Pathways of karyological differentiation in palms (Arecaceae)

Karyological data are given for 56 palm taxa coming from all 6 palm subfamilies. In 11 genera and 17 species, chromosome numbers are reported for the first time. Most chromosome numbers in palms range between 2n = 36 and 2n = 26 in dysploid series. Species of the same genus usually exhibit identical chromosome numbers which additionally may be constant in larger groups of closely related genera (Coryphoideae trib.Corypheae with nearly always 2n = 36,Arecoideae subtribesEuterpeinae andRoystoneinae with 2n = 36,Arecoideae subtrib.Butiinae with mostly 2n = 32). Polyploidy among palms is of minor significance but the endemic Madagascan genusVoanioala (2n = 606 ± 3) is the most striking exception. — With respect to structure of interphase nuclei and longitudinal differentiation of prophase and metaphase chromosomes, the palm family is highly differentiated. Euchromatin types with different prophase condensation properties and fluorochrome and C-banding patterns of heterochromatin permit a discrimination of several subfamilies on the nuclear level (Arecoideae, Ceroxyloideae, Nypoideae, Phytelephantoideae, Calamoideae).Arecoideae andCeroxyloideae, andNypoideae andPhytelephantoideae have some features in common. Subfam.Coryphoideae s. l. is a non-uniform group. — Nuclear characters among palms exclusively found in recentCoryphoideae subtrib.Thrinacinae link palms with other monocotyledons. Most probably, such a nuclear condition represents an ancestral state in the evolution of palm genomes within subfam.Coryphoideae s. l., but also the conspicuous nuclear characters of the other modern palm subfamilies appear to be derived from a similar starting point, since transitional character states are still present in subfam.Calamoideae and some taxa of subfam.Arecoideae. Early karyoevolution in palms obviously did not involve numerical change of the ancient chromosome number of 2n = 36 which started subsequently, as a dysploid reduction in numerous parallel series, independent in subfam.Coryphoideae (2n = 36 to 2n = 28),Calamoideae (2n = 36 to 2n = 26),Ceroxyloideae (2n = 34 to 2n = 26), andArecoideae (2n = 36 to 2n = 28). Possible mechanisms of karyological change are discussed. — Karyological characters are compared to morphological, ecological, taxonomical, and chorological features, and give some new insight into older and more recent phases of palm evolution. (1) Strong deviations in vegetative or floral morphology are often accompanied by major karyological differences, and sometimes the direction of advancement can be traced through intermediate stages. (2) Apart fromCoryphoideae subtrib.Thrinacinae, the strongest concentration of apparently original karyological traits is found in the more basal members of each subfamily. (3) The most successful and actively radiating colonizers of the forest floors in evergreen tropical forests which belong to completely different subfamilies (Old WorldLicuala, New WorldChamaedorea andGeonoma), appear to be very advanced karyologically.     

Cytogenetics and cytotaxonomy ofVelloziaceae

Chromosome number and other cytological features are reported from 35 species ofVelloziaceae, including several African and Brazilian populations. All analyzed species show areticulate interphase nuclei and prophase/prometaphase chromosomes with proximal early condensation. Most heteropycnotic blocks do not seem to correspond to heterochromatin since, at least inVellozia patens, they do not stain differentially after C-banding procedures. Regarding the chromosome number, three main groups could be identified. The first comprised diploid species of the generaNanuza, Vellozia and the Brazilian species ofXerophyta with 2n = 14 or 16; the second comprised tetraploid species with 2n = 34, and included all Brazilian species of subfam.Barbacenioideae; the third group, of hexaploid species, comprised the African representatives of the genusXerophyta. A single population ofVellozia, possibly of hybrid origin, had 2n 32. A basic number of x = 8 is proposed for the family. The karyological information supports the hypothesis that theVelloziaceae originated on the South American, rather than on the African continent.     

