A systematic study of the Linum tenuifolium group (Linaceae)

NICHOLLS, M. S., 1985. A systematic study of the Linum tenuifolium group (Linaceae) . A new classification of the Linum tenuifolium group is presented based upon detailed systematic study of 74 populations in nature. Previous confusion over taxonomy and nomenclature is resolved. Linum salsoloides, L. appressum and L. suffruticosum are treated as subspecies of L. tenuifolium . The morphological variation within each subspecies is discussed with reference to its distribution.     

A new subspecies of Linum gyaricum (Linaceae) from Greece

Christodoulakis, D. 1995. A new subspecies of Linum gyaricum (Linaceae) from Greece. - Nord. J. Bot. 15: 145–147. Copenhagen. ISSN 0107–055X.
A new subspecies, Linum gyaricum ssp. icaricum from Ikaria island (E Aegean, Greece), is described and illustrated.     

Infra-specific genetic and morphological diversity in Linum album (Linaceae)

The genus Linum L. (Lineacea) has over 15 species, subspecies or ecotypes in Iran. These species show extensive geographical distribution and form many local populations throughout the country. Linum album is herbaceous medicinal plant containing important lignans such as podophyllotoxin (PTOX) and 6-methoxy podophyllotoxin (MPTOX), which have antiviral and anticancer properties. Studying the genetic and morphological diversity of different geographical populations produces detailed knowledge about population divergence and identification of the infra-species taxa if at all they are present. Moreover, the populations that differ in their genetic content and structure may also differ in their chemical and medicinal properties. The present study considers morphological and genetic diversity analyses of 20 L. album geographical populations by using nuclear ISSR markers, genome size, and cytogenetic characteristics. These populations differed significantly in many of their quantitative morphological characters and in some of their qualitative features. They also differed significantly in their molecular characteristics and genome size. Details of morphological and molecular variations are reported and discussed.     

A new species of Linum (Linaceae) from west Anatolia, Turkey

A new species, Linum vuralianum Yılmaz & Kaynak, is described and illustrated from west Anatolia, Turkey. It is closest to L. flavum L., but differs by its annual or biennial life form, the presence of rosette leaves at anthesis, and the absence of staminodes. The World Conservation Union (IUCN) threat category and observations on the population are noted. The geographical distribution of the new species and its related taxon is given.  © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society , 2008, 156 , 459–462.     

Intraspecific 5S rRNA gene variation in flax,Linum usitatissimum (Linaceae)

A relatively large proportion of the flax genome (~ 3%) is comprised of 5S rRNA genes. This study focuses on the intraspecific sequence variation among five distinct groups of 5S rRNA genes. The results indicate that group 1 and 2 5S rRNA genes most closely resemble other angiosperm 5S genes, while groups 3–5 are highly divergent. Sequence variation is higher in the spacer region compared to the transcribed region for all pairwise comparisons. The large degree of sequence variation observed in this study is discussed with respect to genome organization and proposed models for repetitive sequence maintenance.     

Life history variation in blue flax (Linum perenne: Linaceae): seed germination phenology

Linum perenne L. is a nonclonal perennial herb widely distributed across a range of habitats. Variation in seed germination patterns was examined for 21 Intermountain collections and for the commercial cultivar ‘Appar’. Collections from sites with long, snowy winters were largely dormant at harvest and responded positively to chilling. Collections from middle elevation sagebrush-grassland sites were generally nondormant and contained a fraction induced into secondary dormancy by chilling. Collections from palouse prairie and pinyon-juniper sites were generally nondormant and unaffected by chilling, as was ‘Appar’. When seeds of contrasting populations were planted in seed retrieval experiments at low, middle, and high elevation sites, their field germination phenology was predictable from laboratory experiments. In common garden experiments, there were significant among-sibship differences in germination for each garden-grown wild accession but not for ‘Appar’, suggesting that differences both among and within populations may be genetically based. Garden-grown seeds were generally less dormant than wild-collected seeds, possibly because of selection during propagation. Results demonstrated the existence of ecologically relevant among-population and within-population variation in germination phenology for blue flax. Seeds of high-montane populations are spring-emerging and have little provision for between-year carryover. Populations from lower elevation habitats with less predictable weather have seeds with contrasting germination patterns, allowing for emergence in both fall and spring as well as for seed bank carryover. ‘Appar’ seeds lack these predictive and carryover mechanisms.     

