Karyotypic studies in Solanum section Lasiocarpa (Solanaceae)

Mitotic chromosomes of 13 species included in Solanum sect. Lasiocarpa were studied. All species have 2n = 24. The chromosome numbers of S. stagnale, S. felinum, and S. repandum are reported for the first time. Statistical analyses of chromosomes, genome length, and centromere position yielded estimates of karyotype composition and asymmetry. A generalized karyotype of the section shows that most of the chromosomes are metacentric (73%) or submetacentric (25.6%). Two pairs of subtelocentric chromosomes are present only in S. sessiliflorum. Satellites are quite common, characterizing chromosomes for ten species and are attached to the short arms of either m or sm chromosomes. They could be seen in 10% to 70% of the cells. Only S. pseudolulo bears two satellited chromosomes, one of them attached to a long chromosome arm. Although the section is chromosomally quite homogeneous, species can be distinguished by karyotype formula, presence of satellites in a particular chromosome pair, and total chromosome length. There are no indications of major chromosomal rearrangements within the section. Using cytological data exclusively, cluster analysis indicates S. sessiliflorum is isolated in the section. Solanum candidum and S. vestissimum are somewhat isolated as well and have unique karyological traits. Solanum pectinatum has a peculiar karyotype, but in the phenogram it is not particularly separated. Karyotype features suggest that morphological differentiation was not always followed by chromosomal divergence. The origin of the domesticated S. quitoense is no further elucidated by our data.     

In vitro organogenesis and plant regeneration from leaves of Solanum candidum Lindl,S. quitoense Lam. (naranjilla) and S. sessiliflorum Dunal

Adventitious shoots and roots were regenerated from leaf segments of 3 Solanum species: S. candidum Lindl., S. quitoense Lam. and S. sessiliflorum Dunal. Leaf explants differentiated shoots on modified MS medium supplemented with 23–163 M kinetin and 0–5.7 µM indoleacetic acid (IAA). Excised shoots were induced to form roots by transfer to media with benzyladenine (BA) and naphthaleneacetic acid (NAA) at 0.09 and 0.11 µM respectively for S. quitoense and 0.01 µM NAA for S. candidum and S. sessiliflorum. Adventitious roots were produced directly from leaf explants with 0–140 µM kinetin and 0–5.7 µM IAA in combination. Rooted plants were successfully established in the greenhouse.     

Diversity and evolution of rice progenitors in Australia

In the thousands of years of rice domestication in Asia, many useful genes have been lost from the gene pool. Wild rice is a key source of diversity for domesticated rice. Genome sequencing has suggested that the wild rice populations in northern Australia may include novel taxa, within the AA genome group of close (interfertile) wild relatives of domesticated rice that have evolved independently due to geographic separation and been isolated from the loss of diversity associated with gene flow from the large populations of domesticated rice in Asia. Australian wild rice was collected from 27 sites from Townsville to the northern tip of Cape York. Whole chloroplast genome sequences and 4,555 nuclear gene sequences (more than 8 Mbp) were used to explore genetic relationships between these populations and other wild and domesticated rices. Analysis of the chloroplast and nuclear data showed very clear evidence of distinctness from other AA genome Oryza species with significant divergence between Australian populations. Phylogenetic analysis suggested the Australian populations represent the earliest‐branching AA genome lineages and may be critical resources for global rice food security. Nuclear genome analysis demonstrated that the diverse O. meridionalis populations were sister to all other AA genome taxa while the Australian O. rufipogon‐like populations were associated with the clade that included domesticated rice. Populations of apparent hybrids between the taxa were also identified suggesting ongoing dynamic evolution of wild rice in Australia. These introgressions model events similar to those likely to have been involved in the domestication of rice.     

