Fractious chromosomes: Hybrid disruption and the origin of selfish genetic elements

Supernumerary B chromosomes are dispensable elements of the genome which can be retained in populations at high frequencies, despite being deleterious, through the ability to undergo non-Mendelian inheritance. Their mode of origin is, however, obscure. Recent work on gynogenetic fish has demonstrated the incorporation of small, unstable, centromere-containing microchromosomes, probably of interspecific derivation, into an asexual lineage⁽¹⁾. That these resemble B chromosomes both in structure and behaviour is consistent with the proposal that hybridisation between closely related species may be a significant mode of origin for such selfish genetic elements. Additional work on the B chromosome of a parasitoid wasp and observations on patterns of chromosome breakage from somatic cell hybrids also support this hypothesis.     

Distribution of a supernumerary chromosome system and aspects of karyotypic evolution in the genus Prochilodus (Pisces,Prochilodontidae)

A cytogenetic study on eight species of the genus Prochilodus, family Prochilodontidae, from different hydrographic basins of Brazil is reported. All species had a diploid number of 2n=54 and NF=108. Supernumerary heterochromatic chromosomes of variable frequency were detected in Prochilodus scrofa and in P. cearensis. To determine the occurrence and frequency of the supernumerary chromosomes in P. scrofa, specimens from different localities belonging to the basin of the Paraná river were analysed. The high karyotypic similarity detected demonstrated that, at least in these eight species, evolution occurred without marked changes in chromosome macrostructure.     

Chromosomal location of genes controlling seed proteins in species related to wheat

Summary The seed proteins of Chinese Spring wheat stocks which possess single chromosomes from other plant species related to wheat have been separated by gel electrophoresis in the presence of sodium dodecyl sulphate. Marker protein bands have been detected for both arms of barley chromosome 5, chromosome E (= 1R) and B (= 2R) of rye, chromosomes A,B (= 1C^u) and C (= 5C^u) of Aegilops umbellulata and chromosomes I and III of Agropyron elongatum. These studies, and previous findings, indicate that chromosome 5 of barley, chromosome 1R of rye, chromosome I of Ag. elongatum and possibly chromosome 1C^u of Ae. umbellulata are similar to chromosomes 1A, 1B and 1D in hexaploid wheat in that they carry genes controlling prolamins on their short arms and genes controlling high-molecular-weight (apparent molecular weight greater than 86,000) seed protein species on their long arms. These findings support the idea that all these chromosomes are derived from a common ancestral chromosome and that they have maintained their integrity since their derivation from that ancestral chromosome.     

Chromosomal Mapping of Repetitive DNAs in the Grasshopper Abracris flavolineata Reveal Possible Ancestry of the B Chromosome and H3 Histone Spreading

Supernumerary chromosomes (B chromosomes) occur in approximately 15% of eukaryote species. Although these chromosomes have been extensively studied, knowledge concerning their specific molecular composition is lacking in most cases. The accumulation of repetitive DNAs is one remarkable characteristic of B chromosomes, and the occurrence of distinct types of multigene families, satellite DNAs and some transposable elements have been reported. Here, we describe the organization of repetitive DNAs in the A complement and B chromosome system in the grasshopper species Abracris flavolineata using classical cytogenetic techniques and FISH analysis using probes for five multigene families, telomeric repeats and repetitive C₀t-1 DNA fractions. The 18S rRNA and H3 histone multigene families are highly variable and well distributed in A. flavolineata chromosomes, which contrasts with the conservation of U snRNA genes and less variable distribution of 5S rDNA sequences. The H3 histone gene was an extensively distributed with clusters occurring in all chromosomes. Repetitive DNAs were concentrated in C-positive regions, including the pericentromeric region and small chromosomal arms, with some occurrence in C-negative regions, but abundance was low in the B chromosome. Finally, the first demonstration of the U2 snRNA gene in B chromosomes in A. flavolineata may shed light on its possible origin. These results provide new information regarding chromosomal variability for repetitive DNAs in grasshoppers and the specific molecular composition of B chromosomes.     

