The influence of B-chromosomes on chiasma frequencies in Black Mexican Sweet Corn

The mean chiasma frequency at first metaphase of meiosis in pollen mother cells of a maize variety, Black Mexican Sweet Corn, increases with increasing number ofB-chromosomes. The increased chiasma frequency is also reflected by a decrease in the number of rod bivalents at first metaphase. There is some indication that theB-chromosomes also influence the distribution of chiasmata between pollen mother cells. The increase in chiasma frequency due toBs is closely analogous to the increase in crossing-over carlier attributed toBs between marker genes located on chromosome III. Together the evidence confirm the role ofBs in controlling genetic recombination at meiosis.     

The effect of X-irradiation on male meiosis in Schistocerca gregaria (Forskål)

The pattern of response of chiasma frequency to X-irradiation has been studied in germ line cells of male imagines of Schistocerca gregaria. A correlation has been established between the observed changes in chiasma frequency of the L and M type bivalents and the time in the meiotic cycle at which the treatment is given. Two radio-sensitive periods have been identified in meiosis itself. At one (meiotic DNA synthesis) X-irradiation produces a decrease in chiasma frequency while at the other (leptotene-early zygotene) the treatment leads to an increase in chiasma frequency. Small bivalents however do not respond to treatment and form a single chiasma under all conditions.     

Metaphase I bound arms and crossing over frequency in rye

Using a Giemsa C-banding procedure it has been possible to identify at meiosis three chromosome pairs of a local Spanish rye cultivar. Two of these chromosomes (3 and 5) were heterozygous for an interstitial C-band in the long arm and the other (chromosome 7) was heterozygous for a telomeric C-band, also in the long arm. From the frequency of being bound at metaphase I and the frequency of recombined chromatids at anaphase I in the arms considered, estimates of actual chiasma frequencies have been derived. The results have been compared with those obtained in a Fl between two inbred lines. It is concluded that: (i) Although the frequency of bound arms analyzed was similar in all cases, the chiasma frequency was higher in the cultivar than in the Fl plants. Cultivar plants showed a variation in chiasma frequency for the bivalent arms studied which was correlated with the frequency of bound arms per cell, indicating that the estimation for chiasma frequency by means of bound arm frequency has an error that increases with increasing number of bound arms per cell, (ii) Evidence of chiasma terminalization has not been found, (iii) It is suggested that the different rye chromosomes have different chiasma localization patterns, which, in turn, are related with the chiasma frequency.     

Sex differences in recombination

One of the stronger empirical generalizations to emerge from the study of genetic systems is that achiasmate meiosis, which has evolved 25–30 times, is always restricted to the heterogametic sex in dioecious species, usually the male. Here we collate data on quantitative sex differences in chiasma frequency from 54 species (4 hermaphroditic flatworms, 18 dioecious insects and vertebrates and 32 hermaphroditic plants) to test whether similar trends hold. Though significant sex differences have been observed within many species, only the Liliaceae show a significant sexual dimorphism in chiasma frequency across species, with more crossing over in embryo mother cells than in pollen mother cells; chiasma frequencies are unrelated to sex and gamety in all other higher taxa studied. Further, the magnitude of sexual dimorphism, independent of sign, does not differ among the three main ecological groups (dioecious animals, plants, and hermaphroditic animals), contrary to what would be expected if it reflected sex-specific selection on recombination. These results indicate that the strong trends for achiasmate meiosis do not apply to quantitative sex differences in recombination, and contradict theories of sex-specific costs and benefits. An alternative hypothesis suggests that sex differences may be more-or-less neutral, selection determining only the mean rate of recombination. While male and female chiasma frequencies are more similar than would be expected under complete neutrality, a less absolute form of the hypothesis is more difficult to falsify. In female mice the sex bivalent has more chiasmata for its length than the autosomes, perhaps compensating for the absence of recombination in males. Finally, we observe that chiasma frequencies in males and females are positively correlated across species, validating the use of only one sex in comparative studies of recombination.     

Occurrence and behaviour of B chromosomes in Artemisia frigida Willd.

In A. frigida Willd., B-chromosomes are reported for the first time. Their number was limited to 2 in all the cells studied. Analysis of chromosome behaviour at meiosis showed that B-chromosomes reduced the mean chiasma frequency and modified the phenotypic and biochemical characteristics of the plants.     

