Chiasmata and chromosome breakages are related to crossing over in Drosophila ananassae males.

A cytogenetic analysis of male crossing over in Drosophila ananassae revealed that cytological exchanges resulted in genetic crossing over, and that chiasma frequency and the genetic recombination correlated positively in chromosomes 2 and 3. Furthermore, the frequency of chromosome breakages correlated positively with chiasma frequency. Paracentric inversion heterozygosity had no detectable influence on the chromosome pairing or exchange events within the inversion loop at meiosis. Scoring of the chiasma demonstrated that males homozygous for the previously mapped enhancers of male crossing over had low frequencies of chiasmata, whereas higher frequencies of chiasmata were observed in males heterozygous for enhancers. The results presented here indicate that the genetic factors controlling male crossing over are involved in the origin of chromosome breakages and in exchange events.     

Chiasmata and variability in Lolium and Festuca populations

There are significant differences in mean pollen mother cell chiasma frequencies between populations within Lolium perenne, L. multiflorum and Festuca pratensis. The differences are genotypically controlled. With low chiasma frequencies the chiasmata are distally located. With increasing chiasma frequency the frequency of chiasmata in interstitial segments increases. Shorter lived populations have higher chiasma frequencies than the more perennial. — The higher the chiasma frequency of a population the lower the phenotypic and genetic variance for characters under polygenic control, such as flowering time, and the less effective also is the response to selection for such characters. These observations are interpreted on the premise that high chiasma frequencies are instrumental in the breaking up of supergene sequences in interstitial chromosome segments.     

The quantitative analysis of chromosome pairing and chiasma formation based on the relative frequencies of M I configurations. IV. Interchange heterozygotes

The quadrivalent of an interchange heterozygote can be divided into six regions, each with its own probability of being bound by at least one chiasma. This gives rise to 2⁶ or 64 different possible combinations of bound and not bound regions and as many M I configurations, several of which are isomorph. Sixteen different types of configuration occur, 10 of which involve more than two chromosomes and can thus be distinguished from the non-interchange bivalents. As each combination of bound and not bound regions has a certain probability of occurring depending on the probability of each region to be bound or not, a frequency expectation value can be estimated for each configuration. Also, when the relative frequencies of the different configurations are known, the probabilities of being bound can be estimated for each of the six regions of a quadrivalent, and from these the crossing-over potentials. One translocation heterozygote was studied, involving the long arm of the satellite chromosome of rye (Secale cereale). Crossing-over was reduced, perhaps mainly by partner exchange. The terminal segments retained a relatively high chiasma frequency. The interstitial regions had low chiasma frequencies even when relatively long.     

The B-chromosomes of Locusta migratoria I. Detection of negative correlation between mean chiasma frequency and the rate of accumulation of the B's; a reanalysis of the available data about the transmission of these B-chromosomes

B-chromosomes were studied in two Spanish populations of Locusta migratoria. Both exhibit a high frequency of B-carrying individuals (75% and 94%). In both the B-chromosomes are mitotically unstable and they accumulate in the male germ line. The mean rate of accumulation is 28.5% in Baños de la Encina and 31.9% in Carboneras. There are no significant differences in mean cell chiasma frequency and between-cell variance of chiasmata between follicles with different number of B's within individual males. Likewise, there are no significant differences in mean cell chiasma frequency and between-cell variance between males with and without supernumeraries. However, a significant negative correlation exists between mean chiasma frequency and the rate of accumulation.     

Son-sire-regression based heritability estimates of chiasma frequency, using T70H mouse translocation heterozygotes, and the relation between univalence, chiasma frequency and sperm production.

