THE DISTRIBUTION OF ANTHOCHLOR FLORAL PIGMENTS IN NORTH AMERICAN COREOPSIS (COMPOSITAE): TAXONOMIC AND PHYLETIC INTERPRETATIONS

Anthochlors (chalcones and aurones) occur in floral tissues of all members of North American Coreopsis. The genus in North America consists of 46 species distributed in 11 sections, and a total of five chalcone aglycones are found in it. The chalcone butein occurs in all sections, and is the only anthochlor constituent in the closely related woody Mexican sections Electra and Anathysana. These two sections are viewed as primitive. The morphologically distinct but putatively unspecialized woody Mexican section Pseudo-Agarista sequesters both okanin and butein; the same two anthochlors also characterize two additional sections of herbaceous perennials, Silphidium and Tuckermannia. Sections Euleptosyne and Pugiopappus, comprised of annuals restricted primarily to California, exhibit the rare chalcone stillopsidin in addition to butein and okanin. The remaining four sections, Palmatae, Coreopsis, Calliopsis and Eublepharis are among the more specialized in North America, and are sometimes considered to represent a distinct phyletic line or two related lines. All or some of the members of these sections sequester the very rare chalcone lanceolatin in floral tissues in addition to okanin. Butein is also present in some or all species in each section, and isoliquiritigenin occurs rarely in two sections. Those sections considered more specialized have additional anthochlors relative to primitive sections, and any compound found in a more primitive section also occurs in a more advanced one. Thus, the trend in floral anthochlors in North American Coreopsis is toward increased number of compounds rather than reduction.     

INTERSPECIFIC HYBRIDIZATION IN HAPLOPAPPUS AND ITS BEARING ON CHROMOSOME EVOLUTION IN THE BLEPHARODON SECTION

Jackson , R. C. (U. Kansas, Lawrence.) Interspecific hybridization in Haplopappus and its bearing on chromosome evolution in the Blepharodon section. Amer. Jour. Bot. 49 (2) : 119–132. Illus. 1962.—Cytological analyses of interspecific hybrids between H. gracilis (n = 2) and H. spinulosus ssp. australis (n = 8) indicate that ssp. australis is a segmental allotetraploid, derived from past hybridization between 2 taxa with chromosome numbers of n = 4. Analysis of hybrids between H. gracilis (n = 2) and H. ravenii (n = 4), a previously undescribed species, has shown that the chromosome segments of these 2 species are almost completely homologous. Differential contraction is suggested as the explanation for the disappearance in late pachytene of presumed non-homologous segments which were evident in some cells at early pachytene. The pairing relationship of gracilis and ravenii chromosomes at pachytene and later prophase I stages of meiosis indicates that gracilis has evolved from ravenii by an aneuploid reduction process similar to that described for Crepis. The close morphological relationship of the 2 species adds further support to this proposition. Data from the cytological analysis of both interspecific hybrids indicate that x = 4 is the basic chromosome number for the Blepharodon section of Haplopappus.     

Quantitative nuclear DNA differences associated with genome evolution in Guizotia (Compositae)

The present communication deals with 2C nuclear genome size variation in a fairly small genus Guizotia. Twenty-four accessions belonging to six species, out of seven known, were analysed in order to elucidate the extent of DNA variation both at an intra—as well as interspecific level. At the intraspecific level none of the species exhibited significant differences in their genome size. Between the species, the 2C DNA amounts ranged from 3.61 pg in G. reptans to 11.37 pg in G. zavattarii; over three-fold DNA variation is evident. Apparently these interspecific DNA differences have been achieved independent of the numerical chromosomal change(s), as all the Guizotias share a common chromosome number 2n=2x=30. The cultivated oilseed crop, G. abyssinica (7.57 pg), has accommodated nearly 78% extra DNA in its chromosome complement during the evolutionary time scale of its origin and domestication from the wild progenitor G. schimperi (4.25 pg). The extent of genomic DNA difference(s) between the species has been discussed in the light of their interrelationships and diversity.     

