Interaction among different heterocromatic variants in the grasshopper Dichroplus elongatus

Simultaneous chromosome polymorphisms for supernumerary elements allow us to analyse the relationships among different forms of heterochromatic variation in nature. We report simultaneous variation patterns for supernumerary segments in chromosomes S10 (SS10), S9 (SS9) and S6 (SS6) and B chromosomes in nine populations of the grasshopper Dichroplus elongatus from two biogeographic provinces from east Argentina. Our results show spatial chromosome differentiation for three out of four supernumerary heterochromatic variants (B chromosomes, SS6 and SS10). The incidence of B chromosomes was negatively correlated with the SS10 frequency. The distribution pattern analysis shows different degree of differentiation among populations for each supernumerary heterochromatic variant suggesting that the detected chromosome variation cannot be explained by interaction between migration and genetic drift. Moreover, the observed population chromosome differentiation was not in agreement with the hierarchical analysis of molecular of heterogeneity at mitochondrial DNA level (mtDNA). The present results point out the importance of the interaction among heterochromatic variants in the chromosome intraspecific variation in east Argentina natural populations of the grasshopper D. elongatus.     

Mitotically unstable B chromosome polymorphism in the grasshopper Dichroplus elongatus

Dichroplus elongatus, a widespread South American phytophagous grasshopper, exhibits polymorphisms for supernumerary chromosomes and segments (SS) in natural populations in Argentina. In this paper we review the available information on B chromosome polymorphism in D. elongatus related to geographic distribution, patterns of chromosome variation and influence on sperm formation. In D. elongatus the different forms of supernumerary variants are not independent. The proportion of B-carrying individuals (B prevalence) is negatively correlated with SS10 and positively with SS6 frequencies. The analysis of population structure considering the different supernumerary variants would suggest that the patterns of chromosome variation can not be explained only by random factors. Geographic distribution was analyzed scoring the prevalence of B chromosomes in 13 natural populations collected in three different biogeographical provinces from Northwest (Las Yungas province) and East (Espinal and Pampeana provinces) of Argentina. The detected heterogeneity may be explained by significant differentiation between Northwest and East regions and among populations within Las Yungas and Pampeana provinces. Correlation analysis suggested that B chromosome prevalence is associated with maximum temperature and with latitude. Additional information about the nature of the patterns of B chromosome variation was obtained comparing them with those obtained at the mitochondrial DNA level. The hierarchical analysis of molecular differentiation revealed discrepancy with respect to chromosome differentiation and also suggested that the pattern of B chromosomes may not be explained by historical factors. We also discussed the probable influence on fertility of carriers considering the production of abnormal sperm formation (macro and microspermatids) in relation to the number of Bs per follicle.     

Undertransmission of a supernumerary chromosome segment through heterozygous females possessing B chromosomes in the grasshopper Eyprepocnemis plorans.

The transmission ratio (ks) for a supernumerary chromosome segment was studied in a total of 54 heterozygous females collected from two Spanish natural populations of the grasshopper Eyprepocnemis plorans. Our analysis clearly demonstrated that ks is negatively dependent on the number of B chromosomes in the female. The possible mechanisms by which B chromosomes may cause undertransmission of the supernumerary segment, and the implications of this phenomenon for the maintenance of this extra chromosome segment, are discussed.     

Meiotic self-pairing of the Psalidodon (Characiformes,Characidae) iso-B chromosome: A successful perpetuation mechanism

B chromosomes are non-essential additional genomic elements present in several animal and plant species. In fishes, species of the genus Psalidodon (Characiformes, Characidae) harbor great karyotype diversity, and multiple populations carry different types of non-essential B chromosomes. This study analyzed how the dispensable supernumerary B chromosome of Psalidodon paranae behaves during meiosis to overcome checkpoints and express its own meiosis-specific genes. We visualized the synaptonemal complexes of P. paranae individuals with zero, one, or two B chromosomes using immunodetection with anti-medaka SYCP3 antibody and fluorescence in situ hybridization with a (CA)₁₅ microsatellite probe. Our results showed that B chromosomes self-pair in cells containing only one B chromosome. In cells with two identical B chromosomes, these elements remain as separate synaptonemal complexes or close self-paired elements in the nucleus territory. Overall, we reveal that B chromosomes can escape meiotic silencing of unsynapsed chromatin through a self-pairing process, allowing expression of their own genes to facilitate regular meiosis resulting in fertile individuals. This behavior, also seen in other congeneric species, might be related to their maintenance throughout the evolutionary history of Psalidodon.     

