Sex ratio variation and spatial segregation of the sexes in populations ofRumex acetosa andR. acetosella (Polygonaceae)

Populations ofRumex acetosa andR. acetosella were studied during two growth seasons. The ramet sex ratios ofR. acetosa were always female-biased. InR. acetosella the sex ratios expressed more variation but were mostly female-biased. In both species the sex ratios commonly varied between subpopulations reflecting a partial spatial segregation of the sexes. No marked differences between sexes in vegetative vigour were detected in either species. Interactions between sex ratios, various soil characteristics and population densities were determined. Possible mechanisms for causing biased sex ratios and partial spatial segregation of the sexes are discussed.     

In vitro organogenesis in two dioecious species,Rumex acetosella L. and R. acetosa L. (Polygonaceae)

Callus cultures were established from dioecious plant species Rumex acetosella and R. acetosa, using cotyledons, hypocotyls and stem tips of aseptically germinated seedlings as primary explants. Cultures were also established from male and female R. acetosella adult plants, starting from vegetative lateral buds. Cell division was induced using a high 2,4-D concentration, while bud induction and multiplication were stimulated on a medium with high BAP/IAA ratio. Cotyledon fragments of both species produced only rhizogenic calli. Hypocotyl-derived calli of R. acetosella produced buds, while those of R. acetosa showed no bud forming response under these conditions. Bud multiplication occurred in stem tip cultures of both species and in lateral bud cultures of R. acetosella. Calli derived from male plants produced more buds than those from female. Shoots were easily rooted using IBA, and plantlets were effectively transferred to soil. Flowering was not induced in culture. The sex of regenerated male and female plants was not altered by the culture conditions.     

The effects of seed weight on growth and competitive ability of Rumex acetosella from two successional old-fields

Summary The effects of seed size on growth, biomass allocation and competitive ability of Rumex acetosella plants grown either individually or in competition were studied in two populations (6 months and 15 years old respectively) sampled from a postcultivation successional gradient. For plants grown individually there were highly significant effects of seed weight on growth after 43 days, with a higher relative growth rate (RGR) observed for plants raised from heavier seeds. However at the end of the experiment, seedlings developed from lighter seeds had a RGR 2 times greater than those from heavier seeds. Final biomass of the two types was not significantly different after 73 days of growth. When plants were grown individually, there were only slight differences between populations, but when grown in monocultures of 4 plants per pot, plants from the old population had higher root and leaf biomass per pot whereas those from the young population had a higher reproductive effort per pot. This suggests that a trade-off between allocation to sexual and vegetative reproduction occurs over successional time. In mixtures of light and heavy seeds, plants from light seeds were shorter, had fewer leaves and lower biomass than plants from heavy seeds, which were also taller and produced more dry matter than plants grown from heavy seeds in monoculture. The significant effects of seed weight and population on biomass parameters persisted unit the end of the experiment. Seedlings from heavy seeds were strong competitors: those from the young population grew better in the presence of neighbors than in monoculture and those from the late successional population suppressed the more the growth of their partners. Seedlings from light seeds were subordinate competitors. These results suggest that seedlings from seeds of different sizes benefit from contrasting ecological conditions and that selection acts on reproductive output along successional gradients.     

Variation and covariation among life-history traits in Rumex acetosella from a successional old-field gradient