New chromosome counts in the genus Centaurea (Asteraceae) from Turkey

Twenty‐two chromosome counts are reported in 16 species, four subspecies and two varieties of the genus Centaurea. These are mostly Turkish local endemics of section Cheirolepis, a complicated group from the Eastern clade of the Jacea group. Twenty‐one reports are new. Prevalence of the basic chromosome number x = 9 among the eastern sections of the Jacea group is confirmed. © 2009 The Linnean Society of London, Botanical Journal of the Linnean Society, 2009, 159 , 280–286.     

POLLEN TUBE GROWTH AND INCOMPATIBILITY FOLLOWING INTRA- AND INTER-SPECIFIC POLLINATIONS IN LILIUM LONGIFLORUM

The growth of pollen tubes of a sampling of Lilium species in styles of L. longiflorum incubated at 24 C for 48 hr after pollination indicated two types of interspecific incompatibility. Pollen tubes of two self-compatible species of section Leucolirion, the section including L. longiflorum, stopped growth abruptly upon reaching the stylar canal, were of abnormal morphology, and were incapable of continued growth with longer incubation. Pollen tubes of self-compatible or self-incompatible species of sections Sinomartagon, Pseudolirium, Liriotypus, and Daurolirion approached but did not exceed the length of intraspecific incompatible pollen tubes in styles of L. longiflorum. Pollen tube morphology was normal and tubes were capable of continued growth with additional incubation. Unilateral interspecific incompatibility occurred in reciprocal crosses between self-incompatible L. longiflorum and self-compatible L. regale and L. formosanum, but exceptions occurred in Aurelian hybrids. Incubation of interspecifically pollinated L. longiflorum styles at 39 C, which removes the self-incompatibility reaction, had no effect on interspecific incompatibility.     

Phylogenetic relationships in Carex, subgenus Vignea (Cyperaceae), based on ITS sequences

To evaluate the sectional classification in Carex, subgenus Vignea, the ITS region of 58 species of 20 sections was analyzed with Neighbor Joining (NJ) and Markov chain Monte Carlo (MCMC) methods. Sections Dioicae, Physodeae and Ovales are found to be monophyletic, with C. bohemica well integrated in the section Ovales. Section Heleonastes turns out to be monophyletic, if C. canescens is treated separately in section Canescentes. Section Elongatae is monophyletic, but C. remota is placed in section Remotae and C. bromoides in section Deweyanae. In both analyses, six representatives of section Arenariae cluster together in a terminal group, whereas C. disticha, C. repens and C. siccata form a basal cluster. C. maritima, as the only member of section Incurvae, shares this basal position. C. chordorrhiza is ascribed to section Chordorrhizeae and not ascribed to the paraphyletic section Divisae. C. vulpina and C. otrubae are assigned to section Vulpinae and separated from the heterogeneous section Stenorhynchae. The other members of sections Divisae, Muehlenbergianae, Multiflorae and Stenorhynchae are scattered throughout the trees. The representatives of section Foetidae are dispersed in both analyses, section Paniculatae appears to be non-monophyletic in the molecular results as well. The subgenus appears subdivided in at least four larger subgroups in all analyses. Whereas these subgroups are strongly supported, the relationships between these subgroups remain only poorly resolved.     

Nested insertions of short mobile sequences in Drosophila P elements

A potentially full-sized P element isolated from the genome of Drosophila ambigua by polymerase chain reaction amplification was completely sequenced. It has a length of 3329 bp and the termini are formed by 33 bp inverted repeats. Sequence comparisons show that it can be classified as a member of the T-type P element subfamily. The translational reading frames of all four exons are interrupted by stop codons and frameshift mutations. At the 3′ end of exon 3 a 687 bp insertion sequence (IS-amb-P) is found that also occurs in the form of dispersed copies (IS-amb) in the genome in D. ambigua. At the interspecific level it shows homology to mobile sequences of other species of the obscura group. Although variable in length, these IS elements are characterized by conserved sections without coding function and by 14 bp inverted repeats, one at a terminal, the other at a subterminal position. In situ hybridization revealed that P elements in D. ambigua are restricted to only two euchromatic sites on chromosome elements A and E. This situation resembles that found in Drosophila guanche and Drosophila subobscura where P homologs are clustered at a single site on chromosome element E and where the section corresponding to exon 3 of P elements carries an IS element. The gene sik-hom, which is located at the 5′ side of the D. guanche cluster of P homologs, was used as a marker to examine whether the P element sites on chromosome element E of D. guanche and D. ambigua are homologous. The results suggest that the nested insertions of IS elements into P elements must have occurred independently in the two different lineages. Received: 13 October 1997; in revised form: 11 December 1997 / Accepted: 12 December 1997     