THE RELATIONSHIPS BETWEEN MALE AND FEMALE FERTILITY AND AMONG TAXA IN DIPLOID WHEATS

Pollen stainability appears to be a reliable indication of the ultimate seed set in diploid interspecific hybrid and backcross populations in Triticum L. The correlation between percent pollen stained and number of seeds set is positive and highly significant (r = 0.92). Estimates of male and female fertility in the hybrids and backcrosses are interpreted to indicate that the domesticated diploid Triticum monococcum L. and wild diploid T. boeoticum Boiss. em. Schiem, are one and the same species, and that T. urartu Tum. is not a variety of monococcum or boeoticum, but rather a separate species. The F₁ hybrids and backcrosses between monococcum and boeoticum are normally male and female fertile. The F₁ hybrids between monococcum and urartu are completely sterile and complete to partial sterility exists in backcrosses.     

Extensive Hybridization and Introgression between Melastoma candidum and M. sanguineum

Natural hybridization can lead to various evolutionary outcomes in plants, including hybrid speciation and interspecific gene transfer. It can also cause taxonomic problems, especially in plant genera containing multiple species. In this study, the hybrid status of Melastoma affine, the most widespread taxon in this genus, and introgression between its putative parental species, M. candidum and M. sanguineum, were assessed on two sites, Hainan and Guangdong, using 13 SSR markers and sequences of a chloroplast intergenic spacer. Bayesian-based STRUCTURE analysis detected two most likely distinct clusters for the three taxa, and 76.0% and 73.9% of the morphologically identified individuals of M. candidum and M. sanguineum were correctly assigned, respectively. 74.5% of the M. affine individuals had a membership coefficient to either parental species between 0.1 and 0.9, suggesting admixture between M. candidum and M. sanguineum. Furthermore, NewHybrids analysis suggested that most individuals of M. affine were F2 hybrids or backcross hybrids to M. candidum, and that there was extensive introgression between M. candidum and M. sanguineum. These SSR data thus provides convincing evidence for hybrid origin of M. affine and extensive introgression between M. candidum and M. sanguineum. Chloroplast DNA results were consistent with this conclusion. Much higher hybrid frequency on the more disturbed Guangdong site suggests that human disturbance might offer suitable habitats for the survival of hybrids, a hypothesis that is in need of further testing.     

Application of interspecific sesquiploidy to introgression of PLRV resistance from non-tuber-bearing Solanum etuberosum to cultivated potato germplasm

Summary Hybridization of synthetic allotetraploids of S. pinnatisectum with S. etuberosum (4x-EP) with S. acaule (2n = 4x = 48) resulted in two individuals that were highly fertile, in contrast to all other progenies. The unique individuals are hexaploids, 2n = 72, while the other progenies are tetraploids, 2n = 48. They are thought to be the products of a union between 2n eggs of S. acaule and normal 1n microspores of 4x-EP. The fertile hexaploids (designated 6x-AEP) produced abundant selfed seed and viable hybrids with cultivated diploid potato, S. phureja, when developing embryos were rescued from berries and cultured before transplanting to pot culture. The extreme variability in chromosome constitution of the hybrids with S. phureja and selfed progenies indicates that addition and substitution lines of etb chromosomes bearing genes of interest to breeders could easily be produced from this material. The production of sesquiploids, as the 6x-AEP hybrids are called, is discussed as a useful bridging step in the introduction of alien genes from genomes that share little homology with the cultivated genome.This work was part of the requirement for a Ph.D. degree of the senior author R. Chavez at The University of Birmingham, UK     

Interspecific hybridization inRibes

The genusRibes, 2n=16, consisting of over 150 shrubby species has been divided byRehder into 15 sections grouped into four subgenera. Data on interspecies crosses within and between these sections and subgenera made by previous workers and at East Malling are tabulated and discussed.Species within a section are morphologically similar and will usually cross, forming fertile hybrids; 33 such hybrids in 6 of the 15 sections are known. Inter-sectional and inter-subgeneric crosses between more unlike species often fail, but matur flowering plants have been obtained from crosses involving the following combinations: –Symphocalyx x Calobotrya, Symphocalyx x Eucoreosma, Calobotrya x Eucoreosma, Eucoreosma x Heritiera, Eucoreosma x Ribesia, Eugrossularia x Robsonia, Euberisia x Eugrossularia, Symphocalyx x Eugrossularia, Calobotrya x Eugrossularia, Eucoreosma x Eugrossularia, andRibesia x Eugrossularia. Of these, only the combinationsEugrossularia x Robsonia andCalobotrya x Eucoreosma show any fertility.On the basis of these results, it is suggested that a more natural classification ofRibes would be to retainRehder's subgeneraBerisia, Grossularia andGrossularioides, while creating from his sevenRibesia sections, six new subgenera, viz.Microsperma, Symphocalyx, Calobotrya (comprising the sectionsCalobotrya andEucoreosma), Cerophyllum, Heritiera, andRibesia. With this classification, in the light of present knowledge, all intra-sectional hybrids are fertile, intra-subgeneric hybrids at least partially fertile, and such inter-subgeneric hybrids as can be raised are sterile.     

