Chromosome evolution in fish: sex chromosome variability in Eigenmannia virescens (Gymnotiformes: Sternopygidae)

New data are presented on the sex chromosomes of the fish species Eigenmannia virescens (Gymnotiformes, Sternopygidae). A new finding, involving the occurrence of ZZ/ZW sex chromosomes, is described in specimens sampled from the S?o Francisco and Amazon river basins in Brazil. All individuals had a chromosome number of 2n = 38. The homologs of the sex chromosome pair from the S?o Francisco river basin sample differed only in their morphology, while those from the Amazonian sample differed both in morphology and heterochromatin pattern. A possible model for the evolution of the sex chromosomes in E. virescens is proposed, including data from populations from the Paraná (Brazil) river basin, in which male heterogamety has already been described. The occurrence of different sex chromosome systems in species and populations of the neotropical freshwater fish fauna is discussed.     

Complex sex chromosome system in Eigenmannia sp. (Pisces,Gymnotiformes)

Chromosomes of Eigenmannia sp. (7 males and 15 females) collected from the Tietê River in Botucatu (SP, Brazil) were examined from gill, kidney and testicular cells. The diploid chromosome number in males was 2n=31 and in females, 2n=32. In both sexes the number of chromosomal arms was 40. The difference in diploid number was due to the fusion of two acrocentrics. Mitotic and meiotic studies suggested that one of the fused acrocentrics was the Y chromosome. The sex-determining mechanism in Eigenmannia sp. could therefore be XX, AA in the female and X, \-YA A in the males. One of the males presented 2n=30 chromosomes due to the occurrence of another fusion of acrocentrics. C-banding analysis of the mitotic chromosomes revealed constitutive heterochromatin in the centromeric regions of all acrocentrics. However, small metacentrics were C-band negative. The YA chromosome is C-band negative except for a small amount of heterochromatin in the centromeric region. The nucleolar organizer region as identified by Ag-staining is present in the interstitial region of chromosome pair No. 10.     

Non-homologous sex chromosomes in two species of the genus Eigenmannia (Teleostei: Gymnotiformes)

The Neotropical genus Eigenmannia is a fish group with unknown species diversity where representatives possess a broad range of chromosomal sex determining systems namely XY/XX, X(1)X(2)Y/X(1)X(1)X(2)X(2), ZZ/ZW as well as homomorphic sex chromosomes. To test the homology of two heteromorphic XY sex chromosome systems present in two sympatric populations, reciprocal cross-species FISH experiments were performed using probes derived by microdissection of X and Y chromosomes present in analyzed specimens of Eigenmannia virescens and Eigenmannia sp.2, respectively. While X and Y paint probes hybridized to species-specific sex chromosomes, in reciprocal cross-FISH both probes hybridized exclusively to autosomes. The result suggests multiple independent origins of the XY systems in the analyzed populations.     

Microgeographic diversity in the neotropical knife-fish Eigenmannia macrops (Gymnotiformes,Sternopygidae)

Synopsis Examination of 681 individuals of electric knife-fishes, Eigenmannia macrops, from the Orinoco River, Venezuela, revealed a pattern of morphological diversity which covaried with habitat. Fishes from the caños (small stream tributaries) are on average smaller than those from offshore habitats and lagoons. Even at the same body size the tails of caño fishes are relatively shorter, their bodies are deeper, and their heads are relatively wider than those from other habitats. Caño fishes are more different from fishes of the deep channels than from those of the shallow channels, beaches, or lagoons. Several factors are examined for their possible relationship to the origin of the observed diversity including electrophysiology, locomotory adaptation, growth rate and ecology. The problems of genetic versus physiological bases of morphological variation are addressed.     

Nine polymorphic microsatellite loci in the Neotropical electric eel Eigenmannia (Teleostei: Gymnotiformes)

Ten microsatellite loci were isolated from a species of the Neotropical electric eel, Eigenmannia, a genus of freshwater fish characterized by small populations and low vagility. Nine microsatellites were polymorphic, the number of alleles ranging from seven to 27, and values of observed heterozygosity ranging from 0.327 to 0.741. These loci were developed for population genetic studies of Eigenmannia sp. 2, however, cross‐amplification carried out with other species of this genus as well as other Gymnotiformes genera indicate that these molecular markers are also potentially useful for population‐level studies in closely related species.     

