Turtles demonstrate variability in sex determination and, hence, constitute an excellent model for the evolution of sex chromosomes. Notably, the sex determination of the freshwater turtles from the family Chelidae, a species-rich group with wide geographical distribution in the southern hemisphere, is still poorly explored. Here we documented the presence of an XX/XY sex determination system in seven species of the Australasian chelid genera Chelodina, Emydura, and Elseya by conventional (karyogram reconstruction, C-banding) and molecular cytogenetic methods (comparative genome hybridization, in situ hybridization with probes specific for GATA microsatellite motif, the rDNA loci, and the telomeric repeats). The sex chromosomes are microchromosomes in all examined species of the genus Chelodina. In contrast, the sex chromosomes are the 4th largest pair of macrochromosomes in the genera Emydura and Elseya. Their X chromosomes are submetacentric, while their Y chromosomes are metacentric. The chelid Y chromosomes contain a substantial male-specific genomic region with an accumulation of the GATA microsatellite motif, and occasionally, of the rDNA loci and telomeric repeats. Despite morphological differences between sex chromosomes, we conclude that male heterogamety was likely already present in the common ancestor of Chelodina, Emydura and Elseya in the Mesozoic period.
- MeSH
- chromozom X genetika MeSH
- chromozom Y genetika MeSH
- genom * MeSH
- karyotyp MeSH
- mikrosatelitní repetice MeSH
- molekulární evoluce * MeSH
- pohlavní chromozomy genetika MeSH
- procesy určující pohlaví MeSH
- želvy MeSH
- zvířata MeSH
- Check Tag
- mužské pohlaví MeSH
- ženské pohlaví MeSH
- zvířata MeSH
- Publikační typ
- časopisecké články MeSH
- práce podpořená grantem MeSH
Squamate reptiles are considered to exhibit indeterminate growth. Nevertheless, current literature disputes the available definitions of this growth type, presents new theoretical models, and questions its universality in cold-blooded vertebrates. We have followed up on our previous research employing micro-CT to explore growth plate cartilage (GPC) in the epiphysis of long bones, which is responsible for longitudinal skeletal growth by the endochondral ossification process. We focused on numerous and highly diversified group of the Iguania clade comprising Acrodonta (agamas and chameleons) and Pleurodonta ("iguanas"). We recorded the absence of GPC in most of the examined adult Pleurodonta specimens and interpret it as an irreversible arrest of skeletal growth. This finding clearly rejects the universality of indeterminate growth in lizards. On the other hand, we found apparent GPC preservation in most of the adult specimens belonging to Acrodonta. This suggests a preserved ability to continue body growth throughout most of their life. We discuss the uncovered disparity between Acrodonta and Pleurodonta and emphasize the importance of GPC degradation timing.
- MeSH
- chrupavka * diagnostické zobrazování růst a vývoj MeSH
- fylogeneze * MeSH
- ještěři metabolismus MeSH
- rentgenová mikrotomografie * MeSH
- růstová ploténka * diagnostické zobrazování růst a vývoj MeSH
- zvířata MeSH
- Check Tag
- zvířata MeSH
- Publikační typ
- časopisecké články MeSH
- práce podpořená grantem MeSH
Chameleons are well-known, highly distinctive lizards characterized by unique morphological and physiological traits, but their karyotypes and sex determination system have remained poorly studied. We studied karyotypes in six species of Madagascan chameleons of the genus Furcifer by classical (conventional stain, C-banding) and molecular (comparative genomic hybridization, in situ hybridization with rDNA, microsatellite, and telomeric sequences) cytogenetic approaches. In contrast to most sauropsid lineages, the chameleons of the genus Furcifer show chromosomal variability even among closely related species, with diploid chromosome numbers varying from 2n = 22 to 2n = 28. We identified female heterogamety with cytogenetically distinct Z and W sex chromosomes in all studied species. Notably, multiple neo-sex chromosomes in the form Z1Z1Z2Z2/Z1Z2W were uncovered in four species of the genus (F. bifidus, F. verrucosus, F. willsii, and previously studied F. pardalis). Phylogenetic distribution and morphology of sex chromosomes suggest that multiple sex chromosomes, which are generally very rare among vertebrates with female heterogamety, possibly evolved several times within the genus Furcifer. Although acrodontan lizards (chameleons and dragon lizards) demonstrate otherwise notable variability in sex determination, it seems that female heterogamety with differentiated sex chromosomes remained stable in the chameleons of the genus Furcifer for about 30 million years.
