Inheritance of Chromosome-Length Polymorphisms in Coprinus Cinereus

We have investigated the inheritance of chromosome-length polymorphisms in the basidiomycete Coprinus cinereus. The electrophoretic karyotypes of interfertile strains of C. cinereus are strikingly different, and crosses between strains with different karyotypes yield progeny with chromosomes of new sizes. Repeated backcrossing of a mutant to one parent often stabilizes the mutant chromosome at a unique size; this then becomes a chromosome-length polymorphism marker for that mutant gene. A comparison of mutant strains, their wild-type progenitor, and backcrossed strains revealed that these marker chromosomes are not caused by the initial mutagenic treatment and are found only in progeny of crosses between strains with polymorphic chromosomes. Thus, they are most likely formed by meiotic recombination. For the rad12 gene, the marker chromosome can further recombine to become the size of the homolog of the backcross parent. For the rad3 gene, both ectopic and homologous recombination events are likely involved in the generation of the marker chromosomes. As predicted by a recombination model, a cross to a new wild-type parent can change the size of a mutant marker chromosome. Therefore, changes in chromosome length are a common and prominent feature of the genome of this sexual fungus, and a variety of karyotypes is tolerated by the organism.     

SUPPRESSION OF SEX-RATIO MEIOTIC DRIVE AND THE MAINTENANCE OF Y-CHROMOSOME POLYMORPHISM IN DROSOPHILA

Like several other species of Drosophila, D. quinaria is polymorphic for X-chromosome meiotic drive; matings involving males that carry a “sex-ratio” X chromosome (X^SR) result in the production of strongly female-biased offspring sex ratios (Jaenike 1996). A survey of isofemale lines of D. quinaria from several populations reveals that there is genetic variation for partial suppression of this meiotic drive. Crossing experiments show that there is Y-linked, and probably autosomal, variation for suppression of drive. Y-linked suppressors of X-chromosome drive have now been described in several species of Diptera. I develop a simple model for the maintenance of Y-chromosome polymorphism in species polymorphic for X-linked meiotic drive. One interesting feature of this model is that, if there is a stable Y-chromosome polymorphism, then the equilibrium frequency of the standard and sex-ratio X chromosomes is determined solely by Y-chromosome parameters, not by the fitness effects of the different X chromosomes on their carriers. This model suggests that Y-chromosome polymorphism may be easier to maintain than previously thought, and I hypothesize that karyotypic variation in Y chromosomes will be found to be associated with suppression of sex-ratio meiotic drive in other species of Drosophila.     

The odd-even effect in mitotically unstable B chromosomes in grasshoppers

The odd-even effect, by which B chromosomes are more detrimental in odd numbers, has been reported in plants and animals. In grasshoppers, there are only a few reports of this effect and all were referred to as traits related to the formation of aberrant meiotic products (AMPs). Here we review the existing information about B chromosome effects on AMPs, chiasma frequency and the number of active nucleolus organizer regions (NORs) per cell. Polysomy for A chromosomes and B chromosomes are two kinds of chromosome polymorphism frequently found in grasshoppers. In some aspects, e.g. meiotic behaviour and mitotic instability leading to individual mosaicism (in the case of mitotically unstable Bs), polysomic As show similar characteristics to B chromosomes. In fact, polysomy is regarded as one of the main mechanisms for B chromosome origin. Here we review some features of meiotic behaviour in known cases of polysomy and mitotically unstable Bs in grasshoppers, in looking for possible causes for the odd-even effect. In all these traits, the odd-even effect was apparent, although its appearance was not universal in any case, with variation among species or populations within the same species. The equational division and lagging of the extra chromosomes, when univalents, could favour the appearance of abnormal meiotic products, and the formation of bivalents, when there are two or more extra chromosomes, inhibits this process. Therefore, the odd-even effect might be a consequence of the concomitant operation of both aspects of extra chromosome meiotic behaviour. The possibility that the odd-even effect might result from an increase in cell stress generated by odd numbers is suggested.     