Comparative analysis of female and male meiosis in three meiotic mutants of tomato.

Female meiosis was analysed in squash preparations of ovules from three meiotic mutants and wild-type plants of tomato. In the completely asynaptic mutant as6, chromosome pairing and chiasma formation were virtually absent in both sexes. In the partially asynaptic mutant asb, with intermediate levels of chromosome pairing at pachytene, there were a higher number of chiasmate chromosome arms in female meiosis than in male meiosis, whereas in the desynaptic mutant as5 there were normal levels of chromosome pairing at pachytene and a similar reduction in chiasma frequency in the two sexes. In wild-type tomato, we found slightly higher numbers of chiasmate chromosome arms in female meiosis than in male meiosis. We propose that the higher female chiasma frequencies in mutant asb and wild-type tomato result from a longer duration of female meiotic prophase. This would allow chromosomes more time to pair and recombine. It is possible that a longer duration of prophase I does not affect mutants as5 and as6, either because the meiotic defect acts before the pairing process begins (in as6) or because it acts at a later stage and involves chiasma maintenance (in as5).     

The control of chiasma distribution in the locust,Schistocerca gregaria (Forskål)

Chiasma distribution at diplotene in Schistocerca gregaria males can be taken to indicate the positions at which crossing over occurred prior to diplotene since chiasma terminalisation is entirely absent. Analysis of chiasma frequency and position leads to a model for the mechanism controlling distribution which has three main components. — i) The bivalents vary in length between cells due to unknown factors and increase in bivalent length leads to an increase in chiasma frequency. — ii) Within bivalents chiasma initiation is sequential from telomere to centromere with the first chiasma usually forming close to the telomere. — iii) Interference operates with the same polarity and determines a distance within which crossing over is precluded.     

Behavior of meiotic chromosomes in Pinus wallichiana, P. strobus and their hybrid and nrDNA localization in pollen mother cells of the hybrid by using FISH

The complete process of meiosis was investigated in Pinus wallichiana, P. strobus and their artificial hybrid （F1） using microsporocytes. It is revealed that there were slightly lower chiasma frequency, lower ring bivalent frequency, lower meiotic index and distinctly higher frequency of aberrance （chromosomal bridges, fragments or micronuclei） in pollen mother cells （PMCs） of the hybrid （F1） than those of the parental species, which showed a certain degree of differentiation between homologous chromosomes of the two parents. However, relatively higher frequency of ring bivalents and higher meiotic index in all the three entities indicate the great stability of genomes of parental species, and the differentiation of genomes between the two parents must have been slight. Total nineteen signal loci of 18S rDNA were observed in nine bivalents of the hybrid （F1）, among which one bivalent bears two loci, while the others have only one. It is suggested that distinct differentiation at genetic level existed in homologous chromosomes of the two parental species, whereas only slight differentiation at karyotypic and genomic levels take place between the parent species.     

Resolution of chiasmata in oocytes requires separase-mediated proteolysis

In yeast, resolution of chiasmata in meiosis I requires proteolytic cleavage along chromosome arms of cohesin's Rec8 subunit by separase. Since activation of separase by the anaphase-promoting complex (APC/C) is supposedly not required for meiosis I in Xenopus oocytes, it has been suggested that animal cells might resolve chiasmata by a separase-independent mechanism related to the so-called "prophase pathway" that removes cohesin from chromosome arms during mitosis. By expressing Cre recombinase from a zona pellucida promoter, we have deleted a floxed allele of separase specifically in mouse oocytes. This prevents removal of Rec8 from chromosome arms and resolution of chiasmata. It also hinders extrusion of the first polar body (PBE) and causes female sterility. mRNA encoding wild-type but not catalytically inactive separase restores chiasma resolution. Both types of mRNA restore PBE. Proteolytic activity of separase is therefore essential for Rec8's removal from chromosome arms and for chiasma resolution but not for PBE.     

