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PubMed1984

The morphological sequence of XY pairing in the Norway rat Rattus norvegicus.

The sequence of XY pairing at meiotic prophase in the Norway rat, Rattus norvegicus, has been studied in spread preparations of spermatocytes obtained from pubertal males. As in most mammals, sex chromosome pairing is delayed in relation to that of the autosomes. At one stage in pachytene, the Y is fully paired in synaptonemal complex association with about one-third of the X. Observation in spread preparations at pachytene and diplotene and in air-dried metaphase I preparations indicates that the long arm of the Y pairs with the short arm of the X. Pairing of the Y with both ends of the X is seen in about 4% of pachytene spermatocytes. The possibility that XY pairing in the rat may be nonhomologous (Ashley 1983) is considered, and the view is expressed that the XY synaptonemal complex may be incomplete in fine structural detail, thus not providing for the effective pairing required in true reciprocal recombination. The same mechanism that excludes crossing over from heterochromatic regions of autosomes may also operate to minimize or prevent crossing over in the sex pair of mammals.

Animals↗
PubMed1986

Ultrastructural analysis of meiotic recombination and chiasma formation.

Meiotic recombination appears to be mediated by recombination nodules, small electron dense spheres, associated with the central region of the synaptonemal complex. These structures are a prerequisite for, not the result of crossing over and they undergo complex structural, numerical and distributional changes before the final distribution of crossovers is achieved. In the organisms studied the nodules are placed randomly along and among the bivalents at zygotene. Subsequently, the distribution is modified to insure that each bivalent and generally each bivalent arm attain a nodule and that close spacing of nodules is minimized. Analysis of the number and location of recombination nodules at pachytene provides knowledge of the number and physical distribution of crossovers in the individual bivalents. Chiasmata originate from recombination nodules. At early diplotene the chiasma consists of a retained segment of synaptonemal complex containing a derivative of a recombination nodule. At late meiotic prophase the segment is eliminated after the formation of a chromatin chiasma.

Animals↗
PubMed1976

Ultrastructural study on the meiotic prophase nucleus of rat oocytes.

Rat oocytes in the meiotic prophase are studied by means of classical techniques of electron microscopy, preferential staining methods for DNA and RNA and specific enzymatic hydrolysis. The axial cores in leptotene and the lateral arms in the pachytene synaptonemal complex are composed by fibrils that keep a positive contrast after the application of the ethylenediaminetetraacetic acid staining method. They disappear with RNAse treatment, which reveals the presence of chromatin fibrils in the zone occupied by the cores. Preferential staining for DNA corroborates this evidence. Medial arm and lateral-medial fibrils are formed by ribonucleoproteic filaments that form bridges between pairing homologues in the zygotene. In the advanced pachytene stage, the RNA becomes scarce in these structures. No DNA can be detected either in the lateral-medial fibrils or in the medial arm. During diplotene the synaptonemal complex loses its individually and the synaptic space becomes wider and irregular. At the same time, loss of chromatin and a large increase of RNA-containing particles occur. These processes lead to the typical interphasic arrangement of nuclear components seen in the dictyate stage.

Animals↗
PubMed1990

The HOP1 gene encodes a meiosis-specific component of yeast chromosomes.

The HOP1 gene in Saccharomyces cerevisiae is important for meiotic chromosomal pairing, because hop1 diploids fail to form synaptonemal complex during meiosis and are defective in crossing over between, but not within, chromosomes. We demonstrate here that the HOP1 gene is transcriptionally regulated during sporulation and that the HOP1 protein is situated along the lengths of meiotic chromosomes. Furthermore, the HOP1 protein contains a Cys2/Cys2 zinc finger motif. A mutation within this motif that changes a cysteine to serine results in the hop1 phenotype, consistent with the possibility that the HOP1 gene product acts in chromosome synapsis by directly interacting with DNA. These observations demonstrate that HOP1 encodes a component of meiotic chromosomes, perhaps serving as a constituent of the synaptonemal complex.

Amino Acid Sequence↗
PubMed1987

Age-related changes in the meiotic chromosomes of the nematode Caenorhabditis elegans.

