Genotype-dependent effect of B-chromosomes on chiasma frequency in Eyprepocnemis plorans (Acrididae: Orthoptera)

In order to analyze the effect of B-chromosomes on chiasma frequency, the offspring of different females of the grasshopper Eyprepocnemis plorans have been studied. From the comparison between individuals of the different families having a given number of B's and between individuals with different numbers of B-chromosomes within the same family, it can be concluded that the effect of the genetic background on chiasma frequency is greater than that produced by the presence of B's, and that there is a between-families variation in the effect of B-chromosomes which, in addition, is negatively correlated with the chiasma frequency of related individuals without B's. This genotype-dependent effect of B-chromosomes is discussed.     

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.     

Genetic and environmental components of chiasma control

Variation in chiasma frequency within and between individuals has been investigated in Schistocerca gregaria and Stethophyma grossum. By taking sequential samples of the same testis in S. gregaria it has been demonstrated that there is considerable variation in chiasma frequency between times, within individuals, and that this variation is not evenly distributed between individuals of the same population. The response and recovery patterns after exposure to successive temperature treatments also indicate a differential sensitivity of individuals within the experimental population. — An analysis of interfollicular variation in chiasma frequency has revealed significant differences between follicles within individuals both in S. gregaria and a Spanish population of S. grossum. A comparable analysis on individuals of S. grossum carrying supernumerary segments shows that the presence of such segments increases the amount of variation between follicles within individuals and also between individuals within the population. — The distribution of cell chiasma frequencies in S. gregaria is normal whereas in Stethophyma cell frequencies approximate to a Poisson distribution. The structurally different supernumerary segments, present in Austrian and Spanish populations of S. grossum, both increase mean chiasma frequency and they modify the between cell variance in different ways — the former interchromosomally and the latter intrachromosomally. — The differences in chiasma frequency between follicles and between times within individuals and the differential reaction to heat shock reflect differences in the pattern of genotype-environment interaction. Similarly it can be argued that the presence of polymorphisms involving supernumerary material may play an equivalent role by increasing the between cell or between bivalent variance within individuals, a unique form of genotype-environment interaction at the endophenotypic level. — Such variation in genotype sensitivity to environmental modification has important adaptive value especially in organisms which are subject to recurrent, often random, changes in their environment.     

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.     

Population cytology of the genus Phaulacridium

Samples were taken from twenty three Australian mainland populations of the shorthorned grasshopper, Phaulacridium vittatum. Five of these populations were found to be polymorphic for B-chromosomes. The anaphase I segregation of the B-chromosome appeared to be random with respect to the X (contra Jackson and Cheung, 1967). The B-chromosomes did not appear to influence chiasma frequency in the population in which they were found.     

Presence and behaviour of B-chromosomes in Acanthophyllum laxiusculum (Caryophyllaceae)

Two karyotypic variants were recognized in populations of Acanthophyllum laxiusculum Schiman-Czeika. Variant A had 2N = 30 and variant B showed the presence of 0 to 3 B-chromosomes in addition to 2N = 30 chromosomes. Analysis of chromosome behaviour at meiosis showed that the presence of B-chromosomes increases chiasma frequency in A-chromosomes; this effect was higher for plants with odd numbers of B-chromosomes compared with plants with even numbers of B-chromosomes. Comparisons of variants A and B, suggests that B-chromosomes have an effect pollen stainability and seed production. It seems that the presence of B-chromosomes may increase pollen stainability and seed production in variant B.     

Segregation-distortion in the B-chromosome system ofTettigidea lateralis (say) (Orthoptera: Tetrigidae)

A marginal population ofTettigidea lateralis was found to be polymorphic with respect to a large, mitotically stable supernumerary (B) chromosome. Male and female individuals may carry one or two B-chromosomes. In the male sex the frequency of individuals with one B was 33.8% whereas that of 2B-carriers was 2.9 %. A comparison with a small sample of female individuals suggests similar frequencies of B-chromosome carriers in the two sexes. The pycnocity cycle of the B's is virtually identical to that of the X chromosome which is always distinguishable by virtue of its larger size and other structural details. Persistent heterochromatic associations between the B and the X, which may last until metaphaseanaphase I, lead to a preferential migration of the B with the X to the same pole in male carriers of a single supernumerary. This distortional segregation of the B-chromosome may produce a differential transmission of the supernumerary to the two sexes if the various types of gametes are equally functional. Achiasmate, persistent B-B associations in 2B individuals can also cause segregation-distortion. The two supernumeraries segregate to the same pole in approximately 1/3 of the spermatocytes, but their poleward movement relative to that of the X is random. Both –B and +B individuals show only a single chiasma per individual bivalent. However, the presence of a single B raises very significantly the frequency at which the chiasma forms at the extreme distal ends of the L₁-L₂ and M₃-M₄ autosomes. The effect on recombination exerted by the supernumeraries and the possible implications of the segregation-distortion system ofT. lateralis are discussed in the light of recent studies on comparable B-chromosome polymorphisms.Research supported by N.R.C. of Canada.     

