Chiasmata distribution in the normal karyotype of mice

The number of chiasmata per cell and variance of chiasmata numbers were studied, as well as the recombinational interaction between different bivalents in CBA/Lac mice male line. No competition of bivalents for chiasmata was discovered in mice; at the same time, the chiasmata within one arm of the chromosome interfere with each other. The number of chiasmata per bivalent is estimated for each chromosome independently. The number of chiasmata per chromosome is limited both from below (minimum one chiasma independently of its size) and from above (positive interference of chiasmata).     

Fitting of a binomial distribution to the number of chiasmata per bivalent

Statistical tests on the distribution of the number of chiasmata per chromosome, collected from literatures, showed that they can be approximated by binomial distributions with one obligatory chiasma, i.e., B(N-1, p). N is proportional to the average number of chiasmata, while p is nearly constant for the species tested.     

Ecological correlates of chiasma frequency and recombination index of plants

This paper tests four theories of the maintenance of genetic recombination using published chiasma frequencies of 194 plant species representing 15 families of angiosperms. The theories are that recombination is favoured by environmental unpredictability, by fluctuating selection, by mutational load or by sib-competition. The level of genetic recombination was approximated by recombination index and by the number of excess chiasmata. Both measures were higher in animal-dispersed than in other species and increased as dispersal distance decreased. This association was found at all taxonomic levels. Chiasma frequency was not associated with life span of the species. Recombination index was lower in perennial than in annual species, but the reverse trend was observed among genera. Number of chiasmata per bivalent tended to decrease as the number of chromosomes increased, but this association was statistically insignificant at all taxonomic levels. No association was found between number of chiasmata per cell and nuclear DNA content.
These patterns did not support the theories that environmental unpredictability or fluctuating selection due to parasite pressure favour recombination, and they offered only very little support for the theory that recombination is favoured so that mutational load can be decreased. The only theory that was consistent with the observed data is the sib-competition theory, which suggests that genetic recombination is maintained so that the intensity of competition with sibs is reduced.     

Effects of supernumerary heterochromatin on chiasma condition in two species of Acrididae (Orthoptera)

Amblytropidia australis and Dichroplus elongatus were found to be polymorphic for supernumerary heterochromatin. In both, basic karyotypes are 2n=22+XO in males.Mitotically unstable extra chromosomes were detected in a population of A. australis. The Bs are telocentric and their number varies from O to 2 within individuals. Mean frequencies of interstitial and total chiasmata at diplotene were compared between individuals with and without Bs. The mean frequency of interstitial chiasmata increases with the number of Bs per cell.A supernumerary terminal segment in S₁₀ pair was observed in a heterozygous condition in several individuals of D. elongatus. The localization and frequency of chiasmata at diplotene were studied. The segment has an intrachromosomal effect since it modifies the location of chiasmata in the bivalent involved.Fellow of the Consejo Nacional de Investigaciones Científicas y Técnicas (CONICET).     

Lampbrush chromosomes in the japanese quail Coturnix coturnix japonica: cytological maps of macro chromosomes and meiotic crossover frequency in females

Cytological maps of lampbrush macrobivalents of the Japanese quail (Coturnix coturnix japonica) were constructed. Investigation of chiasmata allowed determination of the meiotic frequency of reciprocal genetic recombination (crossing over) in Japanese quail females. The total chiasma number in bivalents of Japanese quail oocyte nuclei was determined to be 53-58. Macrobivalents 1-5 and Z of the Japanese quail had on average 3.3 chiasmata per bivalent, and microbivalents, 1.0-1.1 chiasmata per bivalent. The chiasmata (crossover) frequency in Japanese quail females was lower than in chicks. In macrochromosomes of Japanese quail females, one crossover occurred per 43.9 Mb, and in chicken, per 30.0 Mb. Judging from chiasma frequency, the genetic length of the Japanese quail genome is likely to be 2650-2900 cM. Crossover frequency in the species was 0.023 per Mb in macrobivalents and 0.07-0.08 Mb in microbivalents and for the total genome, 0.041 crossovers per Mb. The genetic length of one Mb (theta) in female Japanese quails was 1.14 cM in macrochromosomes, 3.60-4.12 cM in microchromosomes, and about 1.96-2.15 cM averaged over the genome.     