Metabolism of Hydroxy Fatty Acids in Developing Seeds in the Genera Lesquerella (Brassicaceae) and Linum (Linaceae)

Members of the genus Lesquerella produce seed oil that contains a high proportion of hydroxy fatty acids (HFAs). There are three groups of Lesquerella species that are distinguished by their most abundant seed oil fatty acid: lesquerolic acid (20:1OH; e.g. Lesquerella fendleri), densipolic acid (18:2OH; e.g. Lesquerella kathryn), and auricolic acid (20:2OH; e.g. Lesquerella auriculata). To investigate the biochemistry of HFA production in Lesquerella species, the conversion of putative radiolabeled intermediates of HFA biosynthesis, including 18:1, 20:1,18:1OH, 18:2OH, and 20:1OH, was examined in developing embryos of L. fendleri, L.kathryn, and L. auriculata. The results are consistent with (a) 18:1OH formation by hydroxylation of 18:1, (b) elongation and desaturation of 18:1OH to produce 20:1OH and 18:2OH, respectively, and (c) desaturation of 20:1OH to produce 20:2OH. The desaturation of 20:1OH was also found to occur in developing embryos of high, but not low, linolenic acid flax. This suggests that the desaturation is catalyzed by the extraplastidial linoleate desaturase. Confirming this suggestion, it was notable that 18:1OH and 18:2OH were found in low and high linolenic flax (Linum usitatissimum) seeds, respectively, at levels of 0.2 to 1%.     

Pollen flow, population composition, and the adaptive significance of distyly in Linum tenuifolium L. (Linaceae)

The function of distyly as a device promoting inter-morph pollination was proposed by Darwin over a century ago, though only recently have experimental studies been undertaken to test this hypothesis. These studies are unrealistic since they assume regular spatial distribution of morphs within a population and fail to consider intra-flower pollination in assessing distyly's effects. Pollen flow in two populations of Linum tenuifolium L. was investigated using a fluorescent-dye marker technique. One population was dimorphic and self-incompatible; the other was monomorphic and self-compatible. In both populations pollen dispersal was leptokurtic with over 75% of grains detected within 5 m from the source. Intra-flower pollinations comprised a large proportion of individual stigma loads (42% for the dimorphic form; 49% for the monomorphic form). Total stigmatic loads for the dimorphic population showed equal proportions of pin and thrum pollen on both long-style and short-style stigmas, though variation in individual loads was marked. Values ranging from 34.2 to 61.5% legitimate pollen were recorded. The irregular distribution of morphs observed at the dimorphic site will reduce the efficiency of heterostyly as a device promoting intermorph pollination. The reduction in intra-flower pollination suggests adaptive significance of this floral dimorphism in increasing the efficiency of pollen dispersal to the individual.     

山地五月茶的蝇类传粉研究

观测了山地五月茶Antidesma montanum的开花物候、开花动态、访花者种类和访花行为,并对其繁育系统、花粉组织化学、花粉胚珠比（P／O）、花粉活力进行了检测。结果表明：山地五月茶是雌雄异株植物;其总状花序花期可长达7天;雄花／雌花花序数目为140．33±27．79／208．33±33．65（n=6）,雌雄花的颜色为很淡的黄绿色小型花;雄花单花花期为2天,花药开裂当天有活力,花粉为非淀粉型,花粉胚珠比为3333．33±607．18;单花花蜜量可达0．34±0．03μL,花蜜含糖量3．69％±0．30％。花的结构和开花式样适合蝇类传粉。芳香的气味是吸引蝇类的直接物质：花蜜是传粉者的报酬。主要的传粉昆虫为双翅目Diptera丽蝇科Calliphoridae的Chrysomya megacephala、Chrysomyasp．、寄蝇科Tachinidae的Drino sp．和蝇科Muscidae的Spilogona sp．、Mitroplatia sp．。套网不授粉的处理不结实,表明山地五月茶不存在无融合生殖,人工辅助授粉的坐果率（39．1％）略高于自然坐果率（36．7％）,二者无显著差异,表明其坐果率主要受自身资源分配限制。还讨论了蝇类传粉与雌雄异株性系统以及蝇类传粉与热带林中小型黄绿色花植物的相关性。     

Fly pollination and duodichogamy in Bridelia stipularis and Cleistanthus sumatranus (Phyllanthaceae)