Delimiting the Origin of a B Chromosome by FISH Mapping,Chromosome Painting and DNA Sequence Analysis in Astyanax paranae (Teleostei,Characiformes)

Supernumerary (B) chromosomes have been shown to contain a wide variety of repetitive sequences. For this reason, fluorescent in situ hybridisation (FISH) is a useful tool for ascertaining the origin of these genomic elements, especially when combined with painting from microdissected B chromosomes. In order to investigate the origin of B chromosomes in the fish species Astyanax paranae, these two approaches were used along with PCR amplification of specific DNA sequences obtained from the B chromosomes and its comparison with those residing in the A chromosomes. Remarkably, chromosome painting with the one-arm metacentric B chromosome probe showed hybridization signals on entire B chromosome, while FISH mapping revealed the presence of H1 histone and 18S rDNA genes symmetrically placed in both arms of the B chromosome. These results support the hypothesis that the B chromosome of A. paranae is an isochromosome. Additionally, the chromosome pairs Nos. 2 or 23 are considered the possible B chromosome ancestors since both contain syntenic H1 and 18S rRNA sequences. The analysis of DNA sequence fragments of the histone and rRNA genes obtained from the microdissected B chromosomes showed high similarity with those obtained from 0B individuals, which supports the intraspecific origin of B chromosomes in A. paranae. Finally, the population hereby analysed showed a female-biased B chromosome presence suggesting that B chromosomes in this species could influence sex determinism.     

The modification of crossing over in maize by extraneous chromosomal elements

Summary Five regions of the maize genome were tested for their response to endogenous factors influencing recombination. These included heterochromatic B chromosomes and abnormal chromosome 10 as well as the sex in which recombination occurred.The frequency of recombination in the proximal A ₂-Bt and Bt-Pr segments of chromosome 5 was increased in the presence of B chromosomes, with the male meiocytes showing a greater response than the female meiocytes. In addition, experiments involving 0, 1, 2 and 4 B's revealed a dosage effect of B chromosomes on crossing over in chromosome 5. Recombination in the proximal Wx-Gl ₁₅ interval of chromosome 9 was found to be slightly higher than normal in male flowers when two B chromosomes were present. This increase was accompanied by a decrease in the adjacent Sh-Wx segment. Crossing over in the distal C-Sh segment and in the C-Sh-Wx-Gl ₁₅ regions of female flowers was unaffected by B's.Comparisons of plants heterozygous for abnormal chromosome 10 (K10 k10) and homozygous for the standard chromosome 10 (k10 k10) showed that abnormal 10 greatly enhances crossing over in the A ₂-Bt and Bt-Pr segments of chromosome 5. In contrast to the finding with B's, the effect is greater in female than in male sporocytes. K10 showed no significant effect on recombination in the C-Sh-Wx-Gl ₁₅ region of chromosome 9 except in male sporocytes, where there was a slight increase in the Sh-Wx region of 0 B K10 k10 plants and a possible interaction with B chromosomes to raise the level of recombination between Wx and Gl ₁₅. The fact that the regions adjacent to the centromere of chromosome 9 show little or no response to the presence of K10 indicates that the proximal heterochromatin of this chromosome differs qualitatively from that of other maize chromosomes. This conclusion is supported by a comparison of the effects of B chromosomes, K10 and sex on crossing over in chromosomes 5 and 9.Dedicated to Dr. M. M. Rhoades on the occasion of his seventieth birthday.     

Molecular Characterization of a Maize B Chromosome Centric Sequence

Supernumerary chromosomes are widespread in the plant kingdom but little is known of their molecular nature or mechanism of origin. We report here the initial cloning of sequences from the maize B chromosome. Our analysis suggests that many sequences are highly repetitive and shared with the normal A chromosomes. However, all clones selected for B-specificity contain at least one copy of a particular repeat. Cytological mapping using B chromosome derivatives and in situ hybridization show that the B specific repeats are derived from the centric region of the chromosome. Sequence analysis of this repeat shows homology to motifs mapped to various plant and animal centromeres and to the maize neocentromere. A precise localization of these sequences among breakpoints within the B centromere and an homology to a facultative centromere, suggest a role for this sequence in centromere function.     

