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Light microscopic observations on the behavior of silver-stained trivalents in pachytene cells of Sigmodon fulviventer (Rodentia, Muridae) heterozygous for centric fusion.

Silver-stained pachytene cells of male cotton rats, Sigmodon fulviventer, heterozygous for a naturally occurring Robertsonian fusion, were examined by light microscopy. Silver staining clearly revealed synaptonemal complexes of all autosomal bivalents and of the translocation trivalent, as well as X-Y conformations within the sex vesicle. The synaptonemal complex of the trivalent conforms closely in morphology to that observed by electron microscopy in hybrids of other species. In all cases, the proximal telomeric knobs of the two acrocentric elements were paired and projected in the same direction from the metacentric element. The geometry of trivalent pairing in pachytene may predispose to normal disjunction and, therefore, account for the high fertility observed in S. fulviventer heterozygotes.

Animal Population Groups↗

Robertsonian polymorphism in chromosomes of Oryzomys subflavus (Rodentia, Cricetidae).

Eighty-four specimens of Oryzomys subflavus, collected in the State of Pernambuco, Brazil, were studied. A Robertsonian chromosome polymorphism, characterized by a varying diploid number of 50, 49, 48, and 46, was found. All specimens showed a chromosome arm number of 56. G-banding patterns in somatic cells allowed identification of the chromosome pairs (2, 3, 5, and 7) involved in centric fusion. C-banding revealed the presence of constitutive heterochromatin near the centromere the X chromosome and those of the autosomes. The Y chromosome presented a large heterochromatic block in the distal portion of its long arm.

Animals↗

An assessment of the nature of chromosomal rearrangements in 18 species of Peromyscus (Rodentia: Cricetidae).

G- and C-banded karyotypes from 9 previously unstudied species of Peromyscus are described and compared to the 9 species described in the literature. Additional new data are presented for P. eremicus. A hypothetical evolutionary tree for the 18 species was constructed based on the assumption that, where possible, shared chromosomal rearrangements became established in a common ancestor. A minimum of 60 chromosomal rearrangements (34 heterochromatic additions and 26 pericentric inversions) were needed to construct the most parsimonious tree. Seven of the 18 species have heterochromatin in noncentromeric areas. Three times as many inversions have been identified in the 11 largest pairs as have been observed in the 12 smallest pairs. Peromyscine rodents are characterized by a pattern of chromosomal variation quite distinct from that described for other rodent genera, such as Sigmodon and Mus. Karyotypic orthoselection in rodents appears to be relatively common in closely related species.

Animals↗

Chromosomal variability of sex chromosomes and NOR's in Trichomys apereoides (Rodentia, Echimyidae).

G-banding, C-banding, and silver staining techniques were used to analyze the chromosomes of Trichomys apereoides. In a sample of 22 specimens the diploid number was 2n = 30, with several pairs of biarmed chromosomes and a large metacentric (pair 1) with a secondary constriction, or nucleolar organizing region (NOR), in the short arm. One male had a heteromorphic pair 1, with only one NOR-bearing homolog. C-bands showed constitutive heterochromatin in the centromeric region of almost all autosomes. The X has a large block in the proximal region and an interstitial small band in the long arm. The Y is totally heterochromatic. One heteromorphic female with an increased C-band block in one of the X chromosomes was observed. The Y is a small submetacentric; one male showed a metacentric Y, resulting probably from a pericentric inversion. The sexual bivalent and the synaptonemal complexes of all autosomal bivalents were revealed by silver staining in pachytene cells.

Animals↗

Complex evolution of sex chromosomes in Gerbillidae (Rodentia).

The sex chromosomes of 11 species of Gerbillidae (Meriones tristrami, M. crassus, M. libycus, M. persicus, M. unguiculatus, Gerbillus hesperinus, G. nigeriae, G. gerbillus, G. campestris, Gerbillurus tytonis, and Taterillus gracilis) are reported. A very complex evolution of the X chromosome and, to a lesser degree, of the Y chromosome was observed, including several inversions, translocations with autosomes, and an increase in heterochromatin. The role of constitutive heterochromatin, isolating autosome and gonosome segments in translocations (thus preventing any position effect), is discussed. The sequence of rearrangements affecting the sex chromosomes is considered with a view toward establishing the phylogenetic relationship of the species studied. The tendency to accumulate the same rate type of rearrangement, namely, gonosome-autosome translocations, is another demonstration that chromosomal evolution is not random in a given group of species.

Animals↗

Nucleolar organizer regions in Akodon arviculoides (cricetidae, rodentia): evidence for the activity of rDNA genes in both X chromosomes of females.