T(1;13)70H/+ translocation heretozygous mice were used for assessing heritability values for chiasma frequencies and the epididymal sperm count. The chiasma frequency estimates were based on 15 son-sire pairs, the translocation heterozygotes being maintained in a Swiss random-bred genetic background. The chiasma frequencies were scored separately for the T70H/+ derived multivalent, specific pairing segments within the multivalent and the remaining bivalents. Chiasma counts within these specified parts of the genome were positively correlated. The heritability estimates, significantly greater than zero, ranged from 0.78-0.98, depending on the chromosome segments included. These results indicate a strong genetic control on a cellular basis for the formation of chiasmata in the mouse. Despite significantly positive correlations and regressions between the various chiasma frequencies and the sperm count (for which 29 pairs of observations were available), no significant heritability estimate for the sperm count was obtained. The relation between the chiasma frequency and the sperm count was weakest when the chiasma count was confined to a region of the translocation-caused multivalent in which the absence of a chiasma almost always resulted in the production of an univalent. This indicates that in the translocation heterozygotes used, the overall chiasma frequency has a greater predictive value for the sperm count than autosomal univalence alone.     

The calculation of the map length of interchange segments from meiotic configuration frequencies

Standard mapping functions permit the calculation of map length from the frequency by which a chromosome segment has at least one chiasma. In interchange heterozygotes, approximately exact estimates of such frequencies can be obtained from the relative frequencies of specific groups of meiotic metaphase I configurations. Examples are given for two interchanges inSecale cereale.     

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 quantitative analysis of chromosome pairing and chiasma formation based on the relative frequencies of M I configurations

In autotetraploids, chromosome pairing may be in the form of quadrivalents or bivalent pairs. Whether or not the quadrivalents are maintained until first meiotic metaphase depends on the formation of chiasmata. The relative frequencies of M I configurations thus contain information both on pairing and on chiasma formation. With distal chiasma localisation six configurations can be recognised and their relative frequencies determined: ring quadrivalents, chain quadrivalents, trivalents (with univalent), ring bivalents, open (rod) bivalents, univalent pairs. These represent five degrees of freedom permitting five parameters to be estimated: the frequency (f) of quadrivalent pairing; the frequencies of chiasmate association of the two ends (arms in metacentrics), a′, b′, after quadrivalent pairing, and a, b after bivalent pairing. — The appropriate formulae have been derived and applied to observations on Tradescantia virginiana (4n=24) which has pronounced distal chiasma localisation. Slight modifications make the model applicable to autotetraploids with interstitial in addition to distal chiasmata. In T. virginiana, chromosome pairing appeared to be random between homologues (65.8% quadrivalent pairing; 55.4% observed at M I). After quadrivalent pairing chiasmate association is frequent in the “average long” arm (95.0%) and much less so in the other arm (60.5%). This is attributed to partner exchange. After bivalent pairing chiasma frequencies are still different for the two arms (93.8% and 83.5% association respectively) but much less pronounced. Various complications are discussed.     

The effects of ageing on the meiotic chromosomes of male and female mice

The effects of age on the chiasma frequencies, chiasma position and numbers of univalents at MI in males and females of three strains of mouse were examined. Males showed a slight but non significant rise in chiasma frequency in age due to an increase in bivalents with two chiasmata at the expense of single chiasmata bivalents. In contrast, females exhibited a significant decrease in chiasma frequency with age due to the loss of two chiasma bivalents with a corresponding increase in single terminal chiasmata bivalents. In both males and females there was no significant increase in univalents with age in the strains studied. Of interest was the finding of a greater degree of contraction of the MI chromosomes in the oocytes of old relative to young females, a differential contraction that was independent of culture time. This finding is discussed with regard to the production line theory and non disjunction at Anaphase in other strains of mice.     

Male and female meiosis in grasshoppers II. Chorthippus brunneus

The frequencies and distribution patterns of chiasmata in the autosomal bivalents of Chorthippus brunneus are very similar in male and female meiosis. The mean cell chiasma frequency of oocytes is however significantly higher than that of spermatocytes, but this difference is almost certainly due to the incremental effect of the X bivalent on the chiasma frequencies of oocytes. The implications of these findings in terms of chiasma control are briefly discussed.     

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.     

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.     