Molecular cytogenetic analysis of Brassica rapa-Brassica oleracea var. alboglabra monosomic addition lines

Interspecific alien chromosome addition lines can be very useful for gene mapping and studying chromosome homoeology between closely related species. In this study we demonstrate a simple but robust manner of identifying individual C-genome chromosomes (C5, C8 and C9) in the A-genome background through the simultaneous use of 5S and 25S ribosomal probes on mitotic and meiotic chromosomes of three different Brassica rapa-B. oleracea var. alboglabra monosomic addition lines. Sequential silver staining and fluorescence in situ hybridisation indicated that 18S-5.8S-25S rRNA genes on the additional chromosome C9 are expressed in the A-genome background. Meiotic behaviour of the additional chromosomes was studied in pollen mother cells at diakinesis and metaphase I. In all of the addition lines the alien chromosome was most frequently observed as a univalent. The alien chromosome C5, which carries an intercalary 5S rDNA locus, occasionally formed trivalents that involved either rDNA- or non rDNA-carrying chromosomes from the A genome. In the case of chromosomes C8 and C9, the most frequently observed intergenomic associations involved the regions occupied by 18S-5.8S-25S ribosomal RNA genes. It is possible that not all such associations represent true pairing but are remnants of nucleolar associations from the preceding interphase. Variations in the numbers and distribution of 5S and 25S rDNA sites between cultivars of B. oleracea, B. oleracea var. alboglabra and B. rapa are discussed.This revised version was published online in April 2005 with corrections to Fig. 2.     

The chromosomes of Puschkinia libanotica during mitosis

The chromosome complement of Puschkinia libanotica is described. In addition to five pairs of A chromosomes plants may possess up to 7 B chromosomes. Part of the long arm of the B chromosome gives rise to a heterochromatic mass in interphase nuclei and this can be seen to be a double structure in G₁ nuclei and a quadruple structure in G₂ nuclei. It is believed that these configurations represent the pre- and post-replication forms of subchromatids in the heterochromatic segment of the B chromosome. Microdensitometry of metaphase chromosomes shows that the segment of the B chromosome that is heterochromatic during interphase has no more DNA per unit volume than any of the euchromatic A chromosomes.     

The banding pattern of polytene chromosomes of Drosophila guanche compared with that of D. subobscura

A detailed map of the salivary gland chromosomes of Drosophila guanche is presented and compared to the standard gene arrangements of D. subobscura. Generally, the polytene chromosomc banding patterns of the two species show a high degrce of homology. Only Segment I of the sex chromosome (Chromosome A) shows marked differences. The banding pattern proposed for this segment in D. guanche could have originated from a cluster of overlapping inversions including A₁ arrangement.     

The cytology of Brachycome lineariloba

A study of approximately 300 plants in the taxonomic species B. lineariloba is reported. Five biological species differing in chromosome number exist in this species complex. The species with the lowest number (species A, n = 2) often carries B chromosomes, which may be large, or minute. It also exists in three racial forms which differ in karyotype. Observations on naturally occurring hybrids in zones of overlap show that two of the races differ by an unequal interchange. — The species with n = 8 (species C) is probably of amphidiploid origin from the cross A X B. Species B, E and D, with n = 6, 5 and 4 respectively, may represent a series of reductions in chromosome number. They show close karyotypic relationships. The relationship of species A with D, E and B is obscure.     

Macro- and microcolinearity between the genomic region of wheat chromosome 5B containing the Tsn1 gene and the rice genome

The Tsn1 gene in wheat confers sensitivity to a proteinaceous host-selective toxin (Ptr ToxA) produced by the tan spot fungus (Pyrenophora tritici-repentis) and lies within a gene-rich region of chromosome 5B. To use the rice genome sequence information for the map-based cloning of Tsn1, colinearity between the wheat genomic region containing Tsn1 and the rice genome was determined at the macro- and microlevels. Macrocolinearity was determined by testing 28 expressed sequence markers (ESMs) spanning a 25.5-cM segment and encompassing Tsn1 for similarity to rice sequences. Twelve ESMs had no similarity to rice sequences, and 16 had similarity to sequences on seven different rice chromosomes. Segments of colinearity with rice chromosomes 3 and 9 were identified, but frequent rearrangements and disruptions occurred. Microcolinearity was determined by testing the sequences of 26 putative genes identified from BAC contigs of 205 and 548 kb in length and flanking Tsn1 for similarity to rice genomic sequences. Fourteen of the predicted genes detected orthologous sequences on six different rice chromosomes, whereas the remaining 12 had no similarity with rice sequences. Four genes were colinear on rice chromosome 9, but multiple disruptions, rearrangements, and duplications were observed in wheat relative to rice. The data reported provide a detailed analysis of a region of wheat chromosome 5B that is highly rearranged relative to rice.     