Cytogenetic studies in Eigenmannia virescens (Sternopygidae, Gymnotiformes) and new inferences on the origin of sex chromosomes in the Eigenmannia genus

Background

Previous studies suggested that multiple domestication events in South and South-East Asia (Yunnan and surrounding areas) and India have led to the genesis of modern domestic chickens. Ha Giang province is a northern Vietnamese region, where local chickens, such as the H'mong breed, and wild junglefowl coexist. The assumption was made that hybridisation between wild junglefowl and Ha Giang chickens may have occurred and led to the high genetic diversity previously observed. The objectives of this study were i) to clarify the genetic structure of the chicken population within the Ha Giang province and ii) to give evidence of admixture with G. gallus. A large survey of the molecular polymorphism for 18 microsatellite markers was conducted on 1082 chickens from 30 communes of the Ha Giang province (HG chickens). This dataset was combined with a previous dataset of Asian breeds, commercial lines and samples of Red junglefowl from Thailand and Vietnam (Ha Noï). Measurements of genetic diversity were estimated both within-population and between populations, and a step-by-step Bayesian approach was performed on the global data set.

Results

The highest value for expected heterozygosity (> 0.60) was found in HG chickens and in the wild junglefowl populations from Thailand. HG chickens exhibited the highest allelic richness (mean A = 2.9). No significant genetic subdivisions of the chicken population within the Ha Giang province were found. As compared to other breeds, HG chickens clustered with wild populations. Furthermore, the neighbornet tree and the Bayesian clustering analysis showed that chickens from 4 communes were closely related to the wild ones and showed an admixture pattern.

Conclusion

In the absence of any population structuring within the province, the H'mong chicken, identified from its black phenotype, shared a common gene pool with other chickens from the Ha Giang population. The large number of alleles shared exclusively between Ha Giang chickens and junglefowl, as well as the results of a Bayesian clustering analysis, suggest that gene flow has been taking place from junglefowl to Ha Giang chickens.     

FMRFamide-like immunoreactivity in the brain and pituitary of the teleost. Eigenmannia lineata (Gymnotiformes)

The distribution of cells immunoreactive for the molluscan tetrapeptide FMRFamide in the brain and the pituitary of Eigenmannia was investigated immunohistochemically by the use of the peroxidase-antiperoxidase (PAP) technique and unlabelled antibodies. FMRFi neurons were located in the ganglion of the nervus terminalis at the rostroventral side of the bulbus olfactorius. FMRFi perikarya were also found in a dorsomedial diencephalic nucleus, in the nucleus ventromedialis, in some liquor-contacting neurons of the nucleus lateralis tuberis and of the nucleus recessus lateralis and posterior. The perikarya of the midbrain pre-pacemaker nucleus were only weakly immunoreactive for FMRFamide while large FMRFi neurons (T-cells) occurred in lamina VI of the torus semicircularis, in the brain stem, in dorsal and medial layers of the lobus lineae lateralis posterior (LLLp) and in the medullary electric organ pacemaker nucleus (pm). FMRFi fibers and nerve endings were found in the bulbus olfactorius, in medial areas of the telencephalon, and rather densely in the rostral diencephalon. Ventrocaudally to most of the hypothalamic nuclei the occurrence of immunoreactive fibres increased; many coursed to the pituitary through the pituitary stalk. FMRFi fibres also appeared in the deep layers of the tectum opticum, in the torus semicircularis, in the medial and lateral medulla and below the pacemaker nucleus. Wherever FMRFamide-immunoreactivity occurred fibres and nerve endings could be found in close contact with blood vessels.     

Sex differences in the electrocommunication signals of Sternarchogiton nattereri (Gymnotiformes: Apteronotidae)

In this study we examined electrocommunication behavior in Sternarchogiton nattereri (Apteronotidae), a weakly electric fish from South America. We focused on variation between females and males lacking external dentition and used playbacks of simulated conspecifics to elicit chirps (modulations of their electric organ discharge, EOD). Chirp responses were not affected by the frequency of the playback stimulus. EOD frequency, chirp rate, and chirp duration were not sexually dimorphic; however, the amount of chirp frequency modulation was significantly greater in toothless males than in females. These results reinforce that sex differences in chirp structure are highly diverse and widespread in the Apteronotidae.     