In this study morphological variation and the potential for competition to affect biomass and seedling selection of the families of five populations of Rumex acetosella L. sampled along a successional old-field gradient have been investigated. Seeds from 25 families were submitted to four competitive regimes: no competition (one plant per pot), medium competition (two plants/ pot taking plants from the same population), high within-population competition (four individuals from the same population in a pot) and high between-population competition (four individuals from two different populations in a pot). Eight traits were analysed after 3 months of growth for variation among families within populations. A significant difference among families within the two older populations was recorded for sexual biomass and related components. High sensitivity of these traits to density was observed in all populations except the youngest, suggesting specialization to particular environmental conditions in late successional populations, and a good adaptive capacity to buffer environmental variation in the pioneer population. Little significant interaction between competitive regimes and families within populations was found, i.e. genotypes within each population showed little variation in their response to environmental variation. Genotypic variance decreased with increasing competitive conditions for the majority of the traits. However, the percentage of variance in sexual reproduction explained by family was stable among treatments. Tradeoffs between vegetative reproduction and sexual reproduction were recorded at the population level along the successional gradient, with increasing competitive conditions. As succession proceeds, we observed a decrease in sexual reproduction and an increase in vegetative reproduction. At the family level, correlation among traits were similar when plants were grown in the absence of competition and at high density, with a significant negative correlation between sexual reproduction and vegetative reproduction. For both sprout number and sexual biomass, the performance of families grown under all the treatments was positively correlated. Together these results indicate allocational constraints on the reproductive biology of R. acetosella that may be favoured by natural selection and have influenced population differentiation along the successional gradient. However, they also revealed that the potential exists for evolutionary specialization through plasticity, in response to variation in environmental conditions.     

Sexual dimorphism in clonal growth forms and ramet distribution patterns in <Emphasis Type="Italic">Rumex acetosella</Emphasis> (Polygonaceae)

We investigated clonal traits in the dioecious herb Rumex acetosella to characterize sexual dimorphism in clonal forms and to correlate below-ground clonal patterns and above-ground ramet distributions. We recorded creeping root length, branching patterns, ramet and clump (caespitose ramets from the same position on the root) sprouting patterns, and biomass allocations in three females and males. We also estimated the patch size of flowering ramets within a quadrat. No sexual dimorphism was detected in the frequencies of branches and flowering ramets per root length. Male plants allocated proportionally more biomass to below-ground organs. Total root length did not differ between the sexes. Females sprouted more clumps with fewer flowering ramets per root length than males, which sprouted fewer clumps with more flowering ramets, which meant that clump sprouting patterns were phalanx-like in females and guerrilla-like in males. Flowering ramets were aggregately distributed in both females and males and patch sizes were similar between sexes, indicating that the spreader propagations were not found in the guerrilla-like males. We assumed that sexual dimorphism occurred in response to physiological integration for higher reproductive effort in females.     

Rumex subgen.Rumex sect.Axillares (Polygonaceae) in South America

Rumex sect.Axillares first described by the author in 1937 is known to be represented in N. America by 20 endemic species. From S. America only five species, all endemic, were previously known; this number is raised in the present revision to 15. Outside America there exist small groups of endemic species in S. Africa and in the islands of Hawaii as well as in Australia and scattered single species in C. Africa, on the remote Atlantic islands of Tristan de Cunha and Gough Island, and in eastern Asia.—In Europe the section is completely absent, except for three American species which occur as rare aliens. TheAxillares are not only morphologically well-characterized by their growth—no leaf rosettes, flowering period practically unlimited by formation of later flowering axillary shoots, flowers often monoecious—but also by their genetic structure expressed by a different behaviour in hybridization. Hybrids are frequent between species of sect.Rumex (syn.Simplices) and are easy to recognize by their sterility which results in a certain peculiarity in habit. In contrast, no spontaneous hybrids are known with certainty among species of sect.Axillares. Most artificial hybrids are fairly fertile. So far only three hybrids between species of the two sections have been known. Three more have been found by the author in Chile, all being highly sterile and showing the growth characteristics resulting from sterility as occur in hybrids between species of sect.Rumex.

