Heterochromatin Content Research Articles

The Chromatin Assembly Factor Subunit FASCIATA1 Is Involved in Homologous Recombination in Plants

DNA replication in cycling eukaryotic cells necessitates the reestablishment of chromatin after nucleosome redistribution from the parental to the two daughter DNA strands. Chromatin assembly factor 1 (CAF-1), a heterotrimeric complex consisting of three subunits (p150/p60/p48), is one of the replication-coupled assembly factors involved in the reconstitution of S-phase chromatin. CAF-1 is required in vitro for nucleosome assembly onto newly replicated chromatin in human cells and Arabidopsis thaliana, and defects in yeast (Saccharomyces cerevisiae) affect DNA damage repair processes, predominantly those involved in genome stability. However, in vivo chromatin defects of caf-1 mutants in higher eukaryotes are poorly characterized. Here, we show that fasciata1-4 (fas1-4), a new allele of the Arabidopsis fas1 mutant defective in the p150 subunit of CAF-1, has a severe developmental phenotype, reduced heterochromatin content, and a more open conformation of euchromatin. Most importantly, homologous recombination (HR), a process involved in maintaining genome stability, is increased dramatically in fas1-4, as indicated by a 96-fold stimulation of intrachromosomal HR. Together with the open conformation of chromatin and the nearly normal expression levels of HR genes in the mutant, this result suggests that chromatin is a major factor restricting HR in plants.

DNA methylation in stingless bees with low and high heterochromatin contents as assessed by restriction enzyme digestion and image analysis

The stingless bee genus Melipona has been divided into two groups, based on their heterochromatin content. Melipona quadrifasciata and Melipona rufiventris have low and high levels of heterochromatin, respectively. Since condensed chromatin may be rich in methylated DNA sequences, M. quadrifasciata and M. rufiventris nuclei may contain different amounts of methylated CpG. These differences could be assessed by comparing Feulgen-DNA values obtained by image analysis of cells treated with the restriction enzymes Msp I and Hpa II that distinguish between methylated and unmethylated DNA. Msp I and Hpa II cleave the sequence -CCGG-, but there is no cleavage by Hpa II if the cytosine of the central CG dinucleotide is methylated. Malpighian tubules of M. quadrifasciata and M. rufiventris were treated with Msp I and Hpa II prior to the Feulgen reaction, and analyzed by automatic scanning microspectrophotometry. The Feulgen-DNA values for the heterochromatin of M. rufiventris and for the small heterochromatin and some euchromatin domains of M. quadrifasciata mostly decreased after treatment with Msp I, but were unchanged after treatment with Hpa II. CpG methylation, although detected in diverse chromatin compartments in different bee species, may induce silencing effects required for the same cell physiology.

Karyotype variation, evolution and phylogeny in Borago (Boraginaceae), with emphasis on subgenus Buglossites in the Corso-Sardinian system.

Karyological variation in the Mediterranean genus Borago and cytogeography of subgenus Buglossites in Corsica, Sardinia and the Tuscan Archipelago were investigated in combination with a molecular phylogenetic analysis aimed at elucidating relationships between subgenera and taxa with different chromosome features. Karyotype analysis was performed on population samples of B. pygmaea, B. morisiana, B. trabutii and B. officinalis. Phylogenetic analyses were based on ITS1 nrDNA and matK cpDNA sequences. Four base numbers were found, x = 6, 8, 9 and 15, and three ploidy levels based on x = 8. In subgenus Buglossites the Sardinian endemic B. morisiana is diploid with 2n = 18, while B. pygmaea includes three allopatric cytotypes with 2n = 30 (Sardinia), 2n = 32 (southern Corsica) and 2n = 48 (central northern Corsica and Capraia). In subgenus Borago, the Moroccan endemic B. trabutii and the widespread B. officinalis have 2n = 12 and 2n = 16, respectively. Molecular data support the monophyly of Borago, while relationships in subgenus Borago remain unclear. Borago trabutii appears as the earliest divergent lineage and is sister to a clade with B. officinalis, B. morisiana and B. pygmaea. Subgenus Buglossites is also monophyletic, but no correspondence between ITS1 phylogeny and B. pygmaea cytotypes occurs. Chromosome variation in Borago is wider than previously known. Two base numbers may represent the ancestral condition in this small genus, x = 6 or x = 8. An increase in chromosome number and karyotype asymmetry, a decrease in chromosome size and heterochromatin content, and the appearance of polyploidy are the most significant karyological changes associated with the divergence of the Buglossites clade. High ITS1 variation in the tetra- and hypotetraploid races of B. pygmaea suggests a multiple origin, while the lower polymorphism of the hexaploid race and its allopatric distribution in the northernmost part of the range is better explained with a single origin via union of unreduced and reduced gametes.