Genetic diversity and population structure of Buddleja crispa Bentham in the Himalaya-Hengduan Mountains region revealed by AFLP

Buddleja crispa Benth. is one of the most morphologically variable species in genus Buddleja, and it is widely distributed in the Himalaya-Hengduan Mountains (HHM) region. This study used AFLPs as a tool to examine the genetic variation among and within 25 populations of B. crispa. Analysis of population genetics of the species aimed to clarify morphological variation, current distribution patterns, strong adaptability to habitats, and the effects of geological factors in the HHM region. The genetic structure results, based on PCoA and NJ cluster analyses, revealed that the populations of B. crispa were divided into two genetic groups. Furthermore, the peripheral populations had lower genetic diversity than the populations in the center of the distribution areas (Three Parallel Rivers). We conclude that the gene flow (predominantly seed and pollen flow) and the population differentiation of B. crispa might be more affected by the barriers formed by rivers and mountains than by geographic distance.     

Population Genetic Structure of the Grasshopper Eyprepocnemis plorans in the South and East of the Iberian Peninsula

The grasshopper Eyprepocnemis plorans subsp. plorans harbors a very widespread polymorphism for supernumerary (B) chromosomes which appear to have arisen recently. These chromosomes behave as genomic parasites because they are harmful for the individuals carrying them and show meiotic drive in the initial stages of population invasion. The rapid increase in B chromosome frequency at intrapopulation level is thus granted by meiotic drive, but its spread among populations most likely depends on interpopulation gene flow. We analyze here the population genetic structure in 10 natural populations from two regions (in the south and east) of the Iberian Peninsula. The southern populations were coastal whereas the eastern ones were inland populations located at 260–655 m altitude. The analysis of 97 ISSR markers revealed significant genetic differentiation among populations (average G_ST =  0.129), and the Structure software and AMOVA indicated a significant genetic differentiation between southern and eastern populations. There was also significant isolation by distance (IBD) between populations. Remarkably, these results were roughly similar to those found when only the markers showing low or no dropout were included, suggesting that allelic dropout had negligible effects on population genetic analysis. We conclude that high gene flow helped this parasitic B chromosome to spread through most of the geographical range of the subspecies E. plorans plorans.     

A chromosomal study on Greek populations of the genusApodemus (Rodentia, Murinae) reveals new data on B chromosome distribution

During this study, 94 specimens (51 males, 43 females) of the wood mouseApodemus sylvaticus (Linnaeus, 1758), the yellow-necked mouseA. flavicollis (Melchior, 1834) and the western broad-toothed mouseA. epimelas (Nehring, 1902) from 20 localities of Greece were karyologically examined. The first two species were found to be widely distributed and their otherwise very similar karyotype (2n=48, FN=48) could be clearly distinguished, based on C-banding pattern. The third species appeared to have a more limited geographical distribution and its karyotype was distinguished from that of the previous two species, since it contained two pairs of small metacentric autosomes (2n=48, FN=52). The chromosomal study further revealed that nine individuals ofA. flavicollis possessed supernumerary B chromosomes (2n=49–50, FN=49–50). Meiotic chromosome preparations revealed that in contrast to sex chromosomes and autosomes, B chromosomes do not participate in bivalent formation. On the other hand, no supernumerary chromosomes were found in the studiedA. sylvaticus andA. epimelas material.     

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 transmission of rye B chromosomes in natural pollination

Summary This paper examines the transmission of B chromosomes in natural (but controlled) pollination, in order to obtain results which can be applied to natural populations of rye. The frequencies of the female gametes in both 2n= 14+1 and 2n=14+2 rye plants have been estimated with reference to their chromatid constitution. From the results obtained on the offspring, it seems that preferential distribution takes place during female meiosis of 2n= 14+2 plants. It has been demonstrated that pollen carrying B chromosomes formed in plants of 2n=14+2 was more competitive than normal pollen. On the contrary, when it was formed from plants 2n=14+1, B chromosome elimination by pollen was total. This process may be considered as sporophytic determination. The genetic significance of the presence of B chromosomes in natural populations is discussed. It is proposed that B chromosomes may be the cytological expression of a complex evolutionary system which results in conservation of population genetic variability.     