Variation in chiasma frequency among eight accessions of Arabidopsis thaliana

Natural variation in meiotic recombination frequency in Arabidopsis thaliana has been assessed by analyzing chiasma frequency variation among a range of geographically and ecologically diverse accessions. Fifty pollen mother cells at metaphase I of meiosis were analyzed from each of eight accessions and fluorescence in situ hybridization was applied to enable identification of all 10 chromosome arms. There was no significant variation in mean chiasma frequency between plants within accessions, but there was significant variation between accessions. Further analysis confirmed this finding and identified two particular accessions, Cvi and Ler, as having chiasma frequencies significantly lower than those of the other accessions. The analysis also revealed that the pattern of chiasma distribution between arms and among chromosomes is not consistent over accessions. Further detailed analyses were conducted on each individual chromosome (1-5) in turn, revealing that chromosome 4, one of the acrocentric chromosomes, is the least variable while the other acrocentric chromosome (2) is the most variable. These findings indicate the existence of recombination regulatory elements in Arabidopsis and we conclude that it may be possible in the future to identify these elements and determine their mode of action. The practical implications of such developments are considerable.     

The effect of B chromosomes on homoeologous pairing in species hybrids

The effect of B chromosomes on meiosis is described in the diploid and tetraploid interspecific hybrid Lolium multiflorum x Lolium perenne. Although the parental species are very closely related, the presence of B chromosomes in the diploid hybrid reduced both chiasma frequency and the number of bivalents at meiosis by a small but significant amount. However at the tetraploid level the presence of B chromosomes did not seem to alter the pairing pattern and chiasma frequency in any way. The use of B chromosomes to stabilize meiosis in amphiploids of this type between closely related outbreeding species is therefore ruled out.     

Male and female meiosis in diploid and colchitetraploid Phlox drummondii Hook. (Polemoniaceae)

Comparative studies on male and female meiosis in diploid (2n = 2x = 14) and colchitetraploid (2N = 4x = 28) ornamental Phlox drummondii reveal higher chiasma frequency in embryo-sac mother cells as compared with pollen mother cells in the diploid, and significant differences in the pairing properties of chromosomes in the pollen mother cells and embryo-sac mother cells of the colchitetraploid. Male meiosis in the colchicine-induced autotetraploid was abnormal with 51.70% of chromosomes associated in quadrivalents and tridents in 96% of the cells. This was followed by discordant anaphase I disjunction. In the female meiocytes the chromosomes formed 14 bivalents in 86.66% of the cells. Quadrivalents (1–3) appeared only in 13.34% of the cells. It is concluded that meiosis in male and female cells of the colchitetraploid is governed and regulated by different controlling systems of gene(s).     

The cytogenetic structure of Tasmanian populations of Phaulacridium vittatum

In 8 out of 20 Tasmanian populations of Phaulacridium vittatum from 0.3-11.0 percent of the males carried a single supernumerary chromosome. In such males the X and B univalents are both heteropycnotic at first prophase of male meiosis and associate with another in a non-homolgous manner in about two-thirds of the diplotene cells examined. In all 56 B-containing individuals studied, however, these associations lapse by first metaphase and the X and the B move at random with respect to one another at first anaphase. The supernumerary in this species is stable and only 5 of the 56 individuals with supernumeraries showed evidence of B-chromosome non-disjunction in the pre-meiotic mitoses. Since there was no other evidence for the loss or gain of supernumeraries in the male line it is clear that B-transmission is regular in the males of this species. There were significant differences between population with respect to mean cell chiasma frequency but there was no significant effect of the B-chromosome on this metric. Additionally a comparison of mean cell chiasma frequency in follicles with and without supernumeraries from a mosaic individual shows no significant difference.     

Male and female meiosis in grasshoppers

Male meiosis in the grasshopper Stethophyma grossum is well known as an example of proximal chiasma localisation. An investigation of female meiosis in oocytes of this species shows that both the frequency and distribution of chiasmata are quite different from the male situation. Mean chiasma frequency per cell (14.98) in considerably higher in females than in males (11.28) which agrees with the trend established in other comparative studies of male and female meiosis. More strikingly, males and females also show not only different but quite opposite patterns of chiasma distribution. In spermatocytes of males, chiasmata are strictly localised proximally in most bivalents, but in oocytes of females very few chiasmata form in proximal regions and nearly all chiasmata form either in distal or interstitial regions. The genetical significance of these findings is considered.     