Pairing and recombination at meiosis of Brassica rapa (AA) × Brassica napus (AACC) hybrids

Interspecific crosses contribute significantly to plant evolution enabling gene exchanges between species. The efficiency of interspecific crosses depends on the similarity between the implicated genomes as high levels of genome similarity are required to ensure appropriate chromosome pairing and genetic recombination. Brassica napus (AACC) is an allopolyploid, resulting from natural hybridization between Brassica rapa (AA) and Brassica oleracea (CC), both being diploid species derived from a common ancestor. To study the relationships between genomes of these Brassica species, we have determined simultaneously the pairing and recombination pattern of A and C chromosomes during meiosis of AAC triploid hybrids, which result from the interspecific cross between natural B. napus and B. rapa. Different AAC triploid hybrids and their progenies have been analysed using cytogenetic, BAC-FISH, and molecular techniques. In 71% of the pollen mother cells, homologous A chromosomes paired regularly, and usually one chromosome of each pair was transmitted to the progeny. C chromosomes remained mainly univalent, but were involved in homoeologous pairing in 21.5% of the cells, and 13% of the transmitted C chromosomes were either recombined or broken. The rate of transmission of C chromosomes depended on the identity of the particular chromosome and on the way the hybrid was crossed, as the male or as the female parent, to B. napus or to B. rapa. Gene transfers in triploid hybrids are favoured between A genomes of B. rapa and B. napus, but also occur between A and C genomes though at lower rates.     

Relationships among pansies (Viola section Melanium) investigated using ITS and ISSR markers

Sequences of the nuclear region ITS and the variable molecular markers ISSR were used to estimate the phylogeny of the section Melanium of the genus Viola. We confirm that the so-called pansies form a derived and monophyletic group. Two floral characters, the upturned side petals and the large size of pollen grains appear to be synapomorphies in Melanium. The Melanium species are very closely related, as shown by the reduced genetic variation compared to the other sections of Viola. Our analyses suggest x=5 or x=7 as the possible base chromosome number of the section Melanium. Polyploidy and hybridization would have played an important role in the evolutionary history of this clade resulting in a wide range of chromosome number. The low genetic differentiation and the complex cytological evolution suggest that diversification in Melanium is the result of a reticulate evolution and rapid radiation in Europe and Northern Africa.     

A three-locus system of interspecific incompatibility underlies male inviability in hybrids between Drosophila buzzatii and D. koepferae

In hybrids between the sibling species D. buzzatii and D. koepferae, both sexes are more or less equally viable in the F₁: However, backcross males to D. buzzatii are frequently inviable, apparently because of interspecific genetic incompatibilities that are cryptic in the F₁. We have performed a genetic dissection of the effects of the X chromosome from D. koepferae. We found only two cytological regions, termed hmi-1 and hmi-2, altogether representing 9% of the whole chromosome, which when introgressed into D. buzzatii cause inviability of hybrid males. Observation of the pattern of asynapsis of polytene chromosomes (incomplete pairing, marking introgressed material) in females and segregation analyses were the technique used to infer the X chromosome regions responsible for this hybrid male inviability. The comparison of these results with those previously obtained with the same technique for hybrid male sterility in this same species pair indicate that in the X chromosome of D. koepferae there are at least seven times more regions that produce hybrid male sterility than hybrid male inviability. We have also found that the inviability brought about by the introgression of hmi-1 is suppressed by the cointrogression of two autosomal sections from D. koepferae. Apparently, these three regions conform to a system of species-specific complementary factors involved in an X-autosome interaction that, when disrupted in backcross hybrids by recombination with the genome of its sibling D. buzzatii, brings about hybrid male inviability.     