- MeSH
- fylogeneze * MeSH
- ještěři klasifikace genetika MeSH
- karyotyp * MeSH
- molekulární evoluce * MeSH
- pohlavní chromozomy * MeSH
- procesy určující pohlaví * MeSH
- zvířata MeSH
- Check Tag
- mužské pohlaví MeSH
- ženské pohlaví MeSH
- zvířata MeSH
- Publikační typ
- časopisecké články MeSH
- práce podpořená grantem MeSH
Sex determination in varanids, Gila monsters, beaded lizards, and other anguimorphan lizards is still poorly understood. Sex chromosomes were reported only in a few species based solely on cytogenetics, which precluded assessment of their homology. We uncovered Z-chromosome-specific genes in varanids from their transcriptomes. Comparison of differences in gene copy numbers between sexes across anguimorphan lizards and outgroups revealed that homologous differentiated ZZ/ZW sex chromosomes are present in Gila monsters, beaded lizards, alligator lizards, and a wide phylogenetic spectrum of varanids. However, these sex chromosomes are not homologous to those known in other amniotes. We conclude that differentiated sex chromosomes were already present in the common ancestor of Anguimorpha living in the early Cretaceous or even in the Jurassic Period, 115-180 Ma, placing anguimorphan sex chromosomes among the oldest known in vertebrates. The analysis of transcriptomes of Komodo dragon (Varanus komodoensis) showed that the expression levels of genes linked to anguimorphan sex chromosomes are not balanced between sexes. Besides expanding our knowledge on vertebrate sex chromosome evolution, our study has important practical relevance for breeding and ecological studies. We introduce the first, widely applicable technique of molecular sexing in varanids, Gila monsters, and beaded lizards, where reliable determination of sex based on external morphology is dubious even in adults.
Despite their long history with the basal split dating back to the Eocene, all species of monitor lizards (family Varanidae) studied so far share the same chromosome number of 2n = 40. However, there are differences in the morphology of the macrochromosome pairs 5-8. Further, sex determination, which revealed ZZ/ZW sex microchromosomes, was studied only in a few varanid species and only with techniques that did not test their homology. The aim of this study was to (i) test if cryptic interchromosomal rearrangements of larger chromosomal blocks occurred during the karyotype evolution of this group, (ii) contribute to the reconstruction of the varanid ancestral karyotype, and (iii) test homology of sex chromosomes among varanids. We investigated these issues by hybridizing flow sorted chromosome paints from Varanus komodoensis to metaphases of nine species of monitor lizards. The results show that differences in the morphology of the chromosome pairs 5-8 can be attributed to intrachromosomal rearrangements, which led to transitions between acrocentric and metacentric chromosomes in both directions. We also documented the first case of spontaneous triploidy among varanids in Varanus albigularis. The triploid individual was fully grown, which demonstrates that polyploidization is compatible with life in this lineage. We found that the W chromosome differs between species in size and heterochromatin content. The varanid Z chromosome is clearly conserved in all the analyzed species. Varanids, in addition to iguanas, caenophidian snakes, and lacertid lizards, are another squamate group with highly conserved sex chromosomes over a long evolutionary time.
- MeSH
- heterochromatin genetika MeSH
- ještěři genetika MeSH
- karyotyp MeSH
- karyotypizace metody MeSH
- molekulární evoluce MeSH
- pohlavní chromozomy genetika MeSH
- zvířata MeSH
- Check Tag
- zvířata MeSH
- Publikační typ
- časopisecké články MeSH
- práce podpořená grantem MeSH
The reconstruction of the evolutionary dynamics of karyotypes and sex determining systems in squamate reptiles is precluded by the lack of data in many groups including most chameleons (Squamata: Acrodonta: Chamaeleonidae). We performed cytogenetic analysis in 16 species of chameleons from 8 genera covering the phylogenetic diversity of the family and also phylogenetic reconstruction of karyotype evolution in this group. In comparison to other squamates, chameleons demonstrate rather variable karyotypes, differing in chromosome number, morphology and presence of interstitial telomeric signal (ITS). On the other hand, the location of rDNA is quite conserved among chameleon species. Phylogenetic analysis combining our new results and previously published data tentatively suggests that the ancestral chromosome number for chameleons is 2n = 36, which is the same as assumed for other lineages of the clade Iguania, i.e., agamids and iguanas. In general, we observed a tendency for the reduction of chromosome number during the evolution of chameleons, however, in Rieppeleon brevicaudatus, we uncovered a chromosome number of 2n = 62, very unusual among squamates, originating from a number of chromosome splits. Despite the presence of the highly differentiated ZZ/ZW sex chromosomes in the genus Furcifer, we did not detect any unequivocal sexual differences in the karyotypes of any other studied species of chameleons tested using differential staining and comparative genomic hybridization, suggesting that sex chromosomes in most chameleons are only poorly differentiated.