Synaptonemal complex study in some species of Gerbillidae without heterochromatin interposition

Synaptonemal complexes were studied in Gerbillus campestris, Meriones libycus, M. shawi, M. crassus, and in two hybrids M. shawi x M. libycus (Gerbillidae, Rodentia). In both the pure species and hybrids, there was no pairing of X and Y chromosomes, as was previously observed in Psammomys obesus and other Gerbillidae species with gonosome-autosome translocations. A pair of autosomes was also located in proximity to the sex chromosomes in pachytene and showed unusual meiotic behavior with no, incomplete, or much delayed pairing. This chromosome pair, composed of late replicating heterochromatin, exists in most Gerbillidae species and is constant in number, but variable in size across the species. Both meiotic and mitotic characteristics indicate that this pair may correspond to a new type of chromosome which is different from B chromosomes. We do not know if there is a relationship between the presence of this chromosome and the unusual behavior of the sex chromosomes. In Gerbillidae species, the lack of pairing of both sex and heterochromatic chromosomes obviously does not prevent their correct meiotic segregation.     

Meiosis Drives Extraordinary Genome Plasticity in the Haploid Fungal Plant Pathogen Mycosphaerella graminicola

Meiosis in the haploid plant-pathogenic fungus Mycosphaerella graminicola results in eight ascospores due to a mitotic division following the two meiotic divisions. The transient diploid phase allows for recombination among homologous chromosomes. However, some chromosomes of M. graminicola lack homologs and do not pair during meiosis. Because these chromosomes are not present universally in the genome of the organism they can be considered to be dispensable. To analyze the meiotic transmission of unequal chromosome numbers, two segregating populations were generated by crossing genetically unrelated parent isolates originating from Algeria and The Netherlands that had pathogenicity towards durum or bread wheat, respectively. Detailed genetic analyses of these progenies using high-density mapping (1793 DArT, 258 AFLP and 25 SSR markers) and graphical genotyping revealed that M. graminicola has up to eight dispensable chromosomes, the highest number reported in filamentous fungi. These chromosomes vary from 0.39 to 0.77 Mb in size, and represent up to 38% of the chromosomal complement. Chromosome numbers among progeny isolates varied widely, with some progeny missing up to three chromosomes, while other strains were disomic for one or more chromosomes. Between 15–20% of the progeny isolates lacked one or more chromosomes that were present in both parents. The two high-density maps showed no recombination of dispensable chromosomes and hence, their meiotic processing may require distributive disjunction, a phenomenon that is rarely observed in fungi. The maps also enabled the identification of individual twin isolates from a single ascus that shared the same missing or doubled chromosomes indicating that the chromosomal polymorphisms were mitotically stable and originated from nondisjunction during the second division and, less frequently, during the first division of fungal meiosis. High genome plasticity could be among the strategies enabling this versatile pathogen to quickly overcome adverse biotic and abiotic conditions in wheat fields.     

Evolutionary dynamics of a satellite DNA in the tiger beetle species pair Cicindela campestris and C. maroccana.

Satellite repeat elements are an abundant component of eukaryotic genomes, but not enough is known about their evolutionary dynamics and their involvement in karyotype and species differentiation. We report the nucleotide sequence, chromosomal localization, and evolutionary dynamics of a repetitive DNA element of the tiger beetle species pair Cicindela maroccana and Cicindela campestris. The element was detected after restriction digest of C. maroccana total genomic DNA with EcoRI as a single band and its multimers on agarose gels. Cloning and sequencing of several isolates revealed a consensus sequence of 383 bp with no internal repeat structure and no detectable similarity to any entry in GenBank. Hybridization of the satellite unit to C. maroccana mitotic and meiotic chromosomes revealed the presence of this repetitive DNA in the centromeres of all chromosomes except the Y chromosome, which exhibited only a very weak signal in its short arm. PCR-based tests for this satellite in related species revealed its presence in the sister species C. campestris, but not in other closely related species. Phylogenetic analysis of PCR products revealed well-supported clades that generally separate copies from each species. Because both species exhibit the multiple X chromosome karyotypic system common to Cicindela, but differ in their X chromosome numbers (four in C. maroccana vs. three in C. campestris), structural differences could also be investigated with regard to the position of satellites in a newly arisen X chromosome. We find the satellite in a centromeric position in all X chromosomes of C. maroccana, suggesting that the origin of the additional X chromosome involves multiple karyotypic rearrangements.     