Double insertion of homogeneously staining regions in chromosome 1 of wild Mus musculus musculus: effects on chromosome pairing and recombination

An examination of the meiotic pattern of chromosome 1 isolated from a feral mouse population and containing a double insertion (Is) of homogeneously staining regions (HSRs) was carried out. The region delineated by the proximal breakpoint of Is(HSR;1C5) 1Icg and the distal breakpoint of Is(HSR;1E3)2Icg is desynapsed during the early pachytene stage and heterosynapsed at the midpachytene, as shown by electron microscopic analysis of synaptonemal complexes. The HSRs have no effect on the segregation of chromosome 1 in heterozygous mice. The lack of homosynapsis in the region under study causes chiasmata redistribution in heteromorphic bivalents. In normal males, single chiasmata are located in the medial part of the chromosome. In heterozygotes, this segment is heterosynapsed and unavailable for recombination. This leads to a significant decrease in the frequency of bivalents bearing single chiasmata. The total number of chiasmata per bivalent is much higher in heterozygous males than in normal ones. The recombination frequency between proximal markers fz and In also is higher in heterozygous animals. The increase in the total chiasma number in the heteromorphic bivalent is due to the addition of double chiasmata located mostly at precentromeric and pretelomeric regions of the chromosome.     

The effect of colchicine on meiosis in Lilium speciosum cv. “Rosemede”

Chromosome pairing and chiasma frequency were studied in meiocytes at diakinesis of Lilium speciosum cv. Rosemede fixed up to 21 days after the start of either continuous or 3 day pulse colchicine treatment. The two treatments gave similar results. In pulse treated pollen mother cells (PMCs) the mean chiasma frequency per cell fell from 26.4 in controls to 8.5 after fourteen days while the mean number of univalents per cell increased from 0.05 to 17.58. There was a negative correlation between mean chiasma frequency per bivalent and per PMC in colchicine treated buds; univalents were preferentially induced in bivalents with one chiasma, and preferentially excluded in bivalents with 4 chiasmata. Some chiasmata were redistributed to surviving bivalents despite the concurrent reduction in chiasma frequency per meiocyte. — Colchicine sensitivity began in premeiotic interphase and extended to mid or late zygotene in PMCs; ongoing synapsis was unaffected. However, susceptibility to univalency was asynchronous between bivalents occurring at zygotene in short chromosomes but at late premeiotic interphase in the longest chromosomes. The number of chiasmata per bivalent could be altered by colchicine without inducing univalents, but the ultimate effect was to reduce the number of chiasmata per bivalent (or per chromosome arm) directly to zero. The major factors determining the order and extent of reduced pairing and chiasma number were total chromosome length and arm length. Pairing and chiasma formation in embryo sac mother cells were less sensitive to colchicine than in PMCs, but their behavior was otherwise similar.     

Lampbrush Chromosomes in the Japanese Quail Coturnix coturnix japonica: Cytological Map of Macrochromosomes and Meiotic Crossing-over Frequency in Females

Cytological map of lampbrush macrobivalents of the Japanese quail (Coturnix coturnix japonica) were constructed. Investigation of chiasmata allowed to estimate the frequency of reciprocal genetic recombination (crossing over) in Japanese quail female meiosis. The total chiasma number in bivalents of Japanese quail oocyte nuclei was determined to be 53–58. Macrobivalents 1–5 and Z of the Japanese quail had on average 3.3 chiasmata per bivalent, and microbivalents, 1.0–1.1 chiasmata per bivalent. The chiasmata (crossover) frequency in Japanese quail females was lower than in chicken. In macrochromosomes of Japanese quail females, one crossover occurred per 43.9 Mb, and in chicken, per 30.0 Mb. Judging from chiasma frequency, the genetic length of the Japanese quail genome is likely to be 2650–2900 cM. Crossover frequency in the species was 0.023 per Mb in macrobivalents and 0.07–0.08 Mb in microbivalents and for the total genome, 0.041 crossing over per Mb. The genetic length of one Mb (recombination rate ) in female Japanese quails was 1.14 cM in macrochromosomes, 3.60–4.12 cM in microchromosomes, and about 1.96–2.15 cM averaged over the genome.     

The chiasmatic systems in populations of two spiny leaf beetles (Coleoptera,Chrysomelidae)

The number and distribution of chiasmata per cell at diakinesis has been analyzed in four populations ofDicladispa testacea (L.) and in two ofPolyconia caroli Lepr. both having 2n=16 and a 7+Xy_p male melotic formula. Most bivalents were unichiasmate and the distal chiasma position was the one most frequently encountered. One population ofDicladispa testacea could be distinguished from the remaining ones by the total number of chiasmata per cell. This intraspecific difference is discussed in relation with the degree of isolation between the four populations and other geographical characteristics.     