A role for Drosophila SMC4 in the resolution of sister chromatids in mitosis

BACKGROUND: Faithful segregation of the genome during mitosis requires interphase chromatin to be condensed into well-defined chromosomes. Chromosome condensation involves a multiprotein complex known as condensin that associates with chromatin early in prophase. Until now, genetic analysis of SMC subunits of the condensin complex in higher eukaryotic cells has not been performed, and consequently the detailed contribution of different subunits to the formation of mitotic chromosome morphology is poorly understood. RESULTS: We show that the SMC4 subunit of condensin is encoded by the essential gluon locus in Drosophila. DmSMC4 contains all the conserved domains present in other members of the structural-maintenance-of-chromosomes protein family. DmSMC4 is both nuclear and cytoplasmic during interphase, concentrates on chromatin during prophase, and localizes to the axial chromosome core at metaphase and anaphase. During decondensation in telophase, most of the DmSMC4 leaves the chromosomes. An examination of gluon mutations indicates that SMC4 is required for chromosome condensation and segregation during different developmental stages. A detailed analysis of mitotic chromosome structure in mutant cells indicates that although the longitudinal axis can be shortened normally, sister chromatid resolution is strikingly disrupted. This phenotype then leads to severe chromosome segregation defects, chromosome breakage, and apoptosis. CONCLUSIONS: Our results demonstrate that SMC4 is critically important for the resolution of sister chromatids during mitosis prior to anaphase onset.     

Cytology, palynology, and taxonomy of Asteropyrum and four other genera of Ranunculaceae

A karyomorphological study of two species in Asteropyrum (Ranunculaceae) and five in presumably related genera (one each in Caltha , Coptis , and Dichocarpum and two in Thalictrum ) reveals close similarities between Asteropyrum and Caltha , but obvious differences between Asteropyrum and Coptis , Dichocarpum , and Thalictrum , in some karyological features, such as the structure of the interphase nuclei, chromosome size, basic chromosome number, and condensation behaviour of chromosomes during the mitotic prophase. The chromosomes of Asteropyrum are categorized as being of the R-type and the basic chromosome number is confirmed as x  = 8, indicating a possible close affinity of Asteropyrum to members in the subfamily Helleboroideae, in particular to Caltha and its allies. Asteropyrum peltatum and Asteropyrum cavaleriei are shown to be more or less differentiated from each other in gross morphology and palynology, and somewhat independent in geographical distribution, albeit with intermediate forms occurring where their distribution zones overlap. It seems justifiable to treat them as two subspecies of A. peltatum .  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 152 , 15–26.     

Reorganization and condensation of chromatin in mitotic prophase nuclei of Allium cepa

This paper studies the process and features of chromosome construction in mitotic prophase cells of Allium cepa. The results showed that a prominent reorganization of chromatin occurred during G₂-early prophase. The 250–400 nm thick compact chromatin threads in G₂ nuclei began to disorganize into about 30, 100 and 220 nm chromatin fibres which constituted the loosely organized chromosome outlines in early prophase before chromosome condensation. In middle prophase, chromosome condensation was characterized by the formation of many condensed regions (aggregates of chromatin), which increased in size (1–1.5 m) when prophase proceeded. Meanwhile, the chromatin threads that constituted and connected the condensed regions became increasingly thicker (120–250 nm). In late prophase adjacent condensed regions fused to form cylinder-shaped chromosomes. Based on these observations, we come to the conclusion that the construction of prophase chromosomes is a two-step process, that is, the reorganization and condensation of chromatin. In addition, we report the study of silver-stained, DNA- and histone-depleted prophase chromosomes, describe morphological features of the non-histone protein (NHP) residue in early, middle and late prophase chromosomes, and discuss the roles of NHPs in chromosome construction.     

Cold-induced Chromosome Regions and Karyosystematics in Sambucus and Viburnum

The genera Viburnum, Sambucus and Lonicera have been investigated for chromosome number and karyomorphology including Giemsa-C-banding, fluorochrome (DAPI/CMA) banding and cold treatment. Cold-induced undercontracted chromosome regions (CIRs) are found in Viburnum and Sambucus for the first time and are apparently identical with larger hc regions, shown by Giemsa C-banding. Certain narrow C-bands are not cold-sensitive. CIRs frequently react brightly CMA-positive in Viburnum and Sambucus, while DAPI fluorescence is virtually ineffective. The occurrence of CIRs within plants is possibly linked to certain nuclear characters such as large chromosomes and continuous condensation behaviour. Cold-induction has possibly also some influence on euchromatin condensation characteristics in prophasic chromosomes. Several karyological characters point to a closer relationship between Viburnum, Sambucus and Adoxa: Relatively large chromosomes, continuous condensation behaviour, reticulate to semireticulate interphase nuclei and presence of CIRs. These genera appear isolated from Lonicera and the Caprifoliaceae s.str., which differ remarkably in karyomorphology.     