GENOME RELATIONSHIPS OF INTRA- AND INTERSECTIONAL SPECIES HYBRIDS OF ONCIDIUM TRIQUETRUM

Chromosome numbers and meiotic behavior of six interspecific hybrids of Oncidium triquetrum were investigated. The somatic chromosome number of O. triquetrum (section Oncidium) x O. variegatum (section Oncidium) was 2n = 42. O. triquetrum x O. splendidum (section Plurituberculata) and O. triquetrum x O. cebolleta (section Cebolletae) were 2n = 39 and O. triquetrum x O. floridanum (section Planilabra), O. triquetrum x O. flexuosum (section Synsepala) and O. triquetrum x O. onustum, (section Onusta) were 2n = 49. The intersectional species hybrid, O. triquetrum x O. variegatum, showed fairly regular meiosis with a mean of 20.4 bivalents, indicating strong homology of parental genomes. In contrast, highly irregular meiosis with poor chromosome pairing was observed in all intersectional hybrids. The mean number of bivalents ranged from 1.8 to 9.5. On the basis of chromosomal affinity at metaphase I, O. triquetrum showed a close relationship to O. variegatum in the same section, but more distant relationships to O. splendidum, O. cebolleta, O. floridanum, O. flexuosum and O. onustum belonging to different sections.     

Molecular and chromosomal organization of two repetitive DNA sequences with intercalary locations in sugar beet and other Beta species

 In a search for repetitive DNA sequences in the sugar beet genome, two sequences with repeat unit lengths of 143 and 434 bp were isolated and characterized. The pSV family showed an unusual conservation of restriction sites reflecting homogenization of the analyzed repeats. Members of the family are organized as tandem repeats as revealed by PCR and sequencing of dimeric units. The pSV satellite occurs in large intercalary arrays which are present on all chromosome arms of sugar beet. The pSV sequence family is present in different abundance in the sections Beta, Corollinae and Nanae but is not detectable by Southern hybridization in the section Procumbentes. The pDRV family is characterized by an interspersed genomic organization. The sequence is detectable in all sections of the genus and is amplified in species of the section Beta but was also detected, although at lower abundance, in the remaining three sections. Fluorescent in situ hybridization has shown that the pDRV sequence family is dispersed over all chromosomes of the sugar beet complement with some regions of clustering and centromeric depletion. Received: 18 March 1998 / Accepted: 31 March 1998     

CHROMOSOME NUMBERS IN CUPHEA (LYTHRACEAE): NEW COUNTS AND A SUMMARY

The American genus Cuphea with ca. 260 species is extremely diverse with respect to chromosome number. Counts are now available for 78 species and/or varieties, or 29% of the genus. Included in this study are first reports for 15 taxa from Brazil, Cuba, Dominican Republic, Mexico, and Venezuela. Twenty-two different numbers are known for the genus, ranging from n = 6 to n = 54. The most common number in the primary center of species diversity in Brazil is n = 8, which is regarded as the base number of the genus. Two numbers are most common in the secondary center in Mexico, n = 10 and n = 12. Species with n = 14 or higher are considered to be of polyploid origin. Polyploids comprise 46% of the total species counted and appear in 9 of the 11 sections for which chromosome numbers have been reported. Aneuploid species comprise ca. 25% of the genus and are known from 7 of the 11 sections. The two subgenera are not characterized by different chromosome numbers or sequences of numbers. None of the 14 sections are circumscribed by a single chromosome number. Morphological and ecological variability in widespread, weedy species is correlated with differing chromosome numbers in some species whereas in others the chromosome number is stable. Summary of chromosome numbers by taxonomic section is presented. Section Euandra, centered in eastern Brazil, and the largest section of the genus, appears to be chromosomally most diverse. In section Trispermum, characterized by difficult, variable species with intermediate forms, two of the four species studied have polyploid races. Section Heterodon, endemic to Mexico and Central America and comprising most of the annual species of the genus, is best known chromosomally. Chromosome numbers have been counted for 25 of 28 species, and 12 different numbers are reported. The most advanced sections, Melvilla and Diploptychia, with numerous species occurring at higher altitudes, are characterized by high polyploids. Apomictic species occur in sect. Diploptycia. The cytoevolution of Cuphea is complex with frequent polyploid and aneuploid events apparently playing a significant role in speciation in both centers of diversity.     