Chiasma frequency and distribution in a sample of human males: chromosomes 1, 2, and 9

Chiasma frequency and the distribution of chiasmata within chromosomes 1, 2, and 9 were investigated in a total of seven males with presumptively normal meiosis. With the exception of one unusual individual there was little variation among the males examined for these characteristics. The exceptional male had a reduced chiasma frequency, which, rather than being considered abnormal, was thought to represent the lower end of the normal range of variability. A statistical analysis of the histograms of chiasma distribution confirmed the impression of overall similarity, and we conclude that genetic maps of these chromosomes may now be constructed without too much disturbance from differences between individuals. The maps will be presented separately in a subsequent paper.     

Higher recombination frequencies in female compared to male meisoses in Brassica oleracea

Linkage maps of the nine chromosomes of Brassica oleracea, based on 75 informative molecular markers, have been compared in first and second backcross progeny from a cross between two doubled haploid lines. The second backcross progeny showed greater recombination frequencies for 75% of the pairs of adjacent markers, but there was no obvious indication that this effect was localised to particular regions of the chromosomes. Four chromosomes increased in genetic length more than twofold, while overall, the total map was 66% longer. The possible causes of this discrepancy are analysed. A sex difference in chiasma distribution and/or frequency at meiosis is thought to be the most likely explanation. The implications of this finding for mapping and map-based applications are discussed.     

Metaphase I bonds,crossing-over frequency,and genetic length of specific chromosome arms of rye

Using a Giemsa C-banding procedure three chromosome pairs (3, 6 and 7) have been identified in meiosis of the F1 of a cross between two rye inbred lines. Two of these chromosome pairs (3 and 7) were heterozygous for a prominent telomeric heterochromatic band. The comparison between the frequencies of the different meiotic configurations at metaphase I, anaphase I and metaphase II presented by these two chromosome pairs has allowed the estimation of the chiasma frequency and the genetic length of the chromosome arms 3 short and 7 long.     

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.     

Genetic and environmental factors affecting chiasma formation in locusts

By means of crosses with the African Migratory locust it has been demonstrated that various strains carry different segregations of genes which quantitatively condition the formation of chiasmata during meiosis. These genes occur in polygenic series, and in one low frequency strain the polygenes show an epistatic effect for low chiasma frequency, while another low frequency strain seems to have attained this property through gamma radiation. In reciprocal crosses between the lastnamed low and a medium frequency strain there appears to be a maternal effect, with heterosis and epistasis respectively in the reciprocal crosses. In one strain two lines were selected for plasticity and non-plasticity of hopper colour change in small populations, and this selection has resulted in multiple changes in that the two lines now diverge in the three main physical criteria of phase transformation, viz. hopper colour, adult morphometric ratios and chiasma frequencies; one line is now solitarious for these characteristics, the other more gregarious. The genotypes selected out could be related to the ability to utilize the gregarization pheromone.A 10° rise in temperature above 22° C has the effect of increasing chiasma frequencies significantly, but increase in the carbon dioxide content of the atmosphere has no effect. Male and female hoppers produce equivalent amounts of the chiasma-inducing pheromone, while solitarised hoppers do not lose the ability to produce this pheromone. The crucial stage for the action of this pheromone on chromosomes is during the fifth nymphal instar. The phase status of parents appears to affect chiasma frequencies in the offspring, more specifically that of the mother, so that there is further evidence of maternal effect on chiasma frequency.     

The Development and Meiotic Behavior of Asymmetrical Isochromosomes in Wheat

To determine which segments of a chromosome arm are responsible for the initiation of chiasmate pairing in meiosis, a series of novel isochromosomes was developed in hexaploid wheat (Triticum aestivum L.). These isochromosomes are deficient for different terminal segments in the two arms. It is proposed to call them ``asymmetrical.' Meiotic metaphase I pairing of these asymmetrical isochromosomes was observed in plants with various doses of normal and deficient arms. The two arms of an asymmetrical isochromosome were bound by a chiasma in only two of the 1134 pollen mother cells analyzed. Pairing was between arms of identical length whenever such were available; otherwise, there was no pairing. However, two arms deficient for the same segment paired with a frequency similar to that of normal arms, indicating that the deficient arms retained normal capacity for pairing. Pairing of arms of different length was prevented not by the deficiency itself, but rather, by the heterozygosity for the deficiency. Whether two arms were connected via a centromere in an isochromosome or were present in two different chromosomes had no effect on pairing. This demonstrates that in the absence of homology in the distal regions of chromosome arms, even if relatively short, very long homologous segments may remain unrecognized in meiosis and will not be involved in chiasmate pairing.