Polymorphism involving heterochromatic segments in Metrioptera brachyptera

A complex pattern of polymorphism involving terminal heterochromatic segments on L₃ and L₄ chromosomes has been uncovered in eight populations of Metrioptera brachyptera. There are individuals in every population which carry reduced segments on one or both L₄'s. In six populations, enlarged heterochromatic segments have been encountered on the L₃ chromosomes in some individuals. The L₄ system is almost certainly stable although the frequency of L₄ karyotypes does not conform to a Hardy-Weinberg distribution in all populations. Stability of the L₃ polymorphism could not be ascertained. A reduction of L₄ heterochromatin leads to a significant rise both in mean cell chiasma frequency and between cell variance. The effect on chiasma frequency is transchromosomal. The normal pattern of strict chiasma localisation tends to be disrupted in germ lines which include modified L₄ chromosomes. There is a reduction in the number of proximal and distal chiasmata and an increase in the frequency of interstitial ones. It is proposed that the standard L₄ heterochromatin may function in conserving heterozygosity and promoting uniformity between parent and offspring. Partial removal may lead to an effective increase in recombination and produce a greater diversity of genotypes for selection to act upon.     

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.     

Spontaneous chromosome mutations in Truxaline grasshoppers

Three distinct mutant conditions are described in single male individuals from three species of short horn grasshopper. Of these, one is an entire germ line mutant of Myrmeleotettix maculatus, heterozygous for a centric fusion between single M₄ and M₅ telocentric chromosomes. In contrast, the remaining two mutants are present in mosaic form. One is heterozygous for an L₁-M₄ interchange in Omocestus viridulus, the other tetrasomic for the M₄ chromosome in Chorthippus parallelus which in addition is characterised by the inclusion of a supernumerary heterochromatic segment on one S₈ homologue. Centric fusion in Myrmehotettix maculatus has neither disturbed the chiasma potential of the elements constituting the fusion multiple nor, has it apparently influenced the production of balanced gametes. The pattern of chiasma formation in the L₁-M₄ interchange multiple lends support to the contention that the process of chiasma formation originates near the distal end of chromosome arms in Omocestus viridulus. There is no interaction between the two mutant conditions of tetrasomy and the presence of supernumerary segments in Chorthippus parallelus. Moreover, because of the precocious nature of two of the four M₄ homologues there is little tendency to form multivalents. The two M₄ bivalents share a similar mean chiasma frequency.     

Parallel polymorphism for supernumerary segments inChorthippus parallelus (zetterstedt)

A complex and parallel pattern of polymorphism for heterochromatic supernumerary segments in the M₇ and S₈ chromosomes has been found in 14 populations of the meadow grasshopperChorthippus parallelus. Nine distinct karyotype classes for these two chromosome pairs occur though they are not equally represented in different populations. Populations differ also with respect to the frequency of supernumerary segments they contain. In all populations the presence of supernumerary segments leads to a significant elevation of mean cell chiasma frequency compared to individuals from the same population lacking such segments. The extent of the effect appears to differ in different populations. The observed frequencies of S₈ karyotypes conform to the expectations of a Hardy-Weinberg distribution. Those of the M₇, however, do not, and in all but one of the 14 populations there is a significant excess of homokaryotypes. In the Ashurst population 26% of the individuals sampled were characterised by germ-line polysomy for the M₄ chromosome, either in the form of entire tetrasomics or more usually as mosaics ranging from tri- to hepta-somics. In all these polysomics the M₄ chromosomes in excess of two were regularly heteropycnotic at first meiotic prophase from zygotene to diakinesis. As a consequence of this multivalents are rare. Extra M₄ chromosomes do not modify the chiasma characteristics of the other chromosomes in the complement. Nor do they modify the action of the supernumerary segments in any way.     

The supernumerary segment system of Stethophyma

Three species of the genus Stethophyma have been cytologically examined and all three show variation both for supernumerary heterochromatic segments and for the distribution of standard heterochromatin among the autosomes. The European species, S. grossum, for example, shows considerable interpopulation variation for standard heterochromatin while two of the populations, from Spain and Austria, show supernumerary segment polymorphism. The segments are located interstitially on the S₁₁ chromosome but occupy different positions in the different populations. — In all species, the presence of the extra heterochromatic segments increases the mean chiasma frequency. Moreover, the influence of the segments upon mean chiasma frequency is different in different populations and in different species. In the Spanish population, the increase is both intra- and interchromosomal whereas in Austria the influence of the segment is completely interchromosomal. — In the American species, S. gracile and S. lineatum, where supernumerary heterochromatic segments are carried on both S₁₀ and S₁₁ chromosomes, the effect on chiasma frequency shows a dosage relationship, an increase in the number of segments per individual being correlated with an increase in mean chiasma frequency. It is suggested that the interstitial segments found in all species have originated by direct duplication of chromosome material. By contrast the terminal segments in S. lineatum and S. gracile may be derived by translocation from a B-chromosome since such a chromosome has been found in one individual of the former species. — The variation in segment structure and the distribution of standard heterochromatin, among the European species of S. grossum suggests that these systems have evolved independently in different populations.On educational leave from the Forest Research Laboratory, Fredericton, N. B. Canada.     