Comparison of pachytene karyotypes from old and young wild-type hermaphrodites and males and the mutant him-5 were made following three-dimensional reconstruction of serial ultrathin sections. Age-related changes included: (1) differential condensation of chromatin with increased variance in length of chromosomes; and (2) increased variation in nuclear and nucleolar volume along with increased density of the nucleoplasm. Synaptonemal complex (SC) fine structure was not altered in the nuclei from older specimens. Attachment of only one end of the SC to the nuclear envelope (NE), common to all nematodes, was present at all ages in the wild-type hermaphrodite and male, however, clustering of the SC ends was present in nuclei from older him-5 hermaphrodites. Condensed chromatin along the SC formed a continuous mass except in those small regions where the chromatin had a granular appearance and was decondensed. Such regions, termed "Disjunction Regulator Regions" (DRR), have been implicated in the regulation of X-chromosome segregation (Goldstein, P., The synaptonemal complexes of Caenorhabditis elegans: Pachytene karyotype analysis of the Dp 1 mutant and disjunction regulator regions. Chromosoma, 93 (1985) 177-182). In the present study, it was observed that the number of DRRs in the nucleus change with aging. In the wild-type hermaphrodite and male, the rate of X-chromosome non-disjunction increases with age which correlates with a decrease in the number of DRRs to the point where they are absent in older males. In him-5, the DRRs increase in number with advanced age, which correlates with an observed decrease in the rate of X-chromosome non-disjunction.

Animals↗
PubMed1977

Extensive pairing of the XY bivalent in mouse spermatocytes as visualized by whole-mount electron microscopy.

Autosomes and sex chromosomes of mouse spermatocytes were examined during zygotene, pachytene, and diplotene by a whole-mount electron-microscope technique after cell dispersion in a detergent solution (Nonidet-P40). Zygotene, pachytene, and diplotene stages can be adequately identified in the preparations. Thus, asynchronous side-by-side pairing of homologous autosomes, some of them displaying attached nucleoli, defines zygotene. Pachytene is identified by complete pairing of homologues. Diplotene is characterized by disjunction of bivalents (autosomes and sex chromosomes), lack of autosomal-attached nucleoli, divergent expansions observed at lateral element endings of disassembled synaptonemal complexes, end-to-end association of the XY pair and well defined outward deformations ('bulges') along sex chromosomal axial cores. X and Y chromosomes display at pachytene an extensive side-by-side pairing segment which decreases in length as meiotic prophase advances. Each sex chromosomal axial core appears double and is formed by close apposition of 2 nearly parallel elements displayed separately along the entire length of the chromosomal core. This double structural feature suggests that each sex chromosomal axial core is presumably composed of 2 chromatid axial cores, each of which, in turn, constitutes the respective lateral elements of short synaptonemal complexes observed at the unpaired segment.

Animals↗
PubMed1977

Homologous chromosome pairing.

Commonly accepted precepts are challenged: (1) that homologous chromosome pairing is normally mediated by nuclear envelope attachment sites; (2) that crossover site establishment awaits synaptic completion; and (3) that it is the function of the synaptonemal complex to hold homologues in register so that equal crossing over can occur, and perhaps to provide machinery for the crossover process. Although these views may eventually be shown to be true, it is felt that currently available evidence does not warrant their full acceptance, and that alternatives should be considered. As examples of alternatives the following ideas, with some supporting evidence, are suggested: (1) homologous chromosome pairing (in non-haplont organisms) may be accomplished by chance meeting of homologue segments (followed by establishment of invisible, elastic connectors) at congression for a mitotic metaphase (in many cases perhaps the premeiotic mitosis); (2) crossover sites may be established before, during, or immediately following initiation of synapsis; and (3) the synaptonemal complex may somehow function in the crossover process at the inception of its formation, but its complete deployment throughout each normal bivalent may serve some other role, such as mediation of the binding of sister chromatids apparently required for chiasma maintenance until anaphase I.

Chromatids↗
PubMed2002

Close, stable homolog juxtaposition during meiosis in budding yeast is dependent on meiotic recombination, occurs independently of synapsis, and is distinct from DSB-independent pairing contacts.

