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.     

The B-chromosome system of Hypochoeris maculata

Populations of Hypochoeris maculata (Compositae) over a wide area of Europe contain a small proportion of individuals with mitoticallystable B-chromosomes. The telocentric Bs are about one sixth the length of the smallest A-chromosome and are heterochromatic. Offspring grown from seeds of natural populations have one or at most two B-chromosomes although plants with up to five Bs have been produced in experiment. The Bs rarely pair during meiotic prophase and the univalents usually undergo centric division at anaphase I. Despite lagging, the Bs are rarely lost during male meiosis. The inheritance of the B-chromosomes differs between the two sides with accumulation in the egg and random inheritance or loss on the pollen side. Although meiotic drive may play some part in the maintenance of the B-polymorphism there is a strict upper limit on the number of Bs in natural populations.     

Genetic control of synapsis and recombination in Lolium amphidiploids

Homologous bivalent formation in amphidiploids of Lolium is promoted during meiosis by diploidising genes carried by A-chromosomes and by supernumerary B-chromosomes. The site and mode of action of these diploidising factors were investigated by comparing the relative frequencies of pairing configurations at meiotic prophase and metaphase I in several different hybrid genotypes. The results indicate that diploidising genes act predominatly by increasing the stringency of synapsis at early stages of meiotic prophase. By contrast, B-chromosomes appear to promote bivalent formation by ensuring that homoeologously paired chromosome segments within multivalents do not crossover. The results show that the additive effects of diploidising genes and B-chromosomes are to a certain extent separable in terms of their mode of action and timing during meiosis.     

Structure,behaviour and repetitive DNA of B-chromosomes in Cestrum nocturnum (Solanaceae)

Abstract

The Cestrum genus is karyotypically exceptional in Solanaceae. It is characterised by a basic number x?=?8, a large chromosomal and genomic size, complex heterochromatin patterns, B-chromosomes (Bs) with particular heterochromatin and distribution of 18–5.8–26S and 5S rDNA. Cestrum nocturnum L. has a diploid number of 2n?=?16 plus a variable number of B-chromosomes. The aims of work was to analyse their numerical variation, structure and behaviour of C. nocturnum B-chromosomes by classical and molecular cytogenetics. The individuals analysed had 2n?=?16?+?0?13 B-chromosomes. All B-chromosomes were metacentric and smaller than A-chromosomes. The number of B-chromosomes showed a great variability between and within individuals, thereby denoting the occurrence of events that promote mitotic and meiotic instability. Cytogenetic techniques made it possible to observe that B-chromosomes are rich in heterochromatin, probably with AT- and GC-rich regions. In addition, molecular techniques allowed to detect homologous sequences of transposable element conserved domains of Ty1-Copia and Ty3-Gypsy superfamilies. These sequences were located by FISH in all B-chromosomes and some A-chromosomes. Our results showed that repetitive DNA could play an important role in chromosomal evolution as well as in the stability of B-chromosomes in C. nocturnum.     

THE CYTOGENETICS OF XANTHISMA TEXANUM DC. (ASTERACEAE) AND ITS B-CHROMOSOMES

The karyotype of A-set and B-chromosomes of Xanthisma texanum DC. are described in detail and measurements are given. The distribution of B-chromosomes in stem tissue is described and the loss of B-chromosomes from primary roots confirmed. The number of B-chromosomes in pollen mother cells was constant within single inflorescence buds and in all buds throughout the flowering period. The behavior of one and two B-chromosomes during meiosis is described in detail. During meiosis, B₁'s divided in 71% of the cases during anaphase II and in 29% of the cases during anaphase I. First anaphase division resulted in some lagging and elimination of B-chromatids during second division. During anaphase I, B₁₁'s divided into chromosomes and in anaphase II these divided into chromatids in 95% of the cases. In the other 5 % non-disjunction of the B-chromosomes or chromatids and A-set chromosome abnormalities occurred. Post meiotic preferential distribution during pollen mitosis resulting in a doubling of the number of B-chromosomes passed to progeny via pollen occurred, but no doubling occurred in the female line. Crosses involving plants with two B-chromosomes each yielded fewer plants with three B-chromosomes than expected. Polyploidy is extremely rare in the species. The evolutionary significance of B-chromosome behavior is discussed in relation to its origin and survival.     

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.     

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.     

Evidence of B-chromosomes in the karyotype of Barypeithes pellucidus Boheman 1834 (Coleoptera, Curculionidae, Entiminae) from Central Europe

B-chromosomes were observed in spermatogonial mitotic metaphases, meiotic metaphases I and II of Barpeithes pellucidus from one population in Slovakia. The number of B-chromosomes ranged from one to six per cell and they paired with the sex heterochromosomes in the first meiotic metaphase and rarely with the autosomes. In metaphase I one B-chromosome was always associated with X chromosome forming a tripartite complex. The XyBp was easily recognizable as a complex of three chromosomes in a parachute association The size of the B-chromosomes was approximately the same or a little smaller than that of the y heterochromosome which was the smallest element of the regular chromosome set. Their staining intensity seems to be similar to that of the autosomes and sex chromosomes, respectively. The behaviour of B-chromosomes during mitosis and meiosis in weevils is briefly discussed.     

