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"Goldfish."
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Functional Divergence of Multiple Duplicated Foxl2 Homeologs and Alleles in a Recurrent Polyploid Fish
Evolutionary fates of duplicated genes have been widely investigated in many polyploid plants and animals, but research is scarce in recurrent polyploids. In this study, we focused on foxl2, a central player in ovary, and elaborated the functional divergence in gibel carp (Carassius gibelio), a recurrent auto-allo-hexaploid fish. First, we identified three divergent foxl2 homeologs (Cgfoxl2a-B, Cgfoxl2b-A, and Cgfoxl2b-B), each of them possessing three highly conserved alleles and revealed their biased retention/loss. Then, their abundant sexual dimorphism and biased expression were uncovered in hypothalamic–pituitary–gonadal axis. Significantly, granulosa cells and three subpopulations of thecal cells were distinguished by cellular localization of CgFoxl2a and CgFoxl2b, and the functional roles and the involved process were traced in folliculogenesis. Finally, we successfully edited multiple foxl2 homeologs and/or alleles by using CRISPR/Cas9. Cgfoxl2a-B deficiency led to ovary development arrest or complete sex reversal, whereas complete disruption of Cgfoxl2b-A and Cgfoxl2b-B resulted in the depletion of germ cells. Taken together, the detailed cellular localization and functional differences indicate that Cgfoxl2a and Cgfoxl2b have subfunctionalized and cooperated to regulate folliculogenesis and gonad differentiation, and Cgfoxl2b has evolved a new function in oogenesis. Therefore, the current study provides a typical case of homeolog/allele diversification, retention/loss, biased expression, and sub-/neofunctionalization in the evolution of duplicated genes driven by polyploidy and subsequent diploidization from the recurrent polyploid fish.
Journal Article
Molecular and immunohistochemical characterization of intestinal macrophages subsets in goldfish
Macrophage-colony stimulating factor (M-CSF) plays a crucial role in the proliferation and differentiation of the monocyte-macrophage lineage across vertebrates. In this study, we identified and characterized two CSF1 paralogues and their corresponding receptor genes in the intestine of the goldfish (
Carassius auratus
), showing sequence homology with known teleost species. Immunohistochemical analysis confirmed the presence of CSF, its receptor CSF-R1, and bone morphogenetic protein 2 (BMP2) in intestinal macrophages. These macrophages were localized within mucosal, submucosal, and muscularis layers, suggesting distinct functional subtypes. Quantitative PCR analysis revealed differential gene expression patterns, with
csf1a
,
csf1b
, and
csf1ra
highly expressed in the brain, while
csf1rb
transcripts were predominant in the intestine. Immunophenotypic characterization using CD14 and CD86 markers further demonstrated macrophage heterogeneity. Additionally, BMP2-expressing macrophages were observed in the muscularis externa, implying a potential role in neuromuscular regulation. These findings provide novel insights into the molecular and immunohistochemical profiles of goldfish intestinal macrophages, highlighting their potential role in immune responses and gut homeostasis.
Journal Article
Goldfish on vacation
Three goldfish live in a small bowl, in an apartment building, in the middle of a big city, until one summer they get to go on vacation--in a fountain, with lily pads, and reeds, and other neighborhood goldfish.
Book
Sex chromosome and sex locus characterization in goldfish, Carassius auratus (Linnaeus, 1758)
Background
Goldfish is an important model for various areas of research, including neural development and behavior and a species of significant importance in aquaculture, especially as an ornamental species. It has a male heterogametic (XX/XY) sex determination system that relies on both genetic and environmental factors, with high temperatures being able to produce female-to-male sex reversal. Little, however, is currently known on the molecular basis of genetic sex determination in this important cyprinid model. Here we used sequencing approaches to better characterize sex determination and sex-chromosomes in an experimental strain of goldfish.
Results
Our results confirmed that sex determination in goldfish is a mix of environmental and genetic factors and that its sex determination system is male heterogametic (XX/XY). Using reduced representation (RAD-seq) and whole genome (pool-seq) approaches, we characterized sex-linked polymorphisms and developed male specific genetic markers. These male specific markers were used to distinguish sex-reversed XX neomales from XY males and to demonstrate that XX female-to-male sex reversal could even occur at a relatively low rearing temperature (18 °C), for which sex reversal has been previously shown to be close to zero. We also characterized a relatively large non-recombining region (~ 11.7 Mb) on goldfish linkage group 22 (LG22) that contained a high-density of male-biased genetic polymorphisms. This large LG22 region harbors 373 genes, including a single candidate as a potential master sex gene, i.e., the anti-Mullerian hormone gene (
amh
). However, no sex-linked polymorphisms were detected in the coding DNA sequence of the goldfish
amh
gene.