A site-specific recombination system that probes the relative probabilities that pairs of chromosomal loci collide with one another in living cells of budding yeast was used to explore the relative contributions of pairing, recombination, synaptonemal complex formation, and telomere clustering to the close juxtaposition of homologous chromosome pairs during meiosis. The level of Cre-mediated recombination between a pair of loxP sites located at an allelic position on homologous chromosomes was 13-fold greater than that between a pair of loxP sites located at ectopic positions on nonhomologous chromosomes. Mutations affecting meiotic recombination initiation and the processing of DNA double-strand breaks (DSBs) into single-end invasions (SEIs) reduced the levels of allelic Cre-mediated recombination levels by three- to sixfold. The severity of Cre/loxP phenotypes is presented in contrast to relatively weak DSB-independent pairing defects as assayed using fluorescence in situ hybridization for these mutants. Mutations affecting synaptonemal complex (SC) formation or crossover control gave wild-type levels of allelic Cre-mediated recombination. A delay in attaining maximum levels of allelic Cre-mediated recombination was observed for a mutant defective in telomere clustering. None of the mutants affected ectopic levels of recombination. These data suggest that stable, close homolog juxtaposition in yeast is distinct from pre-DSB pairing interactions, requires both DSB and SEI formation, but does not depend on crossovers or SC.

Alleles↗
PubMed2000

Structural and functional evidence that a B chromosome in the characid fish Astyanax scabripinnis is an isochromosome.

Astyanax scabripinnis possesses a widespread polymorphism for metacentric B chromosomes as large as the largest chromosome pair in the A complement. On the basis of C-banding pattern, it was hypothesized that these B chromosomes are isochromosomes that have arisen by means of centromere misdivision and chromatid nondisjunction. In the present paper we test this hypothesis by analysing (i) the localization of a repetitive DNA sequence on both B chromosome arms, and (ii) synaptonemal complex formation, in order to test the functional homology of both arms. Genomic DNA digested with KpnI and analysed by gel electrophoresis showed fragments in a ladder-like pattern typical of tandemly repetitive DNA. These fragments were cloned and their tandem organization in the genome was confirmed. A 51-bp long consensus sequence, which was AT-rich (59%) and contained a variable region and two imperfect reverse sequences, was obtained. Fluorescence in situ hybridization (FISH) localized this repetitive DNA into noncentromeric constitutive heterochromatin which encompasses the terminal region of some acrocentric chromosomes, the NOR region, and interstitial polymorphic heterochromatin in chromosome 24. Most remarkably, tandem repeats were almost symmetrically placed in the two arms of the B chromosome, with the exception of two additional small clusters proximally located on the slightly longer arm. Synaptonemal complex (SC) analysis showed 26 completely paired SCs in males with 1B. The ring configuration of the B univalent persisting until metaphase I suggests that the two arms formed chiasmata. All these data provided strong support for the hypothesis that the B chromosome is an isochromosome.

Animals↗
PubMed1978

Meiotic effects of DNA-defective cell division cycle mutations of Saccharomyces cerevisiae.

The meiotic effects of several cell division cycle (cdc) mutations of Saccharomyces cerevisiae have been investigated by electron microscopy and by genetic and biochemical methods. Diploid strains homozygous for cdc mutations known to confer defects on vegetative DNA synthesis were subjected to restrictive conditions during meiosis. Electron microscopy revealed that all four mutants were conditionally arrested in meiosis after duplication of the spindle pole bodies but before spindle formation for the first meiotic division. None of these mutants became committed to a recombination or contained synaptonemal complex at the meiotic arrest.--The mutants differed in their ability to undergo premeiotic DNA synthesis under restrictive conditions. Both cdc8 and cdc21, which are defective in the propagation of vegetative DNA synthesis, also failed to undergo premeiotic DNA synthesis. The arrest of these mutants at the stage before meiosis I spindle formation could be attributed to the failure of DNA synthesis because inhibition of synthesis by hydroxyurea also caused arrest at this stage.--Premeiotic DNA synthesis occurred before the arrest of cdc7, which is defective in the initiation of vegetative DNA synthesis, and of cdc2, which synthesizes vegetative DNA but does so defectively. The meiotic arrest of cdc7 homozygotes was partially reversible. Even if further semiconservative DNA replication was inhibited by the addition of hydroxyurea, released cells rapidly underwent commitment to recombination and formation of synaptonemal complexes. The cdc7 homozygote is therefore reversibly arrested in meiosis after DNA replication, whereas vegetative cultures have previously been shown to be defective only in the initation of DNA synthesis.

Cell Cycle↗
PubMed1974

Transcription sites in spread meiotic prophase chromosomes from mouse spermatocytes.