Chromosome plasticity in Ctenomys (Rodentia Octodontidae): chromosome 1 evolution and heterochromatin variation

A chromosome 1 (Cr1) pericentric inversion is described in six of seven species in the genus Ctenomys (tuco-tucos) from Uruguay. The inversion was inferred from G-band analyses of subtelocentric Cr1 hypothesised to be derived from the ancestral metacentric condition. Cr1 varies across species in heterochromatin amount and localisation including a metacentric chromosome without positive C-bands in C. torquatus, a subtelocentric chromosome with heterochromatic short arms in C. rionegrensis, and a subtelocentric chromosome negative after C-banding in five of the species analysed here. Pachytene chromosomes from C. rionegrensis, a species with the highest heterochromatin content, and C. torquatus, one of the species with the lowest heterochromatin content, were analysed in order to assess possible mechanisms of heterochromatin evolution. This analysis revealed the presence of three heterochromatic chromocenters in C. rionegrensis where bivalents converge, while in C. torquatus only one chromocenter was observed. In both species, highly repetitive DNA was observed, localised in chromocenters after "in situ" hybridisation. Heterochromatin associated protein M31 was localised in chromocenters of both species after immuno-detection. The spread of heterochromatin in Ctenomys chromosomes could be produced by chromatin exchanges at the chromocenter level. We propose the exchange of this DNA associated proteins between non-homologous chromosomes in pachytene to be the responsible for the spread of heterochromatin through the karyotypes of species like C. rionegrensis.

Functional Genomic Analysis of CAF-1 Mutants in Arabidopsis thaliana

Duplication of chromatin following DNA replication requires spatial reorganization of chromatin domains assisted by chromatin assembly factor CAF-1. Here, we tested the genomic consequences of CAF-1 loss and the function of chromatin assembly factor CAF-1 in heterochromatin formation. Genes located in heterochromatic regions are usually silent, and we found that this transcriptional repression persists in the absence of CAF-1 in Arabidopsis. However, using microarrays we observed that genes that are active during late S-phase, when heterochromatin is duplicated, were up-regulated in CAF-1 mutants. Arabidopsis CAF-1 mutants also have reduced cytological heterochromatin content; however, DNA methylation of pericentromeric repeats was normal, demonstrating that CAF-1 is not required for maintenance of DNA methylation. Instead, hypomethylation of the genome, which has only mild effects on the development of wild-type plants, completely arrested development of CAF-1 mutants. These results suggest that CAF-1 functions in heterochromatin formation. CAF-1 and DNA methylation, which is also needed for heterochromatin formation, have partially redundant functions that are essential for cell proliferation. Interestingly, transcriptional repression and heterochromatin compaction can be genetically separated, and CAF-1 is required only for the complete compaction of heterochromatin but not to maintain transcriptional repression of heterochromatic genes.

