The effect of Actinomycin-D on chiasma frequency in Vicia faba

The effect of Actinomycin-D was studied on chiasma frequency in Vicia faba (var. Weibull's Akerabona). A significant decrease in chiasma frequency over control was observed in all cases irrespective of the time of fixation and chemical concentration. However, the maximum decrease was found in the material fixed after a recovery period of 24 hours. The decrease in chiasma frequency showed a linear relationship with time of recovery. Besides reduction in the chiasma frequency, cells showing complete failure of homologous pairing were also observed. These findings are discussed in the light of actinomycin-D action.     

Chiasma frequency variation inCepaea hortensis (Müll.) and a comparison withC. nemoralis (L.)

Chiasma frequency variation has been studied in 12 populations ofCepaea hortensis and the results compared with those previously obtained forC. nemoralis from the same populations. Variation in chiasma frequency is effectively restricted to a single large bivalent. In both species there are significant differences between populations in chiasma frequency. Although mean population mean chiasma frequencies are similar inC. hortensis andC. nemoralis, C. hortensis is less variable thanC. nemoralis. This can be seen, to a greater or lesser extent, at several levels-(i) within cells, (ii) between cells, (iii) between snails, and (iv) between populations-and is most noticeable when interstitial chiasmata are considered separately from terminal chiasmata. Data from 6 populations suggest that chiasma frequency remains stable between successive years. It is suggested that, since sympatric populations were studied, the most likely explanation for differences in chiasma frequency statistics between the two species is thatC. hortensis andC. nemoralis are responding differently to similar selective pressures.     

Genetic Control of Meiosis in Rice, ORYZA SATIVA L. III. Effect of DS Genes on Genetic Recombination

The recombination frequency as influenced by five independent recessive ds genes was measured on three segments of different chromosomes of rice, Oryza sativa L. Each ds gene in the homozygous condition resulted in an almost equally reduced recombination frequency in the three segments. When the mean reduction in recombination frequency was related to the reduction of chiasma frequency, the five ds genes were divided into two types: in one type the reduction of chiasma frequency almost corresponded to the mean reduction of recombination frequency, and in the other the chiasma frequency was greatly reduced in comparison with the mean reduction of recombination frequency. Three of the five ds genes were found to belong to the former group. In both types, normal synaptonemal complexes were observed in pachytene cells homozygous for ds genes. This finding suggests that ds genes do not affect the formation of synaptonemal complexes which are regarded as the prerequisite structure for crossing over.     

Chromosome-specific control of chiasma formation

A desynaptic mutant of Hypochoeris radicata, 2n = 8, has been found in a population from France. The mutant is remarkable in that at metaphase-I over 90% of PMC's have a pair of univalents while two pairs are found in only 1% of cells. Only chromosome IV is affected by the desynapsis which is controlled by a single recessive gene. Bivalent chiasma frequency in cells with a pair of univalents is higher than in cells with complete bivalent formation indicating partial dependence of cell chiasma number on the availability of some factor within the anther. The F₂, produced by full-sib mating, has a raised frequency of univalents in the chromosomes other than pair IV. The higher level results from enforced sib-mating of an obligate outbreeder and is not related to the action of the major gene. It is suggested that chiasma formation is controlled both polygenically and by major genes which operate on a hierarchical system. Some genes affect the behaviour of the entire complement while the action of others is specific to individual chromosomes.     

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.     

Phase transformation and chiasma frequency variation in locusts

The hypothesis, proposed by Nolte, that all locusts undergo an increase in their chiasma frequency when swarming is examined in the Migratory Locust, Locusta migratoria, using (1) animals which have been isolated in the laboratory at hatching, (2) crowded and isolated animals supplied by an independent laboratory and (3) field solitaria material. Animals isolated in the laboratory at hatching did not show a reduced chiasma frequency when compared with crowded controls despite the fact that they did develop a solitaria phenotype. Likewise, crowded and isolated animals supplied by workers at the University of Paris showed no difference in their chiasma frequency. Finally, animals living at very low density on the flood plains of the River Niger in Mali, West Africa, and exhibiting extreme phase solitaria characteristics proved to have very high chiasma frequencies which is the exact antithesis of the hypothesis. — Thus there is no evidence that there is any relationship between phase status or density and chiasma frequency in Locusta migratoria. This is in direct contradiction to Nolte's claims but in agreement with a previous study by the present author on Schistocerca gregaria.     

Asynapsis in Lolium perenne

Eighteen plants displaying varying degrees of asynapsis ranging from weak to very strong were found among four out of six populations of Lolium perenne L. (2n=14) which had been subjected to three cycles of directional phenotypic selection for productivity of green material. No plants were found displaying univalents in the original generation but the incidence increased with cycles of selection, indicating the genetic control and differential distribution of asynaptic genes among these populations. — The analysis of univalents and chiasma frequency of pollen mother cells (PMC) of six partially asynaptic plants chosen for detailed study revealed that univalents occurred throughout all PMC chiasma classes irrespective of chiasma frequency, but the higher the chiasma frequency of any PMC the less the likelihood of univalents occurring. The relationship between chiasma frequency and univalent frequency per PMC per plant was negative. — Mean chiasma frequency per bivalent increased for the asynaptic cells in comparison with the normal in both the weak and medium asynaptic groups which was explained by the availability of additional chiasmata for redistribution.     