CROSSING RELATIONSHIPS AMONG DIPLOID SPECIES OF THE SOLANUM NIGRUM COMPLEX IN NORTH AMERICA

Crosses were made in all combinations of the six diploid species of the Solarium nigrum complex that occur in North America. Some interspecific pollinations failed to yield viable seed; successful crosses gave rise to moderately to highly sterile F₁ hybrids. Results of interspecific crosses suggested phylogenetic relationships that were not completely in accord with those suggested by morphology. Interspecific crosses also gave varied results. All interpopulational crosses within S. interius and S. sarachoides produced fully fertile hybrids. In contrast, hybrids within S. americanum and S. douglasii varied from fully fertile to almost completely sterile. Populations of S. pseudogracile could be divided into two groups which are geographically separated but not morphologically differentiated. Fully fertile hybrids resulted from crosses within a group, whereas crosses between groups gave hybrids with reduced fertility. Four crossing groups were observed within 5. nodiflorum; three of the parental groups are sympatric and are morphologically differentiated. Although hybrid sterility in interspecific crosses is sometimes used to support delimitation of species, the presence of sterility in intraspecific crosses suggests that such an interpretation is unwarranted for the S. nigrum complex. Hybrid sterility, therefore, is not considered to have special taxonomic significance in this complex.     

Distinct protoconchs recognised in three of the larger Mediterranean Cerithium species (Caenogastropoda: Cerithiidae)

The gastropod genus Cerithium includes several polymorphic species which are hardly distinguishable using a morphological approach based on teleoconch characters. Here we show that protoconch characters can be reliably used to identify the larger Mediterranean species (Cerithium alucastrum, C. repandum and C. vulgatum), and to assess their intraspecific variability. Based on a large amount of morphological data, we show that a multispiral, strongly sculptured protoconch (traditionally associated with C. vulgatum) is found in C. alucastrum. This species originated in the Pliocene. A multispiral, weakly sculptured protoconch, not observed previously, is reported for C. vulgatum. A paucispiral protoconch, distinct from those observed previously for C. lividulum and C. renovatum, is reported for C. repandum and can be used for recognising its wide intraspecific variation and understanding its geographical distribution. Cerithium repandum, previously doubtfully recorded outside its type locality, extends back to the mid-Pliocene and is today rather widely distributed in the Mediterranean, although absent from the Aegean Sea and the Alboran Sea. We describe the larval development of C. repandum and C. vulgatum for the first time, confirming their respective non-planktotrophic and planktotrophic development previously inferred from protoconch morphology. Also, we report a possible case of morphological convergence between C. repandum and C. vulgatum occurring in harbour sites, where we found a distinct (possibly Anthropocene) shell form not described previously from other Recent or fossil material.     

Domestication via hybridization of the wild tetraploid oats Avena magna and A. murphyi

The wild tetraploid (2n=28) oat species Avena magna and A. murphyi have been domesticated by having been transferred from the common oat, A sativa (2n=42), the characteristics of non-shedding spikelets glabrous and yellow lemma, and reduced awn formation. Domestication has been achieved by crossing the common oat with either of the tetraploid species and then backcrossing the pentaploid hybrids with pollen of the tetraploid wild parent. Among the BC plants obtained only a few produced some seeds. Fertile tetraploids exhibiting the domesticated syndrome have been selected for in the F₂ generation. Although morphologically they were almost indistinguishable from the common oat, they were tetraploids. Wild x domesticated A. magna hybrids were vigorous and fertile. They retained their spikelets at maturity, lemma color and pubescence were intermediate between the parental lines, and awns were formed only on the lower floret of the spikelet. Each of these characteristics segregated in a 31 fashion, indicating single gene control, as in the common oat. These four characteristics formed a linkage group in one F₂ family and two linkage groups in the other two families. The usefulness of the domesticated tetraploids for oat research and production has been discussed. Taxonomically, the domesticated tetraploids were ranked as subspecies: A. magna ssp. domestica, and A. murphyi ssp. rigida.     