The Arabidopsis LHP1 protein is a component of euchromatin

The HP1 family proteins are involved in several aspects of chromatin function and regulation in Drosophila, mammals and the fission yeast. Here we investigate the localization of LHP1, the unique Arabidopsis thaliana HP1 homolog known at present time, to approach its function. A functional LHP1-GFP fusion protein, able to restore the wild-type phenotype in the lhp1 mutant, was used to analyze the subnuclear distribution of LHP1 in both A. thaliana and Nicotiana tabacum. In A. thaliana interphase nuclei, LHP1 was predominantly located outside the heterochromatic chromocenters. No major aberrations were observed in heterochromatin content or chromocenter organization in lhp1 plants. These data indicate that LHP1 is mainly involved in euchromatin organization in A. thaliana. In tobacco BY-2 cells, the LHP1 distribution, although in foci, slightly differed suggesting that LHP1 localization is determined by the underlying genome organization of plant species. Truncated LHP1 proteins expressed in vivo allowed us to determine the function of the different segments in the localization. The in foci distribution is dependent on the presence of the two chromo domains, whereas the hinge region has some nucleolus-targeting properties. Furthermore, like the animal HP1beta and HP1gamma subtypes, LHP1 dissociates from chromosomes during mitosis. In transgenic plants expressing the LHP1-GFP fusion protein, two major localization patterns were observed according to cell types suggesting that localization evolves with age or differentiation states. Our results show conversed characteristics of the A. thaliana HP1 homolog with the mammal HP1gamma isoform, besides specific plant properties.

Variability in Size of the Nuclear Genome in Pygmy Wood Mouse Sylvaemus uralensis (Rodentia, Muridae)

Earlier, in an integral genetic study, the Asian and European races were distinguished within the species Sylvaemus uralensis (pygmy wood mouse) and the European race was divided into the East European and South European forms. Each of these groups differed from the others, in particular, in the quantity of the centromeric heterochromatin in karyotypes of the animals. To establish the pattern of its changes in S. uralensis, in the present study the DNA content in splenocyte nuclei in all races and forms of pygmy wood mice was assessed using DNA flow cytometry. The heterochromatin amount in karyotypes and genome size were shown to be correlated. The East European chromosomal race of S. uralensis (Central Chernozem and Non-Chernozem regions of Russia, Crimea Peninsula, Middle Volga region, and Southern Ural) and the Asian race of this species (East Kazakhstan, Uzbekistan, and East Turkmenistan), which have respectively the highest and the lowest amounts of centromeric heterochromatin in the karyotype, exhibit the greatest difference in the DNA content in the genome. On average, the difference is approximately 8% in males and 6.7% in females; in both cases, the ranges of variability were distinctly different. Against the general background of the trait variation, the Asian race, whose members have the smallest DNA amount in their cells, looks homogeneous. The genome of the South European chromosomal form of S. uralensis (Caucasus, Transcaucasia, Carpathians, and Balkan Peninsula), which exhibits an intermediate content of the centromeric heterochromatin in the karyotype, is smaller that the genome of the East European race (by 3.2% in the group of males and by 1.9%, in the group of females), but larger than that of the Asian race (by 5% in either sex). Thus, the variability of size of centromeric C-blocks in pygmy wood mouse is likely to be associated with elimination (or, conversely, an increase in the amount) of the genetically inert chromatin. It is suggested that a significant contribution to the variability of genome size in S. uralensis is made by heterochromosomes, or, more precisely, their variable regions, which seem to be largely heterochromatic.