The globally distributed family Phyllanthaceae comprises 1745 species, all of which produce unisexual flowers, and most of the species occur in the tropics. The few species in which the pollination system has been studied are pollinated by flies, wasps, or specialized moths, and at least two species are duodichogamous. Here we report on the flowering phenology and pollination of the Chinese species Bridelia stipularis and Cleistanthus sumatranus, which were studied on the tropical Hainan Island. Both species are duodichogamous and indicated to be fly‐pollinated. The calliphorid species Chrysomya megacephala, Isomyia isomyia, Pierretia sp., Hemipyrellia sp., and Achoetandrus tufifacies were the most frequent flower visitors to B. stipularis; some of these species together with syrphids (Meliscaeva cinctella, Dideopsis sp., and Eristalis sp.) were the most abundant visitors to Cl. sumatranus flowers. Fruit set in hand‐pollinated flowers was higher than in open‐pollinated flowers, but because of natural fruit abortion the difference was not significant, indicating that fly pollinators do not limit reproduction of these Phyllanthaceae species. This study provides an addition to the limited Asian literature on pollination by flies and the even more limited literature on duodichogamy. It could help to broaden our understanding for analysis of the evolution of duodichogamy in future research, but the specific ecological factors behind the evolution of this rare sexual system are still difficult to generalize.     

Effects of cadmium and mycorrhizal fungi on growth, fitness, and cadmium accumulation in flax (Linum usitatissimum; Linaceae)

? Premise of the study: Agricultural soils have become contaminated with a variety of heavy metals, including cadmium. The degree to which soil contaminants affect plants may depend on symbiotic relationships between plant roots and soil microorganisms. We examined (1) whether mycorrhizal fungi counteract the potentially negative effects of cadmium on the growth and fitness of flax (Linum usitatissimum) and (2) whether mycorrhizal fungi affect the accumulation of cadmium within plant parts. ? Methods: Two flax cultivars (Linott and Omega) were grown in three soil cadmium environments (0, 5, and 15 ppm). Within each cadmium environment, plants were grown in either the presence or absence of mycorrhizal fungi. Upon senescence, we measured growth and fitness and quantified the concentration of cadmium within plants. ? Key results: Soil cadmium significantly decreased plant fitness, but did not affect plant growth. Mycorrhizal fungi, which were able to colonize roots of plants growing in all cadmium levels, significantly increased plant growth and fitness. Although mycorrhizal fungi counteracted the negative effects of cadmium on fruit and seed production, they also enhanced the concentration of cadmium within roots, fruits, and seeds. ? Conclusions: The degree to which soil cadmium affects plant fitness and the accumulation of cadmium within plants depended on the ability of plants to form symbiotic relationships with mycorrhizal fungi. The use of mycorrhizal fungi in contaminated agricultural soils may offset the negative effects of metals on the quantity of seeds produced, but exacerbate the accumulation of these metals in our food supply.     

Variation and evolution inLinum tenuifolium (Linaceae)

The phenetic variation inL. tenuifolium s.l. was assessed using multivariate analyses of 27 characters found to be variable within the species. These results are discussed and interpreted in the context of the reproductive biology, chromosome number and ecological responses of the taxa. Evidence suggests that both the self-compatible and tetraploid races were derived independently from a self-incompatible, diploid type. Although no taxonomic scheme is compatible with all of the evidence presented, the study supports a recent decision to recognize four taxa at subspecific level.     

Fly pollination in Ceropegia (Apocynaceae: Asclepiadoideae): biogeographic and phylogenetic perspectives

Background and Aims

Ceropegia (Apocynaceae subfamily Asclepiadoideae) is a large, Old World genus of >180 species, all of which possess distinctive flask-shaped flowers that temporarily trap pollinators. The taxonomic diversity of pollinators, biogeographic and phylogenetic patterns of pollinator exploitation, and the level of specificity of interactions were assessed in order to begin to understand the role of pollinators in promoting diversification within the genus.

Methods

Flower visitor and pollinator data for approx. 60 Ceropegia taxa were analysed with reference to the main centres of diversity of the genus and to a cpDNA–nrDNA molecular phylogeny of the genus.

Key Results

Ceropegia spp. interact with flower-visiting Diptera from at least 26 genera in 20 families, of which 11 genera and 11 families are pollinators. Size range of flies was 0·5–4·0 mm and approx. 94 % were females. Ceropegia from particular regions do not use specific fly genera or families, though Arabian Peninsula species are pollinated by a wider range of Diptera families than those in other regions. The basal-most clade interacts with the highest diversity of Diptera families and genera, largely due to one hyper-generalist taxon, C. aristolochioides subsp. deflersiana. Species in the more-derived clades interact with a smaller diversity of Diptera. Approximately 60 % of taxa are so far recorded as interacting with only a single genus of pollinators, the remaining 40 % being less conservative in their interactions. Ceropegia spp. can therefore be ecological specialists or generalists.