Sex differences in meiosis between Viciafaba L. and its close wild relatives

Comparison of chiasma frequency was made in male and female meiocytes of the widely cultivated Vicia faba L. and its close wild allies in the Vwia narbonensis species complex. Contrary to most of the earlier reports of higher chiasma frequency in female sex cells, a higher chiasma frequency was observed in male sex cells. This pattern is restricted to the species representing the Narbonensis complex, since Vicia faba shows identical chiasma frequency in male and female sex cells. The genetic significance of these findings is that chiasma formation in the sex cells of the putative allies is governed and regulated by similar controlling systems of genes, but that of V. faba indicates a different kind of system.     

Population cytology of the genus Phaulacridium

Twenty seven populations of the grasshopper Phaulacridium marginale were sampled from South Island, New Zealand. An analysis of the chiasma frequency data showed that populations differed significantly for this metric. The populations appeared to fall into two distinct size groups. Morphological measurements confirmed this and a reanalysis of the chiasma frequency data revealed that although the two groups differed significantly for both mean cell chiasma frequency and between cell variances, there was no difference between populations within a group. Analysis of climatological data suggests that the “larger” individuals show different environmental (habitat) preferences from the smaller, which are restricted to one region of South Island. The taxonomic status of Ph. marginale as a single species is called into question.     

Influence of B-chromosomes on meiosis in pearl millet

B-chromosome behaviour and the effects of B's on the endophenotype were studied in the third back-cross progeny of a cross between non-B and B-carrying parents ofPennisetum typhoides Stapf et Hubb. In the experimental material a regular increase in B-chromosome chiasma frequency and an increase in the variation of mean chiasma frequency with increased number of B's was observed. When four or more B-chromosomes were present quadrivalents occurred more frequently. In these respects the experimental material differed from the B-carrying parent. B's had no effect on the mean chiasma frequency of A-chromosomes when present in numbers of up to and including four but when present in numbers of more than four had a depressing effect. The variation of the mean chiasma frequency of the A-chromosomes increased with increasing number of B's. No correlation was observed between A-chromosome chiasma frequencies and B-chromosome chiasma frequencies. In the experimental material B's when present in larger numbers i.e. more than four had deleterious effects on A-chromosome behaviour and on fertility. In the effects of B's on the endophenotypic characters the experimental material differed from the B-carrying parent. It is suggested that the behaviour and effects of B's on the endophenotypic characters are the result of interaction between the A-chromosome genotype and the B-chromosomes.     

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.     

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.     

The Relationship between Chiasmata and Crossing over in TRITICUM AESTIVUM

Telocentrics for the β arm of chromosome 4A and the long arm of 6B were used as cytological markers for the determination of chiasma frequency. In concomitant studies of recombination, terminal segments of rye and T. umbellulatum chromatin carrying Hp (Hairy peduncle) and Lr9 (Leaf-rust resistance), respectively, marked 4A and 6B. Two temperatures, 21° and 32°, were used for both the 4A and 6B experiments.—Only one chiasma was observed in each heteromorphic bivalent. Because there was a substantial reduction in pairing between diakinesis and metaphase I, all determinations of chiasma frequency were made at diakinesis. In the 21° experiments, agreement was good between genetic recombination and cytological prediction on the basis of the partial chiasmatypy hypothesis that each chiasma represents a crossover. At 32° both chiasma frequency and crossing over, but particularly the latter, were strongly reduced. The fewer crossovers than expected are explained in part by stickiness of chromosomes at the high temperature, sometimes resulting in adjacent chromosomes being wrongly scored as having a chiasma, and in part by premetaphase disjunction of some recombined bivalents and subsequent independent behavior of the two resulting univalents.—Male transmission of the 4A telocentric from the heteromorphic bivalent was unusually high: 51% at 21° and 31% at 32°.     

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.     

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.     

Variation in chiasma frequency during tulip anther development

In tulip anthers meiosis begins first in the pollen mother cells next to the point of entry of the vascular bundle at the base. It then proceeds in a wave to the free end of the anther. The first cells to pass through meiosis have the lowest chiasma frequency and cells with a lower cell chiasma frequency have a lower total bivalent length. It is suggested that some factor which passes through the vascular bundle may cause the developmental variations in chiasma frequency possibly by first altering chromosome length (Summary see p. 179.)     

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.     

Chromosomal diversity and morphological dimorphism in Moroccan wild oat, Avena agadiriana

Chromosomal diversity and morphological dimorphism were examined by breeding the intraspecific hybrids among six accessions of Avena agadiriana Baum et Fedak. The accession M55 occasionally showed a quadrivalent, which indicated that it was heterogeneous for a reciprocal translocation. The chromosome pairings in the intra-early-flowering ecotype hybrids were almost normal with high chiasma frequency at metaphase I. However, the intra-late-flowering ecotype hybrids showed a quadrivalent or a trivalent with a univalent, similar to the inter-ecotype hybrids. The marginal population of the northeastern end, M74, always showed a quadrivalent or a trivalent in all the hybrids due to a unique large reciprocal translocation. This result was consistent with the late-flowering ecotype. There was a genocline of the chiasma frequency resulting from chromosomal rearrangements between adjacent populations. There was also a significant negative correlation between the mean chiasma frequency and the annual rainfall in each collection site (r = ?0.880). The two distinct ecotypes were characterized by the different magnitude and number of chromosomal rearrangements, reciprocal translocation and loss of satellite chromosomes, and they were adapted to the wide range of environments along the Atlantic coast of Morocco.     

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.     

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.