Chromosome numbers in Heterotheca,including Chrysopsis (Compositae: Astereae), with Phylogenetic interpretations

A summation of both previously reported and original data on chromosome numbers is presented for species ofHeterotheca sens. lat. Chromosome numbers are listed for 24 taxa, including 20 different species. Chromosome numbers for the genus aren = 4, 5, 9, 12, and 18. The most common and possibly basic number for the genus isn = 9, with then = 18 taxa considered to be tetraploids, and those withn = 4 and 5 as probable aneuploid derivatives ; alternativelyn = 4 and 5 could be considered the basic numbers withn = 9 the result of polyploidy. Several nomenclatural changes have been made including the following new combinations:Heterotheca bolanderi (Gray) Harms,H. fastigiata (Greene) Harms, andH. villosa var.hispida (Hook.) Harms.     

Evolution and systematic relationships in theTriticeae (Poaceae)

The evolution and taxonomic relationships in theTriticeae are discussed with the view to highlight aspects of this agronomically important group of plants, which may be of interest to molecular biology. Some of these aspects are addressed in more detail in adjoining papers in which specific genomic loci have been examined at the DNA sequence or isozyme level. Aspects discussed include the systematics and geographic distribution of theTriticeae species, isozyme and chromosome pairing studies on some of the species as well as more recent developments in DNA analyses. A survey of the systematics of theTriticeae indicated that the genomic system ofLöve is probably the most useful starting point for interpreting molecular data even though the system has many problems from a taxonomic point of view. The geographical distribution ofTriticeae species, using both published and unpublished data, suggested that information of this type taken together with the theory of continental drift provides a broad time-span for considering data from DNA sequence studies. The significance, and modes of analyses, of isozyme studies were assessed because they often provide valuable characters in determining relationships between species. The main character underlyingLöve's andDewey's analyses of theTriticeae, namely chromosome pairing, is discussed with particular reference to isozyme studies to show that in some cases, such as species ofHordeum sensu lato, consistent relationships are obtained. Finally, new developments in understanding chromosome structure are considered in relation to the above variables in the taxonomy and evolution of theTriticeae.     

The nature of polyploidy inReseda sect.Leucoreseda (Resedaceae)

We have found two chromosome levels (n = 10 and n = 20) in Spanish species ofReseda sect.Leucoreseda, four species on the first (R. undata, R. paui, R. suffruticosa, R. barrelieri), only one on the second level (R. alba). As already the species with n = 10 apparently behave as polyploids, we propose x = 5 as the original basic chromosome number for this section. The seed protein profile ofR. alba (2n = 40) reveals close relationships withR. undata (2n = 20), while the esterase isozymes suggest affinities withR. paui (2n = 20). Thus,R. alba can be regarded either as an autopolyploid fromR. paui or more likely as an allopolyploid fromR. paui andR. undata. Finally, the DNA values suggest a diploidization process inR. alba since its origin.     

Chromosome numbers of Panamanian Lecythidaceae and their use in subfamilial classification

Nine species of Lecythidaceae subfamily Lecythidoideae in four genera whose chromosome numbers were previously unknown, have 17 as their basic chromosome number:Eschweilera pittieri, three other unidentified species ofEschweilera, Grias cauliflora, Gustavia dubia, G. superba, Lecythis minor, andL. tuyrana. All are diploid exceptGustavia superba, which is tetraploid.Couroupita guianensis, which was previously—and probably incorrectly—reported to have a gametic chromosome number of 18, also hasn = 17. The known chromosome numbers support recognizing at least three of Niedenzu’s subfamilies: Planchonioideae withx = 13, Napoleonaeoideae withx = 16, and Lecythidoideae withx = 17. His fourth subfamily, Foetidioideae, with one genus of five species, has not been counted. Cytological data have been and probably will be useful in indicating to what subfamily problematic genera belong and in showing interesting phytogeographic patterns within the family. On the other hand, cytological data provide no recognizable clues relating the Lecythidaceae to other families.     