Karyomorphology, heterochromatin patterns and evolution in the genus Ophrys (Orchidaceae)

Karyotype structures and heterochromatin distribution in representative taxa of the genus Ophrys are compared, based on Feulgen-stained and banded somatic metaphase chromosomes. The karyotypes of Ophrys iricolor, O. lupercalis, O. caesiella, O. lutea, O. lunulata, O. x. tardans, O. apifera, O. praecox, O. lacaitae and O. insectifera are described for the first time. The karyological analyses indicate the relationships among the species with respect to asymmetry indices and heterochromatin content. Chromosomal differences have been helpful in clarifying the taxonomic position of Ophrys species that do not have clear affinities. The representative species of Araniferae, Fuciflorae and Ophrys sections exhibited the most asymmetrical karyotypes, while chromosome complements of the O. fusca–O. lutea group, of O. tenthredinifera and of O. bombyliflora proved to be less asymmetrical. Weakly heterochromatic chromosomes, with heterochromatin present mostly in thin centromeric bands, characterize Ophrys C-banded karyotypes. Chromomycin A3 (CMA) staining revealed that the analysed species exhibit a weak pattern of CMA+ bands at centromeric, intercalary or telomeric regions. No DAPI bright blocks were observed. The significance of the karyological data is discussed with regard to the relationships between the analysed species. © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society, 2005, 148, 87–99.

Giemsa C-Banded Karyotypes of two Cytotypes (2x, 4x) of Psathyrostachys Lanuginosa (Poaceae; Triticeae)

The karyotypes of the diploid (2n = 2x = 14) and tetraploid (2n = 4x = 28) cytotypes of the outbreeding P. lanuginosa collected from one population in China were investigated by Giemsa C- and N-banding and AgNO3 staining. The complement of the diploid consisted of 14 large chromosomes: 8 metacentrics and 6 SAT-chromosomes, one metacentric and 2 submetacentric pairs, with small to minute, polymorphic, completely heterochromatic satellites. Except that only eight SAT-chromosomes were observed, the complement of the tetraploid matched a doubling of that of the diploid and indicated an autoploid origin. The existence of 12 chromosomes with nucleolus organizers was established through AgNO3 staining demonstrating nucleoli. C-banding patterns comprised from zero to three (2x), or four (4x) small bands per chromosome giving a low content of constitutive heterochromatin (< 3 percnt;). The level of banding pattern polymorphism was low. N-banding produced no bands. Variation in the size of nucleoli demonstrated a differential nucleolus forming capacity of nucleolus organizers among SAT-chromosomes. The basic karyotype was similar to those of P. fragilis, P. juncea and P. stoloniformis, indicating a close relationship and the presence of genome N. Meiotic analysis suggested the presence of a chromosomal rearrangement in the diploid. The frequency of interstitial chiasmata was high. Differences in pollen diameters differentiated the two cytotypes.

Cloning, characterization and localization of Chinese hamster HP1 isoforms.

The Chinese hamster is one of the few mammalian species that are characterized by relatively poor heterochromatin content. It was intriguing to test whether or not the lack of large blocks of heterochromatin in the hamster chromosomes could be correlated with the absence or species-specific differences of the HP1 proteins, the main structural components of heterochromatin. To address this, we attempted to clone HP1 from the Chinese hamster. It is shown here that all three isoforms of HP1 known in mammals are present in hamster, and the amino acid sequences deduced from the cDNAs of the isoforms are 97-100% identical to those of the known mammalian homologues. All three isoforms are localized mainly in heterochromatic regions in the native chromosomes and nuclei. The hamster HP1 alpha gene was cloned, sequenced and mapped to the short arm of hamster chromosome 2. These data indicate that the Chinese hamster has all the HP1 components necessary for the establishment of heterochromatin. The limited amount of heterochromatin in hamster cells may probably be attributed to the unusual satellite DNA content of the hamster genome.