Looking for natural variation in chiasma frequency in Arabidopsis thaliana

Information concerning natural variation either in chiasma frequency or in the genetic basis of any such variation is a valuable tool to characterize phenotypic traits and their genetic control. Here meiotic recombination frequencies are analysed in nine geographically and ecologically diverse accessions of Arabidopsis thaliana, and a comparative study was carried out incorporating previous data from another eight accessions. Chiasma frequencies, estimated by counting rod and ring bivalents at metaphase I, varied up to 22% among accessions. However, no differences were found among plants of the same accession. There was a relationship, which does not necessarily imply direct proportionality, between the size of the chromosomes and their mean chiasma frequency. Chiasma frequency and distribution between arms and among chromosomes were not consistent over accessions. These findings indicate the existence of genetic factors controlling meiotic recombination both throughout the whole genome and at the whole chromosome level. The reliability of chiasma scoring as an indicator of reciprocal recombination events is also discussed.     

Chromosome pairing and chiasma formation in autopolyploids of different Lathyrus species.

Chromosome pairing and chiasma formation were studied in natural and induced tetraploids (2n = 28) of Lathyrus odoratus (induced), Lathyrus pratensis (natural and induced), Lathyrus sativus (induced), and Lathyrus venosus (natural), as well as in triploids of L. pratensis and diploids of L. odoratus, L. pratensis, and L. sativus. All natural tetraploids appeared to be autotetraploids and their meiotic metaphase I behaviour was very similar to that of the induced autotetraploids, with average numbers of pairing partner switches exceeding 4 or even 5. Multivalent frequencies were high, but the numbers of chiasmata were not much higher than necessary to maintain the configurations. Interstitial chiasmata were common, but not predominant. Fertility was reduced, but sufficient for predominantly vegetatively reproducing species. The triploids of L. pratensis had an even higher multivalent frequency than the tetraploids, but still produced some viable progeny at or close to the tetraploid level, suggesting that in mixed populations of diploids and tetraploids, triploids can contribute to gene flow between the ploidy levels. There was no significant correlation between chiasma frequency and ring bivalent frequency in the diploids and multivalent frequency in the corresponding tetraploids. In the tetraploids, chiasma frequency and multivalent frequency were negatively correlated.     

Cytology of inbreds and f 1 hybrids of pearl millet

Summary The meiotic behaviour of chromosomes was studied in four inbred lines and their F ₁ hybrids of P. typhoides. The inbred lines showed a decrease in mean chiasma frequency when compared with the population plants, but differed from one another in their mean chiasma frequencies. Together with the decrease in mean chiasma frequency the inbreds showed variation in mean chiasma frequencies. The inbred lines showed a number of meiotic abnormalities such as extra chromosomes, extra fragments, desynapsis, persistent nucleoli and differential condensation of chromosomes. The F ₁ hybrids of these inbreds exhibited heterosis for chiasma frequency. All the F ₁'s had mean chiasma frequencies higher than the means of the respective participating parents. The F ₁'s, however, differed in the degree of heterosis exhibited. In the F ₁ hybrids, the variation in mean chiasma frequency, both between plants and between PMC's within plants, was significantly lower than that of the inbred lines. The effect of environment was studied in the inbred lines and their F ₁ hybrids. The mean chiasma frequencies of the inbred lines were significantly lower, and the variation in mean chiasma frequencies was greater, in the stress season. The mean chiasma frequencies of F ₁'s did not show any significant differences between the two seasons. The F ₁'s exhibited less variation in mean chiasma frequency than the inbred lines, showing that F ₁'s were developmentally more stable. The F ₁'s did not show any meiotic abnormalities in either season.     

The effect of B chromosome number on chiasma frequency within and between individuals of Gibasis linearis (Commelinaceae)

Meiotic analysis of 21 plants from a single population of Gibasis linearis which contained from 0 to 6 apparently identical B chromosomes showed that the mean chiasma frequencies were significantly higher with increasing numbers of Bs. It was also found that those meiotic cells of a 5B plant which contained fewer than five B chromosomes showed a marked fall in chiasma frequency, demonstrating that the influence of B chromosomes on chiasma formation in A chromosomes acts at a cellular rather than a whole-anther level.     