Evolutionary breakpoint regions and chromosomal remodeling in Harttia (Siluriformes: Loricariidae) species diversification

The Neotropical armored catfish genus Harttia presents a wide variation of chromosomal rearrangements among its representatives. Studies indicate that translocation and Robertsonian rearrangements have triggered the karyotype evolution in the genus, including differentiation of sex chromosome systems. However, few studies used powerful tools, such as comparative whole chromosome painting, to clarify this highly diversified scenario. Here, we isolated probes from the X₁ (a 5S rDNA carrier) and the X₂ (a 45S rDNA carrier) chromosomes of Harttia punctata, which displays an X₁X₁X₂X₂/X₁X₂Y multiple sex chromosome system. Those probes were applied in other Harttia species to evidence homeologous chromosome blocks. The resulting data reinforce that translocation events played a role in the origin of the X₁X₂Y sex chromosome system in H. punctata. The repositioning of homologous chromosomal blocks carrying rDNA sites among ten Harttia species has also been demonstrated. Anchored to phylogenetic data it was possible to evidence some events of the karyotype diversification of the studied species and to prove an independent origin for the two types of multiple sex chromosomes, XX/XY₁Y₂ and X₁X₁X₂X₂/X₁X₂Y, that occur in Harttia species. The results point to evolutionary breakpoint regions in the genomes within or adjacent to rDNA sites that were widely reused in Harttia chromosome remodeling.     

Origin of the X1X1X2X2/X1X2Y sex chromosome system of Harttia punctata (Siluriformes,Loricariidae) inferred from chromosome painting and FISH with ribosomal DNA markers

Harttia is a genus in the subfamily Loricariinae that accommodates fishes popularly known as armored catfishes. They show extensive karyotypic diversity regarding interspecific numerical/structural variation of the karyotypes, with the presence of the XX/XY₁Y₂ multiple sex chromosome system, as found in H. carvalhoi. In this context, this study aimed to characterize Harttia punctata chromosomally, for the first time, and to infer the rearrangements that originated the X₁X₁X₂X₂/X₁X₂Y multiple sex chromosome system present in this species. The data obtained in this study, with classical (Giemsa, C-banding and AgNORs) and molecular methodologies (fluorescence in situ hybridization) and chromosome microdissection, indicated that a translocation between distinct acrocentric chromosomes bearing rRNA genes, accompanied by deletions in both chromosomes, might have originated the neo-Y chromosome in this species. The data also suggest that the multiple sex chromosome systems present in H. carvalhoi and H. punctata had an independent origin, evidencing the recurrence of chromosome alterations in species from this genus.     

ISOZYME NUMBER IN SUBTRIBE ONCIDIINAE (ORCHIDACEAE): AN EVALUATION OF POLYPLOIDY

The Oncidiinae has attracted attention because of the variation it exhibits in chromosome number, n = 5–30, which is greater than the range in the rest of the Orchidaceae. The genus Psygmorchis, with n = 5 and 7, has been a particular focus of controversy, and many authors have suggested that 5 and 7 are the base numbers for the subtribe. The other taxa in the subtribe presumably evolved through hybridization and polyploidy. Other workers have found that the lowest counts correlate with derived morphological conditions and have hypothesized that these low numbers result from aneuploid reductions, while higher numbers are associated with ancestral morphologies and are not the result of polyploidy. These two hypotheses were evaluated by determining isozyme numbers for 13 enzymes in species that span the chromosomal range known for the Oncidiinae (n = 5–30). Isozyme number has been shown to be a reliable indicator of polyploidy in angiosperms because polyploids display isozyme multiplicity relative to diploids. This analysis revealed no differences among species in isozyme number for the enzymes examined. Therefore, our data reject the hypothesis that species with higher chromosome numbers are polyploid.     

Isozymic gene linkage map of the tomato: Applications in genetics and breeding

Summary New linkage data are presented for the situation of five previously unlocated isozymic loci of the tomato and closely related species with homosequential chromosomes.Prx-1 lies on chromosome 1, where it is also linked withSkdh-1; Aps-2 is linked withGot-4 on chromosome 8;Tpi-2 has been allocated to chromosome 4; and a linkage has been detected betweenPgi-1 andEst-4, whose respective chromosome has not yet been determined. These and previously published data have been summarized in the form of an isozyme linkage map. Twenty-two loci have thus been mapped on nine of the twelve tomato chromosomes. We discuss some new applications of mapped isozymic genes. In certain types of segregations, isozymic genes are far more efficient than morphological markers in providing linkage information. They greatly expedite the cytogenetic investigation of species hybrids and can be utilized to facilitate backcross transfers of genes from wild to cultivated taxa.     