Biased distribution of repetitive elements: a landmark for neo-Y chromosome evolution in Drosophila miranda

It is generally assumed that the sex chromosomes developed from a pair of homologs. Over evolution, the proto-Y chromosome, with a very short differential segment, matured in its final stage into a heterochromatic and, for the most part, genetically eroded Y chromosome. The constraints on the evolution of the proto-Y chromosome have been speculated upon since the sex chromosomes were discovered. Several models have been suggested. Drosophila miranda has proved to be a unique and potent model system to study Y-chromosome evolution. We use selected test genes distributed along the neo-Y chromosome as entry gates to analyze the molecular mechanisms involved in the process of Y-chromosome evolution. Here, we report our findings on the Krüppel gene (Kr), which is located distally on the neo-sex chromosome pair.     

Sex chromosome evolution in non-mammalian vertebrates

Birds, reptiles, amphibia and fish have an enormous variety of chromosomal sex determination mechanisms that apparently do not follow any phylogenetic or taxonomic scheme. A similar picture is now emerging at the molecular level. Most genes that function downstream of the mammalian master sex-determining gene, Sry, have been found in non-mammalian vertebrates. Although the components of the machinery that determines sex seem to be conserved, their interaction and most importantly the initial trigger is not the same in all vertebrates. This variety is the consequence of the extremely dynamic process of the evolution of sex determination mechanisms and sex chromosomes, which is prone to create differences rather than uniformity.     

Phylogenetic systematics and historical biogeography of the Neotropical electric fish Sternopygus (Teleostei: Gymnotiformes)

Interrelationships among 10 extant species of the Neotropical electric fish Sternopygus are inferred from phylogenetic analysis of 66 morphological characters, including features of pigmentation, body proportions, meristics and osteology. A total of 287 lots containing 677 specimens were examined. The important findings of this study are: (1) S. branco is the most basal species unique among congeners in being restricted to whitewater rivers in the Central Amazon Basin, (2) S. sp. ‘cau’ from the Rio Caura of Venezuela is the sister taxon to (S. obtusirostris + S. astrabes), (3) S. castroi is a junior synonym of S. astrabes, (4) S. macrurus is the sister taxon to (S. arenatus + S. xingu + S. aequilabiatus species group) and (5) S. arenatus is the sister taxon to (S. xingu + S. aequilabiatus species group). A key to the adults of Sternopygus species is provided. Several features of S. astrabes previously thought to be plesiomorphic are now considered derived, including: short body cavity, paedomorphic cranial osteology, and the habitat restriction to terra firme streams. Sternopygus species assemblages in the Pacific (trans‐Andean) and Atlantic (cis‐Andean) slopes of northwestern South America are not monophyletic and do not result exclusively from local or regional radiations. The clade composed of S. macrurus, S. arenatus, S. xingu and the S. aequilabiatus species group is inferred to predate the Middle Miocene uplift of the Eastern Cordillera (c. 11.8–12.2 Ma). As currently recognized S. macrurus is the most widely distributed and most eurytopic gymnotiform species, inhabiting all hydrogeographical regions of tropical South America and most lowland aquatic habitats. Other Sternopygus species have much more restricted geographic and ecological distributions. Perceptions of phylogenetic patterns in Sternopygus are shown to be highly sensitive to taxon sampling.     