Gametogenesis and nucleotypic effects in the tetraploid red vizcacha rat, Tympanoctomys barrerae (Rodentia, Octodontidae)

Nucleotypic effects link DNA content with nuclear size and cell dimensions of reproductive cells in polyploid organisms. We studied the gametogenesis of the allotetraploid rodent Tympanoctomys barrerae, aiming to determine these effects in reproductive cells. The species' cofamily members, Octodon degus and Spalacopus cyanus were used as control. Spermatogenesis and oogenesis in T. barrerae follows the pattern of differentiation and sequence of events of the control species, but varied nucleotypic effects were observed. Exceedingly large, spatulated spermatozoa with a submedially attached flagellum are characteristic of male T. barrerae. The diameter of the nuclei of primordial and growing follicles as well as those of the Graaff follicles, of the granulose, and of luteal cells are significantly larger and heavily heterochromatic. Moreover, the width of the pellucid zone is 108% thicker in T. barrerae than in S. cyanus. Binucleation was recorded in 26% of luteal bodies examined whereas no binucleated cells are detected in the diploid control. Likewise, large heterochromatic nucleoli were observed in the follicle cells but not in S. cyanus. This finding and the high heterochromatin content of reproductive cells in the red vizcacha rat is probably associated with its genome complexity so that redundant genetic information is silenced through heterochromatinization.

Sex-chromosome heterochromatin variation in the wood mouse, Apodemus sylvaticus

Variation in heterochromatin content, as revealed by G- and C-banding, was studied in the sex chromosomes of the wood mouse, Apodemus sylvaticus. The sex-chromosome heterochromatin was also characterized by DAPI staining. Variation in sex chromatin was recorded in extremely large (giant) sex chromosomes in certain individuals and populations. In some individuals, the Y chromosome was the largest element of the complement. Different variants of both the X and Y chromosomes were found within a single population. The variation is therefore a type of population polymorphism and should not be used for taxonomic discrimination.

Karyotype morphology and cytogeography in Brunnera and Cynoglottis (Boraginaceae)

A comparative study of karyotype morphology and heterochromatin patterns in Brunnera andCynoglottis (Boraginaceae) was carried out with traditional methods and Giemsa C-banding. Two polymorphic species ofCynoglottis , each with two subspecies, and two of Brunnera were investigated using native population samples from the central-eastern Mediterranean and Middle East. Pollen size of these samples was measured to investigate relationships with ploidy level. C. barrelieri subsp. barrelieri and subsp. serpentinicola are characterized by 2n =18 and smaller pollen grains in contrast to C. chetikiana subsp. chetikiana and subsp.paphlagonica , which are fundamentally tetraploid with 2 n=36. The occurrence of cytotypes with 2n =12 and 2 n=24 in both subspecies of C. chetikiana, however, would suggestx =6 as the original haploid number and x=9 as a derived one. Furthermore, the finding of a hypoploid cytotype with 2 n=16 in C. barrelieri ssp. barrelieri was consistent with previous reports and suggested relationships with Anchusa. Karyoevolutionary processes possibly associated with such a wide chromosome variation in Cynoglottis are discussed. Brunnera macrophylla and B. orientalis share a complement of 2 n=12 and an apparently identical karyotype, which differs from Cynoglottis in terms of asymmetry, chromosome size and morphology. A basic C-banding style was present in Brunnera and Cynoglottis, but heterochromatin content increased from the former to the latter. The parallel increase in chromosome number, heterochromatin content and size of the pollen from Brunnera to Cynoglottis may reflect an evolutionary progression, and is consistent with the supposed ancient origin of Brunnera.

Karyotype morphology and cytogeography in Brunnera and Cynoglottis (Boraginaceae)