A multiple sex-chromosome system in Antarctic ice-fishes

Summary We have studied the chromosomes of 11 of the 15 known species of the notothenioid family Channichthyidae, the specialized whiteblooded Teleosts endemic to the Southern Ocean (ice-fishes). In the female sex, all studied species have the same diploid number of forty eight mostly acrocentric (uniarmed) chromosomes; however there is an interspecific variability in the chromosome morphology, type and quantity of repetitious DNA (usually seen as heterochromatin) localization of silver-stained nucleolar organizers. At least five of the studied species show a multiple sex-chromosome system possibly originated by the translocation of an autosome on an early Y gonosome morphologically similar to the X: the digametic males (2n = 47) show a X₁Y X₂ and the homogametic females (2n = 48) a X₁X₁X₂X₂ gonosomic constitution. This peculiar sex determining mechanism, otherwise rare in Teleosts, can be considered apomorphic in the same way as other morphofunctional characters usually interpreted as adaptive in these fishes.Some of the data presented here were collected during the European Polarstern Study (EPOS) sponsored by the European Science Foundation     

Multiple sex chromosome system of X1X1X2X2/X1X2)Y type in lutjanid fish, Lutjanus quinquelineatus (Perciformes)

The karyotype and other chromosomal markers as revealed by C-banding and Ag-staining were studied in Lutjanus quinquelineatus and L. kasmira (Lutjanidae, Perciformes). While in latter species, the karyotype was invariably composed of 48 acrocentric chromosomes in both sexes, in L. quinquelineatus the female karyotype had exclusively 48 acrocentric chromosomes (2n = 48) but that of the male consisted of one large metacentric and 46 acrocentric chromosomes (2n = 47). The chromosomes in the first meiotic division in males showed 22 bivalents and one trivalent, which was formed by an end-to-end association and a chiasmatic association. Multiple sex chromosome system of X₁X₁X₂X₂/X₁X₂Y type resulting from single Robertsonian fusion between the original Y chromosome and an autosome was hypothesized to produce neo-Y sex chromosome. The multiple sex chromosome system of L. quinquelineatus appears to be at the early stage of the differentiation. The positive C-banded heterochromatin was situated exclusively in centromeric regions of all chromosomes in both species. Similarly, nucleolus organizer region sites were identified in the pericentromeric region of one middle-sized pair of chromosomes in both species. The cellular DNA contents were the same (3.3 pg) between the sexes and among this species and related species.     

A bubaline-derived satellite DNA probe uncovers generic affinities of gaur with other bovids

DNA typing using genome derived cloned probes may be conducted for ascertaining genetic affinities of closely related species. We analysed gaurBos gaurus, cattleBos indicus, buffaloBubalus bubalis, sheepOvis aries and goatCapra hircus DNA using buffalo derived cloned probe pDS5 carrying an array ofBamHI satellite fraction of 1378 base residues to uncover its genomic organization. Zoo-blot analysis showed that pDS5 does not cross hybridize with non-bovid animals and surprisingly with female gaur genomic DNA. The presence of pDS5 sequences in the gaur males suggests their possible location on the Y chromosome. Genotyping of pDS5 withBamHI enzyme detected mostly monomorphic bands in the bubaline samples and polymorphic ones in cattle and gaur giving rise to clad specific pattern. Similar typing withRsaI enzyme also revealed clad specific band pattern detecting more number of bands in buffalo and fewer in sheep, goat and gaur samples. Copy number variation was found to be prominent in cattle and gaur withRsaI typing. Our data based on matched band profiles (MBP) suggest that gaur is genetically closer to cattle than buffalo contradicting the age-old notion held by some that gaur is a wild buffalo. The pDS5 clone has a potential for estimating the generic and genetic relationship amongst closely related bovid species.     

Ploidy determination in Sphagnum samples from Svalbard,Arctic Norway,by DNA image cytometry

Abstract

We measured DNA content of cell nuclei, stained with the Feulgen method, using branch tips of 11 species of Sphagnum from Svalbard, Arctic Norway, as an alternative to chromosome counting. Nine species were haploid and two were diploid, with no intraspecific variation in ploidy level. The results conformed to known chromosome numbers and/or to expectations from isozyme studies. Ploidy levels were determined for the first time in S. tundrae and S. fimbriatum ssp. concinnum (haploid) and S. arcticum and S. olafii (diploid). No mitotic divisions were observed, but unreplicated interphase nuclei still allowed precise ploidy determinations. Basic DNA contents of all Sphagnum species were very similar, and measurement of a few nuclei proved sufficient to ascertain ploidy level despite very low nuclear DNA content. Advantages of the DNA image cytometry method are: mitotic or meiotic cells are not required to be found, and only a small amount of material is required.     