An analysis of frequencies of chromosome configurations in wheat and wheat hybrids

Presynaptic association of homologous chromosomes is a prerequisite to the sequence of events that lead to chiasma formation. This association of homologous chromosomes, as entire units, occurs with probability a, and chiasma formation occurs independently in opposite chromosome arms with probability c. a and c have been estimated from frequencies of different chromosome configurations at metaphase I of euhexaploid wheat and several derived lines. In the euploid, a is essentially unity and c is of the order of 95%. All changes in the aneuploidy studied involved c rather than a, whereas the change induced by colchicine application primarily involved a.—Observed and expected frequencies of configurations were compared in wheat hybrids in which only homoeologues were present. The expected frequencies of configurations were estimated from the data, based on a being unity for entire groups of homoeologues and c being the probability of chiasma formation between random homoeologous arms. Observed and expected frequences of configurations were in general agreement; however, an excess of observed closed bivalents at the expense of multivalents is interpreted to mean that not all homoeologues are effectively associated in all cells.—In euhexaploid wheat, we suggest that homologues associate with almost certainty, whereas homoeologous pairs of chromosomes associate less efficiently. The aneuploidy examined in this study does not appear to affect the association of chromosomes, but rather the number of chiasmata that eventuate and, in the case of deficiency of chromosome 5B, the distribution of chiasmata within homoelogues, perhaps by way of rendering sites for chiasma formation of homoelogues more similar.     

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.     

Pseudo-multiple formation as a consequence of prolonged non-homologous chromosome association in Metrioptera brachyptera

Multiple configurations involving the L ₃ and L ₄ homologues have been observed in all individuals from a population sample of Metrioptera brachyptera. These associations which persist up to and including metaphase of the first meiotic division are non-homologous and achiasmate in character. They are conditioned by the persistent association of large distally located heterochromatic blocks on the L ₃ and L ₄ chromosomes and are not, as White has proposed, the result of crossing over in translocated terminal duplications. When the L ₃ and L ₄ chromosomes form bivalents the distal heterochromatin restricts crossing over in this region and chiasmata are localised proximally. In other members of the complement chiasmata are localised at either or both the centric and telomeric ends. A relationship is demonstrated between the pattern of chiasma localisation and that of chromosome pairing.     

Studies on trisomics in a Jute hybrid

Fifteen red pigmented trisomics were isolated in the F₂ generation from the cross Corchorus olitorius L. x C. capsularis L. In the F₃ generation a few green trisomics were obtained; more of these were isolated from the backcross generation. A detailed morphological and cytological analysis of the trisomic hybrid populations derived from the F₃ and F₄ generations is presented. The trisomics were scored for several morphologically differentiating characters and most of them were intermediate between the parental species, a few resembling the olitorius parent more. Cytological studies showed the formation of abnormal sporads in trisomics with different frequencies indicating a misdivision at meiosis. This imbalance at meiosis results in a higher percentage of pollen sterility in the trisomics as compared with the parents. Analysis of M₁ of meiosis showed that there were: differences in the frequencies of the various chromosome configurations between the two categories of trisomics; significantly higher trivalent frequencies per PMC in the green trisomics; in contrast significantly hihger univalent frequencies per PMC in the red trisomics. No significant difference in chiasma formation was observed between red and green trisomics, nor between trisomics and their parental species. It appears that segmental homology in the parental chromosomes has probably resulted in varying degrees of preferential pairing in the trisomic hybrid.     

