Giemsa C-banded karyotypes of diploid and tetraploidHordeum bulbosum (Poaceae)

The similar-looking basic genomes ofHordeum bulbosum (2x and 4x) have five rather similar metacentric, one submetacentric, and one satellited choromosome. C-banding patterns are characterized by one or two centromeric, or juxtacentromeric, small to larger bands in most chromosomes, by bands at the nucleolar organizers, by small or very small telomeric bands, and by the nearly complete lack of intercalary bands. Banding pattern polymorphism is widespread. Banding patterns supported by chromosome morphology enable identification of homologues, and discrimination between non-homologues inH. bulbosum (2x). The C-banded karyotype ofH. bulbosum (4x) supports an autopolyploid origin, but it was possible to identify only homologues of submetacentrics and SAT-chromosomes.     

Pollen evolution and systematics inAnnonaceae with special reference to the disulcate Australian endemic genera

All genera ofAnnonaceae endemic in Australia (Ancana, Fitzalania, Haplostichanthus) show almost exactly the same type of disulcate (disulculate) pollen with intact exine extending over the sulci. Tetrad stages inHaplostichanthus andAncana reveal a latudinal subequatiorial orientation of the two sulci at the proximal hemisphere. Sometimes they fuse into a ±zonosulcate aperture.Fissistigma pollen grains are ±globose and have a flattened pole with a central elevation and a concentric groove, covered by a somewhat reduced exine. This palynological characters give further support for separating the generaAncana andFissistigma. Germination was observed inHaplostichanthus where the pollen tube emerges at one of the two sulci and inFissistigma where the flattened part breaks up during germination. The aperture types described here are obviously transitional stages between aperturate and inaperturate pollen grains and are discussed in regard to pollen evolution.     

Comments on flower evolution and beetle pollination in the generaAnnona andRollinia (Annonaceae)

Dynastid scarab beetle pollination appears basic within the genusAnnona. Those species ofAnnona which are more morphologically derived, as well as allRollinia spp. possess reduced floral chambers and attract small beetles likeNitidulidae orStaphylinidae. Pollination of the primitive species ofAnnona byDynastinae would imply that the genus had not evolved before the Tertiary. The fossil record is in congruence with this hypothesis. Once again it is stressed that the cantharophilous syndrome, as it is found in theAnnonaceae, Magnoliaceae, Eupomatiaceae andCalycanthaceae, with beetles being exclusive pollinators, is a secondary and derived condition and obviously different from the expected basic entomophily of the original angiosperms.     

C-banding inLiriodendron tulipifera (Magnoliaceae): Some karyological and systematic implications

InLiriodendron tulipifera the Giemsa C-banding pattern differs remarkably from the distribution of condensed chromatin in prophase as revealed by conventional methods. The nature of proximal bands in metaphase chromosomes is interpreted to be different from the terminal ones. A comparison ofLiriodendron with some C-banded karyotypes of ± related taxa indicates a relatively high degree of karyomorphological differentiation in some woodyPolycarpicae. This has to be considered in karyosystematic comparisons.     

Ultrastruktur und systematische Bedeutung des Pollens beiBocageopsis,Ephedranthus, Malmea undUnonopsis (Annonaceae)

The four genera investigated show solitary sulcate pollen grains. The structure and sculpture of the sporoderm is very similar inBocageopsis andUnonopsis, and supports the close relationship of both genera; their flower morphology also is very similar. In contrast, the sporoderm ofEphedranthus, and especially that ofMalmea, is different in some characters and suggests more remote relationships; this is also confirmed by differences in flower morphology. Within this group of genera a distinct exine progression can be recognized from non-columellate (Malmea) to granular (Unonopsis guatterioides), further to somewhat irregularly (Bocageopsis, someUnonopsis species), and finally to very regularly columellate (Ephedranthus). The sculpture of the tectum varies from a reticulum with large lumina (Malmea) to one with small performations (Ephedranthus). Within theAnnonaceae the genusMalmea is among the most primitive in respect to pollen structure. The sulcus of the four genera is very large and runs over 1/3 of the pollen grain. It is characterized by a reduction of the exine and a bulgy thickening of the intine.

     

Infraspecific variation in C-banded karyotype and chiasma frequency inCucumis sativus (Cucurbitaceae)

Infraspecific cytogenetical variation was studied in a diverse collection of five non-cultivated and cultivatedCucumis sativus accessions. The individual chromosomes of different accessions could be identified by the C-banding pattern and chromosome measurements. About 40–50% of the genomic area are made up of heterochromatin inC. sativus. The non-cultivated accessions exhibit more heterochromatin and lower chiasma frequencies per pollen mother cell than cultivated accessions. There is infraspecific variation in C-banding pattern, karyomorphology and multinucleolate cells. The use of C-banding in infraspecific classification is discussed.     