New insights into sex chromosome evolution in anole lizards (Reptilia,Dactyloidae)

Anoles are a clade of iguanian lizards that underwent an extensive radiation between 125 and 65 million years ago. Their karyotypes show wide variation in diploid number spanning from 26 (Anolis evermanni) to 44 (A. insolitus). This chromosomal variation involves their sex chromosomes, ranging from simple systems (XX/XY), with heterochromosomes represented by either micro- or macrochromosomes, to multiple systems (X₁X₁X₂X₂/X₁X₂Y). Here, for the first time, the homology relationships of sex chromosomes have been investigated in nine anole lizards at the whole chromosome level. Cross-species chromosome painting using sex chromosome paints from A. carolinensis, Ctenonotus pogus and Norops sagrei and gene mapping of X-linked genes demonstrated that the anole ancestral sex chromosome system constituted by microchromosomes is retained in all the species with the ancestral karyotype (2n?=?36, 12 macro- and 24 microchromosomes). On the contrary, species with a derived karyotype, namely those belonging to genera Ctenonotus and Norops, show a series of rearrangements (fusions/fissions) involving autosomes/microchromosomes that led to the formation of their current sex chromosome systems. These results demonstrate that different autosomes were involved in translocations with sex chromosomes in closely related lineages of anole lizards and that several sequential microautosome/sex chromosome fusions lead to a remarkable increase in size of Norops sagrei sex chromosomes.     

Chromosome numbers in callirhoe (Malvaceae)

Forty-four chromosome counts from the North American genusCallirhoe are reported, including first-recorded counts ofC. alcaeoides, C. digitata, C. bushii, C. triangulata, andC. involucrata var.tenuissima. Information concerning chromosome numbers from present and previous reports is summarized and evaluated. Observations are presented on meiosis for the genus in general, for male sterile individuals of gynodioecious species, and for selected F₁ hybrids. Pollen fertility data are recorded for the latter. Chromosomally three species groups are recognized: an exinvolucellate, diploid species group withn=14; an involucellate, tetraploid and octoploid species pair,n=28 and 56; and an involucellate, diploid and tetraploid species group withn=15 and 30. Intrageneric relationships and those within the tribe Malveae subtribe Malvinae are discussed.     

Chromosomal segregational mechanisms in ant-lions (Myrmeleontidae,Neuroptera)

In a single male specimen of Myrmeleon mexicanum Banks the sex chromosomes, normally X and Y, were replaced by what appeared to be X¹X² and Y. These segregated as expected on that interpretation in only half of the spermatocytes — in the other half, one X and the Y segregated from the other X. This atypical segregation is explicable on the assumption that one of the supposed Xs is a supernumerary, not a sex chromosome, and the diploid complement of the male comprises six pairs of autosomes plus a supernumerary and the X and Y sex chromosomes. The orientation of the X chromosomes at first metaphase was variable: kinetochoric activity may be localized midway the length of the chromosome, as in gonial mitosis, or terminally. Comparative study of three congeneric species, seven of Brachynemurus, one of Psammoleon, and one of Vella showed normal segregation in all, and no evidence for secondary kinetochoric activity. In nine of the species studied one pair of autosomes was unconjoined at first metaphase in 0.3%–1.2% of primary spermatocytes. These autosomes segregated precociously with the sex chromosomes in the central unit of the spindle. In one exceptional male of Brachynemurus hubbardi Currie all first meiotic metaphases showed this behavior, and a compound X¹X²/Y¹Y² system was thus simulated. Bivalent formation replaced distance segregation of sex chromosomes in 0.4%–3.2% of the spermatocytes in seven of the thirteen species studied. These sex-bivalents frequently displayed partial or complete failure in congression.     