CYTOTAXONOMIC STUDIES IN CHRYSOTHAMNUS (ASTEREAE,COMPOSITAE)

The basic chromosome number for Chrysothamnus is x = 9 as determined by 234 original counts representing all species. Polyploidy is frequent only in C. viscidiflorus where diploids, tetraploids, and hexaploids are found. Pollen size in the genus is basically correlated with style length rather than ploidy level. Most aspects of meiosis appear normal except for the tendency for nucleolar fragmentation. To reduce chromosome clumping during meiosis, an ice-water pretreatment was utilized. Karyotypes in Chrysothamnus reflect similar trends of specialization noted in chromosome structure for other Compositae. The taxa are predominantly self-fertilized; however, one instance of interspecific hybridization is noted. Taxonomic significance of the chromosome numbers, karyotypes, and breeding system in Chrysothamnus is discussed.     

Carex, subgenus Carex (Cyperaceae) – A phylogenetic approach using ITS sequences

To evaluate the sectional classification in Carex, subgenus Carex, the ITS region of 117 species belonging to 32 sections was analyzed with Neighbor Joining (NJ) and Markov chain Monte Carlo (MCMC) methods. In our analyses (1) species of subgenus Indocarex appear as a statistically well supported group within subgenus Carex. (2) The representatives of sections Vesicariae, Hirtae, Pseudocypereae, Ceratocystis, Spirostachyae, Bicolores, Paniceae, Trachychlaenae, Scirpinae, Atratae and Albae group in statistically supported clades with higher support in MCMC than in NJ. (3) C. rariflora clusters with representatives of section Limosae, however only weakly supported. (4) Taxa of section Phacocystis are divided in two statistically supported subclusters that are closely related to a core group of section Hymenochlaenae. (5) Species of sections Montanae, Pachystylae, Digitatae, Phacocystis, Rhomboidales, Careyanae and Frigidae are segregated into two or more clusters each. (6) Five species of section Frigidae cluster together, whereas the seven others are in scattered positions. Based on these results, delimitation of sections is discussed.     

Morphological, molecular and cytological analyses of Carica papaya×C. cauliflora interspecific hybrids

 Morphological, molecular and cytological analyses were performed to assess the hybridity of 120 putative interspecific hybrids of Carica papaya L.×C. cauliflora Jacq. In the putative interspecific hybrids the number of main leaf veins was intermediate between the two parents while the hermaphrodite flower sex form and the low vigour were distinctive features of these hybrids. Petiole length, stem diameter, leaf length, leaf width and flower colour were similar to C. papaya, whereas leaf shape, type, serration, venation, petiole hairiness and flower shape were similar to C. cauliflora. Markers generated by the polymerase chain reaction using 72 10-mer primers (random amplified polymorphic DNA) revealed a high level of polymorphism (64%) between C. papaya and C. cauliflora. Seventeen of these primers yielded reliable and easily scorable polymorphic banding patterns that were further screened to reveal hybrids. A range of 1–5 RAPD primers consistently confirmed that all 120 plants were genetic hybrids, with all of them containing at least one band from the male parent. Cytological analysis revealed that 7–48% of the cells in many of the interspecific hybrids were aneuploid suggesting that chromosome elimination was occurring. The frequency of aneuploid cells was negatively associated (r=0.88) with the number of bands from the male parent integrated into the hybrid. Pollen fertility of the hybrids was from 0.5 to 14.0% while C. papaya and C. cauliflora had 88.0–99.0% and 90.0–97.0% fertile pollen, respectively. Received: 28 February 1996 / Accepted: 27 September 1996     

The genotypic control of homoeologous chromosome association in Lolium temulentum × Lolium perenne interspecific hybrids

Two contrasting genotypes of Lolium perenne and two inbred lines of L. temulentum were examined with regard to their effect on homoeologous chromosome pairing in interspecific hybrids derived from them. Substantial differences in chiasma frequency were observed between the hybrid progeny of the different parental types. The background genes involved were found to operate in the presence and in the absence of B chromosomes. The combination of A chromosome genes present in some of the 0B hybrids was found to result in a considerable suppression of chiasma formation at the diploid level, and the restriction of pairing to strict homologues at the tetraploid level. It appears, therefore, that genes are present within the diploid species of the genus Lolium which are capable of performing a function similar to that of the Ph locus in wheat.     