Morphology and Meiotic/Mitotic Behavior of B Cromosomes in a Japanese Harvestman, Metagagrella tenuipes (Arachnida: Opiliones): No Evidence for B Accumulation Mechanisms

Earlier, it has been demonstrated that wild populations of a Japanese harvestman Metagagrella tenuipes (Arachnida: Opiliones) are polymorphic for B chromosomes. In this paper, we present results of a study of the morphology and mitotic and meiotic behavior of the Bs. The B chromosomes varied considerably in size and proportion of eu- and heterochromatin. The single nucleolus organizing region, found in males, was located on a chromosome of the A complement. Some intercell variation in number of Bs may be explained by accidental chromosome losses during chromosome preparation. We also found no intertissue variation in number of Bs. There were also no differences in mean number of B chromosomes per individual among males and females, adult and subadult harvestmen. Segregation of Bs in mitotic and meiotic divisions was nonrandom; B chromosomes tended to segregate equally between daughter cells. The results obtained provide no support for the hypothesis of existence of B accumulation mechanism in this species.     

Condensin I reveals new insights on mouse meiotic chromosome structure and dynamics

Chromosome shaping and individualization are necessary requisites to warrant the correct segregation of genomes in either mitotic or meiotic cell divisions. These processes are mainly prompted in vertebrates by three multiprotein complexes termed cohesin and condensin I and II. In the present study we have analyzed by immunostaining the appearance and subcellular distribution of condensin I in mouse mitotic and meiotic chromosomes. Our results demonstrate that in either mitotically or meiotically dividing cells, condensin I is loaded onto chromosomes by prometaphase. Condensin I is detectable as a fuzzy axial structure running inside chromatids of condensed chromosomes. The distribution of condensin I along the chromosome length is not uniform, since it preferentially accumulates close to the chromosome ends. Interestingly, these round accumulations found at the condensin I axes termini colocalized with telomere complexes. Additionally, we present the relative distribution of the condensin I and cohesin complexes in metaphase I bivalents. All these new data have allowed us to propose a comprehensive model for meiotic chromosome structure.     

两种髭蟾的Ag—NORs,C—带及核型的研究

本文叙述和比较了峨眉髭蟾和哀牢髭蟾的核型,C-带和Ag-NORs,结果表明两者有下列相同方面:2n=26(6+7)、除No.3为SM外,其余诸对概为M,次缢痕和Ag-NoRs位于6q,并都表现出异形现象,No.1长臂有长度异形,但所增染色体片断不呈现C-带正染,C-带正染主要是在各对染色体的着丝点区域,另外No.2短臂近着丝区域亦为正染。这些表明二者之间有很大的核带型同源性。但是二者间在核型的某些对应染色体之间,在相对长度(6对)和臂比值(2对)方面有显著性差异,故可推测其机制是相互易位和臂间倒位。另外,哀牢髭蟾未发现与性别相关的异形性染色体。     

Sex and speciation: The paradox that non-recombining DNA promotes recombination

The benefits of sexual reproduction that outweigh its costs have long puzzled biologists. Increased genetic diversity generated by new allelic combinations, as enhanced by recombination during meiosis, is considered a primary benefit of sex. Sex-determining systems have evolved independently on numerous occasions. One of the most familiar is the use of sex chromosomes in vertebrates. Other eukaryotic groups also use sex chromosomes or smaller sex-determining regions within their chromosomes, such as the mating type loci in the fungi. In these organisms, sexual reproduction and its associated meiotic recombination are controlled by regions of the genome that are themselves blocked in recombination. Non-recombining DNA that is essential for recombination presents a paradox. One hypothesis is that sex-determination requires or leads to highly diverse alleles, establishing this block in recombination. A second hypothesis to account for the common occurrence of these types of sex-determining systems is that they combine mechanisms for recombination suppression and reproductive isolation, thereby promoting the evolution of new species. The fungal kingdom represents the ideal eukaryotic lineage to elucidate the functions of non-recombining regions in sex-determination and speciation.     