A comparative study of karyotype morphology and heterochromatin patterns in Brunnera andCynoglottis (Boraginaceae) was carried out with traditional methods and Giemsa C-banding. Two polymorphic species ofCynoglottis , each with two subspecies, and two of Brunnera were investigated using native population samples from the central-eastern Mediterranean and Middle East. Pollen size of these samples was measured to investigate relationships with ploidy level. C. barrelieri subsp. barrelieri and subsp. serpentinicola are characterized by 2n =18 and smaller pollen grains in contrast to C. chetikiana subsp. chetikiana and subsp.paphlagonica , which are fundamentally tetraploid with 2 n=36. The occurrence of cytotypes with 2n =12 and 2 n=24 in both subspecies of C. chetikiana, however, would suggestx =6 as the original haploid number and x=9 as a derived one. Furthermore, the finding of a hypoploid cytotype with 2 n=16 in C. barrelieri ssp. barrelieri was consistent with previous reports and suggested relationships with Anchusa. Karyoevolutionary processes possibly associated with such a wide chromosome variation in Cynoglottis are discussed. Brunnera macrophylla and B. orientalis share a complement of 2 n=12 and an apparently identical karyotype, which differs from Cynoglottis in terms of asymmetry, chromosome size and morphology. A basic C-banding style was present in Brunnera and Cynoglottis, but heterochromatin content increased from the former to the latter. The parallel increase in chromosome number, heterochromatin content and size of the pollen from Brunnera to Cynoglottis may reflect an evolutionary progression, and is consistent with the supposed ancient origin of Brunnera.

NOR regions of polychaete worms of the genus Ophryotrocha studied by chromosome banding techniques and FISH.

This article reports the results of cytogenetic analyses carried out on 10 species of polychaete worms belonging to the genus Ophryotrocha (Dorvilleidae). Nucleolar organizer regions (NORs) were characterized by Ag staining, C-banding, CMA3 staining, and ribosomal fluorescent in situ hybridization (rDNA FISH). Extensive intraspecific variation in NOR number and distribution were observed in O. costlowi, O. sp. macrovifera, O. notoglandulata, O.l. labronica, O. l. pacifica (2n = 6), O. p. puerilis, O. diadema (2n = 8), O. hartmanni, O. gracilis (2n = 10). In O. sp. robusta (2n = 10), Ag-NORs were always located on a single chromosome pair. CMA3 staining suggests a possible trend toward a GC-rich rDNA compartmentalization. In O.l. labronica, O. p. puerilis, O. diadema, and O. sp. robusta rDNA FISH shows that Ag and FISH signals coincide. Results from C-banding seem to indicate that the increased genome size (GS) observed in O. sp. macrovifera (0.8 pg) and O. hartmanni (1.16 pg) compared to the base GS value of the genus (0.4 pg) cannot be attributed to variation in the heterochromatin content.

Karyomorphology and Evolution in Italian Populations of Three Neottieae Species (Orchidaceae).

A karyological analysis on Italian populations of 3 species, belonging to the genera Cephalanthera and Listera (Orchidaceae : Neottieae tribe) and characterized by asymmetrical karyotypes, was undertaken. Karyomorphological and C-banding data strongly suggest that in 2 species, namely C. damasonium and L. cordata, chromosomal rearrangements involving the large and medium-large chromosomes constituted the principal mechanism of chromosome evolution. This processis particularly evident in L. cordata, where karyotype evolution occurred by means of centric fission along with an increase in chromosome number and in the overall chromosomal heterochromatin content. Altogether the present observation on Italian specimens of the Neottieae corroborate previous proposals of a central role of Robertsonian rearrangements and quantitative heterochromatin varia-tion in karyotype reorganization.

Genome size variations are related to X‐chromosome heterochromatin polymorphism in Arvicanthis sp. from Benin (West Africa)

The African grass rat, Arvicanthis sp., from Benin shows sex‐chromosome polymorphism for the presence in the karyotype of three X‐chromosome forms: subtelocentric (Xst), submetacentric (Xsm), and metacentric (Xm). Here we report statistically significant differences of diploid genome size (g.s.) among individuals, from 6.22 ± 0.06 pg to 6.80 ± 0.10 pg, depending on the X‐chromosome form. The evaluation of the total and X‐chromosome heterochromatin content alone (Xst: 2.16 ± 0.22%; Xsm: 5.02 ± 0.22%; Xm: 6.54 ± 0.28%), supports this suggestion. DNA base‐pair preferential staining revealed that heterochromatin of both autosome and sex chromosomes is AT‐rich. Digestion of genomic DNA with sixteen restriction enzymes produced DNA bands representing the repetitive fraction of the genome. When these were used as probes for fluorescence in situ hybridization (FISH), two hybridization patterns were found: a) FISH positive signals localized in the pericentromeric areas of every chromosome of the complement and b) chromosome specific signals co‐localizing with both the pericentromeric regions of every chromosome and the heterochromatic regions of Xsm and Xm, but not on the Xst chromosome. FISH performed with a (TTAGGG)n probe revealed a telomere‐only pattern.