Effects of centric-shift polymorphisms on chiasma conditions inTrimerotropis pallidipennis (Oedipodinae:Acrididae)

Two populations ofTrimerotropis pallidipennis from Mendoza (MA) and Uspallata (UA) (Argentina), polymorphic with respect to centric shifts and B-chromosomes, were cytologically analysed. The medium-sized chromosomes from both populations were polymorphic for 7 (MA) and 5 (UA) inversions respectively. A clear tendency towards chiasma localisation was evident in both populations although they differ significantly with respect to mean frequencies of both total and interstitial chiasmata. The analysis of these frequencies in relation to the polymorphisms showed that in UA, the inversions do not affect chiasma formation while in MA, total chiasma frequency is negatively correlated in a significant fashion with the number of heteromorphic bivalents. This decrease is related, in non-B-carriers, to a significant increase of interstitial chiasmata. Therefore a redistribution of chiasmata, which could increase genetic recombination, is evident. The successful maintenance of polymorphisms in each population might depend, in part, on their effects on recombination, which could be related with the local requirements of genetic variability.     

Pseudomultiple production in Ageneotettix deorum deorum

Meiosis in Ageneotettix deorum deorum is characterised by extensive pseudomultiple formation during prophase. The association of non-homologous chromosomes takes place prior to pairing and chiasma formation and occurs to a varying degree in all prophase cells. These pseudomultiples originate during interphase as a consequence of the association of heterochromatin. All autosomes carry procentric heterochromatic segments of variable size and the L₁, L₃ and M₅ chromosomes also possess terminal heterochromatic regions. The association of non-homologous chromosomes during zygotene and pachytene does not appear to impede pairing or the frequency and distribution of chiasmata. — A majority of the pseudomultiples dissociate after diakinesis, during orientation and congression on the spindle. However in 4% of the cells examined, associations, mainly quadrivalents, persist through metaphase. — Heterochromatic associations of non-homologous chromosomes are again evident during second prophase, though here they involve only the centric heterochromatic regions; 9% of these associations persist through second metaphase. — The nature and behaviour of the pseudomultiples in Ageneotettix are pertinent to the interpretation of terminal associations in other Orthoptera and provide evidence that persistent associations can arise following a non-chiasmate association.On educational leave from the Forest Research Laboratory, Fredericton, N.B. Canada.     

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.     

Unstable B chromosomes in Silene maritima With. (Caryophyllaceae)

Unstable B chromosomes in Silene maritima With. (Carophyllaceae). B chromosomes have been found in 7 out of 39 populations of Silene maritima With. (Carophyllaceae) studied. These 7 populations are all from the Norfolk coast. The B chromosome is small, 0.8 μm long, telocentric and appears euchromatic. Within plants the B chromosome is highly unstable with different numbers in cells of the same root or pollen mother cells (pmcs) of an anther. The transmission of the Bs is efficient and in controlled crosses progeny plants usually have higher mean B frequencies than the B-containing parents. At metaphase I the pairing behaviour of the Bs is variable and they often appear as univalents. In general the presence of Bs in pmcs results in an increase in chiasma frequency/cell and populations with Bs have higher mean chiasma frequencies than those without Bs. Cloned plants grown under stress conditions induced by the addition of NaCl to a nutrient solution showed no difference in mean B number/cell or distribution when compared with controls grown in nutrient solution only.     

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 B-chromosome system of myrmeleotettix maculatus (thunb.)

It is established that in populations of Myrmeleotettix maculatus with B-chromosomes these chromosomes occur at stable frequencies and are present to the same extent in both the males and the females of the same population. It is also established that the B-chromosome content of a population is positively correlated with its chiasma frequency and that, within a population, individuals with single B-chromosomes tend to have higher chiasma frequencies than individuals lacking B-chromosomes. Since this effect is not increased by the addition of further supernumeraries it is argued that selection operates in favour of individuals with single B-chromosomes. Finally it is shown that the level of B-chromosomes in a population is related to temperature and especially to rainfall so that B-chromosomes are absent from populations in climatically stringent environments.     

Incipient Genome Differentiation in Gossypium. I. Chromosomes 14, 15, 16, 19 and 20 Assessed in G. HIRSUTUM, G. RAIMONDII and G. LOBATUM by Means of Seven a-D Translocations

The genus Gossypium is favorable for study of genome divergence at several levels. Early stages of divergence have been studied among four D genomes by comparing chiasma frequencies (reciprocal exchanges) between pairs of genomes and between individual counterpart chromosomes marked by heterozygous translocations. D₅ (G. raimondii) shows barely detectable differentiation from from D_h (G. hirsutum), whereas D₇ (G. lobatum) is considerably less closely related to D_h than is D₅. Fragmentary data suggest that D_2–2 (G. harknessii) falls between D₅ and D₇ in its relationship to D_h. Since chiasma frequencies in individual chromosomes and marked regions exhibit the same order of relationships as their corresponding whole genomes, it is concluded that the genome differentiation is generalized (i.e., nucleus-wide) rather than localized in specific chromosomes or chromosome regions. Estimates of relationships based on reciprocal exchange frequencies agree with those based upon preferential synapsis in allohexaploids reported previously. Since preferential synapsis and reciprocal exchange frequencies reveal the same order of relationships, it is concluded that to some extent they reflect common underlying changes in chromosome properties, despite recent evidence that synapsis and crossing over are under independent genetic control.     

