The relation between pairing preference and chiasma frequency in tetrasomics of rye

The association pattern of marked tetrasomes of Secale chromosome 1R at meiotic first metaphase was analyzed. Two of the four chromosomes were identical with terminal C-bands at both arms; the other two were also identical but lacked C-bands and were homologous or homeologous with the first two. Four different types of heterozygotes for 1R were studied: (i) autotetraploid hybrids between genetic variants within Secale cereale subsp. cereale, (ii) tetraploid hybrids between subspecies of Secale cereale, (iii) tetraploid hybrids between species of Secale, and (iv) autotetrasomes of S. cereale in a wheat background. Earlier observations that heterozygous associations (banded with unbanded) had consistently higher chiasma frequencies than homozygous associations were extended and confirmed. To analyze this phenomenon more closely, the possible relations between this correlation and several other meiotic phenomena were studied. For this analysis, three genetically different autotetraploid hybrids within S. cereale were selected that differed with respect to the relation between pairing type and chiasma frequency. Special attention was given to different patterns of interference and other meiotic phenomena in the two chromosome arms of chromosome 1R. No relations between such phenomena and the relation between pairing type and chiasma frequency could be established. A hypothesis is formulated assuming that long-distance homologue attraction is concentrated in a limited number of sites and that in different genotypes, different patterns of active sites are present. Moderately weak attraction sites can pair with strong homologous sites under favorable genetic conditions, but two weak sites cannot. Then, heterozygotes have more effective pairing initiation and consequently chiasma formation than homozygotes. Under less favorable conditions, only strong sites are effective, and then, homozygotes pair better, but the chiasma frequency is lower. A model of the forces involved in homologue attraction is presented.Key words: autopolyploids, preferential pairing, chiasma frequency, homologue attraction, Secale.

The pattern of homoeologous recombination in triploid hybrids of Lolium multiflorum with Festuca pratensis

Homoeologous chromosomes of Lolium-Festuca hybrids are capable of frequent meiotic pairing and recombination. The frequency and distribution of recombination was studied by genomic in situ hybridization in backcross progenies of reciprocal triploid hybrids of Lolium multiflorum with Festuca pratensis. Significant differences in the male transmission of the parental and translocated chromosomes were observed depending on the cytoplasm of the F1 hybrids and the ploidy level of the female test cross partner. The frequency of intergeneric translocations in the progeny indicated that, on average, there must have been at least 4.5 homoeologous arms paired in the F1 hybrids; the actual frequency might have been higher because of pre- or post-zygotic selection against the F. pratensis chromatin, which probably eliminated certain gametes with Festuca-Lolium translocations. Both parental species are known for localized distal chiasmata, but the intergeneric translocation breakpoints were distributed along the entire lengths of the chromosome arms. The change in the distribution of homoeologous recombination might have been related to different pairing initiation of homologues and homoeologues. It probably resulted from allocation of additional chiasmata to chromosome arms and produced a net increase in recombination.Key words: homoeologous exchanges, Lolium-Festuca, translocations, recombination.

Cytogenetics of Lathyrus palustris, a natural autohexaploid

American as well as British forms of the perennial plant Lathyrus palustris have 2n = 42 chromosomes with one group of 6 long submetacentric chromosomes, two groups of 6 medium-long subacrocentric chromosomes, three groups of 6 medium-long to medium-short submetacentric to subacrocentric chromosomes, and one group of 6 medium-short metacentric chromosomes. One haploid complement measures 45.8 μm at somatic metaphase. At meiotic first metaphase both forms show extensive multivalent formation. In 50 metaphase I cells of one plant of the British form there were 51 ring hexavalents, 80 chain and open-branched hexavalents of various forms, 2 quinquivalents, 41 ring quadrivalents, 58 chain quadrivalents, 20 branched quadrivalents of various forms, 1 trivalent, 128 ring bivalents, 280 open bivalents, and 9 univalents. It is concluded that L. palustris is most likely a natural autohexaploid with frequent multivalent pairing, many points of pairing initiation, several partner exchanges per multivalent, and many interstitial chiasmata. It may well be the first fully documented true natural autohexaploid reported. Pollen fertility was 56.2% and seed set 37%, apparently sufficient for a perennial plant species.Key words: Lathyrus, natural, autohexaploid, meiosis, multivalents.