Mouse spermatocytes at pachytene stage have been examined by whole-mount electron microscope techniques complemented with autoradiography as an approach for visualizing their transcriptive activity. Structural elements of meiotic bivalents, such as synaptonemal complexes and chromatin fibers, have been satisfactorily displayed in the total set of autosomal and sexual bivalents in single spermatocytes. Adequate preservation of the entire set of bivalents has provided a basis for recognition of sites where presumptive preribosomal RNA and heterogeneous nuclear RNA species are being transcribed at different segments of autosomal bivalents. Nucleoli attached to the basal knob region where nucleolar organizer cistrons are assumed to be located and ribonucleoprotein fibrils associated with distinct chromatin loops have been recognized. These structural findings have been correlated with display of [(3)H]uridine incorporation sites in thin-section and whole-mount electron microscopy autoradiographic preparations. A low transcriptive activity of the sexual bivalent contrasted with extensive gene expression in autosomal bivalents. Each sex chromosome shows a double axial core. A short region of pairing with a synaptonemal complex joins the two chromosomes at one end. We conclude that variations in the rate of RNA synthesis throughout meiotic prophase stages in the mouse are expressed as fluctuations in the amount and distribution of distinct RNA species at specific segments of the bivalents.

Animals↗
PubMed1982

[Karyosphere and nuclear envelope ultrastructure of oocytes of the common frog Rana temporaria after nuclease treatment and extraction of chromatin and alkali-soluble proteins].

The late oocyte karyosphere of Rana temporaria consists mainly of fibrillar nucleoli and micronucleoli, Fibrillar material containing the pore complexes (pseudomembranes) and modified synaptonemal complexes. Nuclease treatment of the karyosphere reveals regularly repeating bands in the nuclei and micronuclei, the fibrillar material associated with the nuclei being attenuated. After a subsequent treatment with DNase-1, RNase A and 1.5 M Nacl and karyosphere is disintegrated to involve remains of extracted nucleoli and fibrillar material containing pore complexes. 0.025 N NaOH extracted the bulk of the karyosphere and left only the islets if fibrillar material. The nuclear envelope proved to be far more stable to the above treatments. It retained its structure and was loosened partly only after 1.5 M NaCl or 0.025 N NaOH extraction, the pore complexes disappearing only after alkaline extraction. It is assumed that the karyosphere represents a skeletal structure (matrix) of the oocyte nucleus.

Animals↗
PubMed2000

Synaptomenal complex analysis of four breeds of Bos taurus taurus x B. taurus indicus hybrids.

The synaptonemal complex (SC) was analyzed in four F1 hybrids of Bos taurus taurus and B. taurus indicus including Gyr-Simmental (G-S), Nelore Simmental (N-S), Gyr-Holstein-Friesian (G-H) and Nelore-Piemontese (N-P). We analysed the frequency of various types of SC abnormalities and the frequency of cells with SC abnormalities. The results were compared with similar observations made on purebred animals. All the animals studied possessed 29 autosomal and one sex bivalent. The frequency of cells with abnormalities in the hybrids were 28.0% in the N-P, 29.1% in the G-S, 33.3% in the N-S and 40.0% in the G-H. The frequency of cells with abnormalities in the four hybrids was 31.5%; 57.9% of these abnormalities occurred in zygotene and 42.0% occurred in pachytene. The comparisons among the hybrids and among the hybrids and their parental breeds showed that the only significant difference was between Gyr and Gyr-Holstein-Friesian animals. Some aspects of the relationship between the frequency of cells with anomalies and the fertility of hybrids are discussed.

Animals↗
PubMed1997

Meiotic cells monitor the status of the interhomolog recombination complex.

During meiosis, mutations that cause defects at intermediate stages in the recombination process confer arrest at the end of prophase (e.g., pachytene). In yeast, mutations of this type include rad50S, dmc1, rad51, and zip1. Rad50 is likely part of a recombination initiation complex. DMC1, RAD51, and ZIP1 encode two RecA homologs and a synaptonemal complex protein, respectively. We report here the effects of mutations in two other (meiosis-specific) genes, RED1 and MEK1/MRE4, that encode a chromosome structure component and a protein kinase, respectively. A red1 or mek1/mre4 mutation alleviates completely rad50S, dmc1, rad51, and zip1 arrest. Furthermore, the red1 and mek1/mre4 mutations define a unique, previously unrecognized aspect of recombination imposed very early in the process, during DSB formation. Finally, the red1 and mek1/mre4 mutations appear to alleviate prophase arrest directly rather than by eliminating, or permitting bypass of, the rad50S, dmc1, rad51, or zip1 defects. These and other observations suggest that a meiosis-specific regulatory surveillance process monitors the status of the protein/DNA interhomolog recombination machinery as an integral entity, in its proper chromosomal context, and dependent upon its appropriate Red1 and Mek1/Mre4-promoted development. We speculate that a properly developed recombination complex emits an inhibitory signal to delay progression of meiotic cells out of prophase until or unless the recombination process has progressed, at least past certain critical steps, and perhaps to completion.