Cryptic sex in fungi

It is generally accepted that genetic recombination in eukaryotes is mostly driven by meiotic divisions as part of the sexual cycle. The maintenance of the sexual cycle that combines beneficial genes in sexual offspring from two parents provides an advantage for a species in novel or changing environments. Sex thus preserves beneficial and removes deleterious mutations. However, some eukaryotes, including many fungi lack sex entirely, and thus, it was assumed that recombination occurring during mitotic (somatic) divisions is the main force to shape the genome of these asexually propagating microbes. However, several recent reports of a sexual cycle in asexually propagating fungi put this concept in question. Here, we summarize the accumulating evidence for the general occurrence of cryptic sex in filamentous fungi in which sexual reproduction has not been previously reported.     

Chromosome polymorphism and C-banding variation of the brachypterous grasshopper Podisma sapporensis Shir. (Orthoptera, Acrididae) in Hokkaido, northern Japan

The grasshopper Podisma sapporensis consists of two main chromosome races in Hokkaido. The western group of populations of P. sapporensis, belonging to the XO race, has a diploid number of chromosomes 2n = 23 in the male and 2n = 24 in the female (sex determination XO male/XX female). The eastern group of populations of this species, belonging to the XY race, differs from the western one as a result of Robertsonian translocation between the originally acrocentric X chromosome and M5 autosome in homozygous state, having resulted in the forming of chromosome sex determination neo-XY male/neo-XX female (2n = 22). These races are geographically isolated by the mountainous system consisting of the Mts Daisetsu and Hidaka range, occupying the central part of the island. The hybrid zones between the races have not so far been discovered. Various levels of polymorphism for the pericentric inversions and C-banding variation exist in different chromosomes throughout populations in both chromosome races. In some solitary populations (the population at the summit of Mt Yotei, populations in the vicinity of Naganuma, Oketo, and Tanno) pericentric inversions are fixed in some pairs of chromosomes, which enables marking of the discrete karyomorphes. In the Mt Daisengen population all chromosomes are two-armed as a result of fixing the pericentric inversions. These facts contradict karyotypical conservatism of the tribe Podismini. The level of diversity of P. sapporensis karyotypes could provide a new perspective on the evolutionary process of different karyotype in Orthoptera. The considerable occurrence of polymorphism in chromosomes suggests that karyotypic diversification is undergoing in P. sapporensis. The authors also proposed that P. sapporensis would be divided into four chromosome subraces in the XO chromosome race and two chromosome subraces in the XY race, on the basis of karyotypic features. These races may have been established by fundamental climatic changes during the glacial epoch.     

C-bands on chromosomes of 32 beetle species (Coleoptera: Elateridae, Cantharidae, Oedemeridae, Cerambycidae, Anthicidae, Chrysomelidae, Attelabidae and Curculionidae)

C-banding patterns of 32 beetle species from the families Elateridae, Cantharidae, Oedemeridae, Cerambycidae, Anthicidae, Chrysomelidae, Attelabidae and Curculionidae were studied using the C-banding technique. Mitotic and meiotic chromosomes were previously described for 14 species. From among 18 species that had never been cytogenetically studied, we determined the diploid and haploid chromosome numbers and the sex determination system for 12 beetles. The karyotype for 6 species is not described because of a lack of mitotic and meiotic metaphases. Results confirm that most of the beetle species possess a small amount of heterochromatin and C-positive segments are weakly visible in pachytene stages and weakly or imperceptible in mitotic and meiotic metaphases. In some species with a large amount of heterochromatin, C-bands were observed in the centromeric region in all autosomes and the X chromosome. The Y chromosome does not show C-bands with the exception of Oedemera viridis in which it possesses a small band of heterochromatin.     