Silver-stained cells from 13 specimens of Akodon arviculoides (2n = 14 and 15) were analyzed for the location and frequency of Ag-NOR's. They were located terminally on four pairs of chromosomes, 3q, 4q, 5p, and Xp. In these 13 individuals, the number of Ag-NOR's per cell varied between two and eight, and each chromosome had a characteristic pattern of frequency of NOR's in each individual. Since in A. arviculoides a chromosomal polymorphism resulting from three autosomal pericentric inversions in chromosomes 2, 3, and 5 and a complex rearrangement involving chromosome 1 exist, it was possible to determine the frequency of Ag-staining in each homologue of these pairs in heterozygous individuals. Both X chromosomes had active NOR's in the females. An Ag-NOR was also present on the single X chromosome of the males.

Animals↗

The origin of multiple sex chromosomes in the gerbil Gerbillus gerbillus (Rodentia: Gerbillinae).

The sex chromosomes of the partly sympatric species of gerbils Gerbillus pyramidum and G. gerbillus (Mammalia: Gerbillinae) were investigated by a variety of light- and electron-microscope methods, including DNA replication banding and synaptonemal complex (SC) techniques. The sex-chromosome mechanism of G. pyramidum is of the maleXY:femaleXX type, whereas that of G. gerbillus is of the less common maleXY1Y2:femaleXX system. The results include the demonstration that the X chromosomes of both species are compound. One segment is added to the X chromosome of G. pyramidum, leading to an increase in length from the standard 5% to approximately 7.3%, whereas two different extra segments increase the length of the X chromosome of G. gerbillus to approximately 11% of the length of the haploid genome. In both cases the extra material is autosomal and is also represented in the respective Y chromosomes. Classifying heterochromatin by the variation in staining quality was helpful in elucidating the possible origin of the different chromosome segments, including the pericentromeric regions. Observations on meiotic chromosome pairing and chiasma formation have confirmed the homologies established by band comparisons. The occurrence of chiasmata between the sex chromosomes supports the autosomal origin of the pairing segments. These and other findings have been interpreted in the framework of a multistep evolutionary model. This sequence starts from a hypothetical pair of sex chromosomes, the X element of which amounts to 5% of the haploid genome, and leads through three translocations involving two pairs of autosomes and one pericentric inversion to the most complex situation of this series, manifested in G. gerbillus. The adaptive value, if any, of autosome incorporation into the sex chromosomes repeatedly occurring here is unknown. It is, however, a remarkable fact that in one species, G. gerbillus, the complex sex-chromosome constitution is conserved over vast geographic distances, and in the other, G. pyramidum, the compound X and Y chromosomes withstand change in the face of extreme autosome restructuring.

Animals↗

Unusual C-band patterns in three karyotypically rearranged forms of Scapteromys (Rodentia, Cricetidae) from Brazil.

Chromosome studies of 30 specimens of the rodent Scapteromys collected at nine localities in southern Brazil revealed the occurrence of three karyotypic taxa with 2n = 36 (one locality), 34 (two localities), and 24 (six localities), although all three had 40 autosomal arms (AN). The G-band analysis indicated that this reduction in diploid number was mainly due to Robertsonian translocations which have occurred along a gradient, possibly in two independent evolutive routes. The C-bands occur on one autosomal pair and on the X and Y in the 2n = 36 and 34 forms and on the X and Y chromosomes only in the 2n = 24 taxon. The broad genomic reorganization which has occurred in this genus, in which the chromosomes do not have large amounts of constitutive heterochromatin, argues against the idea that a large amount of constitutive heterochromatin favors chromosome evolution and speciation.

Animals↗

An X1X1X2X2/X1X2Y mechanism of sex determination in a South American rodent, Deltamys kempi (Rodentia, Cricetidae).

Chromosome studies on 14 specimens of Deltamys kempi disclosed six males with 2n = 37, NF = 38, six females with 2n = 38, NF = 38, and two females with 2n = 37, NF = 38. G- and C-band analyses revealed a Y-autosome translocation in the males leading to a multiple chromosome system of sex determination of the type X1X1X2X2/X1X2Y, this being the second case of such a mechanism described in rodents. At meiosis the males presented a trivalent in which C-banding studies showed an alternate orientation of the sex chromosomes due to end-to-end association of the X1 and Y chromosomes, the Y and the X2 being held together by interstitial chiasmata. At metaphase II both n = 17 + Y and n = 18 + X1 are regularly observed. The two females with 2n = 37, NF = 38, are heterozygous for an autosomal centric fusion involving chromosomes 1 and 13. The product of the Y-autosome translocation constitutes the largest element of the karyotype (9.4% of the haploid set); the X1 chromosome amounts to 7.8% of this set, including a large heterochromatic block. When only its euchromatic region is considered, this percentage decreases to 4.6%. From two to seven NORs were observed at the telomeres, with a mean of 4.4 +/- 1.1 per cell.

