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Chromosome pairing in inter-racial hybrids of the house musk shrew (Suncus murinus, Insectivora, Soricidae).

Two chromosome races of the house shrew Suncus murinus that differ from each other for five Robertsonian translocations (8.17, 9.13, 10.12, 11.16, and 14.15), heterochromatic insertions in chromosomes 7 and X, and multiple rearrangements in the Y chromosome were crossed and then intercrossed in captivity to produce a hybrid stock. Electron-microscopic analysis of synaptonemal complexes in fertile and sterile hybrid males was carried out. Meiosis in sterile males did not progress beyond pachytene and was severely disrupted. Meiotic arrest was not determined by structural heterozygosity: heterozygotes for all variant chromosomes distinguishing two parental races were found in both sterile and fertile male hybrids. Fertile hybrids demonstrated an orderly pairing of all chromosomes. In heterozygotes for Robertsonian fusions, completely paired trivalents were formed between the Robertsonian metacentrics and homologous acrocentrics. In heterozygotes for chromosome 7, bivalents with a small buckle were observed in a small fraction of pachytene cells. No differences were found in the morphology and pairing pattern of sex bivalents, composed of the X and Y chromosomes derived from the same or different parental races. Univalents, multivalents, and associations between X and Y chromosomes and autosomal trivalents, as well as associations of autosomal trivalents with each other, were observed in a small fraction of the pachytene cells of fertile males. Our results indicate that the system controlling male sterility in interracial hybrids of S. murinus is of genic rather than of chromosomal type.

Animals↗

Metal to insulator transitions in clusters.

The strict criterion for metallicity, a finite density of states (DOS) at the Fermi energy (E(F)), cannot be applied to clusters because energy levels are always discrete in a system of finite size. We propose an alternative definition whereby clusters can be considered metallic when the gap between occupied and unoccupied states at E(F) is consistently smaller than or equal to the Kubo band gap delta. We use the experimental findings of photoelectron spectroscopy of anionic clusters to analyze band gaps of various cluster families. Monovalent clusters (alkali and noble metals) grossly follow the shell structure pattern, producing band gaps smaller than delta for most cluster sizes, with some exceptional sizes exhibiting electronic shell closure or symmetry-induced band gaps. Among the bivalent metals, only mercury shows consistent band gap closure with increasing cluster size, that is a simple insulator-metal transition. Other bivalent elements such as Zn and Mg exhibit a much more complicated behavior. We also briefly discuss complex cluster families such as aluminum and transition metals.

Aluminum↗

Meiotic chromosome pairing in fetal oocytes of trisomy 21 human females.

The influence of trisomy on meiotic chromosome association and synapsis was studied in oocytes of two trisomy 21 fetuses. The patterns of association of the three chromosomes 21 were determined by analysis of late zygotene to early diplotene fetal oocytes after immunofluorescent staining of synaptonemal complexes. The identity of chromosome 21 was confirmed using FISH with either a whole chromosome 21 paint or an alpha-satellite DNA repeat probe. In both fetuses, a wide variety of configurations was present at pachytene. The most common configurations were a trivalent (35.5% and 51.6% of analyzable cells) and a bivalent plus univalent (62.9% and 45.2%). These different frequencies between the fetuses were not significant. Trivalents showed either triple synapsis or double synapsis with pairing-partner switches. The extent of triple synapsis varied from a short segment, either terminal or interstitial, to the whole chromosome length. Through use of immunofluorescent staining of the centromeres, we identified novel types of abnormal chromosome behavior in trisomy 21 fetal oocytes. Thus, we found that 6/41 trivalents had one of the chromosomes associated "out of register," i.e., in a nonhomologous fashion, with its two homologs. Likewise, we found three cells with bivalent plus univalent configurations, in which the univalent showed self-synapsis. The presence of three copies of chromosome 21 therefore results not only in the formation of complex and highly variable synaptic associations but also causes a significant increase in the occurrence of nonhomologous synapsis in human fetal oocytes.

Abortion, Induced↗

Meiotic mutations in rye Secale cereale L.

