B-Chromosomes inAllium flavum (Liliaceae) and some related species

B-Chromosomes ofAllium flavum, A. stamineum andA. carinatum were C-banded. InA. flavum different types of B's were found, one of them possessing a nucleolus organizer. In the B's ofA. flavum andA. stamineum the banding patterns resemble those found in the standard chromosomes. The B inA. carinatum is only terminally banded. InA. flavum chiasma localization in the B's appears to be dependent on C-band location, just as is the case in the A-chromosomes. An increase in B's was found to cause a small (but significant) increase in chiasma frequency in the PMC's. This may result from an alteration in the nucleotype by the B's.     

A survey of C-band patterns in chromosomes ofLilium (Liliaceae)

C-band patterns are described for 20Lilium spp. distributed across six sections. All species have a similar basic karyotype (n = 12) but C-bands differ markedly between them. The patterns are characterized by a dispersed scattering of thin intercalary bands as well as centric and NOR bands. Only one species,L. canadense, shows a clear equilocal pattern with intercalary C-bands occurring proximally in all of the longer chromosome arms. Comparing species, similar patterns are revealed forL. regale andL. sulphureum, forL. formosanum andL. longiflorum (all in sect.Leucolirion) and to a lesser extent forL. hansonii, L. martagon, andL. tsingtauense (sect.Martagon). The pattern forL. henryi (previously classed in sect.Sinomartagon) matches those ofL. regale andL. sulphureum quite well and its transfer to sect.Leucolirion is proposed. This is consistent with results from interspecies hybrids betweenL. henryi andL. regale (and related species) which are reportedly fertile. No other clear similarities in C-band patterns were seen across species. It seems that C-band patterns change rapidly inLilium and hence their usefulness in classification will be restricted to identifying closely related species.Dedicated to Prof.D. G. Catcheside on the 80th anniversary of his birth.     

A new species of Allium (Liliaceae)

Allium stoloniferum, a new species from southern Mexico, belongs to theA. kunthii alliance on the basis of the appearance of the perianth segments and the presence of long slender rhizomes, but differs by the presence of bulbils.     

Distance-dependent performance of asexual progeny in Allium vineale (Liliaceae)

Local adaptation within and among populations may have an impact on processes ranging from speciation to the evolution of mixed breeding systems and dispersal strategies. It is also one potential factor that could favor the production of asexual over sexual propagules. This field experiment tested whether asexually produced bulbils of Allium vineale demonstrate local adaptation to the parental microsite at the scale of natural dispersal from the parent (5, 25, 50, 100, and 1000 cm). Both "home' and randomly chosen "away' genotypes were planted at each location to determine the relative performance of the "home' genotype. Overall, bulbil performance declined with distance from the parent. In particular, "home' bulbils outperformed "away' bulbils at a distance of 25 cm from the parent, indicating that local adaptation has occurred at the scale of natural dispersal in this species. The variance in propagule performance also increased at farther distances from the parent, indicating that the predictability of offspring performance decreases with distance. Fine-scale local adaptation within the range of seed dispersal in this population may be one factor favoring asexual reproduction in Allium vineale.     

Phylogeny of Chinese Allium (Liliaceae) using PCR-RFLP analysis

Eighteen representative species were selected from all the nine sections of Chinese Allium on the basis of the classification of morphology and cytotaxonomy. The trnK and rpL16 gene fragments of chloroplast DNA were amplified from 18 species by PCR method. The two cpDNA fragments were digested by 26 restriction enzymes, and 303 polymorphic restriction sites were found, of which 163 were informative. The restriction site data were analyzed with PAUP (version 3.1.1) and MEGA (version 1.01) as well as PHYLIP. As a result, the genus Allium could be classified into six subgenera. The recognition of Sect. Anguinum in the Flora of China is reasonable, Sect. Rhizirideum, Sect. Haplostemon and Sect. Cepa are not monophyletic. The infrageneric system of this genus was also discussed.     

Taxonomy and distribution of Allium ursinum (Liliaceae) in Poland and adjacent countries

The paper presents distribution data on Allium ursinum L. in Poland, with additional records from the other European countries. The study comprised 367 herbarium specimens, drawn from most of Polish herbaria and individuals obtained during field investigations. Two subspecies have been recorded: A. ursinum L. subsp. ursinum and A. ursinum subsp. ucrainicum Kleop. et Oxner. The study also documents the presence of intermediate populations at the border of the distribution ranges for both subspecies. In addition, the paper presents the pedicel structure and pollen grains ultrasculpture of each subspecies under a scanning electron microscope (SEM). Finally, questions in respect of the distribution data and habitat preferences are discussed.     