Effect of a rye B chromosome and its segments on homoeologous pairing in hybrids between common wheat and Aegilops variabilis

Rye B chromosomes, which are supernumerary chromosomes dispensable for the host but increase in number by non-disjunction after meiosis, have been reported to affect meiotic homoeologous pairing in wheat-rye hybrids. The effect of a rye B chromosome (B) and its segments (B-9 and B-10) on homoeologous pairing was studied in hybrids between common wheat (2n=42) and Aegilops variabilis (2n=28), with reference to the Ph1 gene located on wheat chromosome 5B. The B-9 and B-10 chromosomes are derived from reciprocal translocations between a wheat and the B chromosomes, and the former had the B pericentromeric segment and the latter had the B distal segment. Both the B and B-9 chromosomes suppressed homoeologous pairing when chromosome 5B was absent. On the other hand, the B-9 and B-10 chromosomes promoted homoeologous pairing when 5B was present. On pairing suppression, B-9 had a greater effect in one dose than in two doses, and B-9 had a greater effect than B-10 had in one dose. These results suggested that the effect of the B chromosomes on homoeologous pairing was not confined to a specific region and that the intensity of the effect varied depending on the presence or absence of 5B and also on the segment and dose of the B chromosome. The mean chiasma frequency (10.23) in a hybrid (2n=36) possessing 5B and one B-9 was considerably higher than that (2.78) of a hybrid (2n=35) possessing 5B alone, and was comparable with that (14.09) of a hybrid (2n=34) lacking 5B. This fact suggested that the B chromosome or its segment can be used in introducing alien genes into wheat by inducing homoeologous pairing between wheat and alien chromosome.     

B chromosomes in Sternorrhyncha (Hemiptera, Insecta)

In the hemipteroid insects of the suborder Sternorrhyncha, B chromosomes are relatively common in comparison with other suborders of Hemiptera. However, the occurrence of supernumerary chromosomes is restricted, in most cases, to several genera or closely related species. At least in some species of Psylloidea with the XY sex determination system, a mitotically stable B chromosome integrated into an achiasmatic segregation system with the X, and became fixed as a Y chromosome. In some Aphidoidea with a multiple X system of sex determination, B chromosomes appear to be in fact non-functional X chromosomes. Supernumerary chromosomes thus probably play an important role in the evolution of sex determination systems in Sternorrhyncha.     

A repetitive DNA sequence common to the different B chromosomes of the genus Brachycome

Dot-like micro B chromosomes of Brachycome dichromosomatica were analysed for their sequence composition. Southern hybridization patterns of a total micro B probe to genomic DNA from plants with and without micro Bs demonstrated that the micro Bs shared sequences with the A chromosomes. In addition to telomere, rDNA and common A and B chromosome sequences, a new B-specific, highly methylated tandem repeat (Bdm29) was detected. After in situ hybridization with Bdm29 the entire micro B chromosome was labelled and clustering of the condensed micro Bs could be observed at interphase. A high number of Bdm29-like sequences were also found in the larger B chromosomes of B. dichromosomatica and in other Bs within the genus Brachycome. Received: 30 May 1997; in revised form: 20 August 1997 / Accepted: 20 August 1997     

Heat-shock DNA homology in distantly related species of Drosophila

Polytene chromosomes of D. melanogaster and D. virilis were hybridized in situ with ¹²⁵I labeled mRNA isolated from polysomes of D. melanogaster tissue culture cells incubated at 37° C. ¹²⁵I mRNA hybridized preferentially with subdivisions 87A and 87Cl of the D. melanogaster 3R chromosome; grains were also observed at regions 93D, 95D and over the chromocenter. A considerable cross hybridization of this mRNA with D. virilis polytene chromosomes was observed. The 29C region of the D. virilis second chromosome was the main site of hybridization. Significant grain numbers also appeared in region 20F of the same chromosome. The two regions mentioned belong to heat shock loci in the latter species. Based on label intensity we conclude that region 29C of D. virilis contains DNA sequences retaining molecular homology with those at subdivisions 87A and 87Cl of D. melanogaster. SDS-polyacrylamide gel electrophoresis revealed similar distributions of heat shock proteins in the two species studied.     