Conclusions
These results show that our goldfish strain has a relatively large sex locus on LG22, which is likely the Y chromosome of this experimental population. The presence of a few XX males even at low temperature also suggests that other environmental factors in addition to temperature could trigger female-to-male sex reversal. Finally, we also developed sex-linked genetic markers, which will be important tools for future research on sex determination in our experimental goldfish population. However, additional work would be needed to explore whether this sex locus is conserved in other populations of goldfish.
Journal Article
Gilbert Goldfish wants a pet
Gilbert has everything a goldfish could want except a pet of his own, but none of the animals who come near his fishbowl seem quite right until Fluffy, with his long tail and whiskers, appears.
Book
Is there direct photoentrainment in the goldfish liver? Wavelength-dependent regulation of clock genes and investigation of the opsin 7 family
Widespread direct photoentrainment in zebrafish peripheral tissues is linked to diverse non-visual opsins. To explore whether this broadly distributed photosensitivity is specific to zebrafish or is a general teleost feature, we investigated hepatic photosynchronization in goldfish. First, we focused on the opsin 7 family (OPN7, a key peripheral novel opsin in zebrafish), investigating its presence in the goldfish liver. Subsequently, we studied whether light can directly entrain the goldfish liver and retina clocks. Silico analysis revealed seven OPN7 paralogs from four gene families, suggesting expansion through whole-genome and tandem duplications. The paralogs of families OPN7a, OPN7b, and OPN7d were mainly localized in neural tissues, while OPN7c paralogs were more abundant in peripheral tissues—including the liver—suggesting divergent roles. Light (independently of the wavelength employed) directly induced the
per2a
clock gene in the retina both in vivo and in vitro, confirming expected photoentrainment. However, in the liver, photoinduction of
per1a
and
cry1a
only occurred in vivo, not in vitro. These results suggest an indirect light-entrainment mechanism of the goldfish hepatic clock, possibly mediated by other oscillators or photosensitive organs. Our findings challenge the assumption of widespread direct photosensitivity in the peripheral tissues of teleosts. Further research is needed to understand the role of tissue-specific photoentrainment and non-visual opsins in diverse teleost species.
Journal Article
SOS water
\"With humour and a touch of poetry, this picture book features a sailor named Lalo, and Rosa, a goldfish, dealing with the pollution caused by plastic water bottles. Lalo wants to find a safe and clean place for his fish friend, but everywhere they go has been invaded by plastic water bottles.\"-- Provided by publisher.
CHILDBOOK
Chronic hypoxia induces alternative splicing of transcripts in the goldfish brain
Several species evolved mechanisms to tolerate periods of severe environmental hypoxia and anoxia. Among them, goldfish are unique as they do not enter a comatose state under such conditions. Taking advantage of the recently published and annotated goldfish genome, we had previously profiled the transcriptomic response of the goldfish brain under normoxic (21 kPa oxygen saturation, N) and hypoxic conditions (2.1 kPa oxygen saturation) after 1 and 4 weeks (1WH, 4WH). Using the RNA-Seq data, we report the occurrence of alternative mRNA splicing (skipped exon, retained intron, alternative 3′ or 5′ splice sites, and mutually exclusive exons). At 1WH/N, 1004 significant alternative splicing events on 769 gene loci were identified, increasing to 1187 on 963 loci at 4WH/N. There were 305 loci with alternatively spliced transcripts common to both 1WH/N and 4WH/N, 221 of which exhibited the same precise location and splicing mechanism. Specific gene transcripts affected by alternative splicing events were almost entirely different from previously identified differentially expressed genes under chronic hypoxia. GO-term enrichment analyses of gene loci of alternatively spliced transcripts, however, did include similar pathways as previously identified for DEGs. These include epigenetic machinery, ion channel activity (1WH/N), glutamate signaling (4WH/N), endothelial cell function, and ATP hydrolyzation pathways (1WH/N + 4WH/N). We describe selected examples of alternatively spliced transcripts to discuss possible functional relevance in the goldfish brain response to chronic hypoxia. Together, our data identified an additional layer of regulation in brain pathways relevant to hypoxia tolerance in goldfish, which complement previously reported gene expression changes.
Journal Article