Heteropycnosis of an underreplicating chromosome

Drosophila nasutoides has an extraordinary genome since 62% of its DNA resides in chromosome4. This element mainly consists of constitutive heterochromatin which does not polytenize. Earlier studies of heterochromatin attributed little attention to the fact that condensed chromosomes often vary in condensation. This paper reports that chromosomes of the same complement display different degrees and kinetics of condensation. InD. nasutoides, even sex specific differences can be observed. The results of a comparative microphotometric study on neuroblast metaphases in both sexes revealed the following picture. The process of chromosome condensation is not restricted to mitotic prophase but continues into the metaphase. The mean condensation is not equal for all chromosomes. In the metaphase of the female, Feulgen density increases from theX chromosome, via3 and2, to chromosome4. In the male, the order isX, 2, 3, Y, and4. During the metaphase of the male, chromosomes condense with similar kinetics. In contrast, chromosomes of the female display asynchrony as monitored by area and length determinations. TheX chromosomes of the female probably have enhanced shortening during prophase. This would explain the metaphase of the female where theX chromosomes shorten less than the autosomes, and why each of theX chromosomes is 15% shorter than theX chromosome in the metaphase of the male. Further differences were observed in the longitudinal and lateral compaction of the chromosomes in males and females. The sex chromosomes and chromosome3 condense by shortening, while chromosomes2 and4 preferentially reduce their diameter. The large amount of DNA engaged in heteropycnosis and the isochromosome nature allow the identification of chromosome4 during interphase. At this stage, a new category of extreme DNA packaging was detected. The interphase density of chromosome4 can exceed that of metaphase by a factor of up to 8. Two events account for this high degree of condensation:(1) the homologues are particularly associated due to somatic pairing and (2) the arms are further tightened as a result of pericentric folding. The features of the isochromosome suggest that the interaction of chromatids during interphase is essentially caused by specific DNA sequences. The data confirm that heteropycnosis not only interferes with gene expression but also strongly inhibits DNA synthesis in endocycles.     

Pericarp structure and phylogeny of theLamiaceae-Verbenaceae-complex

Pericarp structure was investigated in 158 species of the familiesLamiaceae andVerbenaceae. Data from 221 out of 262 genera ofLamiaceae s.l. and a few ofVerbenaceae s.str. were collected in a table. A cladistic analysis was performed on the basis of pericarp characters only. The abandonment of subfam.Pogostemonoideae as a taxonomic unit is considered. Examples of groups given additional support by similarities in pericarp characters are: (1) the gynobasic-styled labiates (subfamiliesPogostemonoideae, Lamioideae, Nepetoideae); (2) aLamioideae-Pogostemonoideae-group; (3)Nepetoideae; (4) aWestringia-Hemigenia-Hemiandra-Microcorys group (in subfam.Chloranthoideae); (5) aLepechinia-Chaunostoma-group (inNepetoideae); (6) aPrunella-Cleonia-group (inNepetoideae).     

Karyomorphological studies in some parasitic species of the Scrophulariaceae,I

A karyomorphological comparison of four genera of the tribe Buchnereae and two genera of the tribe Euphrasieae, root parasites of the subfamily Rhinanthoideae of the Scrophulariaceae, is presented along with 12 new chromosome counts. The interphase nuclei are of the simple chromocenter type. Macranthera and Aureolaria have more subtelocentric chromosomes than those of the other genera studied, which indicates higher asymmetric variation in their karyotypes. In all the karyotypes decrease in size from the longest to the shortest chromosomes is gradual. All the species studied show karyotypic heteromorphology and heteromorphic homologous chromosomes.     