Blotting, Southern↗
PubMed1992

Chromosomal synapsis and the meiotic process in male mesquite lizards, Sceloporus grammicus complex.

Meiosis in males of the F5 cytotype of Sceloporus grammicus was examined through the analysis of synaptonemal complexes (SCs), diakinetic (metaphase I) nuclei, and secondary spermatocytes (metaphase II configurations). These data allowed the establishment of criteria for substaging of zygonema and pachynema, morphological characterization of the SC complement, and comparison of the orientation and segregation of the autosomes and sex chromosomes. The analysis of nuclei from all stages of meiotic prophase I (leptonema through diakinesis) provided a useful means of partitioning the temporal sequence of early meiotic events. Three substages of zygonema (Z1-Z3) were established, based on the extent of synapsis of the microchromosomal and macrochromosomal elements. Synaptic initiation of the autosomes and sex chromosomes was synchronous. Two patterns of macrochromosomal synapsis were observed. Whereas synapsis of the biarmed elements was biterminal (i.e., progressing from both ends of the homologs), synapsis of the acrocentric elements was uniterminal involving only the distal (noncentromeric) ends of the homologs. Unique sex-chromosomal characteristics were not observed in S. grammicus and, therefore, the substaging of pachynema was based on subjective criteria. Examination of diakinesis--metaphase I and metaphase II configurations indicated low levels of diakinetic irregularities with balanced segregation of the autosomal bivalents and the sex-chromosomal trivalent.

Animals↗
PubMed2003

Chromatin dynamics in the male meiotic prophase.

During the male meiotic prophase in mouse and man, pairing and recombination of homologous chromosomes is accompanied by changes in chromatin structure. In this review, the dynamics of assembly and disassembly of the chromatin-associated complexes that mediate sister chromatid cohesion (cohesin) and maintain chromosome pairing (the synaptonemal complex) are described. Special features of the meiotic S phase are discussed, and also the dynamics of several key players that act together after the S phase at sites of meiotic double-strand break DNA repair. Current knowledge on histone modifications that occur during the male meiotic prophase is discussed, with special attention for the inactive chromatin of the X and Y chromosomes that constitutes the sex body. Finally, it is discussed that in the future, it will be possible to view the true chromatin dynamics during male meiosis in time, in living cells, through analysis of fluorescent-tagged proteins expressed in transgenic mice, using advanced fluorescent microscopy techniques.

Animals↗
PubMed1992

Synapsis, recombination, and meiotic segregation in the mesquite lizard, Sceloporus grammicus, complex. II. Fission heteromorphism of the FM2 cytotype and evolution of chromosome 2.

Somatic and meiotic chromosomal and synaptonemal complex techniques were used to characterize the chromosomal complement and to study the fission heteromorphism of chromosome 4 in the FM2 cytotype of Sceloporus grammicus. Analysis of silver-stained somatic metaphases revealed that the nucleolar organizer region in this cytotype is located at the distal end of a pair of medium-sized acrocentric chromosomes, rather than on the largest acrocentric chromosomal pair, as previously reported. This condition is hypothesized to be the result of at least two sequential rearrangements. Analysis of surface-spread zygotene and pachytene nuclei indicated that the components of the chromosome 4 trivalent initiated synapsis at their distal telomeric regions. Although synapsis of the fission trivalent was synchronous with that of the homomorphic autosomal pairs, completion of synapsis was delayed in the trivalent. Associations between the fission trivalent and other autosomal or sex-chromosomal elements occurred in approximately one third of the pachytene nuclei examined. Analysis of secondary spermatocytes (metaphase II configurations) revealed low levels of nondisjunction in fission heterozygotes. These analyses indicate that FM2 individuals heterozygous for the fission rearrangement of chromosome 4 suffer no meiotic deficit.

Animals↗