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)     

Recombination nodules in the oocytes of the chicken, Gallus domesticus

Chicken oocytes at pachytene were processed with the microspreading technique (Moses, 1977), and their synaptonemal complex (SC) complements were analyzed by electron microscopy. Ellipsoidal nodules, 140 X 120 nm in diameter, were associated with the central space of synaptonemal complexes. The average number of nodules per pachytene oocyte was 57.5. The number of nodules per bivalent showed a clear linear relationship with SC length, except for the microchromosomes, which showed a single obligatory nodule. The distribution of nodules along the 10 longest SCs was nonrandom, with low frequencies in the vicinity of kinetochores and high frequencies near the telomeres. The microchromosomes showed a single nodule whose average location was 1.21 micron from the kinetochore. In the ZW pair there was a single nodule whose average location was 0.31 micron from the paired telomeres and not more than 0.65 micron from them. The total number of nodules per cell and the number of nodules in each of the five major bivalents showed good agreement with the total number of chiasmata and the number of chiasmata of the major bivalents of roosters. Thus, these nodules share the characteristics of recombination nodules described in other organisms. The single, obligatory, strictly localized recombination nodule found in the pairing end of the ZW pair strongly suggests that recombination between the Z and W chromosomes in the female chicken is a regular process that may be similar to the obligatory recombination between the pairing ends of the human X and Y chromosomes that was recently described in studies using DNA probes.     

The relationship between chromatid interference and the mapping function.

A formula for a general mapping function involving only the number of chiasmata between two points on the chromosome and the average excess of four-stranded double exchanges over two-stranded double exchanges is derived. A formula is also given for deriving the proportion of three-stranded double exchanges after any given number of chiasmata; this is related to mapping for linkage with the centromere. It is shown that this proportion tends to a limit of 2/3 after a large number of chiasmata, whether or not chromatid interference is operating.     

Competition for chiasmata in diploid and teisomic maize

Summary The addition of an extra chromosome, in particular the short arm of chromosome V, inZea Mays is shown to lead to (a) less competition for chiasmata between the chromosomes, (b) an increase in the number of chiasmata formed. These effects are in accordance with the hypothesis of control by an effective upper limit to the number of chiasmata which may be formed in any nucleus. The addition of an extra chromosome raises the upper limit of the number of possible chiasmata. This is further supported by the fact that the increase in chiasma formation is not confined to the trisomic chromosome but is, as far as can be judged, shared by all the bivalents. The bearing of these findings on chiasma frequency studies of polyploid series is briefly discussed.     

A mathematical framework for examining whether a minimum number of chiasmata is required per metacentric chromosome or chromosome arm in human

We introduce a piecewise linear regression called "hockey stick regression" to model the relationship between genetic and physical lengths of chromosomes in a genome. This piecewise linear regression is an extension of the two-parameter linear regression we proposed earlier [W. Li and J. Freudenberg, Two-parameter characterization of chromosome-scale recombination rate, Genome Res., 19 (2009) 2300-2307]. We use this, as well as the one-piece regression with a fixed y-intercept, to compare the two competing hypotheses concerning the minimum number of required chiasmata for meiosis: minimum one chiasma per chromosome (PC) and per chromosome arm (PA). Using statistical model selection and testing, we show that for human genome data, one-piece PC (PC1) is often in a statistical tie with two-piece PA model (PA2). If an upper bound for the segmentation point in two-piece regression is imposed, PC is usually the preferred model. This indicates that a presence of more than one chiasmata is rather caused by the relationship between chromosome size and chiasma formation than by cytogenetic constraints.     

Chromosome association and pollen fertility of parental and interspecific hybrids of Lycopersicon esculentum X L. hirsutum and L. pennellii

Cytological studies were carried out on two wild species (L. hirsutum and L. pennellii) and the cultivated species (L. esculentum) of tomato and their F1 hybrids. Both parents and hybrids show a diploid chromosome number of 2n=24. The meiotic behaviour of the cultivated species showed a high degree of chromosome homology resulting in a high level of chiasmata frequency per bivalent. In contrast, the two wild species showed a slight increase in uniyalent frequency and a decrease in bivalent formation and chiasmata frequency. The meiotic behaviour of the hybrids showed a high level of univalents and low levels of bivalents as well as trivalents. Highly significant decreases in chiasmata frequency and increases in meiotic abnormalities, especially in the L. esculentum X L. pennellii hybrid, also were detected. The high meiotic irregularity and low chiasmata frequency recorded in the second hybrid indicated the disharmony and difference between its parental genomes and also served to predict its sterility. With regard to degree of pairing recorded in the hybrids, there is a possibility that sterility in such cases may refer to genetic factors in addition to the previously mentioned reasons. Pollen fertility showed no great difference between L. esculentum and L. hirsutum and their F1 hybrid, but a significant decrease was recorded in the L. esculentum X L. pennellii hybrid, which was clearly associated with high meiotic irregularity, low chiasmata frequency and chromosome association.     