Localization of chromosome regions in potoroo nuclei (<Emphasis Type="Italic"> Potorous tridactylus </Emphasis>Marsupialia: Potoroinae)

Chromosome paints of the rat kangaroo ( Aepyprymnus rufuscens, 2n =32) were used to define chromosome regions in the long nosed potoroo ( Potorous tridactylus, 2n =12 female, 13 male) karyotype and localize these regions in three-dimensionally preserved nuclei of the potoroo to test the hypothesis that marsupial chromosomes have a radial distribution. In human nuclei chromosomes are distributed in a proposed radial fashion. Gene-rich chromosomes in the human interphase nucleus are preferentially located in the central area while gene-poor chromosomes are found more at the periphery of the nucleus; this feature is conserved in primates and chicken. Chromosome ordering in nuclei of P. tridactylus is related to their size and centromere position. Its relationship with replication patterns in interphase nuclei and metaphase was studied. In addition it was observed that the nucleus was not a smooth entity but had projections occupied by specific chromosome regions. Edited by: R. Allshire     

Nuclear DNA Amounts in Palms (Arecaceae)

Nuclear DNA amounts are reported for 83 species and 53 genera of palms, covering all of the six subfamilies. 4C DNA contents range between 3.89 and 55.62 pg in diploids, showing an approximate 14.3-fold variation in genome size. Polyploids have DNA contents of up to 156.40 pg/4c which demonstrates a 40.2-fold variation. Diploids with high DNA contents occur in three subfamilies of palms (Coryphoideae, Calamoideae, Arecoideae), and seem to be further restricted to particular tribes or subtribes (Thrinacinae, Borasseae, Lepidocaryeae, Caryoteae, some subtribes of Areceae). Palms from the subfamilies Nypoideae and Phytelephantoideae have the lowest DNA amounts, followed by the Phoeniceae and the Corypheae: Livistoninae from the subfamily Coryphoideae. Although DNA amounts in some genera and subtribes are usually constant, e.g., in Phoenix, Phytelephas, the Livistoninae, Dypsidinae, diploid Butiinae), considerable variation occurs at the diploid level in some large and apparently actively evolving genera such as Chamaedorea, Pinanga, Cenoma and possibly Bactris. Formaldehyde fixation is recommended for palms, as conventional ethanol-acetic acid fixation has proved to be unsuitable for DNA estimation of Feulgen-stained nuclei by microdensitometry, since it can lead to errors up to 2.5-fold in extent. Chromosome counts are reported for 72 of the species studied, of which 42 are new.     

Dynamics of chromosome compaction during mitosis

We have quantitatively studied the space-time dynamics of mitotic chromosome compaction in cultured amphibian cells. After collecting digital phase-contrast images we have done digital image analysis to study spatial correlations in density. We find a characteristic distance at which the strongest correlations occur, which provides a quantitative measure of the size of patches of dense chromatin during interphase and early prophase. Later in mitosis, this length corresponds to the thickness of prophase and metaphase chromosomes. We find that during interphase strong correlations exist at a few-micrometer length; during prophase this correlation length progressively drops as the chromosomes are compacted. Our data are explained by a model based on assembly of chromatin loops onto already fiberlike interphase chromosomes. To test this model we have microinjected cobalt hexamine trichloride into interphase nuclei and have observed the rapid condensation of the interphase chromatin into thick fibers with a spacing similar to the native-state interphase correlation length determined from our image analysis.     

Meiosis-like reduction during somatic embryogenesis of Arabidopsis thaliana

Summary Somatic meiosis-like reduction was observed in some cells of the embryogenic callus of Arabidopsis thaliana. Two types were identified. One type was somatic chromosome reductional grouping, in wich the chromesomes in a cell were separated direetly at either prophase or metaphase. Chromosome reductional grouping happened more frequently in polyploid cells, and the morphology of the chromosomes did not show the role of the spindle fibers. The other type was somatic meiosis which was analogous to the process of gametogenesis, characterized by the pairing and synapsis of homologous chromosomes. The roles of somatic meiosis-like reduction in somatic embryogenesis and somaclonal variations are discussed     

Stygiinae Yakovlev,subfam. n., a new subfamily of palaearctic carpenter-moths (Lepidoptera,Cossidae)

A new subfamily of carpenter moths, Stygiinae Yakovlev, subfam. n. (Lepidoptera, Cossidae) with two genera, Stygia Latreille, [1803] (type genus) and Neostygia Wiltshire, 1980, is described. A brief catalogue of all five species belonging to this subfamily is given.     