Prometaphase I and anaphase I in an interchange heterozygote of Allium triquetrum (Liliaceae)

Four unstable malorientations in a chain quadrivalent, as well as maloriented bivalents, were studied in fixed prometaphase I pollen-mothercells from an interchange heterozygote of Allium triquetrum. The relative frequencies of these malorientations, in cells from pollen sacs of different developmental ages, suggest that maloriented bivalents, on the average, reorient before maloriented quadrivalents; and that, similarly, there are differences in the timing of reorientation amongst the four types of maloriented quadrivalents. — Further, the proportion of anaphase I cells derived from alternate orientation in the interchange quadrivalent is lower than expected in pollen sacs with a high percentage of cells in mid-anaphase; but higher than expected in pollen sacs in which relatively few cells have as yet proceeded into anaphase, and also in those in which most cells have already passed through anaphase. Arguably, these data are a direct outcome of the differential behavior of unstable maloriented quadrivalents in the preceeding prometaphase.     

Unstable B-chromosomes in a European population of Allium schoenoprasutn L. (Liliaceae)

In Allium schoenoprasum , small, dichromatic B-chromosomes occur in 74% of plants at Lac de Tracens, Pare National des Pyrénées. B-chromosomes from Lac de Tracens were compared with the previously well characterized Bs from British populations of A. schoenoprasum at the River Wye, Powys, and were found to differ in terms of both their morphology and the level of mitotic instability. At Lac de Tracens, two morphological types of B-chromosome were observed: a telocentric of about 1.15 μm in length and a metacentric of about 1.35 μm in length; the most frequent B-type in the River Wye populations (B^-1) was not found. B-chromosomes from the River Wye are essentially stable at root tip mitosis, whereas 25 out of 29 plants from Lac de Tracens (86%) showed between-cell variation in B-number, with non-modal numbers in 330 out of 794 cells (41.6%). Most of the variability occurred within individual roots, rather than between roots of single bulbs or of sister bulbs. No meiotic pairing was observed between the different Bs in hybrids from the geographically isolated populations, suggesting a lack of homology. An independent origin of B-chromosomes in the British and French populations is suggested.     

The Biology of Allium monanthum (Liliaceae) I. Polyploid Complex and Variations in Karyotype

Abstract Plants of Allium monanthum Maxim., whose gender expression are usually dioecious, but rarely hermaphrodite or gynomonoecious, proved to constitute a polyploid complex, consisting of diploid, triploid, and tetraploid individuals. The basic chromosome complement of this species consists of seven metacentric or submetacentric chromosomes and one acrocentric, the latter possessing a satellite on the short arm. Thus, the karyotype formula is expressed as 7V+11 (x=8). The diploid plants (2n = 16) were confined to central Honshu, Japan. Typical female plants possessed the standard karyotype, whereas male plants were heterozygous for two kinds of translocations. The 3x plants (2n=24) are somewhat widely distributed in the areas from the Kanto to Hokuriku district in Honshu. All female triploid plants possessed the standard karyotype. The geographical distribution of 4x plants (2n=32) which express mostly a female phenotype occurred nearly throughout the whole areas investigated; they are geographically isolated from the 2x plants. A majority of 4x plants had the standard karyotype. The remaining tetraploids were of the aberrant type, 4x/51, which has five acrocentric chromosomes, and two aneuploids 4x+1 and 4x-1. Both 3x and 4x forms seem to be of autopolyploid origin. Three kinds of aberrant nucleolar chromosomes with an extra satellite or an inseried secondary constriction were found in the heterozygotes for translocations of 2x plants and also in some plants of the 3x form. These aberrant plants usually form their own homogeneous populations, but were somewhat scattered throughout the range in their distribution. Thus, these individuals are considered to have perpetuated these types of chromosome aberrations which originated in the remote past.     

Microsporogenesis and development of the male gametophyte in Allium senescens L. (Liliaceae) in China

Microsporogenesis and development of the male gametophyte of Allium senescens L. were studied by use of paraffin sections. The anther was found to be four-sporangia. Its wall includes four layers: epidermis, endothecium, middle layer, and secretory tapetum. Cytokinesis during meiosis of microspore mother cells (MMCs) is successive. Most mature pollen grains are two-celled, a few are three-celled. In addition, anomalies were observed during meiosis of MMC and in uninucleate microspores, including laggard chromosomes, lagging chromosomes, anaphase bridges, and micronucleus. These anomalies are possible reasons for abortive pollen.     