Comportement des chromosomes surnuméraires et relation avec le taux de mortalité dans une population africaine de Locusta migratoria migratorioides

Transmission of supernumerary chromosomes is studied in adults and embryos of an African population of Locusta migratoria migratorioides; 34% of the animals show one or two B chromosomes (24% one, 10% two). This percentage is the same in both sexes. During mitosis, B chromosomes are very stable. At meiosis, in some cases they show pairing, in other cases they enter as univalents. The eventuality of synapsis or association is discussed. These two kinds of behaviour could not be explained by the presence of two different B chromosomes as is shown from the study of parthenogenetic progeny. Repartition of B chromosomes in the progeny is different depending on whether the male or the female parent supplies them; so the behaviour of these supernumerary should be conditioned, not by their own structure, but by a connection with the cell. — One B chromosome apparently is neutral. When there are two B chromosomes, in case of synapsis, the number of animals with two B in the progeny is slightly lower than previous; in case of asynapsis, there is a higher lethality of individuals and oocytes with two B chromosomes.     

A correlation between B chromosome frequency and sex ratio in Exochomus quadripustulatus

A survey of natural populations of the British ladybird Exochomus quadripustulatus revealed the presence of a single large, acrocentric, supernumerary (B) chromosome in all sites visited. Studies were confined to male meiosis, where more than one B was never found to accompany the six bivalents and neo-XY sex pair. The percentage of males possessing B chromosomes varied from 6.4% to 28.6% in 14 different populations. The sex ratios present in these populations also varied. In some equal numbers of males and females were present, in others there were significant excesses of females. A linear regression was found between the percentage of B chromosomes and the percentages of males and females in those populations. It is suggested that the B chromosomes are not in themselves responsible for the sex ratio differences found for similar differences in sex ratio have been found in related neo-XY species lacking B chromosomes. It is more likely that those factors affecting sex ratio are also responsible for affecting the frequencies of B chromosomes in different populations.     

B chromosome morphotypes of Apodemus peninsulae (Rodentia) from the Russian Far East

Polymorphism of B chromosome morphotypes in the natural populations of the Korean field mouse Apodemus peninsulae (n = 367) has been studied in five regions of the Russian Far East: Magadan and Amur regions, the Jewish Autonomous region, Khabarovsk and Primorskii regions. Six groups of B chromosome morphotypes have been described on the size and morphological parameters. On the basis of their combinations 78 cell clones have been revealed, 23 of which are found in the animals with a stable karyotype and 76--in mosaics. The mosaics have also more diverse and unique combinations of B chromosome morphotypes. No differences between the geographic and local populations of mice on the studied characteristics have been found. Homogeneous structure of DNA in the B chromosomes of this species at the territory of the Russian Far East, as shown previously, allowed us to combine the numerical and dimensional data for each clone by introduction conditional "mass quantity" of B chromosomes (mB index). The leading role of natural selection in production of "critical" mass of supernumerary chromosomes in individuals with a stable karyotype and the weakening of its role in mosaics has been suggested.     

Temporal frequency stability and absence of effects on mating behaviour for an autosomal supernumerary segment in two natural populations of the grasshopper Eyprepocnemis plorans.

Interannual evolution of a polymorphism for a supernumerary segment in the smallest autosome of the grasshopper Eyprepocnemis plorans has been analysed in two natural populations. The polymorphism seemed to be stable in both populations, despite its undertransmission through heterozygous females carrying B chromosomes. Analyses of the effects of the extra segment on mating behaviour failed to show differential mating success in any sex or consistent effects on mating pattern. These results are discussed in relation to the maintenance of this polymorphism in natural populations.     

A CYTOGENETIC STUDY OF CLARKIA UNGUICULATA II. SUPERNUMERARY CHROMOSOMES

Mooring , John S. (Washington State U., Pullman.) A cytogenetic study of Clarkia unguiculata. II. Supernumerary chromosomes. Amer. Jour. Bot. 47(10): 847–854. Illus. 1960.—Supernumerary chromosomes morphologically comparable to those of the basic complement occur in high frequency in wild populations of this onagraceous annual plant. This paper evaluates the role that these supernumerary chromosomes play in the population dynamics of this species. The results of crossing plants with various numbers of supernumeraries showed that these chromosomes often increased in number in the seed parent. Examination of microsporocytes showed that almost 30% of the 491 plants determined had 1 or more supernumerary chromosomes. Studies of natural populations revealed that the frequency of individuals with supernumeraries varied from 0 to 79% in different populations, and also that in 1 population their frequency varied significantly in different years. It is concluded that supernumerary chromosomes in this species are adaptive under at least certain conditions. It is speculated that they are components of polygenic systems which increase tolerance to environmental extremes.     