The Electric Glass Knifefishes of the Eigenmannia trilineata species‐group (Gymnotiformes: Sternopygidae): monophyly and description of seven new species

Eigenmannia trilineata López and Castello, 1966 (Sternopygidae) was described from the Río de La Plata basin and subsequently cited from most South American river basins. Questions about the limits of this species raise the possibility of the occurrence of undescribed species misidentified as E. trilineata. Herein we propose the Eigenmannia trilineata species group for species that share the presence of the superior medial stripe on the flank. This group comprises: E igenmannia antonioi sp. nov. , from the Rio Anapu, Rio Amazonas basin; E igenmannia desantanai sp. nov. , from the Rio Cuiabá, Rio Paraguay basin; E igenmannia guairaca sp. nov. , from the Riacho Água do Ó, upper Rio Paraná basin; E igenmannia matintapereira sp. nov. , from the Rio Uneiuxi and Rio Urubaxi, Rio Negro basin; Eigenmannia microstoma (Reinhardt, 1852), from the Rio São Francisco basin; E igenmannia muirapinima sp. nov. , from small tributaries of the Rio Amazonas; E igenmannia pavulagem sp. nov. , from the tributaries of Rio Capim, Rio Guamá basin; E. trilineata, from the lower Rio Paraná basin and Río de La Plata basin; Eigenmannia vicentespelaea Triques, 1996, from São Vicente I and II caves, Rio Tocantins basin; and E igenmannia waiwai sp. nov. , from the Rio Trombetas basin. These species can be distinguished from each other by unique sets of meristics, morphometrics, osteological and colour pattern features. © 2015 The Linnean Society of London     

Directional characteristics of tuberous electroreceptors in the weakly electric fish,Hypopomus (Gymnotiformes)

This paper is an electrophysiological study of the directionality of the tuberous electroreceptors of weakly electric fish. We recorded from two classes of tuberous electroreceptors known for pulse gymnotiforms: Burst Duration Coders (BDCs), and Pulse Markers (PMs). Both code for stimulus amplitude, although the dynamic range for BDCs is greater, and both exhibit strong directional preferences. Polar plots of spike number (for BDCs) or spike threshold (for PMs) versus electric field azimuth, are figure-8 shaped with two asymmetrical, elliptical lobes separated by 180°. The best azimuth of these two types of receptors from a given body region correlate with each other and with measures of best azimuth for transepidermal current flow. The shape and asymmetry of the directionality profiles appear to be caused by filter dynamics of the receptors. Pulse Markers are located on the anterior part of the body surface while Burst Duration Coders are located all over. The best directions of receptors in the anterior third of the body vary systematically with location from 0° to 180°. This region is probably critical for determining the direction of local electric fields. Together these receptors provide the CNS with sufficient information to construct a map of horizontal plane electric field directions.Abbreviations BDC Burst Duration Coder - ELL electrosensory lateral line lobe - EOD electric organ discharge - nALL anterior lateral line nerve - PM Pulse Marker     

Phylogenetic interrelationships,taxonomy, and reductive evolution in the Neotropical electric fish genus Hypopygus (Teleostei,Ostariophysi, Gymnotiformes)

A phylogenetic reconstruction of the Neotropical electric fish genus Hypopygus based on 47 parsimony‐informative morphological characters is presented. A series of synapomorphies support the hypothesis of monophyly of Hypopygus, and partially resolve species‐level relationships within the genus. Hypopygus species are recognized here as miniaturized fishes based on two criteria; first, a derived condition of diminutive body size, and; second, the presence of a suite of reductive morphological characters, including partial or total losses, simplifications, and reductions of the anal‐fin rays, scales, cranial bones, and laterosensory canal system. Reductive characters associated with miniaturization comprise 45% of the total number of characters in the phylogenetic reconstruction of the genus. Miniaturization and reductive morphological evolution in Hypopygus are discussed here in the phylogenetic context. A taxonomic revision of Hypopygus is presented, in which five new species are described, two species previously assigned to the genus are redescribed, and a single known species of Stegostenopos is redescribed and included in Hypopygus as a junior synonym. Distribution maps and a key for all eight valid species of Hypopygus are provided, based on the examination of 5014 catalogued museum specimens. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 163 , 1096–1156.     