Localized chiasmata and meiotic nodules in the tetraploid onion Allium porrum.

Allium porrum L. (cultivated leek) (2n = 4x = 32) is a fertile tetraploid that forms bivalents with pericentric chiasmata at metaphase I. To investigate the basis of this unusual behavior for a tetraploid, we describe the karyotype, axial cores, synaptonemal complexes (SCs), and meiotic nodules of A. porrum. The karyotype appears to be autotetraploid. This conclusion is also supported by presynaptic alignment of axial cores in groups of four and partner trades between pairs of SCs. Numerous early nodules are distributed all along axial cores and SCs during zygonema, but they are lost by late zygonema - early pachynema. Late (recombination) nodules (RNs) are present on SCs near kinetochores throughout the remainder of pachynema. This pattern of RNs corresponds to the pattern of pericentric chiasmata. Pachytene quadrivalents usually are resolved into bivalents because partner trades between SC lateral elements rarely occur between RNs on the same segment of SC. Thus, the patterns of crossing-over and partner trades promote balanced disjunction and high fertility in autotetraploid A. porrum. Rare quadrivalents observed at metaphase I must be due to infrequent partner trades between RNs. Polycomplexes, unusual in their number and size, were observed during zygonema. Key words : synaptonemal complex, recombination nodules, localized chiasmata, polycomplex, Allium porrum.     

Allium runemarkii (Liliaceae), a new species from the island of Ewia (W Aegean, Greece)

Allium runemarkii , from the island of Ewia (W Aegean area, Greece), is illustrated and described as a species new to science. This species belongs to Allium sect. Scorodon and is related to the Greek species A. thessalicum and A. erythraeum in the A. obtusiflorum group. It is diploid with 2n=2x=16. Details of its karyotype are given.     

Cytogenetical evidences for hybrid structure and origin of diploid and triploid shallots (Allium cepa var.viviparum,Liliaceae) from Dalmatia (Croatia)

Two viviparous strains of common onion (Allium cepa) denoted asAllium cepa var.viviparum (syn.Allium ×proliferum), traditionally cultivated in the seaside regions of Croatia were found to be diploid (2n = 2x = 16) and triploid (2n = 3x = 24), respectively. The triploid cultivated onion Ljutika has not been previously reported in Europe. Using Feulgen and Giemsa C-banding methods both karyotypes were shown to be of hybrid constitution. Results obtained in this work indicate that Dalmatian viviparous onions arose from spontaneous hybridization ofA. cepa withA. fistulosum.     

A comparative study of chiasmata in male and femaleBettongia penicillata (Marsupialia)

The chiasma frequency and distribution does not differ substantially between the sexes in the marsupialBettongia penicillata. This result contrasts with the observations made on other species of marsupials.     

Agrobacterium tumefaciens-mediated transformation of leek (Allium porrum) and garlic (Allium sativum)

Transgenic leek (Allium porrum) and garlic (Allium sativum) plants have been recovered by the selective culturing of immature leek and garlic embryos via Agrobacterium-mediated transformation using a method similar to that described by Eady et al. (Plant Cell Rep 19:376–381, 2000) for onion transformation. This method involved the use of a binary vector containing the m-gfp-ER reporter gene and nptII selectable marker, and followed the protocol developed previously for the transformation of onions with only minor modifications pertaining to the post-transformation selection procedure which was simplified to have just a single selection regime. Transgenic cultures were selected for their ability to express the m-gfp-ER reporter gene and grown in the presence of geneticin (20 mg/l). The presence of transgenes in the genome of the plants was confirmed using TAIL-PCR and Southern analysis. This is the first report of leek and true seed garlic transformation. It now makes possible the integration of useful agronomic and quality traits into these crops.     

Variation and taxonomy ofFritillaria graeca (Liliaceae) in Greece

The geographic variation inFritillaria graeca, an endemic of Greece, is explored, using multivariate techniques. Eleven morphological features were measured in each of 630 individual plants from 38 localities throughout the range of the species.Scheffe's a posteriori test revealed an almost continuous geographic variation and did not identify groups clearly. Discriminant analysis of the groups derived from an F significance test ofMahalanobis distance resulted in five major groups on the first two discriminant axes representing the subspeciesgraeca, ionica, guicciardii, thessala and its var.othria. A diagnostic key to theF. graeca subspecies is provided.This paper is a part of a doctoral dissertation dealing withFritillaria in Greece, to be submitted to the department of Biology of the Aristotelian University of Thessaloniki.     