Determination of ploidy level and nuclear DNA content in blueberry by flow cytometry

The technique of DNA flow cytometry was used to study variation in DNA content among different ploidy levels, as well as among diploid species, of Vaccinium section Cyanococcus. In a sample of plants of varying ploidy level, the relative fluorescence intensity (RFI) of nuclei stained with propidium iodide was a function of the number of chromosome sets (x), as represented by the linear equation RFI=3.7x-2.3 (r²=95%). The data indicated that DNA flow cytometry could be useful for the determination of ploidy level at the seedling stage in blueberry. They also suggest that conventional polyploid evolution has occurred in this section of the genus Vaccinium with an increase in nuclear DNA content concurrent with the increase in chromosome number. The nuclear DNA content of diploid species of Vaccinium section Cyanococcus was estimated from the relationship of the observed RFI to an internal known DNA standard (trout red blood cells). A nested analysis of variance indicated significant variation among species, as well as among populations within species, in nuclear DNA content, although this variation was small compared to the variation among ploidy levels. The variation in nuclear DNA content corresponded to the phylogenetic relationships among species determined from previous studies.     

Karyotypic conservatism in five species of Prochilodus (Characiformes,Prochilodontidae) disclosed by cytogenetic markers

The family Prochilodontidae is considered a group with well conserved chromosomes characterized by their number, morphology and banding patterns. Thence, our study aimed at accomplishing a cytogenetic analysis with conventional methods (Giemsa staining, silver staining of the nucleolus organizer regions-AgNOR, and C-banding) and fluorescence in situ hybridization (FISH) with 18S and 5S ribosomal DNA probes in five species of the Prochilodus genus (Prochilodus argenteus, Prochilodus brevis, Prochilodus costatus, Prochilodus lineatus and Prochilodus nigricans) collected from different Brazilian hydrographic basins. The results revealed conservatism in chromosome number, morphology, AgNORs 18S and 5S rDNAs location and constitutive heterochromatin distribution patterns. The minor differences observed in this work, such as an Ag-NOR on a P. argenteus chromosome and a distinct C-banding pattern in P. lineatus, are not sufficient to question the conservatism described for this group. Future work using repetitive DNA sequences as probes for FISH will be interesting to further test the cytogenetic conservatism in Prochilodus.     

Cytogenetic characterization of ten araneomorph spiders (Araneae): Karyotypes and meiotic features

Karyotypes, sex chromosome systems and meiotic characteristics are reported for ten spider species belonging to the families Gnaphosidae, Philodromidae, Salticidae, Oxyopidae and Sicariidae by using standard Giemsa staining. The male diploid numbers (2n) and sex chromosome systems are as follows: Berinda hakani 2n = 22 (X₁X₂), Berinda ensigera 2n = 22 (X₁X₂), Trachyzelotes lyonneti 2n = 22 (X₁X₂), Trachyzelotes malkini 2n = 22 (X₁X₂), Zelotes caucasius 2n = 22 (X₁X₂) (Gnaphosidae); Thanatus pictus 2n = 28 (X₁ X₂), Tibellus macellus 2n = 24 (X₁ X₂) (Philodromidae); Neon reticulatus 2n = 21 (X₀) (Salticidae); Peucetia virescens 2n = 28 (X₁X₂) (Oxyopidae) and Loxosceles rufescens 2n = 21 (X₁ X₂Y) (Sicariidae). All species have monoarmed chromosomes with the exception of L. rufescens that has biarmed (metacentric and submetacentric) chromosomes. The obtained data are the first results for the genera Berinda, Trachyzelotes and Neon. Additionally, with the exception of L. rufescens, all species are being chromosomally analyzed for the first time.     

The nature and status of Lysimachia X producta (Primulaceae)—II

Data from chromosome numbers and from morphological variation in wild plants and artificial hybrids further support the hypothesis thatLysimachia Xproducta is of hybrid origin and the taxonomic judgment that it is best treated at the status of interspecific hybrid rather than as a species of hybrid origin. Additional distribution data are presented.     

Genetic divergence among species of the section Strombocarpa,genusProsopis (Leguminosae)

We studied degree of variability and relationships among populations belonging to five species of the section Strombocarpa of the family Leguminosae, namelyProsopis ferox, P. torquata, P. pubescens, P. strombulifera andP. reptans, by isozyme electrophoresis. The genetic similarity betweenP. reptans andP. strombulifera is high enough for the two to be considered subspecies or geographic races ofP. strombulifera instead of good taxonomic species. As expected these species cluster withP. pubescens, all belonging to series Strombocarpae. This group is different isoenzymatically from the remaining species,P. torquata andP. ferox. Comparison of our results with those obtained earlier for species of the section Algarobia showed that genetic variability in the section Strombocarpa is lower than in the section Algarobia, while the species divergence is much higher. These results suggest different adaptive strategies between these sections.