Chiasma frequencies in the meiocytes from the anther diametrical zones in species from the genus <Emphasis Type="Italic">Capsicum</Emphasis> L.

A study of the chiasma frequencies in the meiocytes from three anther diametrical zones (free end, middle, and base) in plants from species of the genus Capsicum was carried out. It was found that the maximum values of the total chiasma frequencies were observed in the meiocytes from the anther free end compared to its middle zone and base. It was concluded that the peculiarities of the distribution of crossover events lengthwise along the anther were the result of its anatomic structure.     

Spermatogenic delay and increased chiasma frequency in T70H/+ male mice with hydroxyurea-Trenimon limited spermatocyte populations

T(1;13)70H/+ male mice were treated with hydroxyurea (HU) and Trenimon (T). This karyotype offers excellent possibilities for estimating number and position of chiasmata and segregation in meiotic anaphase I. By their cell-killing action during spermatogenesis, HU and T produce large gaps in the spermatogenic line. The surviving population between the gaps was analysed at diakinesis--metaphase I and metaphase II. We found by autoradiography a considerable retardation of the development from resting primary spermatocytes (RPS) to metaphase I and II as compared to untreated T70H/+ males. Furthermore we found increased chiasma frequencies in diakinesis--metaphase I (MI) and reduced nondisjunction frequencies at anaphse I as a result of the treatments applied. The latter effect could not be explained by the increased chiasma frequency.     

Dominant enhancer effect of the meiotic mei4 mutant on recombination frequencies restricted to linkage group VI in Podospora anserina.

Summary A mutant which increases second division segregation (SDS) frequency of locus 110 (linkage group VI) was isolated. It was called mei4 because of its meiotic deficiency. The present paper deals with its effect on meiotic recombination when heterozygous. mei4 then only acts on linkage group VI. The SDS frequencies were increased for all markers used, except locus 5 located very close to the centromere. This quasi general enhancement results exclusively in an enlargement of map distance on linkage group VI's proximal part. Crosses involving three mutant genes allowed to check that the distances on the distal part were constant. This is due to a real lack of crossover frequency modification in this region and not to a change of chiasma interference. Among the seven linkage groups of Podospora anserina, group VI exhibits several other particularities concerning meiotic recombination, especially a lower positive chiasma interference and a more regular crossover distribution, suggesting a particular recombination regulation.Laboratoire associé n⁰ 86 du Centre National de la Recherche Scientifique     

Studies on chiasma distribution and chiasma movement in Zoniopoda tarsata (Orthoptera: Romaleidae)

A sample of 27 males of Zoniopoda tarsata from Argentina was studied cytologically. The three largest autosomal pairs and the X were characterized by the presence of interstitial C-bands. Chiasma position relative to the bands was analyzed at diplotene and diakinesis. The frequency of interstitial, terminal and total chiasmata per cell was studied for the whole autosomal bivalent set, analysing the variations between stages and among individuals. The comparison of interstitial chiasma frequencies between stages and among individuals and the study of chiasma position relative to the bands in pairs 1, 2 and 3 indicated that chiasma distribution varied from diplotene to diakinesis. Therefore, terminalization does exist in this species and the movement may occur towards the centromere. The frequency of terminal associations at diplotene showed a high negative correlation (r=-0.89; p<10^-5) with the number of interstitial chiasmata. This correlation would not be expected if the two kinds of association were produced by different (independent) mechanisms. Consequently, terminal associations were considered genuine chiasmata. The correlation between interstitial and total chiasmata was very much lower then the former (r=0.39; p=0.04). This fact, besides the relatively low variation for chiasma number, observed among individuals suggests that in this species the number of interestitial chiasmata, which are the most important in controlling the genetical recombination, is mainly regulated by changes in chiasma distribution, while variations in total chiasma frequency are of much lower magnitude.Member of Carrera del Investigador del Consejo Nacional de Investigaciones Cientificas y Técnicas (CONICET)