Meiotic drive for B-chromosomes in the primary oocytes of Myrmekotettix maculatus (Orthoptera: Acrididae)

Using a modified technique which allowed observation of chromosome orientation in the primary oocyte of grasshoppers at the onset of anaphase, it has been possible to establish that the B-chromosome is distributed preferentially on the egg side of the metaphase plate rather than the polar body side. The frequency of this preferential orientation matches very closely the level of preferential transmission determined from breeding experiments using individuals from the same population. The spindle is asymmetrical in the primary oocyte of this species, and a possible explanation of the meiotic drive is proposed as a result of the conical shape of nucleoplasm surrounding this spindle. The autosomal chiasma frequency of these females is generally lower than comparable males and is increased by the presence of B chromosomes; but the chiasma frequencies of the sexes respond differently to the addition of 1 and 2 B-chromosomes.     

B-chromosome polymorphism and interchromosomal chiasma interference in Eyprepocnemis plorans (Acrididae; Orthoptera)

Using the C-banding technique, the morphology and meiotic behaviour of four different types of B-chromosomes present in several populations of Eyprepocnemis plorans have been studied. The possible relationship between these four types is discussed. The analysis of chiasma frequency in A and B-chromosomes suggests the existence of interchromosomal interference and demonstrates that the chiasma frequency of B-s depends on the genetic background of the individual carrying them.     

Variable transmission rates of a B-chromosome in Myrmeleotettix maculatus (Thunb.) (Acrididae: Orthoptera)

Karyotype comparisons of both parents and progeny from single pair matings in the grasshopper Myrmeleotettix maculatus have shown that there is an accumulation of the large mitotically stable B-chromosome when transmitted through the female. This is presumed to result from a preferential segregation of univalent B-chromosomes at the first division of female meiosis and occurs irrespective of whether the B's are odd or even in number. In the male there is a loss of B-chromosomes. This loss does not appear to be due simply to the lagging and elimination of B-chromosomes in meiosis but probably involves sperm formation or function. When the balance of the gain and loss after one generation is calculated, it shows large overall accumulation in crosses involving individuals from a population in Wales, and a slight loss in individuals from a population in East Anglia. Such differences in transmission rates may be responsible for differences in B-frequency between populations. Since the B-chromosome frequency of these two populations has remained stable over five years, possible forces in the maintenance of the equilibria are examined. Females with B-chromosomes produce more aneuploid embryos than 0B females, but neither this cause of inviability nor general embryo mortality seem sufficient to produce an equilibrium situation. It is necessary to postulate that progeny with more than 2B chromosomes are inviable in order to approach equilibria. The presence of B-chromosomes in females has also led to the formation of polyploid embryos. The possible involvement of repetitive DNA in the formation of unreduced egg nuclei and preferential segregation is discussed.     

Seasonal variation of chiasma frequency in Phyllodactylus marmoratus (Gray) (Gekkonidae — Reptilia)

A study of the male meiotic system in two populations of the gekko Phyllodactylus marmoratus (Gray) has shown that both total and interstitial chiasma frequencies vary cyclically throughout the year. This variation is consistent in each population and was observed over a number of years. The total chiasma frequency (an index of the number of terminal chiasmata) has a different form of cyclic variation than does the interstitial chiasma frequency, and it is argued that they are under independent genetic controls. Reproductive studies suggest that only the sperm with the lowest total chiasma frequencies and greatest range of interstitial frequencies are used for fertilization. An experimental approach has shown that prolonged exposure to low temperature produces a significant increase in total chiasma frequency. It is believed that this environmental cue is responsible for the cyclic nature of total chiasma frequency.     