Comparison of the Giemsa C-banded karyotypes of the three subspecies ofPsathyrostachys fragilis,subspp.villosus (2x),secaliformis (2x, 4x), andfragilis (2x) (Poaceae), with notes on chromosome pairing

The karyotypes of diploidP. fragilis subsp.villosus (2n = 2x = 14) and tetraploid subsp.secaliformis (2n = 4x = 28) were studied by Giemsa C- and N-banding, and AgNO₃ staining and compared with the karyotype of subsp.fragilis (2x). The complements of subsp.villosus and subsp.fragilis were similar, with 8 metacentric and 6 SAT-chromosomes, one metacentric and two submetacentric pairs, with small to minute, polymorphic, heterochromatic satellites. The complement of subsp.secaliformis on the whole agreed with a doubling of the complement of diploidP. fragilis, suggesting autopolyploidy. Only the presence of 12 nucleoli in interphases identified 6 SAT-chromosome pairs. In subsp.villosus one or two extra micronucleoli indicated a chromosome pair with very low nucleolusforming activity, bringing the number of SAT-chromosome pairs to 4. This number may be a characteristc ofPsathyrostachys. Besides very small, inconsistently observed bands, the C-banding pattern consisted of 0–3 small bands per chromosome at intercalary and terminal locations, and at NORs. The level of banding pattern polymorphism was low, but enough to indicate that the taxa are outbreeders. Similarities in chromosome morphology and C-banding patterns identified homology of all chromosomes of subsp.villosus, but for 12 pairs only in subsp.secaliformis. Between plants, reliable identification of homology and homoeology (subsp.secaliformis) was possible only for the SAT-chromosomes and the shortest metacentrics. Chromocentres were very small and the amount of constitutive heterochromatin was low. N-banding stained chromosomes uniformly. The basic karyotypes of theP. fragilis taxa were similar to those ofP. juncea, P. lanuginosa, andP. stoloniformis supporting a close relationship and the presence of a common genome, N. NORs had different nucleolus-forming activities. Meiotic analysis demonstrated a high level of bivalent pairing in the three taxa. A chromosomal rearrangement was suggested in subsp.villosus. The low multivalent frequency in subsp.secaliformis indicates the presence of a pairing regulation mechanism. The majority of chiasmata were interstitial. Pollen grain size discriminated between diploid and tetraploid taxa. The existence of a diploid cytotype of subsp.secaliformis is supported by pollen measurements of herbarium material.     

Genetic diversity and phylogeny in Hystrix (Poaceae, Triticeae) and related genera inferred from Giemsa C-banded karyotypes

The phylogenetic relationships of 15 taxa from Hystrix and the related genera Leymus (NsXm), Elymus (StH), Pseudoroegneria (St), Hordeum (H), Psathyrostachys (Ns), and Thinopyrum (E) were examined by using the Giemsa C-banded karyotype. The Hy. patula C-banding pattern was similar to those of Elymus species, whereas C-banding patterns of the other Hystrix species were similar to those of Leymus species. The results suggest high genetic diversity within Hystrix, and support treating Hy. patula as E. hystrix L., and transferring Hy. coreana, Hy. duthiei ssp. duthiei and Hy. duthiei ssp. longearistata to the genus Leymus. On comparing C-banding patterns of Elymus species with their diploid ancestors (Pseudoroegneria and Hordeum), there are indications that certain chromosomal re-arrangements had previously occurred in the St and H genomes. Furthermore, a comparison of the C-banding patterns of the Hystrix and Leymus species with the potential diploid progenitors (Psathyrostachys and Thinopyrum) suggests that Hy. coreana and some Leymus species are closely related to the Ns genome of Psathyrostachys, whereas Hy. duthiei ssp. duthiei, Hy. duthiei ssp. longearistata and some of the Leymus species have a close relationship with the E genome. The results suggest a multiple origin of the polyploid genera Hystrix and Leymus.     

Different origins of fragile exines within theAnnonaceae

Besides tectate and columellate, 3-layered exine types, in theAnnonaceae, one also finds very fragile, thin exine types. Their single exine layer corresponds either to a former tectum (including infratectal layer) or a former basal layer. The interpretation of the different origin of the remaining layers is based on their different structure and position within the intine. The fact that reduced exine types are obviously not always homologous should be regarded in systematic interpretations.     