Meiosis in allopolyploid Crepis capillaris. II. Autotetraploids

Meiotic chromosome pairing of autotetraploid Crepis capillaris was analysed by electron microscopy of surface-spread prophase I nuclei and compared with light microscopic observations of metaphase I chromosome configurations. Prophase I quadrivalent frequencies are high in all three tetrasomes. (A, D, and C) and partially dependent on chromosome size. At metaphase I quadrivalent frequencies are much lower and strongly dependent on chromosome size. There is no evidence for multivalent elimination during prophase I in this system, and the reduction in multivalent frequency at metaphase I can be explained by an insufficiency of appropriately placed chiasmata. The high frequencies of prophase I quadrivalents far exceed the two-thirds expected on a simple model with two terminal independent pairing initiation sites per tetrasome, suggesting that multiple pairing initiation occurs. Direct observations reveal relatively high frequencies of pairing partner switches (PPSs) at prophase I, which confirms this suggestion. The numbers of PPSs per tetrasome show a good fit to the Poisson distribution, and their positional distribution along chromosomes is random and nonlocalized. These observations favour a model of pairing initiation based on a large number of evenly distributed autonomous pairing sites each with a uniform and low probability of generating a PPS.Key words: autotetraploid, meiosis, Crepis capillaris, multivalent, pairing partner switch.

Correlation between pairing initiation sites, recombination nodules and meiotic recombination in Sordaria macrospora.

The decrease of meiotic exchanges (crossing over and conversion) in two mutants of Sordaria macrospora correlated strongly with a reduction of chiasmata and of both types of "recombination nodules." Serial section reconstruction electron microscopy was used to compare the synapsis pattern of meiotic prophase I in wild type and mutants. First, synapsis occurred but the number of synaptonemal complex initiation sites was reduced in both mutants. Second, this reduction was accompanied by, or resulted in, modifications of the pattern of synapsis. Genetic and synaptonemal complex maps were compared in three regions along one chromosome arm divided into well marked intervals. Reciprocal exchange frequencies and number of recombination nodules correlated in wild type in the three analyzed intervals, but disparity was found between the location of recombination nodules and exchanges in the mutants. Despite the twofold exchange decrease, sections of the genome such as the short arm of chromosome 2 and telomere regions were sheltered from nodule decrease and from pairing modifications. This indicated a certain amount of diversity in the control of these features and suggested that exchange frequency was dependent not only on the amount of effective pairing but also on the localization of the pairing sites, as revealed by the synaptonemal complex progression in the mutants.

Metaphase I pairing of deficient chromosomes and genetic mapping of deficiency breakpoints in common wheat

Metaphase I pairing of deficient chromosomes was analyzed in a set of 'Chinese Spring' (CS) wheat (Triticum aestivum L. em. Thell.) plants with varying lengths of deficiencies in the long arm of chromosome 4A (6, 8, 11, 17, 23, 34, 36, 39, and 50% missing), the long arm of chromosome 5B (49% missing), and the long arm of chromosome 2B (33% missing). Pairing in homologous chromosomes between deficient and complete arms was greatly reduced even by small differences in arm length. In deficiency homozygotes and in an isochromosome derived from a deficient 4AL arm, pairing of the two deficient arms was high and approached that of two complete arms. In plants where deficient and complete arms competed for pairing partners, pairing was exclusively between arms of the same length. These results suggest that in wheat, pairing initiation sites are distributed throughout at least the distal halves of the arms and that the alignment of telomeres may be critical for pairing success. Genetic mapping of the deficiency breakpoints was confounded by misdivision of unpaired chromosomes and abnormal transmission rates. Genetic distances between centromeres and breakpoints appeared to be proportional to metaphase I pairing frequencies.Key words: bread wheat, deficiency, chromosome pairing competition, mapping, telomere, pairing initiation.