The cytology of the genus Briza L. (Gramineae)

Meiosis has been studied in five European and four South American species of Briza. The various species have different chiasma frequencies and different patterns of chiasma localisation. Population studies show that there are significant differences in mean plant chiasma frequency between populations of B. maxima whereas B. media and B. spicata do not show interpopulation differences. B. media is shown to have diploid and autotetraploid races and the distribution of these chromosome races in Europe has been plotted. The South American species studied are all bivalent forming tetraploids and would therefore appear to have an allopolyploid origin. Interchange heterozygosity has been found in B. media and B. elatior, different populations of B. media have been shown to be heterozygous for different interchanges. These topics are discussed in relation to the regulation of recombination.     

The B-chromosome system of the grasshopper Eyprepocnemis plorans subsp. plorans (Charpentier)

Supernumerary chromosomes of two types have been observed in the grasshopper Eyprepocnemis plorans subsp. plorans. One of these (the B-type) is similar in size to an S autosome; the other is smaller (B-type). Both are telocentric and mitotically stable. The frequencies of individuals with the B-type supernumeraries in five natural populations were 56, 56, 70, 71 and 30 per cent respectively. The equivalent levels of the B-type supernumerary were 0, 0, 13, 3 and 0 per cent respectively. Because of the relative infrequency of the B-type only the B-type has been studied in detail. In males with 1B, anaphase I segregation of X and B was random in four populations but non-random in that from Otivar. Here the B was distinctive in having a secondary constriction near the centromere. A study of chiasma frequency among A-chromosomes revealed that the B-type supernumerary increases significantly both the mean chiasma frequency and the between-cell variance. A comparison of body morphometrics failed to reveal any effect of these B-chromosomes on the exophenotype.     

Chiasma frequency effects of structural chromosome change

Three structural chromosome changes in the plant Hypochoeris radicata 2n = 8 have been tested for their effects on chiasma formation: (1) centric fission of chromosome 1, (2) a whole arm exchange between chromosomes 1 and 3, and (3) an interchange between the long arm of chromosome 1 and the short arm of 2 which gives an effectively three-armed pachytene multiple. Mean chiasma frequencies were compared between full-sibs in families segregating for the rearrangements. In each family the chiasma frequency was higher in heterozygotes than basic homozygotes. The size of the chiasma increase is dependant on the number of additional potentially-paired segments in the complement at pachytene. Fission heterozygotes and 1/2 interchange heterozygotes, with one extra pairing region, both form about 0.45 more chiasmata per PMC than full-sib basic homozygotes. The 1/3 exchange, with two additional pairing regions, increases chiasma frequency by twice this, about 0.85 per PMC. Individuals homozygous for the centric fission maintain the raised chiasma level. The chiasma increase appears limited to the chromosome(s) affected by structural change with no detectable interchromosomal effect.     

Chromosome distribution and catenation in Rhoeo spathacea var. concolor

The twelve chromosomes of Rhoeo spathacea variety concolor are arranged in a definite sequence in a ring at meiosis. Identification of all the 12 chromosomes was possible in 119 diakinesis and metaphase I cells. — Pollen viability was measured to be 36.54% by cotton blue staining procedure. Forty five of 56 metaphase I cells (80.36%) had adjacent distribution. Each of the 12 chromosomes was equally likely to be involved in adjacent distribution regardless of their sizes and heterobrachialness. Adjacent distribution occurred randomly at each arm-position in the ring regardless of the lengths of the arm-pairs. — The most frequent chromosome configuration at diakinesis and metaphase I was a chain-of-12 chromosomes (41.18%). Cells with 1 to 4 chains of chromosomes were observed. The observed frequencies of various configurations were in good agreement with the calculated frequencies. The mean number of chiasmata was 10.90 per cell and 0.908 per pair of chromosome arms. The 131 chiasma failures were distributed at random among the 12 arm-positions. Since the lengths of arm-pairs in the ring vary, the randomness may mean that chiasma formation was limited to short terminal segments on all chromosomes.