Spontaneous meiotic mutations of winter rye Secale cereale L. (2n = 14) were revealed in inbred F2 progenies, which were obtained by self-pollination of F1 hybrids resulting from crosses of individual plants of cultivar Vyatka or weedy rye with plants of self-fertile inbred lines. The mutations cause partial or complete sterility, and are maintained in heterozygote condition. Six types of mutations were distinguished as the result of cytological analysis of meiosis and genetic analysis. (1) Plants with nonallelic asynaptic mutations sy1 and sy9 lacked bivalents in 96.8 and 67.0% metaphase I cells, respectively, formed only axial elements but not the mature synaptonemal complex (SC), and had defects in telomere clustering in early prophase I. (2) Weak asynaptic mutant sy3 showed incomplete synapsis at the start of SC degradation at diplotene and lower chiasma number; yet only 2% meiocytes lacked bivalents in MI. (3) Mutations sy2, sy6, sy7, sy8, sy10, and sy19 caused nonhomologous synapsis; i.e., a varying number of univalents and occasional multivalents were observed in MI, which was preceded by switches of pairing partners and fold-back synapsis at mid-prophase I. (4) Mutation mei6 led to the formation of protrusions and minor branched structures of the SC lateral elements. (5) Allelic mutations mei8 and mei8-10 caused irregular chromatin condensation along the chromosome length in prophase I, which was accompanied by chromosome sticking and fragmentation in MI. (6) Allelic mutations mei5 and mei10 determined chromosome supercondensation, caused the disturbance of meiotic spindle assembly, arrested meiosis at various stages but did not affect formation of the pollen wall, thus arrested meiocytes got covered with the pollen wall. Analysis of double mutants revealed recessive epistatic interactions for some mutations; the epistatic group was sy9 > sy1 > sy3 > sy19. This reflects the sequence of meiotic events controlled by the corresponding genes. The expression of sy2 and sy19 proved to be modified by additional genes. Most meiotic mutations found in rye have analogs in other plants.

Chromosome Pairing↗

Heteromorphic sex chromosome system with an exceptionally large Y chromosome in a catfish Steindachneridion sp. (Pimelodidae).

The chromosomes and banding patterns of Steindachneridion sp., a large catfish (Pimelodidae), endemic to the Iguaçu River, Brazil, were analyzed using conventional (C-, G-banding) and restriction enzyme banding methods. The same diploid number (2n = 56) as in other members of the genus and the family was found but the karyotype displayed an XX/XY sex chromosome system. The X chromosome was the smallest submetacentric, while the Y was the largest chromosome in the karyotype. Meiotic analysis showed 27 autosomal bivalents plus one heteromorphic XY bivalent during spermatogenesis. Sex chromosomes had no particular pattern after C-banding but G- and restriction enzyme bandings showed specific banding characteristics. The present finding represents the first report of a well-differentiated and uncommon sex chromosome system in the catfish family Pimelodidae.

Animals↗

The chromosomes of cockerels (Gallus domesticus) during meiosis.

The chromosomes in spermatogenic cells of 10 cockerels were examined. Prior to removing the testes, the birds had been injected with either 2.5 mg vinblastine sulfate or 5 mg colchicine, or were untreated. A cell suspension was made from small pieces of testis and slides were prepared using an air-drying technique. Pretreatments of the birds increased the numbers of spermatogonial and secondary spermatocyte cells at metaphase and rendered them more amenable to analysis. The modal number of macrochromosomes or bivalents was observed in 90.2% of spermatogonia, 98% of primary spermatocytes at diakinesis, and 94.2% of secondary spermatocytes. Polyploid cells at all stages were thought to be largely artifacts of the procedure. Aneuploidy was rare, being found in only 1% of secondary spermatocytes. It is estimated that between 56 and 66 chiasmata occur in the 39 bivalents; of these, an average of 28 are seen in the six macrobivalents. Average numbers were 8.1, 5.9, 4.6, 3.6, 2.4, and 3.5 for macrobivalents 1 through 5 and the Z respectively. The numbers of chiasmata differed significantly among birds but were not differentially affected by pretreatment of the birds.

Animals↗

Nonhomologous associations of C-heterochromatin at human male meitoic prophase.

In human male meiotic prophase, nonhomologous pairings and connections between C-heterochromatic regions were demonstrated by C-banding and G-11 staining. Approximately 40% of the pachytene stages exhibited bivalent associations at regions of C-heterochromatin. Nonhomologous associations were seen between all possible morphological types of bivalents. Evidence is submitted suggesting that nonhomologous associations between regions of C-heterochromatin are a fairly common event in many organisms, including man, and the possible roles of this phenomenon in meiosis are discussed.

Azure Stains↗

Sex vesicle-associated nucleolar organizers in mouse spermatocytes: localization, structure, and function.

Selective silver staining demonstrated that autosomal bivalents containing transcriptively active nucleolar organizers associated with the sex vesicle during pachytene of mouse spermatocytes. Later in pachytene, the nucleolar organizers covered the portion of the sex vesicle furthest from the attachment to the nuclear envelope. Hybridization in situ revealed the presence of rDNA in the silver-positive material. The nucleolus, formed from an autosomal bivalent, exhibited a large fibrillar center surrounded by an electron-opaque fibrillar zone. The nucleolar association with the sex vesicle was studied at early, middle, and late pachytene by hybridization in situ, NOR silver staining, and electron microscopy. These observations enabled us to further define the relationships of the nucleolar components with the X-Y pair.