THE GEOGRAPHICAL DISTRIBUTION OF B-CHROMOSOMES OF XANTHISMA TEXANUM DC. (ASTERACEAE). I. SURVEY OF THE RANGE

The distribution of B-chromosomes throughout the range of Xanthisma texanum has been determined from approximately 750 individuals from 80 wild populations in Texas, Oklahoma, and New Mexico. Counts of meiotic chromosome figures of n = 4_II + 1 B_I and n = 5_II (4_II + IB_II) are reported for the first time for subsp. texanum var. texanum and for subsp. texanum var. orientalis. The wide occurrence of B-chromosomes in natural populations of subsp. drummondii is also documented. B-chromosomes are present in 22 (27.2%) randomly distributed populations whose characteristics are in no way distinct from populations without B-chromosomes. In populations with B-chromosomes present, 19.8% of the plants contained one or two B-chromosomes, but in total only 5.8 % of all plants contained one or two B-chromosomes. All populations in one isolated population group of subsp. drummondii lack B-chromosomes.     

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.     

Chromosome organisation in the Australian plague locust Chortoicetes terminifera

In Chortoicetes terminifera 45 independently-occurring B-chromosomes were tested and 23 distinct banding variants were detected with either G- or C-banding; six types were found more than once. In particular the Type I banding morph was detected 12 times indicating that individuals carrying this type may be under a different regime of selection compared with individuals bearing other types of banding morph; or the Type I may be subjected to a higher rate of meiotic drive in either or both sexes than other types. Also the Type I appeared to be obviously related to four other banding morphs whereas most types were not obviously related to any other banding morphs, but a few were similar in banding pattern to one or two other types. Three types of B-chromosomes were found in three or more different populations. A relatively high frequency of the Type I banding morph was found in one population, which was probably mainly composed of non-migratory individuals, and also in a laboratory-raised population. The most likely mechanisms for small changes in the banding sequence of the B-chromosomes are three-break insertions which are often indistinguishable from inversions. Rearrangements which add or delete bands, or sequences of bands, to or from B-chromosomes are probably the result of exchanges which are now known to take place in rare individuals with two B-chromosomes. The most distal region of all the banding morphs of the B-chromosome in C. terminifera, plus a short interstitial region in some types, is not late-replicating and has the banding characteristics of euchromatin. The rest of the chromatin of the B-chromosomes is heterochromatic and is the latest replicating heterochromatin in the whole genome. It consists of G-bands, which are also deeply stained with C-banding, and alternating G-interbands, which in turn are stained grey with C-banding. Both of these staining combinations are seen in heterochromatin of the normal complement. The heterochromatin of the B-chromosomes is condensed throughout 1st meiotic prophase in the male and in all somatic interphase nuclei where it can be quickly detected using the G-banding technique. The B-chromosome has a relatively constant, acrocentric morphology with a tendency to increase of length of the long arm as band numbers increase. Isochromosomes of the long arm have been seen only in laboratoryraised embryos. From egg pods with significantly fewer than expected B-chromosomes it is strongly suggested that more than one male may fertilize the eggs in a single pod.     

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 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.     

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.     

Cyclic variation of chiasma frequency and distribution in Podarcis sicula (Reptilia: Lacertidae)

The seasonal variations of chiasma frequency and distribution have been studied in the lizard: Podareis sicula. In this species, as in Phyllodactylus (King & Hayman, Chromosoma 69: 131–154, 1978), chiasma frequencies vary following a definite annual cycle, and clearly different trends are shown by interstitial and terminal chiasmata.A comparison between these seasonal chiasma frequency variations and those of environmental temperature shows the existence of a clear correlation between these two parameters. However, this correlation is different in the two types of chiasmata, and may be different within the same type of chiasma depending on the period of the year.A more significant correlation is observed between chiasma cycles and annual variations of the haematic levels of sexual steroid hormones. In particular we observe a highly significant correlation between interstitial chiasma frequencies and testosterone concentration. A less precise correlation between terminal chiasma frequencies and estradiol concentration is also observed.In Podarcis, as in Phyllodactylus, the sperm that will be used for fertilization derive from the spermatocytes showing the highest rate of interstitial chiasmata. This supports the hypothesis that the cyclic variations in interstitial chiasma frequencies represent a mechanism to ensure an adequate level of variability in a given population. The above mentioned correlation between chiasma frequencies and steroia hormone concentrations suggests that the seasonal chiasma cycles are controlled by the same environmental and hormonal factors regulating the spermatogenetic cycle.     

B-chromosome systems in the pocket mouse,Perognathus baileyi: Meiosis and C-band studies

The heterochromatin characteristics and meiotic behavior of the B-chromosome system of the pocket mouse, Perognathus baileyi, are described. B-chromosomes are associated both with a meiotic accumulation mechanism and with an increase in average chiasma frequency in the A-chromosome set in males. Three morphological classes of B-chromosomes are recognizable, and the mechanisms of origin of each are discussed.     

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.