Nuclear DNA changes,genome differentiation and evolution inNicotiana (Solanaceae)

A survey of 51 species fromNicotiana subgg.Tabacum, Rustica andPetunioides has shown that evolution was accompanied by a five-fold variation in nuclear DNA amounts. This variation, however, was not directly correlated with the changes in chromosome number. Drastic rearrangement of karyotypes is characteristic for the evolution ofNicotiana spp. Significant gain or loss in nuclear DNA has often accompanied such changes, but DNA variation has also occurred without significant changes in karyotype arrangements.—The distribution of nuclear DNA is discontinuous inNicotiana, species cluster into DNA groups with consistently regular increments in the mean DNA amounts. The discontinuities are viewed as steady states in terms of genomic balance and biological fitness.—Changes in the amount of nuclear DNA and in the heterochromatin are compared with the morphological, chromosomal and adaptive changes which accompanied speciation in 14 subgeneric sections. The evolutionary significance of DNA variation is discussed.     

Giemsa banded karyotypes,systematics, and evolution inAnacyclus (Asteraceae-Anthemideae)

Giemsa C-banding allows the differentiation of six, otherwise very similar karyotypes from the small genusAnacyclus. Banding style—with stable centromeric and nucleolar bands, and diverse specific banding patterns in distal chromosome segments—contributes significantly to generic demarcation and systematic grouping. The amount of banding corresponds to heterochromatic chromocentres and increases from perennials to annuals. Relationships with other nucleotype parameters and evolutionary mechanisms are discussed.First contribution of a series on Giemsa Banded Karyotypes, Systematics, and Evolution inAnthemideae (Asteraceae).     

Karyomorphological analyses and heterochromatin characteristics disclose phyletic relationships among 2n = 32 and 2n = 36 species ofOrchis (Orchidaceae)

The karyotypes of eight taxa ofOrchis L. with 2n = 32 and 2n = 36 have been investigated using morphometrical measurements following staining with Feulgen, Giemsa (C-banding) and the DNA specific fluorochrome Hoechst 33258. The karyotypes ofO. coriophora subsp.fragrans, andO. papilionacea proved to be the most asymmetrical, whileO. morio andO. longicornu exhibited the most symmetrical karyotypes. Using C-banding and the fluorochrome H33258 only the taxa with high asymmetry indices showed the presence of differentially stained chromatin bands. In most chromosomes heterochromatin bands were present at the telomeric position. The present results seem to indicate that the analysed species do not form a homogeneous group and further subdivisions are possible, which, in turn, do not always correlate with divisions based on morphological characters. Both karyomorphology and heterochromatin distribution coincide in indicating a possible evolutionary pathway.     

Quantitative analyses of C-banded karyotypes,and systematics in the cultivated species of theScilla siberica group (Liliaceae)

Quantified C-band karyograms are presented for the related speciesScilla siberica, S. mordakiae, S. ingridae, S. amoena, andS. mischtschenkoana. Chromosome structure, banding style, and heterochromatin characters suggest a systematic grouping of two more closely related species pairs:S. siberica andS. mordakiae, S. ingridae andS. amoena; they are part of a larger aggregate, well separated fromS. mischtschenkoana. Four different heterochromatin fractions can be recognized inS. siberica and its relatives, but only two inS. mischtschenkoana. C-bands do not replace, but they are added to euchromatin. The particular origin and history of the cultivatedS. amoena and the triploidS. siberica cv. Spring Beauty appear to be responsible for their karyotype constancy, but chromosome conservatism obviously is genuine inS. mischtschenkoana. The banding data support the systematic grouping proposed on a morphological basis, and provide additional evolutionary evidence.Evolution ofScilla and Related Genera, IV.     