Animals↗

Chromosomal homoeologies in hamster species of the genus Phodopus (Rodentia, Cricetinae).

The karyotypes of the two hamster species Phodopus sungorus and P. roborovskii were analyzed with several banding techniques and compared. The homoeologies found between all chromosomes led to the proposal of the most likely common ancestral karyotype of these two species. The differences in the karyotypes were found to be a result of seven independent centric fusions, three inversions, and one possible telomeric fusion, as well as changes in the quantity and DNA-base pair composition of the constitutive heterochromatin.

Animals↗

The chromosomes of Micromys minutus (Rodentia, Murinae). II. Pairing pattern of X and Y chromosomes in meiotic prophase.

Both light and electron microscopy were used to study the pairing behavior of the sex chromosomes of the harvest mouse, Micromys minutus, in surface-spread pachytene spermatocytes. The XY pairing pattern is very exceptional in that the site of synaptic initiation is located interstitially in the short arms of the X and the Y, next to their centromeric regions. From this tiny euchromatic site, synapsis proceeds unidirectionally along the homologous heterochromatic short arms of the X and the Y toward the ends of the chromosomes. After pairing of the short arm is concluded, synapsis begins between the nonhomologous long arms of the X and the Y in the immediate vicinity of the centromeres and progresses unidirectionally toward the end of the long arm of the Y. A synaptic complex develops between the constitutive heterochromatin of the long arm of the Y and the euchromatin of the long arm of the X. Analysis of C-banded and distamycin A/DAPI-stained diakineses revealed a trefoil-like XY bivalent, which was interpreted to be the result of an interstitial chiasma occurring in the paired short arms of the X and the Y. A conspicuous, electron-dense body, about 1 micron in diameter, was found closely associated with the centromeres of the X and the Y in numerous pachytene spermatocytes. A review of the literature showed that comparable XY-associated bodies have been found in only eight other mammals to date.

Animals↗

Synaptonemal complex analysis of B-chromosome behavior in meiotic prophase I in the East-Asiatic mouse Apodemus peninsulae (Muridae, Rodentia).

The mitotic and meiotic chromosomes of four male East-Asiatic mice, Apodemus peninsulae, having three to seven chromosomes in addition to the standard karyotype (2n = 48), were investigated. B-chromosomes were represented by medium-sized metacentric and dotlike chromosomes. Mosaicism of bone marrow cells due to a numerical variation of accessory chromosomes was established for the males examined. Capacity of B-chromosomes to form axial elements and synaptonemal complexes in meiotic prophase I was revealed by electron microscopy. The occurrence of univalents of different morphology, bivalents, and multivalents, corresponding to B-chromosomes, was demonstrated. An increase in the number of B-chromosomes was found in spermatocytes at zygotene-pachytene relative to the number in bone marrow cells, which may be evidence of B-chromosome accumulation in the germ cell line of the East-Asiatic mouse.

Animals↗

XY females in Microtus cabrerae (Rodentia, Microtidae): a case of possibly Y-linked sex reversal.

Microtus cabrerae is a species with blocks of heterochromatin on both sex chromosomes. Polymorphism caused by extensive deletions affecting these heterochromatic segments exists in both X and Y chromosomes. Of a total of 22 males and 22 females studied, four females exhibited sex pair dimorphism similar to that observed in males. C-banding techniques and preliminary data regarding inheritance support the hypothesis that these specimens are XY fertile females and that this sex reversion is most likely inherited through the Y chromosome of these females.

Animals↗

Synaptic adjustment in Peromyscus beatae (Rodentia: Cricetidae) heterozygous for interstitial heterochromatin.

Chromosomal pairing and chiasma formation were studied two individuals of Peromyscus beatae heterozygous for the presence of a large block of interstitial heterochromatin. Although the modified chromosome was of medium size, analysis of C-banded diakinetic configurations revealed that it was the homolog of one of the smallest autosomes. Analysis of silver stained synaptonemal complexes indicated that synapsis was either unidirectional from initiation at one set of telomeres or was bidirectional from initiation at both sets of telomeres. Each pattern resulted in characteristic heteromorphic pairing configurations (interstitial asynapsis or terminally positioned unpaired segments) in early pachynema. These configurations underwent synaptic adjustment and, by mid-pachynema, the lateral elements of the polymorphic bivalent either appeared typical of homomorphic bivalents or exhibited regional heteropycnosis in one or both axes. Synaptonemal complex data for Peromyscus and many other mammalian species reflect an apparent need for fully paired, linear bivalents prior to the end of pachynema.