Size heterogeneity of telomeric DNA in mouse meiotic chromosomes

Heterogeneity for the length of telomeric DNA sequences has been found among different mitotic chromosomes in several mammalian species. However, there are no studies reporting such heterogeneity in meiotic chromosomes. To analyse this heterogeneity we have performed fluorescence in situ hybridization with a telomeric (C(3)TA(2))(3) peptide nucleic acid (PNA) probe on spread metaphase chromosomes during both male mouse meiotic divisions. Our results show that independently of the meiotic division, telomeric DNA signals were always surrounded by DAPI-stained chromatin, even at centromeric regions. Moreover, we have found heterogeneity for the size of telomeric DNA signals among different chromosomes, between homologues, and even within a given chromosome. We discuss the functional significance of the location of telomeric DNA in condensed meiotic chromosomes, and then the possible origin for the different polymorphisms found.     

Karyotype analysis in meiosis: Giemsa banding in the genus Secale L.

Summary Terminal bands of meiotic chromosomes stained by the Giemsa technique are permanent genetic structures of the nucleus during PMC differentiation in 8 samples of wild, primitive, and cultivated species of rye. The characteristic meiotic banding pattern is probably identical with the heterochromatic regions of mitotic chromosomes of root meristem cells (RMC) which have so far been studied. Karyotype analysis can be significantly improved by quantitative studies of the number and size of the bands combined with certain well-known chromosome characters in diplotene and diakinesis. The chromosomes involved in multivalents of some natural and synthetic species hybrids are identified for the first time. The results are discussed both in relation to the problems of chromosome evolution and their significance for marker techniques in cytogenetics.     

Rearrangements of highly polymorphic regions near telomeres of Saccharomyces cerevisiae.

We have examined the mitotic and meiotic properties of telomeric regions in various laboratory strains of yeast. Using a sequence (Y probe) derived from a cloned yeast telomere (J. Szostak and E. Blackburn, Cell 29:245-255, 1982), we found that various strains of Saccharomyces cerevisiae show extensive polymorphisms of restriction endonuclease fragment length. Some of the variation in the lengths of telomeric fragments appears to be under the control of a small number of genes. When DNA from various strains was digested with endonuclease KpnI, nearly all of the fragments homologous to the Y probe were found to be of different size. The pattern of fragments in different strains was extremely variable, with a greater degree of polymorphism than that observed for fragments containing the mobile TY1 element. Tetrad analysis of haploid meiotic segregants from diploids heterozygous for many different Y-homologous KpnI fragments revealed that most of them exhibited Mendelian (2:0) segregation. However, only a small proportion of these fragments displayed the obligate 2:2 parental segregation expected of simple allelic variants at the same chromosome end. From the segregations of these fragments, we concluded that some yeast telomeres lack a Y-homologous sequence and that the chromosome arms containing a Y-homologous sequence are different among various yeast strains. Regions near yeast telomeres frequently undergo rearrangement. Among eight tetrads from three different diploids, we have found three novel Y-homologous restriction fragments that appear to have arisen during meiosis. In all three cases, the appearance of a new fragment was accompanied by the loss of another band. In one of these cases, the rearrangement leading to a novel fragment arose in an isogenic diploid, in which both homologous chromosomes should have been identical. Among these same tetrads we also found examples of apparent mitotic gene conversions and mitotic recombination involving telemetric regions.     