Karyomorphological parameters and C-band distribution suggest phyletic relationships within the subtribeLimodorinae (Orchidaceae)

Karyotype attributes and heterochromatin distribution were used to characterize fourteen taxa of the subtribeLimodorinae (Orchidaceae). The karyotypes were established using morphometrical parameters following Feulgen staining and C-banding. No significant differences in heterochromatin content were found between specimens collected from various sites. Four species of theEpipactis helleborine group possess some chromosome pairs with quite similar heterochromatin patterns; some differences were found inE. distans with respect to other species of this group.Epipactis palustris differed significantly from otherEpipactis species in its different karyotype and its numerous terminal C-bands. The largest differences from the other genera were shown inLimodorum as far as karyomorphology and heterochromatin patterns were concerned. C-band distribution indicated similarity among non-homologous chromosomes, supporting a possible palaeo-polyploid origin for theCephalanthera andEpipactis karyotypes.

Karyotypes and heterochromatin variation (C-bands) in Melipona species (Hymenoptera, Apidae, Meliponinae)

We describe the karyotypes of eight bee species of the genus Melipona and compare them in terms of heterochromatin content and location (C-banding technique). All species had 2n = 18 (females) and n = 9 (males) chromosomes, but a wide variation in heterochromatin content was detected among karyotypes. On the basis of these differences, the species were divided into two functional groups, one of them comprising species with a karyotype having a low heterochromatin content (M. bicolor bicolor, M. quadrifasciata, M. marginata, and M. asilvai), and the other species with a high heterochromatin content (M. seminigra fuscopilosa, M. capixaba, M. scutellaris, and M. captiosa). In the species with high heterochromatin content, heterochromatin occupied practically the entire extent of all chromosomes, with euchromatin being limited to the extremities, a fact that prevented observation of the centromere. In contrast, in the species with karyotypes having a low heterochromatin content, heterochromatin was visualized only in some chromosomes. In the chromosomes in which it was present, heterochromatin was located in the centromere or on the short arm. M. bicolor bicolor had the smallest heterochromatin content with only three chromosome pairs presenting heterochromatin in females. Increased heterochromatin content may be explained by interstitial and pericentromeric growth.

Comparison of the Giemsa C‐banded karyotypes of Hystrix coreana and H. patula (Poaceae, Triticeae)

The karyotypes of Hystrix coreana from eastern USSR and H. patula from USA were investigated by Giemsa C‐banding. Both species are outbreeders and have 2n = 4x = 28. The karyotype of two plants of H. coreana has 10 metacentric, 6 submetacentric, 8 heterobrachial and 4 SAT chromosomes; two plants differed by having 12 metacentric, 4 submetacentric, 8 heterobrachial and 4 SAT‐chromosomes, and 10 metacentric, 4 submetacentric, 9 heterobrachial and 5 SAT‐chromosomes, respectively. The C‐banding pattern had no or few inconspicuous intercalary bands, but conspicuous telomeric C‐bands in one or both arms giving a high content of heterochromatin (16.3–18.2%). The chromosome complement of one plant of H. patula had 8 metacentric, 6 submetacentric, 8 heterobrachial and 6 SAT‐chromosomes. The C‐banding pattern had between 1 and 4 intercalary or centromeric bands and conspicuous telomeric bands on one or both arms giving a high content of constitutive heterochromatin (16.4%).