Loxocaudinae: A new subfamily in the ostracod family loxoconchidae

A new Ostracoda subfamily, Loxocaudinae subfam. nov., separated from the subfamily Loxoconchinae of the family Loxoconchidae with five genera (Loxocauda Schornikov, 1969, Glacioloxoconcha Hartmann, 1990, Phlyctocythere Keij, 1958, Pseudoloxoconcha Müller, 1894 and Sarmatina Stancheva, 1984) is described in the paper. The new subfamily differs from Loxoconchinae Sars, 1926 s. str. by the presence of a compact eye, the absence of an eye tubercle, an originally adont hinge with a tendency to formation of an anterior tooth on the left valve and a posterior one on the right valve, a pronounced caudal process and reduction of fossa-mural sculpture. The volume of the genera that are included in the subfamily is determined. A review of 55 various species and forms mentioned in the literature in open nomenclature, whose taxonomic position has not been determined yet is presented. A total of 45 species close to Loxocauda are attributed to the new subfamily, but proper morphological investigations are necessary for their classification. Ten species that were referred earlier to the genera included in Loxocaudinae are excluded from this subfamily. Issues of functional morphology of Loxocaudinae shells and morphological evolution of their sculpture are considered.     

End-to-end chromosome attachments in mitotic interphase and their possible significance to meiotic chromosome pairing

Cytological studies on telophase and early prophase in roottip cells of several plant species (Allium cepa, 2n=16; four Crepis species, including Crepis capillaris, 2n=6; Callitriche hermaphroditica, 2n=6; Nigella arvensis, 2n=12; Secale cereale, 2n=14) revealed that chromosome ends are attached two by two forming chains of chromosomes (interphase associations). In these chains homologous chromosomes are presumably located adjacent to each other. In Crepis capillaris it was observed that the two nucleolar chromosomes form a separate ring one end attached to the ring of the four remaining chromosomes and the other end attached to the nucleolus. It is proposed that these end-to-end attachments have significance for chromosome pairing in meiosis. The adjacent location of homologous chromosomes in the interphase associations would facilitate rapid and regular synapsis.     

Karyomorphology of four species in Ancylostemon,Briggsiopsis and Lysionotus (Gesneriaceae)

Reported in the present paper are chromosome numbers and karyotypes of three genera of the Gesneriaceae, i.e. Ancylostemon Craib. , Briggsiopsis (Franch.) K. Y. Pan and Lysionotus D. Don. The former two genera are endemic to China. The karyotype of Ancylostemon aureus (Franch.) Burtt is formulated as 2n = 34 = 20m(1sat) + 14sm, with the same chromosome number as its allied species A. convexus Craib. This species is characterized by the interphase nucleus of complex chromocenter type and the proximal type of chromosomes in the mitotic prophase. The chromosome number of the monospecific genus Briggsiopsis is 2n = 34, the same as the lowest chromosome number reported in Briggsia. The karyotype of Briggsiopsis, which is formulated as 2n = 25m + 6sm + 3st, also seems to be primitive among the species of the two genera. Briggsiopsis is characterized by the interphase nucleus of simple-complex chromocenter type and the interstitial-gradient type of chromosomes in the mitotic prophase. The chromosome number of Lysionotus carnosus Hemsl. is the lowest reported in this genus. Its karyotype is formulated as 2n= 30 = 21m + 5sm + 3st + lt. Lysionotus serratus var. pterocaulis, with the karyotype being formulated as 2n= 32 = 2lm + 10sm + lt, has the same chromosome number as var. serratus. These two species show a remarkable differentiation of karyotypes and are characterized by the interphase nuclei of simple-complex chromocenter type and the gradient type of chromosomes in the mitotic prophase. _ .     