- Publikační typ
- časopisecké články MeSH
The Komodo dragon (Varanus komodoensis) is the largest lizard in the world. Surprisingly, it has not yet been cytogenetically examined. Here, we present the very first description of its karyotype and sex chromosomes. The karyotype consists of 2n = 40 chromosomes, 16 macrochromosomes and 24 microchromosomes. Although the chromosome number is constant for all species of monitor lizards (family Varanidae) with the currently reported karyotype, variability in the morphology of the macrochromosomes has been previously documented within the group. We uncovered highly differentiated ZZ/ZW sex microchromosomes with a heterochromatic W chromosome in the Komodo dragon. Sex chromosomes have so far only been described in a few species of varanids including V. varius, the sister species to Komodo dragon, whose W chromosome is notably larger than that of the Komodo dragon. Accumulations of several microsatellite sequences in the W chromosome have recently been detected in 3 species of monitor lizards; however, these accumulations are absent from the W chromosome of the Komodo dragon. In conclusion, although varanids are rather conservative in karyotypes, their W chromosomes exhibit substantial variability at the sequence level, adding further evidence that degenerated sex chromosomes may represent the most dynamic genome part.
Blood cell morphology and count are not uniform across species. Recently, between-species comparisons revealed that the size of red blood cells is associated with body size in some lizard taxa, and this finding was interpreted in the context of the metabolic theory. In the present study, we examined the numbers and the size of blood cells in 2 species of monitor lizards, the mangrove-dwelling monitor (Varanus indicus) and the savannah monitor (V. exanthematicus), and we compared these traits in individuals of different body size. The results revealed that during the course of ontogeny, the size of red blood cells increases with body mass. Because the mass-specific metabolic rate decreases with body size and the cell volume-to-surface ratio decreases with the cell size, changes in the erythrocyte size might be the result of oxygen transport adjustment.
- MeSH
- bazální metabolismus fyziologie MeSH
- druhová specificita MeSH
- erytrocyty cytologie fyziologie MeSH
- ještěři anatomie a histologie klasifikace fyziologie MeSH
- krevní buňky cytologie fyziologie MeSH
- leukocyty cytologie fyziologie MeSH
- spotřeba kyslíku fyziologie MeSH
- velikost buňky * MeSH
- velikost těla * MeSH
- zvířata MeSH
- Check Tag
- mužské pohlaví MeSH
- ženské pohlaví MeSH
- zvířata MeSH
- Publikační typ
- časopisecké články MeSH
- práce podpořená grantem MeSH
- srovnávací studie MeSH
Monitor lizards belong to the largest and the most sexually dimorphic lizards in terms of size, making this group an ideal model for studies analyzing ontogenetic causes of sexual dimorphism. Understanding of these ontogenetic factors is essential to the current discussion concerning patterns of sexual dimorphism in animals. We examined the ontogenetic trajectories of body weight and snout-vent length to analyze the emergence of sexual size dimorphism. Experimental animals were 22 males and 13 females of mangrove-dwelling monitors (Varanus indicus) hatched at the Prague Zoo. They were regularly weighed and measured up to the age of 33-40 months, and subsequently sexed by ultrasonographic imaging. The logistic growth equation was used to describe and analyze the observed growth patterns. Our results confirm considerable sexual size dimorphism in the mangrove monitor. The mean asymptotic body weight of males was nearly three times higher than that of females. As the body size of male and female hatchlings is almost equal, and the growth rate parameter (K) of the logistic growth equation as well as the absolute growth rate up to the age of 12 months do not differ between the sexes, size differences between fully grown males and females should be attributed to timing of the postnatal growth. Males continue to grow several months after they reach the age when the growth of females is already reduced. Therefore, the sexual size dimorphism emerges and sharply increases at this period.
- MeSH
- ještěři růst a vývoj MeSH
- pohlavní dimorfismus MeSH
- stárnutí fyziologie MeSH
- velikost těla fyziologie MeSH
- zvířata MeSH
- Check Tag
- mužské pohlaví MeSH
- ženské pohlaví MeSH
- zvířata MeSH
- Publikační typ
- časopisecké články MeSH
- práce podpořená grantem MeSH