Hybridization drives speciation in Gagea (Liliaceae)

Hybridization seems to play an important role in speciation of Gagea Salisb., a genus which is characterised by polyploid taxa lines and in which diploids (2n = 24) appear only to be common in basal sections. Hybrid detection was applied utilising direct and cloning nrDNA ITS data (ITS1, 5.8S rDNA, ITS2) combined with neighbour and ribotype networks and discussed in connection with previously published cpDNA, morphological and karyological data of the authors. We have evidence of the hybrid origin of taxa within the section Gagea (G. pomeranica, G. megapolitana) and the monophyletic clade of sections Didymobulbos and Fistulosae (G. microfistulosa, G. polidorii, G. cf. bohemica). Morphologically and karyologically differentiated Gagea megapolitana and G. pomeranica, adapted to synanthropic habitats, represent both hybrids of G. pratensis × G. lutea. Gagea microfistulosa represents a hybrid of G. villosa × G. fragifera; Gagea polidorii could represent the reverse hybrid. G. glacialis is also closely related to the latter complex.     

Seed dispersal in Erythronium grandiflorum (Liliaceae)

Primary and secondary seed dispersal was investigated for the glacier lily Erythronium grandiflorum in the Colorado Rocky Mountains. These heavy seeds have no obvious adaptations for biotic or abiotic dispersal, but can be thrown short distances when the dehiscent fruits are shaken by wind. We used sticky traps to measure primary transport of seeds up to 1 m away from individual plants. A seed cafeteria experiment examined the role of ants and rodents in secondary seed transport. Primary dispersal by wind was positively skewed and median transport distances were influenced by variation in plant height. Secondary dispersal was negligible compared to Viola nuttallii, an elaiosome-bearing species. Thus, seed dispersal was highly restricted in E. grandiflorum, and a 1 m radius encompassed the modal section of the seed dispersal curve. The seed dispersal component of gene flow was quantified and combined with previous measurements of pollen flow to yield a more complete estimate of Wright's neighborhood size, N _e, for E. grandiflorum. The lack of a special seed dispersal mechanism in E. grandiflorum is discussed in terms of a source-sink model for seedling establishment with respect to distance from the parental plants.     

Amounts of ribosomal DNA inAllium (Liliaceae)

Eight species ofAllium subgen.Allium sect.Allium have been studied at the cytological level by means of karyological analyses and at the biochemical level with regard to the proportions of ribosomal DNA. All the species have a basic genome of x = 8.A. sativum, A. commutatum, A. ampeloprasum, andA. vineale possess approximately 0.050% rDNA and two nucleolus organizer regions per basic chromosome set.A. sphaerocephalon andA. arvense have two nucleolus organizers, andA. amethystinum three nucleolus organizers per haploid (n = x) genome: the three species possess approximately 0.075% rDNA.A. acutiflorum has five nucleolus organizer regions per haploid genome and 0.121% rDNA. An attempt to relate these differences with functional and ecological characteristics indicates that evolutionary variation of rDNA proportions is not casual. Such data also can help to define systematic affinities and circumscribe infrageneric taxa.     

A Comparison of Leek (Allium porrum) and Onion (Allium cepa) Seed Development

Leek and onion seed dry weight increased exponentially for thefirst 31 days after flowering (DAF) but thereafter the increasein dry weight was slower. Before maximum seed dry weight wasreached at 45 DAF in onion and 59 to 66 DAF in leek, seed moisturecontent, seed oxygen uptake and conductivity of the seed steepwater fell from initially high levels. Although some seeds germinated31 DAF in both species, full germination in both was not reacheduntil 66–80 DAF. Tolerance of the seed to artificial dryingimmediately after harvesting occurred 45 DAF in onion and 74DAF in leek. Free nuclear division continued in the endospermuntil 17–22 DAF in onion and until 31–35 DAF inleek but it was not until 45 DAF in onion and 66 DAF in leekthat the embryo and endosperm filled the cavity formed by thepericarp. After formation of cell walls in the endosperm thepattern of change in cell number in both species was similar.The shrunken appearance of the seed coat in leek, which occurredearly in seed development, was associated with the period offree nuclear division in the endosperm and, in addition, thepericarp was thinner than in onion. There was no evidence thatthe shrunken seed coat early in development was associated withself as opposed to open-pollination. Allium porrum, Allium cepa, seed development, endosperm, embryo, cell number, germination, respiration, seed leachates