B. obscurus clustered with Dactylogyrus species parasitising Gobionidae and Xenocyprididae, including D. squameus , supporting recent suggestions of a paraphyletic origin of the Dactylogyrus genus. In addition to co-introduced parasites, topmouth gudgeon was infected with a local generalist, G. prostae Ergens, 1964, increasing the number of monogeneans acquired in Europe to three species. Nevertheless, monogenean infections were generally lower in non-native host populations, potentially giving an advantage to invading topmouth gudgeon. Les parasites monogènes sont souvent co-introduits avec leurs hôtes poissons dans de nouvelles zones. Cette étude a confirmé la co-introduction de deux Dactylogyridae, Dactylogyrus squameus Gusev, 1955 et Bivaginogyrus obscurus (Gusev, 1955), et d’une espèce de Gyrodactylidae nouvellement décrite, Gyrodactylus pseudorasborae n. sp. en Europe, en même temps que leur hôte poisson, le goujon envahissant Pseudorasbora parva (Temminck & Schlegel) qui vient d’Asie de l’Est. Les trois espèces ont été observées dans les régions du bas Dniepr et du bassin moyen du Danube et avaient des parties sclérifiées haptorales légèrement plus grandes que les mêmes parasites dans leur aire de répartition d’origine. Alors que les Dactylogyridae étaient rares, nous avons enregistré une infection régulière par G. pseudorasborae n. sp. à une prévalence et une abondance relativement élevées. Cette dernière espèce a été observée à la fois dans l’aire de répartition indigène et non indigène du goujon asiatique et ressemble à Gyrodactylus parvae You et al. , 2008 récemment décrit chez P. parva en Chine. Les deux espèces ont été distinguées sur la base de l’analyse génétique de leur séquence ITS de l’ADNr (différence de 6,6 %) et des différences morphométriques dans les crochets marginaux et l’organe copulateur mâle. L’analyse phylogénétique des monogènes Dactylogyridae a montré que B. obscurus se regroupait avec des espèces de Dactylogyrus parasitant les Gobionidae et les Xenocyprididae, y compris D. squameus , soutenant les suggestions récentes d’une origine paraphylétique du genre Dactylogyrus . En plus des parasites co-introduits, le goujon asiatique était infecté par un généraliste local, G. prostae Ergens, 1964, portant le nombre de monogènes acquis en Europe à trois espèces. Néanmoins, les infections par les monogènes étaient généralement plus faibles dans les populations d’hôtes non indigènes, donnant potentiellement un avantage à l’invasion du goujon asiatique.

Parasite-mediated selection is considered one of the potential mechanisms contributing to the coexistence of asexual-sexual complexes. Gibel carp ( ), an invasive fish species in Europe, often forms populations composed of gynogenetic and sexual specimens. The experimental infection was induced in gynogenetic and sexual gibel carp using eye-fluke (Trematoda), and the transcriptome profile of the spleen as a major immune organ in fish was analyzed to reveal the differentially expressed immunity-associated genes related to infection differing between gynogenetic and sexual gibel carp. High parasite infection was found in gynogenetic fish when compared to genetically diverse sexuals. Although metacercariae of are situated in an immune-privileged organ, our results show that eye trematodes may induce a host immune response. We found differential gene expression induced by eye-fluke infection, with various impacts on gynogenetic and sexual hosts, documenting for the majority of DEGs upregulation in sexuals, and downregulation in asexuals. Differences in gene regulation between gynogenetic and sexual gibel carp were evidenced in many immunity-associated genes. GO analyses revealed the importance of genes assigned to the GO terms: immune function, the Notch signaling pathway, MAP kinase tyrosine/threonine/phosphatase activity, and chemokine receptor activity. KEGG analyses revealed the importance of the genes involved in 12 immunity-associated pathways - specifically, FoxO signaling, adipocytokine signaling, TGF-beta signaling, apoptosis, Notch signaling, C-type lectin receptor signaling, efferocytosis, intestinal immune network for IgA production, insulin signaling, virion - human immunodeficiency virus, Toll-like receptor signaling, and phosphatidylinositol signaling system. Our study indicates the limited potential of asexual fish to cope with higher parasite infection (likely a loss of capacity to induce an effective immune response) and highlights the important role of molecular mechanisms associated with immunity for the coexistence of gynogenetic and sexual gibel carp, potentially contributing to its invasiveness.

Non-native species have the potential to alter host community structure, which, in turn, may affect parasite transmission, diversity and distribution, particularly when the introduced host is ecologically or phylogenetically related to its native host(s). In this study, we examine whether the introduction of a competent non-native fish host shifts parasite load and community composition in native fish species. Specifically, we focus on two Eupercarian species: the native Eurasian perch ( Perca fluviatilis ) and the non-native pumpkinseed sunfish ( Lepomis gibbosus ), which was introduced into Europe from North America. Our results show that the parasite community of native perch co-occurring with non-native pumpkinseed (sympatric populations) differ significantly from those of perch inhabiting sites unaffected by pumpkinseed introduction (allopatric populations). Moreover, sympatric perch populations exhibit significantly higher parasite abundance, infracommunity richness and dominance, along with lower equitability, compared to allopatric populations. These findings suggest that the introduction of pumpkinseed may have contributed to shifts in both parasite abundance and diversity in native fish hosts. Additionally, our study indicates a potential parasite spillback of the trematode Bucephalus polymorphus and spill-over of the Asian invasive parasitic copepod Neoergasilus japonicus , both associated with pumpkinseed establishment in the study area. The successful inclusion of non-native fish species into the life cycles of several independently introduced and unrelated invasive parasites underscores the importance of introduced hosts in driving changes within local parasite communities.