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↗

Mid-pachytene chromomere maps of human autosomes.

Using a method of chromosome preparation which yielded pachytene spermatocytes with exceptionally well spread bivalents, we undertook detailed chromomere analysis of each of the 22 autosomal bivalents stained with orcein or quinacrine. A maximum of 442 chromomeres were recognized and mapped at mid-pachytene. The chromomere maps of pachytene chromosomes presented here corresponded well with the high resolution banding maps of somatic chromosomes at the 850 band stage described in the International System for Human Cytogenetic Nomenclature (ISCN, 1981). Our observations of other features of the pachytene chromosomes, such as the parameters, also agree well with previous reports.

Aged↗

Nucleoli, micronucleoli, and nucleolus-like structures in human oocytes at meiotic prophase I studied by the silver-NOR technique.

Meiotic nuclei preparations obtained from human fetal ovaries were studied with the silver-NOR technique. At leptotene, the NOR's were located at the periphery of the nucleoli. The mean number of NOR's per nucleus was 6.4. Moreover, 2-12 micronucleoli, each containing an Ag-positive zone, were observed. At pachytene, each nucleolus was divided into two zones, one of which was Ag-positive. The mean number of Ag-positive zones was 2.1 per nucleus. The subterminal region of the short arm of nucleolar bivalents was in contact with the argyrophilic nucleolar zone. In most cells, micronucleoli were dispersed between the bivalents. A single rounded body which stained heavily with silver was consistently observed. At early diplotene, the NOR's separated, appearing again as distinct rounded Ag-positive structures embedded in the peripheral part of the nucleoli. The silver-stained round body was still visible. Eight to twelve micronucleoli were observed. At a more advanced stage of diplotene, the number of micronucleoli increased, varying from 20 to 50 per nucleus. These observations suggest that the micronucleoli, already visible at the leptotene stage, are the morphological expression of an early transcription of amplified rDNA. The biological significance of the Ag-positive round body remains obscure, but the consistency of its appearance suggests that it has a specific function somehow related to nucleolar proteins.

Cell Nucleolus↗

The meiotic chromosomes of Cebus apella (Cebidae, Platyrrhini).

An analysis of the meiotic chromosomes of Cebus apella (Cebidae, Platyrrhini) has allowed for the visualization of constitutive heterochromatic regions throughout the meiotic cycle and has permitted the identification of four chromosomes in the first and the second divisions. In pachytene nuclei, and in pachytene spreads where synaptonemal complexes were visible, one heterochromatic region was frequently unpaired. No chiasmata were found in heterochromatic regions, and there was no evidence of chiasma terminalization. The role of heterochromatin in homologous pairing and its relationship to chiasma position and number in meiotic configurations were discussed. A comparison was made between bivalent 9 in C. apella and bivalent 21 in man, based on the interspecific homologies between these chromosomes. A similar comparison was made between the Algerian and Asian hedgehogs, where marked differences in the content of constitutive heterochromatin occur in two chromosome pairs.

Animals↗

Chromosome elimination in the Japanese hagfish, Eptatretus burgeri (Agnatha, Cyclostomata).

The modal number of chromosomes in Eptatretus burgeri was 36 in 297 (81.8%) of 363 somatic cells, 52 in 13 (30.2%) of 43 spermatogonia, and 25 (46.8%) or 26 (40.3%) in 162 of 186 first spermatocytes. The relative amount of DNA in a somatic cell to that in a spermatogonium averaged 79.2%. C-band-positive chromatin was observed along almost the entire length of the associated dumbbell-shaped bivalents and part of two other bivalents in the metaphases of first spermatocytes, but was rarely observed in somatic cells. These results indicate that, in E. burgeri, chromosome elimination takes place during the early stages of cleavage, except for the ancestral germ-line cells.

Animals↗

Sex chromosome-autosome translocations in the leaf-nosed bats, family Phyllostomidae. II. Meiotic analyses of the subfamilies Stenodermatinae and Phyllostominae.