Distortional meiotic segregation of a supernumerary chromosome producing differential frequencies in the sexes in the short-horned grasshopper Phaulacridium vittatum

Samples of males reveal the presence of a large, mitotically stable supernumerary chromosome in five populations examined. A stable frequency (11.22±0.3%) of male carriers has been observed over nine years in one population. The supernumerary follows closely the condensation cycle and behaviour of the sex chromosome in spermatogonial mitosis and meiosis. A structure simulating a terminalized chiasma frequently joins the precocious sex and supernumerary chromosomes during meiotic prophase; these two chromosomes move preferentially (70%) to opposite poles during the subsequent stages producing a differential transmission of the supernumerary to the two sexes. It is possible that the stable frequency in the population is maintained by a conserved balance between the two sexes without the need of an elimination system. The evidence supports the interpretation that the supernumerary chromosome is partially homologous with the sex chromosome. The possibility that the supernumerary might evolve into a neo-Y chromosome is suggested.     

B chromosomes in Nierembergia aristata (Solanaceae): nucleolar activity and competition with the A chromosomes

B chromosomes are additional dispensable chromosomes that may be present in some individuals, populations, or species, which have probably arisen from the A chromosomes but follow their own evolutionary pathway. Supposedly, B chromosomes do not contain major genes except for ribosomal DNA (rDNA) sequences that have been mapped on the supernumerary chromosomes of many plants and animals. This paper is a new report of B chromosome occurrence in plants. B chromosomes with nucleolar organizing regions (NORs) were found in a diploid sample of Nierembergiaaristata D. Don (sub nom. N. stricta Miers) (2n = 2x = 16). This is an extreme case in which B chromosomes possess not only strong nucleolar activity, as revealed by conventional staining methods, AgNOR and fluorescence banding, and fluorescent in situ hybridization (FISH), but also show nucleolar competition with the A chromosomes. The observed phenomenon could be analogous to the nucleolar dominance or 'differential amphiplasty' phenomenon that occurs in interspecific hybrids.     

Chromosomal diversification in populations of Characidium cf. gomesi (Teleostei, Crenuchidae)

Comparative cytogenetic studies carried out in two populations of Characidium cf. gomesi from Botucatu region, SP, Brazil, showed a similar karyotypic structure in a diploid number of 50 chromosomes, 32 metacentric and 18 submetacentric chromosomes for males and 31 metacentric and 19 submetacentric chromosomes for females as well as a ZZ-ZW sex chromosome system. Differences between both populations, however, were found in relation to the occurrence of B chromosomes and the distribution of 18S and 5S ribosomal DNA (rDNA) sites. Characidium cf. gomesi from the Alambari Stream, a component of the Tietê River basin, revealed 18S rDNA on Z and W chromosomes, while this gene was located on autosomes in the sample from the Paranapanema River basin. The 5S rDNA sites were observed in a single chromosomal pair (number 25) in the populations from Paranapanema and in two pairs in the specimens from Tietê (numbers 20 and 25). Besides that, in the sample from Paranapanema, both inter and intra-individual variations were found due to the occurrence of up to four heterochromatic supernumerary chromosomes in the cells. The life mode of this fish, restricted to headwaters and subjected to frequent breakdown into sub-populations, may have contributed to the fixation of such chromosomal differences. The karyotypic similarities found in the analysed populations, however, suggest that all are descended from the same ancestor group whereas their differences indicate that they are already existing in reproductively isolated populations.     

Meiotic drive against an autosomal supernumerary segment promoted by the presence of a B chromosome in females of the grasshopper Eyprepocnemis plorans

Twenty-seven out of 50 progeny analyses performed with specimens of the grasshopper Eyprepocnemis plorans were informative about the transmission of a supernumerary heterochromatic chromosome segment. The simultaneous presence of a B chromosome in some of the parents involved in the crosses permitted us to test the relationship between both types of supernumerary heterochromatin with respect to their transmission. The results demonstrated that the supernumerary segment is partly eliminated through females possessing B chromosomes. The implications of this in relation to the occurrence of the extra segment in natural populations are discussed.by S.A. Gerbi