Synaptonemal complex analysis of B-chromosome behavior in meiotic prophase I in the East-Asiatic mouse Apodemus peninsulae (Muridae, Rodentia)

The mitotic and meiotic chromosomes of four male East-Asiatic mice, Apodemus peninsulae, having three to seven chromosomes in addition to the standard karyotype (2n = 48), were investigated. B-chromosomes were represented by medium-sized metacentric and dotlike chromosomes. Mosaicism of bone marrow cells due to a numerical variation of accessory chromosomes was established for the males examined. Capacity of B-chromosomes to form axial elements and synaptonemal complexes in meiotic prophase I was revealed by electron microscopy. The occurrence of univalents of different morphology, bivalents, and multivalents, corresponding to B-chromosomes, was demonstrated. An increase in the number of B-chromosomes was found in spermatocytes at zygotene-pachytene relative to the number in bone marrow cells, which may be evidence of B-chromosome accumulation in the germ cell line of the East-Asiatic mouse.     

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.     

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.     

Chromosomes in a hybrid zone of Israeli mole rars (Spalax, Rodentia)

Chromosomal novelties and the level of meiotic and mitotic abnormalities were studied in a hybrid zone between two chromosomally differentiated Spalax cytotypes of 2n = 58 and 2n = 52. These cytotypes differ by five Rb fusions, four centromeric shifts accompanied by heterochromatin deletion, one paracentric inversion, and the Y-chromosome reorganization. Among 149 specimens studied, 82 were hybrids with 64 different karyotypes ranging in diploid numbers from 2n = 50 to 2n = 60. Nine hybrid specimens were mosaics for the chromosome numbers due to occurrence of cell lines with different Robertsonian chromosome arrangements, and six specimens possessed variable number of B-chromosomes. Mosaicism of B-chromosomes was found also in meiotic cells however chromatid breaks and abnormal chromosome pairing during meiosis occurred very rarely. All these results imply some local genomic instability resulting in the spontaneous process of reversible Rb fusions.     

B-chromosomes in some Himalayan hardwoods

B-chromosomes are reported for the first time for woody angiosperms. Out of six species from the eastern Himalayas in which B-chromosomes were observed, two are polyploid. The number of B-chromosomes varied from 0–5 in the different species. In Sambucus canadensis, Glerodendrum colebrookianum and Neolitsea zeylanica B's were found to pair with each other at Metaphase-I.The financial support for this work was provided by the U. S. Government from PL 480 funds in India (Grant: A7-FS-12).     

Synaptonemal complexes of the A- and B-chromosomes of the spermatocytes from the East Asian mouse Apodemus peninsulae

The analysis of whole-mount preparations of synaptonemal complexes (SCs) from surface-spread spermatocytes of A. peninsulae (2n = 48A + 1, 2, ... 12 B) had revealed SCs of 23 autosomal bivalents, sex bivalent XY, axial cores and SCs of the B-chromosomes. The intercellular and interindividual variability of the number of B-chromosomes varied from 1 to 12 per cell. The SCs of autosomal bivalents were shown to have a typical structure. The structure and behaviour of SCs of sex bivalent throughout meiotic prophase I appeared to be similar to those observed in other species of this order. Mainly B-univalents and less frequently B-bivalents containing SCs were found to be formed in meiotic prophase I. The full homologues appear to be rarely seen among B-chromosomes of the East-Asiatic mouse. A tendency of forming clusters of B-univalents near the sex bivalent was found, in addition to B-bivalents with lateral elements, having the form of bi- and tri-stranded elements with rare synaptic fragments. Besides this, the SCs of the autosomes of pachytene cells were found to contain structures resembling the recombination nodules.     

Chromosomes in a hybrid zone of Israeli mole rars (Spalax, Radentia)

Chromosomal novelties and the level of meiotic and mitotic abnormalities were studied in a hybrid zone between two chromosomally differentiated Spalax cytotypes of 2n = 58 and 2n = 52. These cytotypes differ by five Rb fusions, four centromeric shifts accompanied by heterochromatin deletion, one paracentric inversion, and the Y-chromosome reorganization. Among 149 specimens studied, 82 were hybrids with 64 different karyotypes ranging in diploid numbers from 2n = 50 to 2n = 60. Nine hybrid specimens were mosaics for the chromosome numbers due to occurrence of cell lines with different Robertsonian chromosome arrangements, and six specimens possessed variable number of B-chromosomes. Mosaicism of B-chromosomes was found also in meiotic cells however chromatid breaks and abnormal chromosome pairing during meiosis occurred very rarely. All these results imply some local genomic instability resulting in the spontaneous process of reversible Rb fusions.     