The analysis of exchanges in tritium-labelled meiotic chromosomes

The autoradiographic analysis of exchanges in tritium-labelled meiotic chromosomes is potentially a useful approach to the study of meiotic exchange events since this method differentially labels meiotic chromatids along their entire length. The main problem encountered in earlier autoradiographic studies is that of distinguishing label exchanges generated at chiasmata from label exchanges generated by sister chromatid exchange. This problem was overcome in the present study by the choice of a meiotic system (male meiosis of Stethophyma grossum) where chiasmata are limited to just one proximally localised chiasma in each bivalent. This system allows the positive identification of chiasma-generated label exchanges and demonstrates convincingly the origin of chiasmata through breakage and rejoining of homologous non-sister chromatids. Sister chromatid exchanges are also readily detected in labelled meiotic chromosomes of this species, where they occur with a mean frequency of 0.35 per chromosome. This frequency is similar to that found in mitotic spermatogonial cells and the exchanges are randomly distributed both within and between chromosomes. These features of meiotic sister chromatid exchanges suggest that they are unrelated to non-sister chiasmatic exchanges and they probably have no special meiotic significance.     

Pds5 is required for homologue pairing and inhibits synapsis of sister chromatids during yeast meiosis

During meiosis, homologues become juxtaposed and synapsed along their entire length. Mutations in the cohesin complex disrupt not only sister chromatid cohesion but also homologue pairing and synaptonemal complex formation. In this study, we report that Pds5, a cohesin-associated protein known to regulate sister chromatid cohesion, is required for homologue pairing and synapsis in budding yeast. Pds5 colocalizes with cohesin along the length of meiotic chromosomes. In the absence of Pds5, the meiotic cohesin subunit Rec8 remains bound to chromosomes with only minor defects in sister chromatid cohesion, but sister chromatids synapse instead of homologues. Double-strand breaks (DSBs) are formed but are not repaired efficiently. In addition, meiotic chromosomes undergo hypercondensation. When the mitotic cohesin subunit Mcd1 is substituted for Rec8 in Pds5-depleted cells, chromosomes still hypercondense, but synapsis of sister chromatids is abolished. These data suggest that Pds5 modulates the Rec8 activity to facilitate chromosome morphological changes required for homologue synapsis, DSB repair, and meiotic chromosome segregation.     

Nuclear architecture and chromosome dynamics in the search of the pairing partner in meiosis in plants

The formation of haploid gametes in organisms with sexual reproduction requires regular bivalent chromosome pairing in meiosis. In many species, homologous chromosomes occupy separate territories at the onset of meiosis. To be paired at metaphase I, they need to be brought into a close proximity for interactions that include homology recognition and the establishment of some form of bonds. How homologues find each other is one of the least understood meiotic events. Plant species with large or medium sized genomes, such as wheat or maize, are excellent materials for the cytological analysis of chromosome dynamics at early meiosis, but genes that control meiosis have been identified mainly in small genome species such as Arabidopsis thaliana. This review is focused on the contribution studies on plants are providing to the knowledge of the initial steps of the meiotic process.     

In situ localization of human fibronectin (FN) genes to chromosome regions 2p14----p16, 2q34----q36, and 11q12.1----q13.5 in germ line cells, but to chromosome 2 sites only in somatic cells

The locations of the genes for fibronectin (FN) on chromosomes of human germ line and somatic cells were determined by in situ molecular hybridization with two 3H-labeled DNA probes, one for the region encoding the cell attachment domain of human FN, the other for the 3' noncoding and part of the coding region. Pachytene chromosomes of two males and lymphocyte chromosomes of one of these males and a female were used. Two regions of hybridization on pachytene and somatic chromosome 2 (p14----p16 and q34----q36) were found, but not in all individuals. A third region of hybridization was found at 11q12.1----q13.5 in meiotic, but not with significant frequency in somatic chromosomes. It is not clear if these differences between meiotic and somatic chromosomes, and the large differences between individuals at some of the other hybridization sites, resulted solely from technical factors. The differences between the findings in meiotic and somatic preparations might be due to the presence of four strands in pachytene chromosomes versus only one per somatic chromatid. Individual differences in DNA sequences in the chromosome segment containing the gene, differences in gene locations among individuals, or between meiotic and mitotic chromosomes might account for the other findings. The results confirm some of the earlier studies with cell hybrids that mapped FN genes to chromosomes 2 or 11. The combined findings suggest that some of these loci may be coding for the plasma form of FN and others for the cellular form. The expression of the different FN types by differentiated cells might then depend on the loci that are activated.