Unusual chromosome pairing and B chromosomes inBriza spicata (Poaceae)

One plant from a population ofBriza spicata (Poaceae) was found to have highly irregular meiotic behaviour. It is characterized by having a reduced chiasma frequency, a large between cell variance in chiasma frequency and the formation of multivalents involving pairs of A chromosomes. The B chromosome present in this plant also forms multivalents with a pair of A chromosomes. It is suggested that the normal control of strict bivalent pairing has broken down and homoeologous chromosomes are associating as multivalents. Furthermore, the partial homology of the B chromosome with a pair of A chromosomes is revealed.     

A linkage map of meadow fescue (<Emphasis Type="Italic">Festuca pratensis</Emphasis> Huds.) and comparative mapping with other Poaceae species

A genetic linkage map has been constructed for meadow fescue (Festuca pratensis Huds.) (2n=2x=14) using a full-sib family of a cross between a genotype from a Norwegian population (HF2) and a genotype from a Yugoslavian cultivar (B14). The two-way pseudo-testcross procedure has been used to develop separate maps for each parent, as well as a combined map. A total number of 550 loci have been mapped using homologous and heterologous RFLPs, AFLPs, isozymes and SSRs. The combined map consists of 466 markers, has a total length of 658.8 cM with an average marker density of 1.4 cM/marker. A high degree of orthology and colinearity was observed between meadow fescue and the Triticeae genome(s) for all linkage groups, and the individual linkage groups were designated 1F–7F in accordance with the orthologous Triticeae chromosomes. As expected, the meadow fescue linkage groups were highly orthologous and co-linear with Lolium, and with oat, maize and sorghum, generally in the same manner as the Triticeae chromosomes. It was shown that the evolutionary 4AL/5AL translocation, which characterises some of the Triticeae species, is not present in the meadow fescue genome. A putative insertion of a segment orthologous to Triticeae 2 at the top of 6F, similar to the rearrangement found in the wheat B and the rye R genome, was also observed. In addition, chromosome 4F is completely orthologous to rice chromosome 3 in contrast to the Triticeae where this rice chromosome is distributed over homoeologous group 4 and 5 chromosomes. The meadow fescue genome thus has a more ancestral configuration than any of the Triticeae genomes. The extended meadow fescue map reported here provides the opportunity for beneficial cross-species transfer of genetic knowledge, particularly from the complete genome sequence of rice.Communicated by P. Langridge     

Variation of the chromosome phenotype in zea,solanum and salvia

Summary InSolanum lycopersicum pachytene chromosomes the gradient in chromomere size, originating on both sides of the kinetochore, reveals the following characteristics: 1. a relatively abrupt decrease in size of the large chromomeres, 2. the gradient is related to arm length in 9 of the 12 chromosomes, 3. the gradient is particularly irregular in the short arm of the nucleolar chromosome and in the long arm is not conspicuous, 4. chromosome 6 shows an abrupt interruption in the gradient close to the kinetochore. Salvia viridis andZea mays chromosomes represent intermediate conditions between species with well defined and species without gradients. InSalvia the intermediate condition is manifested by the presence of a very large chromomere on each side of the kinetochore followed by very small chromomeres. In two chromosomes the intermediate condition is particularly apparent. In these chromosomes two chromomeres of intermediate size are present in the proximal region of the long arm. The nucleolar organizing arm has also an irregular pattern in this species.Maize has a less distinct gradient than tomato in all its chromosomes. Chromosomes 3, 4, 5 and 8 are those where the gradient is the least sharp. The nucleolar organizing arm of chromosome 6 has also an irregular pattern.In a translocation between chromosomes 5 and 6 of maize, a segment composed of very small chromomeres from the distal region of 5 which was moved to the right of the kinetochore of chromosome 6, did not change appreciably its phenotype after ten years of cultivation. During the period of cultivation a selection was made for plants where the original phenotype was preserved so that this result cannot be considered as demonstrating an absence of change in chromomere phenotype with changed position.InDrosophila andChironomus salivary gland chromosomes where chromomeres are large, and no selection has been carried out with such a purpose, the pattern and nucleic acid content of the bands is known to change when rearrangements occur within the chromosome.Supported by a grant from the Swedish Natural Science Research Council toA. Lima-De-Faria. This work was partly carried out at the Department of Botany, University of Illinois, U.S.A. during a visit to this department byA. Lima-De-Faria.P. Sarvella's collaboration in this work was done during her stay at the University of Lund.