Chromosomal fusion and meiotic behaviour in Delena cancerides (Araneae: Sparassidae). I. Chromosome pairing and X-chromosome segregation

Surface spreading of meiotic material in Delena cancerides indicates that pairing initiation among metacentric chromosomes with monobrachial homology differs from that of telocentric forms and free metacentric bivalents and results in a star-shaped structure at pachytene. Distance cosegregation of the three X chromosomes in ancestral, telocentric forms is prefaced by a centric association early in prophase I. This centric association of the X chromosomes is conserved in metacentric races despite the presence of an X-autosome fusion.Key words: synaptonemal complex, translocation heterozygote, X chromosome, spider.

Quadrivalent formation in a tetraploid chicken oocyte

Synaptonemal complex analysis of an exceptional tetraploid oocyte from a diploid chicken heterozygous for the MN t (Z;1) rearrangement was performed by electron microscopy of a spread preparation. Ten separate quadrivalents (26% of the chromosomal axes) were analyzed, as well as 50 autosomal bivalents. All the axes less than 2.5 μm in length formed bivalents (38) only, while axes in the 2.5–4.2 μm range formed 5 quadrivalents and 12 bivalents. The longer, separate axes formed quadrivalents only. Partner switches in excess of one were documented. The two identical W chromosomes paired only at the ends of their short arms. Quadrivalent formation may require a threshold length (2.5 μm), at least in this species. The tip of the short arm of the W chromosome may be a pairing initiation point, and it corresponds to the region associated with a localized recombination nodule previously described in diploid oocytes.Key words: quadrivalent, tetraploid, synaptonemal complex, chicken oocyte.

Synaptonemal complex spreading in Allium ursinum: pericentric asynapsis and axial thickenings

In Allium ursinum meiotic pairing of homologues is always incomplete; a proximal region on each bivalent remains regularly unsynapsed even in late pachytene. The spatial correlation of the unsynapsed region with the kinetochore suggests that the kinetochore itself exerts an inhibitory effect on synapsis in its vicinity. This can be interpreted as the cytological basis of the ‘centromere effect’ on recombination in this species. Moreover, the high incidence of a pericentric inversion loop in a heterozygous chromosome pair shows that proximal pairing initiation is possible and that its failure cannot be responsible for pericentric asynapsis. The formation of the inversion loop is complicated by the need for two independent pairing initiation sites because synapsis cannot proceed across the pericentric region. It is proposed that the meiotic bouquet polarization helps in establishing the presynaptic alignment of the homologous sites within the inverted regions and hence to achieve a high rate of inversion loop formation. Thickenings of the axial/lateral elements are not distributed equally along the synaptonemal complex. They are underrepresented in unpaired axes but strikingly abundant at the borders with synapsed regions, suggesting their origin in the pairing forks during the process of synapsis. They are virtually always present at nucleolus-organizing regions and often they appear at corresponding sites on opposite lateral elements. Besides the thickenings several other kinds of axial deformities are present in unpaired axes.

Synaptonemal complex spreading in Allium. II. Tetraploid A. vineale

In tetraploid Allium vineale four homologous axes are closely aligned in unsynapsed regions at early zygotene. This alignment is brought about by intercalary and terminal associations. The intercalary association sites are possibly targets of forces for long distance attraction of homologues and potential pairing initiation sites. The terminal associations are mediated by dense spherules and result possibly from the attachment of the telomeres to the nuclear envelope. In pachytene the alignment is abolished and the four axes are synapsed two by two, resulting in bivalents or, owing to partner switches in the synapsed axes, in quadrivalents. From the number of partner switches per configuration the number of pairing initiation sites is estimated. Homologous alignment and synapsis are discussed, comparing them with the conditions in a triploid species, which were described in a previous paper.Key words: Allium vineale, synaptonemal complex, polyploids, chromosome pairing, meiosis.