Karyosystematics and evolution of AustralianAnnonaceae as compared withEupomatiaceae,Himantandraceae, andAustrobaileyaceae

Chromosome counts are presented for 12 genera and 20 species of AustralianAnnonaceae (all diploid with 2n = 16 or 18; Table 1) and two species ofEupomatiaceae (2n = 20, partly from Papua New Guinea). Detailed studies on interphase nuclear structure, condensing behaviour of chromosomes, and fluorochrome and Giemsa C-banding patterns also includeHimantandraceae (Galbulimima) andAustrobaileyaceae. — Eupomatiaceae completely correspond withAnnonaceae karyologically, their base number 2n = 20 is interpreted to have evolved from 2n = 18 by ascending dysploidy from common ancestors.Eupomatia laurina andE. benettii differ in DNA and constitutive heterochromatin (hc) quantity; their evolution from high to low DNA content probably corresponds to general progressions inMagnoliidae. Austrobaileya has nuclei of the presumably primitive Tetrameranthus type which is closely related to that ofGalbulimima and several other primitive taxa inMagnoliidae. Karyomorphology and other characters support the maintainance of two main branches within theMagnoliidae, Laurales andMagnoliales, withAustrobaileya probably intermediate; theWinteraceae appear more remote.—InAnnonaceae the reestablishment ofAncana is underlined by its chromosome number (2n = 18) the unexpected and specialized disulcate pollen, and various morphological characters which point to a close alliance with the Australian endemic generaFitzalania andHaplostichanthus (also disulcate) and the American genus pairSapranthus/Desmopsis; they are united in the provisionalSapranthus tribe, with a more distant position toFissistigma s. str. (2n = 16). AustralianAnnonaceae exhibit a high generic and a low species diversity; they can be considered as an ± old and partly impoverished outpost of the family with phytogeographical relationships to Asia, Africa and America.—On the base of field observations three main types of floral development inAnnonaceae are proposed, the most elaborated one found in the fly pollinated genusPseuduvaria. The growth form change from shrubs to lianas during the ontogeny ofDesmos andMelodorum, the vegetative propagation of anAncana species and the ecological and evolutionary patterns of the taxa investigated are discussed.     

Heterochromatin and silver banding of rye (Secale cereale,Gramineae) chromosomes

Air-dried chromosomes of rye when stained with aqueous silver nitrate show differential banding patterns. In addition to staining the NOR sites, the silver nitrate stains all regions of constitutive heterochromatin, as identified by Giemsa C-banding, as well as a number of small interstitial regions. However, the heterochromatin on the B chromosome is not stained by the silver method. This is proposed as a rapid and reliable banding method.     

Karyotype constancy and genome size variation in BulgarianCrepis foetida s. l. (Asteraceae)

Ten populations ofCrepis foetida from Bulgaria belonging to the three subspeciesfoetida, rhoeadifolia, andcommutata were analyzed karyologically using haematoxylin staining, Giemsa C-banding, fluorochrome banding, Ag-NOR staining, Feulgen cytophotometry (scanning densitometry and video-based image analysis), and propidium iodide flow cytometry. The quantitatively-evaluated karyotype structure was similar among all populations, with minor variation in a few intercalary sites only and in the amount of NOR-associated heterochromatin (satellites). In contrast to the karyotypic constancy the genome size ofC. foetida subsp.commutata was about 10% lower than those of the other two subspecies, which had similar genome sizes. The genome size measurements using three different methods resulted in highly correlated data. The genome size difference adds some weight to previous taxonomic opinions treatingC. foetida subsp.commutata at species level, asC. commutata. Prof. Dr. Stefan Kozhuharov (4 January 1933–24 August 1997).     

Chromosome banding and pairing behaviour inFestuca andVulpia (Poaceae,Pooideae)

Giemsa C-banding patterns of the grassesFestuca rubra (2n = 6x = 42),Vulpia fasciculata (2n = 4x = 28) and their wild F1 hybrid ×Festulpia hubbardii (2n = 5x = 35) show marked differences between the chromosomes of the two parents that enable unequivocal identification ofFestuca andVulpia chromosomes in the hybrid. Moreover, meiotic banding patterns reveal that both homogenetic (Festuca-Festuca andVulpia-Vulpia) and heterogenetic (Festuca-Vulpia) pairing occurs in the F1. The significance of this in relation to the formation of backcrosses to theFestuca parent and to the evolution of theFestuca polyploid complex in general is discussed.     

Karyologie und ökologisch-morphologische Differenzierung vonPeumus boldus (Monimiaceae,Laurales)

The karyotype analysis ofPeumus boldus reveals a remarkable differentiation of chromosomes in size and form. C-banding pattern and interphase nuclei are discribed and compared with other genera. The great variation in leaf-shape and growth-form is continuous and correlated with the very diverse habitats of the species. The recognition of infraspecific taxa is therefore not recommended. The lower leaf surface is very uniform within the monotypic genus and ± similar with that ofSiparuna. The different evolutionary strategies ofPeumus and related genera are discussed and show a different eco-systematic situation than inMollinedia andSiparuna.