Conclusions

The genus Ceropegia has largely radiated without evolutionary shifts in pollinator functional specialization, maintaining its interactions with small Diptera. Intriguing biogeographic and phylogenetic patterns may reflect processes of regional dispersal, diversification and subsequent specialization onto a narrower range of pollinators, though some of the findings may be caused by inconsistent sampling. Comparisons are made with other plant genera in the Aristolochiaceae and Araceae that have evolved flask-shaped flowers that trap female flies seeking oviposition sites.Key words: Apocynaceae, Asclepiadoideae, Brachystelma, Ceropegia, Diptera, flower evolution, generalization, mutualism, pollination, Riocreuxia, specialization, Stapeliinae     

In vitro polyploidy induction: changes in morphology,podophyllotoxin biosynthesis,and expression of the related genes in Linum album (Linaceae)

Planta - Induction of tetraploidy was performed and podophyllotoxin production increased by upregulating the expression level and enzyme activity of genes related to its biosynthesis in tetraploid...     

The Diversity of Karyotypes and Genomes within Section Syllinum of the Genus Linum (Linaceae) Revealed by Molecular Cytogenetic Markers and RAPD Analysis

The wide variation in chromosome number found in species of the genus Linum (2n = 16, 18, 20, 26, 28, 30, 32, 36, 42, 72, 84) indicates that chromosomal mutations have played an important role in the speciation of this taxon. To contribute to a better understanding of the genetic diversity and species relationships in this genus, comparative studies of karyotypes and genomes of species within section Syllinum Griseb. (2n = 26, 28) were carried out. Elongated with 9-aminoacridine chromosomes of 10 species of section Syllinum were investigated by C- and DAPI/С-banding, CMA and Ag-NOR-staining, FISH with probes of rDNA and of telomere repeats. RAPD analysis was also performed. All the chromosome pairs in karyotypes of the studied species were identified. Chromosome DAPI/C-banding patterns of 28-chromosomal species were highly similar. Two of the species differed from the others in chromosomal location of rDNA sites. B chromosomes were revealed in all the 28-chromosomal species. Chromosomes of Linum nodiflorum L. (2n = 26) and the 28-chromosomal species were similar in DAPI/C-banding pattern and localization of several rDNA sites, but they differed in chromosomal size and number. The karyotype of L. nodiflorum was characterized by an intercalary site of telomere repeat, one additional 26S rDNA site and also by the absence of B chromosomes. Structural similarities between different chromosome pairs in karyotypes of the studied species were found indicating their tetraploid origin. RAPD analysis did not distinguish the species except L. nodiflorum. The species of section Syllinum probably originated from a common tetraploid ancestor. The 28-chromosomal species were closely related, but L. nodiflorum diverged significantly from the rest of the species probably due to chromosomal rearrangements occurring during evolution.     

The systematics ofLinum sect.Linopsis (Linaceae)

SectionLinopsis of the genusLinum is the largest and probably the most widespread of the five commonly recognized sections. Using a number of traits, but especially heterostyly, style union, development of false septa in the fruit, pollen morphology, and chromosome number, together with petal pigmentation, number of floral parts, and stigma shape, the section is divided into five subsections:Dichrolinum, with one series;Halolinum with one;Linopsis, with four;Keniense, with one; andRigida, with two. Of these, subsect.Keniense and two series are new, the remaining four subsections and four series are published in a new rank. The inferred relationships of all of the groups and the general distribution of each are discussed. The section ranges from southern Europe and the Mediterranean region to India, eastern and southern Africa and throughout much of North and South America. The primitive species of the section are believed to be in the Mediterranean region.     

Distyly and pollen dimorphism inLinum suffruticosum (Linaceae)

The Mediterranean distylousLinum suffruticosum has dimorphic pollen. That from short-styled plants has sub-monomorphic surface excrescences with minute papillae; that from long-styled plants has strongly dimorphic excrescences with conspicuous papillae. The differences are much like those described in species of other sections of the genus. Pollen of the related homostylousL. tenuifolium has monomorphic pollen. It combines characteristics of bothL. suffruticosum types but resembles that of the long-styled plants somewhat more closely. Ribbonlike styles of long-styled plants ofL. suffruticosum provide another feature which seems to be controlled by the S supergene thought to be associated with distylic incompatibility. The pollen ofL. suffruticosum andL. tenuifolium is very similar to that of some species of sectionLinastrum and would support their placement in that section.