Relational coiling and chiasma frequency

Conclusions The amount of relational coiling of chromatids during the prophase of mitosis and the average number of chiasmata were determined in the largest chromosome of five species of locusts. The statistical analysis of the data obtained suggests that probably there is a correlation between the amount of relational coiling per unit of chromosome length and the number of chiasmata. A similar suggestion was put forward bySax (1936) andHusted (1938), which was based upon observations made in different plant species. Though the difficulties encountered during the present cytological analysis of the chromosome structures, as well as the small number of species investigated, do not allow us to draw a definite conclusion, nevertheless it seems probable from our data that a greater degree of relational coiling of chromatids during mitosis is accompanied by a higher chiasma frequency during meiosis. The correlation furthermore indicates also that the physical basis of the tendency of chromatids, and that of paired chromosomes, to twist round one another relationally may be the same.Fellow of the Rockefeller Foundation.     

Metaphase I orientation of Robertsonian trivalents in the water-hyacinth grasshopper, Cornops aquaticum (Acrididae, Orthoptera)

Trivalents resulting from polymorphic Robertsonian rearrangements must have a regular orientation in metaphase I if the polymorphisms are to be maintained. It has been argued that redistribution of proximal and interstitial chiasmata to more distal positions is necessary for a convergent orientation, the only one that produces viable gametes. Cornops aquaticum is a South-American grasshopper that lives and feeds on water-hyacinths, and has three polymorphic Robertsonian rearrangements in its southernmost distribution area in Central Argentina and Uruguay. The orientation of trivalents in metaphase I, the formation of abnormal spermatids and the frequency and position of chiasmata in the trivalents, was analysed in a polymorphic population of C. aquaticus. In this study we observed a correlation between the number of trivalents with the frequency of abnormal spermatids; additionally, the number of chiasmata, especially proximal and interstitial ones, was strongly correlated with the frequency of the linear orientation. Therefore we confirmed our previous assumption, based on other evidence, that the chiasmata redistribution in fusion carriers is essential to the maintenance of the polymorphisms.     

Distribution of chiasmas in the 2d and 6th chromosomes involved in Robertsonian translocations in male mice

The distribution of chiasmata in 2 and 6 chromosomes in males homozygous for Rb(2.6)4Iem and Rb(8.17)1Iem was studied. Chiasmata were shown to distribute along chromosomes non-randomly, exchanges occurring in telomeric regions. Chiasmata distribution is substantially different for the cases of one and two chiasmata per bivalent. The main cause for these differences is supposed to be strong positive chiasmata interference (the position of the first chiasma may determine the position of the second one). The centromere blocks this effect, so chiasma in one arm does not interfere with that in the second arm. It has been shown that the frequency of double exchanges depended on not only the distance between markers under study, but also on marker position in the chromosome.     

Nucleotypic influences on chromosome-specific chiasma variation in Crepis capillaris. I. Responses to early colchicine treatment and chromosome doubling.

Additive and multiplicative effects of colchicine treatment at the seedling stage and of chromosome doubling on chromosome-specific chiasma frequency at metaphase I have been measured in comparisons between C0 and untreated diploids and between C0 autotetraploids and C0 diploids. Early colchicine treatment increases the frequency of chromosome C univalents to 1.8% but has no similar effect on chromosomes A and D. Colchicine treatment has little net effect on mean chiasma frequency, deducting an average of 0.204 chiasmata per set but otherwise multiplying the mean by a factor of 1.182. These additive and multiplicative effects represent averages of six phenotypes. Chromosome doubling in a tetraploid-diploid chimaera subtracts an average of 0.265 chiasmata per set but otherwise doubles the numbers of chiasmata at the diploid level (x 2.134). Comparison of six diploids and tetraploids reveals modest average additive (+ 1.103) and multiplicative effects (x 1.190). The implications of these findings are discussed in the light of new analyses of previously published data. Key words : chiasmata, Crepis, colchicine, meiosis, polyploidy.     

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.