Unstable B-chromosomes in a European population of Allium schoenoprasutn L. (Liliaceae)

In Allium schoenoprasum , small, dichromatic B-chromosomes occur in 74% of plants at Lac de Tracens, Pare National des Pyrénées. B-chromosomes from Lac de Tracens were compared with the previously well characterized Bs from British populations of A. schoenoprasum at the River Wye, Powys, and were found to differ in terms of both their morphology and the level of mitotic instability. At Lac de Tracens, two morphological types of B-chromosome were observed: a telocentric of about 1.15 μm in length and a metacentric of about 1.35 μm in length; the most frequent B-type in the River Wye populations (B^-1) was not found. B-chromosomes from the River Wye are essentially stable at root tip mitosis, whereas 25 out of 29 plants from Lac de Tracens (86%) showed between-cell variation in B-number, with non-modal numbers in 330 out of 794 cells (41.6%). Most of the variability occurred within individual roots, rather than between roots of single bulbs or of sister bulbs. No meiotic pairing was observed between the different Bs in hybrids from the geographically isolated populations, suggesting a lack of homology. An independent origin of B-chromosomes in the British and French populations is suggested.     

No 5-B compensation by rye B-chromosomes

Summary Chinese Spring mono-5B wheat was crossed with rye plants, with and without B-chromosomes, to produce polyhaploids with and without 5B and with and without rye B-chromosomes. As expected, absence of 5B resulted in a strong increase of homoeologous pairing. It was accompanied by a decrease in chiasmata in the rye B-chromosomes. The rye B-chromosomes were entirely ineffective in compensating for the 5B effect in nulli 5B, 2 rye B types.     

Cytogenetic Analysis of A- and B-Chromosomes of Prochilodus Lineatus (Teleostei, Prochilodontidae) Using Different Restriction Enzyme Banding and Staining Methods

Different cytogenetic techniques were used to analyse the chromosomes of Prochilodus lineatus with the main objective of comparing the base composition of A- and B-chromosomes. The results of digestion of chromosomes with 10 different restriction endonucleases (REs), silver staining, CMA₃ staining and C-banding indicated the existence of different classes of highly repetitive DNA in the A-set and also suggested the existence of compositional differences between the chromatin of A- and B-chromosomes. The 5-BrdU incorporation technique showed a late replicating pattern in all B-chromosomes and in some heterochromatic pericentromeric regions of A-chromosomes. The cleavage with RE BamHI produced a band pattern in all chromosomes of P. lineatus which permitted the tentative pairing of homologues in the karyotype of this species. We concluded that the combined use of the above techniques can contribute to the correct identification of chromosomes and the karyotypic analysis in fishes. On the basis of the results, some aspects of chromosome structure and the origin of the B-chromosomes in P. lineatus are discussed. This revised version was published online in August 2006 with corrections to the Cover Date.     

Occurrence of B-chromosomes in diploid Allium stracheyi Baker and their elimination in polyploids

Summary In a population of Allium stracheyi Baker (2n=14) growing in Darjeeling both diploids and polyploids occur. The diploids contain B-chromosomes varying from 2–10 in number. Polyploids are conspicuous by absence of B-chromosomes. These in diploids are found in the pollen mother cells and also in the pollen, and some are provided with subterminal constriction.Diploid individuals when brought from Darjeeling to Calcutta (i. e. from temperate to tropical regions) became polyploid within a month and the B-chromosomes were simultaneously lost. In order to confirm this unexpected result, the transfer experiment has been repeated thrice with fresh collections in each case and selection of diploid bulbs after cytological observation. In all cases the result has been the same. In rare cases one or two B-chromosomes were found in the polyploid cells which might represent intermediate steps of the disappearance.B-chromosomes in diploids possibly help the individual to compete with polyploids by enlarging the adaptive capacity.The sudden polyploidisation by transfer from the mountains to the plains might have been the result of a shock due to the temperature difference. The high temperature may be deleterious for the reproduction of B-chromosomes, and their degeneration products possibly contribute to cytoplasmic changes and the spindle disturbances which effect polyploidisation.     

Interaction between rye B-chromosomes and wheat genetic systems controlling homoeologous pairing

The interactive effect on homoeologous pairing of rye B-chromosomes with the absence of both pairing suppressor (3A, 3D, 5B) and promotor (3B, 5A, 5D) chromosomes of common wheat (Triticum aestivum L.) is analyzed by comparison of pairing at Metaphase I of 27-, 27+2B, 28- and 28+2B-chromosome plants. These plants were obtained from crosses between the respective wheat monosomics (2n=41) and rye plants (Secale cereale L.) carrying or not carrying two B-chromosomes (2n=14 or 14+2Bs). —The effect of rye B-chromosomes on pairing depends on the function of the wheat chromosome which is absent in the appropriate hybrids, i.e., rye B-chromosomes have a suppressor effect on pairing when the pairing suppressing wheat chromosomes 3A, 3D or 5B are absent, while they behave as promotors when the pairing promoting chromosomes 3B, 5A or 5D are absent.