Marker-assisted breeding of a photoperiod-sensitive male sterile <Emphasis Type="Italic">japonica</Emphasis> rice with high cross-compatibility with <Emphasis Type="Italic">indica</Emphasis> rice

The incomplete fertility of japonica × indica rice hybrids has inhibited breeders’ access to the substantial heterotic potential of these hybrids. As hybrid sterility is caused by an allelic interaction at a small number of loci, it is possible to overcome it by simple introgression at the major sterility loci. Here we report the use of marker-assisted backcrossing to transfer into the elite japonica cv. Zhendao88 a photoperiod-sensitive male sterility gene from cv. Lunhui422S (indica) and the yellow leaf gene from line Yellow249 (indica). The microsatellite markers RM276, RM455, RM141 and RM185 were used to tag the fertility genes S5, S8, S7 and S9, respectively. Line 509S is a true-breeding photoperiod-sensitive male sterile plant, which morphologically closely resembles the japonica type. Genotypic analysis showed that the genome of line 509S comprises about 92% japonica DNA. Nevertheless, hybrids between line 509S and japonica varieties suffer from a level of hybrid sterility, although the line is highly cross-compatible with indica types, with the resulting hybrids expressing a significant degree of heterosis. Together, these results suggest that segment substitution on fertility loci based on known information and marker-assisted selection are an effective approach for utilizing the heterosis of rice inter-subspecies.     

REPRODUCTIVE ISOLATION BARRIERS TO GENE EXCHANGE BETWEEN SOLANUM CHACOENSE AND S. COMMERSONII (SOLANACEAE)

In contrast to the general intercompatibility of diploid species of tuber-bearing potatoes of South America, Solanum chacoense Bitter and S. commersonii Dunal have well-established internal barriers to gene exchange. Pollen tubes of S. commersonii are inhibited in growth down the style of S. chacoense. While S. chacoense pollen tubes are not inhibited in S. commersonii stylar tissue, seed development is abnormal, resulting in production of large numbers of seeds that are not filled out. Even the apparently normal F₁ seed germinated poorly; approximately 0.1 germinable seeds were set per cross as compared with over 30 seeds per cross following intraspecific crosses. Lowland S. chacoense, including populations sympatric with S. commersonii, were not more effectively isolated from S. commersonii than were highland S. chacoense populations, over 500 km from the nearest S. commersonii. The F₁ hybrids produced were vigorous and highly fertile. Crosses between unrelated S. commersonii-chacoense F₁ hybrids produced only 1.3 germinable seeds per cross. This reduction resulted from low numbers of berries set, few seeds per berry, and low seed germination. The F₂ progeny were vigorous, but while many were both male and female fertile there were indications of an increase in sterility over that of the parents and F₁'s. The data were interpreted as indicating that the isolation barriers result from polygenic differences between the parent species and are composed principally of genes that function in several of the processes of seed setting. The accumulation of genes that keep these species isolated was interpreted as not having developed as a result of selection for such isolation in the region of overlap (Wallace effect), but rather as a byproduct of divergence that occurred during their allopatric evolution.     