A human condensin complex containing hCAP-C-hCAP-E and CNAP1, a homolog of Xenopus XCAP-D2, colocalizes with phosphorylated histone H3 during the early stage of mitotic chromosome condensation

Structural maintenance of chromosomes (SMC) family proteins play critical roles in structural changes of chromosomes. Previously, we identified two human SMC family proteins, hCAP-C and hCAP-E, which form a heterodimeric complex (hCAP-C-hCAP-E) in the cell. Based on the sequence conservation and mitotic chromosome localization, hCAP-C-hCAP-E was determined to be the human ortholog of the Xenopus SMC complex, XCAP-C-XCAP-E. XCAP-C-XCAP-E is a component of the multiprotein complex termed condensin, required for mitotic chromosome condensation in vitro. However, presence of such a complex has not been demonstrated in mammalian cells. Coimmunoprecipitation of the endogenous hCAP-C-hCAP-E complex from HeLa extracts identified a 155-kDa protein interacting with hCAP-C-hCAP-E, termed condensation-related SMC-associated protein 1 (CNAP1). CNAP1 associates with mitotic chromosomes and is homologous to Xenopus condensin component XCAP-D2, indicating the presence of a condensin complex in human cells. Chromosome association of human condensin is mitosis specific, and the majority of condensin dissociates from chromosomes and is sequestered in the cytoplasm throughout interphase. However, a subpopulation of the complex was found to remain on chromosomes as foci in the interphase nucleus. During late G(2)/early prophase, the larger nuclear condensin foci colocalize with phosphorylated histone H3 clusters on partially condensed regions of chromosomes. These results suggest that mitosis-specific function of human condensin may be regulated by cell cycle-specific subcellular localization of the complex, and the nuclear condensin that associates with interphase chromosomes is involved in the reinitiation of mitotic chromosome condensation in conjunction with phosphorylation of histone H3.     

TRANSIENT LOCALIZATION OF γ-TUBULIN AROUND THE CENTRIOLES IN THE NUCLEAR DIVISION OF BOERGESENIA FORBESII (SIPHONOCLADALES, CHLOROPHYTA)

Mitosis in Boergesenia forbesii (Harvey) Feldman was studied by immunofluorescence microscopy using anti-β–tubulin, anti-γ–tubulin, and anti-centrin antibodies. In the interphase nucleus, one, two, or rarely three anti-centrin staining spots were located around the nucleus, indicating the existence of centrioles. Microtubules (MTs) elongated randomly from the circumference of the nuclear envelope, but distinct microtubule organizing centers could not be observed. In prophase, MTs located around the interphase nuclei became fragmented and eventually disappeared. Instead, numerous MTs elongated along the nuclear envelope from the discrete anti-centrin staining spots. Anti-centrin staining spots duplicated and migrated to the two mitotic poles. γ–Tubulin was not detected at the centrioles during interphase but began to localize there from prophase onward. The mitotic spindle in B. forbesii was a typical closed type, the nuclear envelope remaining intact during nuclear division. From late prophase, accompanying the chromosome condensation, spindle MTs could be observed within the nuclear envelope. A bipolar mitotic spindle was formed at metaphase, when the most intense staining of γ-tubulin around the centrioles could also be seen. Both spindle MT poles were formed inside the nuclear envelope, independent of the position of the centrioles outside. In early anaphase, MTs between separating daughter chromosomes were not detected. Afterward, characteristic interzonal spindle MTs developed and separated both sets of the daughter chromosomes. From late anaphase to telophase, γ-tubulin could not be detected around the centrioles and MT radiation from the centrioles became diminished at both poles. γ-Tubulin was not detected at the ends of the interzonal spindle fibers. When MTs were depolymerized with amiprophos methyl during mitosis, γ-tubulin localization around the centrioles was clearly confirmed. Moreover, an influx of tubulin molecules into the nucleus for the mitotic spindle occurred at chromosome condensation in mitosis.