Analyses of meiotic pairing and synaptonemal complexes of the composite sex chromosomes of male phyllostomid bats with X-autosome or X- and Y-autosome translocations were performed using Giemsa and silver staining procedures. Typical mammalian sex vesicles were absent in all species analyzed. Stenodermatine species with X-autosome translocations possessed an open ring and tail configuration of the XY1Y2 trivalent. Species with both X- and Y-autosome translocations possessed a closed ring and tail configuration of the neo-XY bivalent. In both cases, the tail represented the autosomal short arm of the X paired with its homologue, either the Y2 in XY1Y2 species or the autosomal arm of the composite Y in neo-XY species. Autosomal pairing of the composite sex bivalent in neo-XY species replaced an association between the original X and Y in late prophase I. The absence of a sex vesicle, the unusual pairing configurations of the composite sex chromosomes, and the presumed absence of meiotic nondisjunction in these species is discussed in light of current hypotheses of sex chromosome behavior in male gametogenesis in mammals.

Animals↗

Synaptonemal complexes in a subfertile man with a pericentric inversion in chromosome 21. Heterosynapsis without previous homosynapsis.

Analysis of surface-spread synaptonemal complexes of zygotene and pachytene spermatocytes was carried out on a human male carrier of a pericentric inversion of chromosome 21 ascertained after four miscarriages. The synaptic behavior of the bivalent, which could be unambiguously identified by its nonaligned kinetochores, was analyzed. All zygotene and pachytene spermatocytes had 22 linearly paired autosomal bivalents, with apparently normal synaptonemal complexes, and no evidence of a loop configuration in the 50 cells analyzed. According to the XY type (classification of Solari), the cells were distributed across zygotene and pachytene stages, not exclusively in the late pachytene to which adjustment is conventionally thought to be confined. It is suggested that inverted segments heterosynapse at early pachytene, without previous homosynapsis. It is expected that this meiotic process leads to failure of crossing-over, reduces the production of unbalanced gametes, and the risk of recombinant offspring, but can increase the incidence of aneuploidy as a result of nondisjunction during meiosis I (a frequent cause of pregnancy wastage).

Adult↗

Precise in situ localization of NCAM, ETS1, and D11S29 on human meiotic chromosomes.

In order to sublocalize NCAM, ETS1, and the anonymous DNA fragment D11S29 within 11q23, in situ hybridization was performed on pachytene bivalents. Analysis of the grain distribution within the band 11q23 indicated that the chromosomal sublocalization of both NCAM and D11S29 was in 11q23.1, whereas ETS1 was found to be localized in 11q23.3. These results clearly demonstrate the usefulness of in situ hybridization applied to pachytene bivalents to obtain accurate gene sublocalization.

Chromosome Mapping↗

Trivalent behavior during prophase I in male mice heterozygous for three Robertsonian translocations: an electron-microscopic study.

A synaptonemal complex (SC) analysis was carried out in male mice heterozygous (CHT/+) for three Robertsonian translocations. All pachytene preparations studied showed the presence of three trivalents. At early pachytene, the nonhomologous centromeric regions of the acrocentric chromosomes were unpaired. Heterosynapsis subsequently took place with complete pairing of the trivalents. Association between one of the three trivalents and the sex vesicle was observed in 30.4% of the nuclei. Association between the unpaired regions of two trivalents was present in 14.4% of the cells, suggesting that the relationship between unpaired regions of structural rearrangements and the X-Y bivalent may simply reflect the tendency of unpaired regions to establish end-to-end associations or heterosynapses among them, which are usually resolved during the pachytene stage of prophase I. Since the sex bivalent always has unpaired regions, these associations often affect the sex chromosomes.

Animals↗

Heterosynapsis and axial equalization of the sex chromosomes of the northern bobwhite quail.

The pairing behavior of the Z and W chromosomes in the female northern bobwhite quail (Colinus virginianus) was analyzed by electron microscopy of silver-stained synaptonemal complexes (SCs). After autosomal pairing was completed, synapsis of the sex chromosomes initiated at the short-arm end of the W chromosome and one end of the Z chromosome. Synapsis then progressed unidirectionally, producing a sex bivalent in which the entire length of the W axis was paired with an equivalent length of the Z axis. Progressive contraction and asymmetrical twisting of the Z axis ultimately resulted in a fully paired configuration with aligned axial ends. Further contraction of the Z axis reduced the extent of asymmetrical twisting such that only the nonaligned centromeric regions distinguished the SC of the ZW bivalent from SCs of similar-sized autosomes in late-pachytene nuclei. Quantitative analyses indicated that the length of the Z axis shortened significantly during the adjustment process, whereas no significant difference occurred in the length of the W axis. The nonalignment of the centromeric regions during transitional stages of ZW synapsis indicates that direct heterosynapsis of nonhomologous segments, followed by axial equalization of the length inequality, is responsible for the length adjustment during synapsis in the sex chromosomes of the bobwhite quail.

Animals↗