HYBRIDIZATION AND CYTOTAXONOMY OF DICENTRA

Biosystematic relationships among species assigned to three subgenera of the genus Dicentra were investigated with respect to hybridization and chromosomal constitution and fertility of the hybrids. Four species of subgenus Dicentra, D. formosa, D. eximia, D. nevadensis, and D. peregrina, were intercrossed in various ways to form diploid, triploid, and tetraploid hybrids. Hybrids at the tetraploid level in this subgenus invariably were highly fertile. Triploid hybrids, as expected, were mostly very sterile. Diploid hybrids varied in this respect, but none was highly fertile. Crosses with two of the remaining four species of subgenus Dicentra produced no hybrids, but abundant seed was obtained in one instance. The two species of the subgenus Chrysocapnos, D. chrysaniha and D. ochroleuca, cross to produce a partially fertile tetraploid hybrid, but cross-pollinations involving these species with those of other subgenera failed. The single species of subgenus Hedycapnos, D. spectablis (diploid) produced no hybrids when cross-pollinated with members of the other two sections. These results are fully concordant with presumed affinities based on morphological similarity In addition, preliminary results of hybridization between the monocotyledonous D. peregrina and a number of dicotyledonous species of Dicentra are reported.     

Downy mildew incidence of pearl millet hybrids with different male-sterility inducing cytoplasms

The use of different sources of cytoplasmic male sterility (CMS) in hybrid seed production of pearl millet [Pennisetum glaucum (L.) R. Br.] is advocated to avoid possible disease epidemics occurring due to cytoplasmic uniformity. The effects of commercially unexploited, but potentially exploitable, sources of CMS, like A₂, A₃ and A₄, on downy mildew [Sclerospora graminicola (Sacc.) Schroet] incidence were studied by using the disease incidence of isonuclear hybrids with male-sterile and fertile cytoplasm. The mean downy mildew incidence of hybrids carrying different male-sterile cytoplasm was similar to that of hybrids retaining the fertile cytoplasm. The cytoplasm accounted for only 0.6% of the total variation and its effect was non-significant; pollinators could explain most of the variation in determining the disease incidence of hybrids. This suggested that these male-sterile cytoplasms are not linked to downy mildew susceptibility and thus can be exploited commercially to broaden the cytoplasmic base of the male-sterile lines and, ultimately, of hybrids.     

Sex at the origin: an Asian population of the rice blast fungus Magnaporthe oryzae reproduces sexually

Sexual reproduction may be cryptic or facultative in fungi and therefore difficult to detect. Magnaporthe oryzae, which causes blast, the most damaging fungal disease of rice, is thought to originate from southeast Asia. It reproduces asexually in all rice‐growing regions. Sexual reproduction has been suspected in limited areas of southeast Asia, but has never been demonstrated in contemporary populations. We characterized several M. oryzae populations worldwide both biologically and genetically, to identify candidate populations for sexual reproduction. The sexual cycle of M. oryzae requires two strains of opposite mating types, at least one of which is female‐fertile, to come into contact. In one Chinese population, the two mating types were found to be present at similar frequencies and almost all strains were female‐fertile. Compatible strains from this population completed the sexual cycle in vitro and produced viable progenies. Genotypic richness and linkage disequilibrium data also supported the existence of sexual reproduction in this population. We resampled this population the following year, and the data obtained confirmed the presence of all the biological and genetic characteristics of sexual reproduction. In particular, a considerable genetic reshuffling of alleles was observed between the 2 years. Computer simulations confirmed that the observed genetic characteristics were unlikely to have arisen in the absence of recombination. We therefore concluded that a contemporary population of M. oryzae, pathogenic on rice, reproduces sexually in natura in southeast Asia. Our findings provide evidence for the loss of sexual reproduction by a fungal plant pathogen outside its centre of origin.     

Geographic isolation and evolution of Mediterranean endemic Cyclamen: insights from chloroplast trnL (UAA) intron sequence variation

 In this study we construct a phylogenetic hypothesis for the relatedness among disjunct subspecies of Cyclamen repandum and its two allopatric congeners, C. creticum and C. balearicum in order to examine the evolutionary divergence of currently isolated populations across the western Mediterranean. The most parsimonious phylogenetic tree obtained from sequencing the cpDNA trnL (UAA) intron suggests a major phylogeographic divide in southern Greece between two clades. The first clade comprises samples of C. repandum subsp. peloponnesiacum (from the Peloponnese) and C. creticum (from Crete). The second comprises samples of C. repandum subsp. repandum (from Croatia, Italy, southern France, Corsica, Sardinia and Sicily), C. repandum subsp. rhodense (from Rhodes and Kos) and C. balearicum (from the Balearic Islands and southern France). These data suggest that C. creticum has evolved in allopatry from C. repandum subsp. peloponnesiacum and that C. balearicum and C. repandum ssp. rhodense have diverged from C. repandum subsp. repandum at its western and eastern distribution limits. At one small site on Corsica, a population of C. repandum may have introgressed with relictual populations of C. balearicum. These divergence patterns illustrate how a phylogenetic perspective can be used to better understand the evolution of endemism in the Mediterranean flora. Received February 19, 2001 Accepted August 22, 2001     