Genetic variation in resistance to Fusarium oxysporum f.sp. lilii in the genus Lilium

Genetic variation in Fusarium resistance in 82 Lilium genotypes was studied. A high level of resistance was found in the Asiatic hybrids (section Sinomartagon) and to a lesser extent in cultivars of L. longiflorum (section Leucolirion). The Oriental hybrids (section Archelirion) showed only moderate resistance. In the sections Sinomartagon, Leucolirion and Martagon species were detected with high levels of resistance. An accession of L. dauricum (section Sinomartagon) was the most resistant source. Possibilities for introgression of Fusarium resistance into cultivars by interspecific hybridisation, using the resistance found in Lilium genotypes, are discussed.     

Molecular analysis of introgressive breeding in coffee (Coffea arabica L.)

Nineteen arabica coffee introgression lines (BC₁F₄) and two accessions derived from a spontaneous interspecific cross (i.e. Timor Hybrid) between Coffea arabica (2n=4x=44) and C. canephora (2n=2x=22) were analysed for the introgression of C. canephora genetic material. The Timor Hybrid-derived genotypes were evaluated by AFLP, using 42 different primer combinations, and compared to 23 accessions of C. arabica and 8 accessions of C. canephora. A total of 1062 polymorphic fragments were scored among the 52 accessions analysed. One hundred and seventy-eight markers consisting of 109 additional bands (i.e. introgressed markers) and 69 missing bands distinguished the group composed of the Timor Hybrid-derived genotypes from the accessions of C. arabica. AFLP therefore seemed to be an extremely efficient technique for DNA marker generation in coffee as well as for the detection of introgression in C. arabica. The genetic diversity observed in the Timor Hybrid-derived genotypes appeared to be approximately double that in C. arabica. Although representing only a small proportion of the genetic diversity available in C. canephora, the Timor Hybrid obviously constitutes a considerable source of genetic diversity for arabica breeding. Analysis of genetic relationships among the Timor Hybrid-derived genotypes suggested that introgression was not restricted to chromosome substitution but also involved chromosome recombinations. Furthermore, the Timor Hybrid-derived genotypes varied considerably in the number of AFLP markers attributable to introgression. In this way, the introgressed markers identified in the analysed arabica coffee introgressed genotypes were estimated to represent from 9% to 29% of the C. canephora genome. Nevertheless, the amount of alien genetic material in the introgression arabica lines remains substantial and should justify the development of adapted breeding strategies. Received: 2 February 1999 / Accepted: 12 May 1999     

Spatial discrimination of pheromones and behavioural antagonists by the tortricid moths Cydia pomonella and Adoxophyes orana

Male moths responding to their species-specific sex pheromone, may cease their upwind flight when pheromone components of sympatric species are added to the mixture. The interspecific interaction between the pheromone response of the tortricid moths Cydia pomonella and Adoxophyes orana was investigated in field-trapping and wind-tunnel studies. Addition of the A. orana pheromone [(Z9)-tetradecenylacetate and (Z11)-tetradecenylacetate] to a source containing the C. pomonella pheromone [(E8, E10)-dodecadienol] resulted in a significant inhibition of attraction by male C. pomonella. It is demonstrated that this behavioural antagonist for C. pomonella must be emitted from the same point source to induce this inhibitory effect. A spatial separation of the two interspecific pheromones (at 14 cm, 5 cm and 0.5 cm crosswind) restored the attraction of the conspecific pheromone for male C. pomonella. In contrast to C. pomonella, male A. orana were not inhibited by point sources releasing both the C. pomonella and A. orana pheromone. We suggest that the discrepancy in the interspecific pheromone interaction between these two tortricids can be explained if we consider the evolutionary ecology of interspecific pheromone communication in C. pomonella and A. orana. Accepted: 18 July 1999