Animals↗

Meiotic behavior of gonosomically variant females of Akodon azarae (Rodentia, Cricetidae).

The meiotic behavior of sex chromosomes has been investigated in variant females of Akodon azarae, both in pachytene oocytes and metaphase I. In somatic cells, these females have a heteromorphic sex pair, in which the minor chromosome has been previously interpreted as a major deletion of the long arm of the X chromosome (dX). After microspreading for synaptonemal complex analysis, pachytene oocytes show two axes of very different lengths (100:17.1), which correspond to the sex chromosomes X and dX. True synapsis is abnormally restricted (43.3%) between these sex chromosomes; on the other hand, self-synapsis of both the X and dX chromosomes is frequent (60%). Single, nonsynapsed axes or axial segments are thickened. Strong chromatin condensation occurs around nonsynapsed axes or axial segments, giving many of these sex pairs an appearance similar to an XY body ("sex vesicle"). The minor gonosome axis differs from that of the Y chromosome of male meiosis, as the former is shorter (relative to the X) and has a different synaptic behavior. In 17 metaphases I from XdX variant females, only heteromorphic, end-to-end joined sex pairs were observed. These variant females differ from the variant females of the wood lemming Myopus schisticolor in several respects, but a similar mechanism seems to be prevalent in other species of the genus Akodon. Self-synapsis of unequal gonosomes in oocytes is assumed as an escape from functional deterioration, following the hypothesis put forward by others.

Animals↗

The sex-determining zinc finger sequences in XY females of Akodon azarae (Rodentia, Cricetidae).

Wild populations of Akodon azarae comprise females with a karyotype indistinguishable from that of males. These individuals were formerly assumed to be Xx, the x being an X chromosome with a deletion of most of its long arm. By using a DNA probe derived from the testis-determining region of the human Y chromosome (comprising a candidate gene for the testis-determining factor, Y-linked zinc finger [ZFY]), we demonstrate that A. azarae gonosomally variant females are XY and not Xx. The ZFY sequences in A. azarae are amplified and located in two different families of EcoRI fragments derived from Y-chromosome DNA. No rearrangement or change in the state of methylation of ZFY or ZFX (X-linked zinc finger) sequences were found in XY females. We propose that sex reversal in A. azarae may be mediated by a gene or genes other than ZFX or ZFY.

Animals↗

The effect of heterochromatin on synapsis of the sex chromosomes of Peromyscus (Rodentia, Cricetidae).

The pairing behavior of the sex chromosomes in male and female individuals representing seven species of Peromyscus was analyzed by electron microscopy of silver-stained zygotene and pachytene configurations. Six species possess submetacentric or metacentric X chromosomes with heterochromatic short arms. Sex-chromosome pairing in these species is initiated during early pachynema at an interstitial position on the X and Y axes. Homologous synapsis then progresses in a unidirectional fashion towards the telomeres of the X short arm and the corresponding arm of the heterochromatic Y chromosome. The distinctive pattern of synaptic initiation allowed a late-synapsing bivalent in fetal oocytes to be tentatively identified as that of the X chromosomes. In contrast to the other species, Peromyscus megalops possesses an acrocentric X chromosome and a very small Y chromosome. Sex-chromosome pairing in this species is initiated at the proximal telomeric region during late zygonema, and then proceeds interstitially towards the distal end of the Y chromosome. These observations suggest that the presence of X short-arm heterochromatin and corresponding Y heterochromatin interferes with late-zygotene alignment of the pairing initiation sites, thereby delaying XY synaptic initiation until early pachynema. The pairing initiation sites are conserved in the vicinity of the X and Y centromeres in Peromyscus, and consequently the addition of heterochromatin during sex-chromosome evolution essentially displaces these sites to an interstitial position.

Animals↗

The synaptic sequence in hydroxyurea-treated spermatocytes of Pitymys duodecimcostatus (Rodentia, Microtidae).

A number of Pitymys duodecimcostatus males from a natural population were treated with hydroxyurea, which produces a gap in the spermatogenic line, to study the synaptic sequence of meiotic prophase. The use of substaging criteria, based on the morphological characteristics of the meiotic cells themselves, gave rise to a classification scheme consistent with the chronological sequence revealed by hydroxyurea treatment, which supports the usefulness of these independent criteria. The characteristics of meiotic prophase in P. duodecimcostatus are also described and discussed.

Animals↗