Infraspecific variation and systematics of cultivatedSetaria italica,foxtail millet (Poaceae)

Foxtail millet (Setaria italica,) is grown as a cereal in southern Europe and in temperate, subtropical, and tropical Asia. Its closest wild relative isS. italica ssp.viridis (green foxtail). Green foxtail is native to temperate Eurasia, but was introduced and became widely established as a weed in temperate and warmer parts of the Americas. Spontaneous and cultivatedS. italica cross to produce fertile hybrids. Derivatives of such crosses, resembling foxtail millet in some inflorescence traits but with efficient natural seed dispersal, accompany the cereal across its range of cultivation. Giant green foxtail of Europe and the American corn belt is a weed of hybrid origin. Foxtail millet was domesticated in the highlands of central China; remains of cultivated foxtail millet are known from the Yang-shao culture period dating back some 5,000 yrs. Comparative morphology suggests that foxtail millet spread to Europe and India as a cereal soon after its domestication. Three cultivated races are recognized. Moharia, from Europe and southwestern Asia, includes cultivars with 5–52 culms, each bearing several, small, more or less erect inflorescences. Cultivars in race maxima are characterized by plants with 1–8 usually unbranched culms that bear large inflorescences; they occur in Transcaucasian Russia and the Far East. Race indica is intermediate in culm number (ave. 6.6) and inflorescence size between races moharia and maxima, and is cultivated in southern Asia.     

Hybridization of Senecio squalidus and S. viscosus and Introgression of Genes from Diploid into Tetraploid Senecio Species

Natural triploid hybrids (Senecio x londinensis Lousley) betweenS. squalidus L. (2n = 20) and 5. viscosus L. (2n = 40) are fairlyfrequently found in Britain. Under glasshouse conditions bothnatural and artificial hybrids displayed very low levels ofseed fertility and gave rise to morphologically diverse F₂ plantsat about the triploid or pentaploid chromosome levels. By theF₄ generation, progeny of a F₂ pentaploid plant had somaticchromosome numbers near to the tetraploid level and considerablyincreased pollen and seed fertilities. Such fertile tetraploidsegregants of S. x londinensis permit the introgression of S.squalidus genes into S. viscosus, and may indicate the courseof introgression into other tetraploid species of Senecio. Senecio, hybridization, introgression     

Homoeologous relationships of Aegilops speltoides chromosomes to bread wheat

 Homoeologous pairing at metaphase I was analysed in the standard-type, ph2b and ph1b hybrids of Triticum aestivum (AABBDD) and Aegilops speltoides (SS). Data from relative pairing affinities were used to predict homoeologous relationships of Ae. speltoides chromosomes to wheat. Chromosomes of both species, and their arms, were identified by C-banding. The Ae. speltoides genotype carried genes that induced a high level of homoeologous pairing in the three types of hybrids analyzed. All arms of the seven chromosomes of the S genome showed normal homoeologous pairing, which implies that no apparent chromosome rearrangements occurred in the evolution of Ae. speltoides relative to wheat. A pattern of preferential pairing of two types, A-D and B-S, confirmed that the S genome is very closely related to the B genome of wheat. Although this pairing pattern was also reported in hybrids of wheat with Ae. longissima and Ae. sharonensis, a different behaviour was found in group 5 chromosomes. In the hybrids of Ae. speltoides, chromosome 5B-5S pairing was much more frequent than 5D-5S, while these chromosome associations reached similar frequencies in the hybrids of Ae. longissima and Ae. sharonensis. These results are in agreement with the hypothesis that the B genome of wheat is derived from Ae. speltoides. Received: 8 January 1998 / Accepted: 4 February 1998