Sex chromosome evolution and speciation in Ficedula flycatchers

Speciation is the combination of evolutionary processes that leads to the reproductive isolation of different populations. We investigate the significance of sex-chromosome evolution on the development of post- and prezygotic isolation in two naturally hybridizing Ficedula flycatcher species. Applying a tag-array-based mini-sequencing assay to genotype single nucleotide polymorphisms (SNPs) and interspecific substitutions, we demonstrate rather extensive hybridization and backcrossing in sympatry. However, gene flow across the partial postzygotic barrier (introgression) is almost exclusively restricted to autosomal loci, suggesting strong selection against introgression of sex-linked genes. In addition to this partial postzygotic barrier, character displacement of male plumage characteristics has previously been shown to reinforce prezygotic isolation in these birds. We show that male plumage traits involved in reinforcing prezygotic isolation are sex linked. These results suggest a major role of sex-chromosome evolution in mediating post- and prezygotic barriers to gene flow and point to a causal link in the development of the two forms of reproductive isolation.     

New species of glass knifefish genus Eigenmannia (Gymnotiformes: Sternopygidae) with comments on the morphology and function of the enlarged cephalic lateral-line canals of Sternopygidae

A new species of Eigenmannia is described from the Rio Paraná (the Grande, Paranapanema and Tietê basins). This new species is distinguished from all congeners by colouration pattern, position of the mouth, relative depth of posterodorsal expansion on infraorbitals 1 + 2, number of teeth, osteological features, number of rows of scales above lateral line (LL) and morphometric data. Comments on the widened cephalic lateral-line canals of Sternopygidae and a dichotomous key to the species of Eigenmannia from the Rio Paraná Basin are provided.     

Sex chromosome variability outlines the pathway to the chromosomal evolution in Microtus thomasi (Rodentia, Arvicolinae)

The underground vole Microtus thomasi , a Balkan endemic, displays remarkable variability in sex chromosome size and morphology. In the present study, we demonstrate this variability in two of its chromosomal races with 2 n  = 44 (i.e. 'thomasi' and 'atticus') with the use of C-banding on a sample of 189 individuals from 50 localities of Greece. In 'thomasi', five different, acrocentric X chromosome variants (X₀–X₄) are described, which differ significantly in size, due to heterochromatin addition. Also, three Y chromosome variants are described (Y₀–Y₂), ranging in size from very small (Y₀) to large (Y₂). The 'atticus' race displays three subtelocentric variants of the X chromosome (X_st0–X_st2), which differ in arm length ratio and heterochromatin content. In Peloponnesus, males of this race exhibit Y₀ and Y₁, whereas, in Attiki (south-east Sterea Ellada), males carry the small metacentric, Y_m. Overall, there is a trend towards sex chromosome size increase in a south to north direction. We propose that the last glaciation must have restricted M. thomasi to a refugium in southern Peloponnesus. During post-glacial colonization, limited northward expansion of its distribution area must have been accompanied by consecutive heterochromatin addition, which is proven today by comparatively larger sex chromosomes in the northern populations of 'thomasi' and 'atticus' in Greece than in their southern populations.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 96 , 685–695.     

Sex chromosome evolution in the clam shrimp,Eulimnadia texana

Chromosomes that determine sex are predicted to evolve differently than autosomes: a lack of recombination on one of the two sex chromosomes is predicted to allow an accumulation of deleterious alleles that eventually leads to reduced functionality and potential physical degradation of the nonrecombining chromosome. Because these changes should occur at an elevated evolutionary rate, it is difficult to find appropriate species in which to test these evolutionary predictions. The unique genetic sex‐determining mechanism of the crustacean Eulimnadia texana prevents major chromosome degeneration because of expression of both ‘proto‐sex’ (i.e. early stage of development) chromosomes in homozygous form (ZZ and WW). Herein, we exploit this unique genetic system to examine the predicted accumulation of deleterious alleles by comparing both homogametic sexual types to their heterogametic counterpart. We report differences in crossing over in a sex‐linked region in the ZW hermaphrodites (～ 3%) relative to the ZZ males (～ 21%), indicative of cross‐over suppression in the ZW hermaphrodites. Additionally, we report that both ZZ and WW genotypes have reduced fitness relative to ZW hermaphrodites, which is consistent with the prediction of harboured recessive mutations embedded on both the Z and the W chromosomes. These results suggest that the proto‐sex chromosomes in E. texana accumulate recessive deleterious alleles. We hypothesize that recessive deleterious alleles of large effect cannot accumulate because of expression in both ZZ and WW individuals, keeping both chromosomes from losing significant function.