---

Resumen Rumex secciónAxillares fue descrito por primera vez por el autor en 1937. Se conoce en América del Norte con 20 especies endémicas. De América del Sur se conocían sólo 5 especies, todas ellas endemicas; este número asciende a 15 en la presente revision. Fuera de las Américas se encuentran pequeños grupos de especies endémicas en Sudáfrica, en las Islas de Hawaii y en Australia; especies aisladas se encuentran en Africa Central, en las lejanas Islas Atlánticas de Tristan de Cunha y Gough Island y en Asia Oriental. La sección no tiene representantes en Europa, con excepción de tres especies americanas introducidas, poco frecuentes. La secciónAxillares se caracteriza bien, tanto morfológicamente por su forma de creciemiento—sin rosetas foliares, período floral prácticamente ilimitado por la formación de inflorescencias axilares parciales, flores a menudo monoicas—como por su estructura genética que se manifiesta en un comportamiento diferente de hibridación. Híbridos son frecuentes en las especies de la secciónRumex (syn.Simplices) y reconocibles por su esterilidad que se expresa en cierta particularidad del habito. Al contrario, no hay seguridad de híbridos espontáneos en la secciónAxillares. La mayor parte de los híbridos artificiales son totalmente fértiles. Hasta ahora se conocían solo tres híbridos entre las especies de ambas secciones. Otros tres han sido encontrados por el autor en Chile. Todos completamente estériles y con las características de hábito típicas resultantes la esterilidad, como ocurre en los híbridos de la secciónRumex.

     

Flavonoid chemistry ofPolygonum sect.Tovara (Polygonaceae): A systematic survey

Polygonum sect.Tovara includes three controversial species;P. virginianum, P. filiforme, andP. neofiliforme. The flavonoid chemistry of these was examined to provide additional information on their delimitation and levels of differentiation. Eight flavonoid compounds were isolated and identified, all of which were 3-O-glycosides of the flavonols kaempferol, quercetin, and myricetin, and their acylated derivatives. Although they exhibit relatively simple flavonoid profiles, the three taxa are readily distinguished by their flavonoid constituents. In addition, they show fundamental differences in flavonol types and glycosylation patterns. These results, in conjunction with evidence from the morphology, strongly suggest thatP. virginianum, P. filiforme, andP. neofiliforme are closely allied but distinct species.     

Isoenzyme diversity in Reynoutria (Polygonaceae) taxa: escape from sterility by hybridization

The genus Reynoutria is represented by four taxa in the Czech Republic – R. japonica var. japonica and compacta, R. sachalinensis and R. × bohemica. Using isoenzyme analysis, we determined the degree of genotype variability in all taxa and compared clones of R. japonica var. japonica from the Czech Republic with those from Great Britain. While the rarely occurring tetraploid variety R. japonica var. compacta possesses low variability, the octoploid female clone of R. japonica var. japonica is genetically uniform in the 93 clones sampled and belongs to the same genotype that is present in the whole Europe. R. japonica var. japonica can be fertilized by the pollen of tetraploid R. sachalinensis and a hexaploid hybrid R. × bohemica is produced. In R. sachalinensis, 16 genotypes were found in the 50 clones sampled. R. × bohemica is genetically the most diverse taxon in the study area, with 33 genotypes recorded among 88 clones sampled.     

Foliar anatomy ofPolygonum (Polygonaceae): Survey of epidermal and selected internal structures

Several anatomical characters in leaves were described, and their distribution determined, for 153 species ofPolygonum, mostly from herbarium specimens. Structures surveyed were epidermal (glandular and nonglandular trichomes, nodules, specialized parenchyma, stomatal apparatus) and internal (cavities, crystals, laticifer-like cells, nodules, subepidermal fibers). Cleared leaves were supplemented by resin-embedded sections and SEM preparations of selected species. No feature defines any taxonomic section, but some features occur only within one section. Laticifer-like cells, epidermal and internal nodules, resin cups, and unique epidermal and subepidermal cavities seem to be unknown elsewhere; other features (invaginated epidermal cells; enlarged crystal cells confined to paraveinal layer) are rare among angiosperms.     

Chromosome number,morphology, pairing,and DNA values of species and hybrids in the genusFallopia (Polygonaceae)

The taxa in this study have in the past been treated together in sect.Tiniaria Meissner ofPolygonum L. s.l., and more recently the large erect rhizomatous herbaceous perennials have been separated from the twining annuals and perennials as the generaReynoutria andFallopia respectively. These taxa range in ploidy level from diploid to octoploid, with base numbers of both 10 and 11 present. The plants are primarily Asiatic in distribution, although many of the large erect perennials are now naturalized in many parts of Europe. Cytological examination has revealed the presence of a number of previously unknown hybrid taxa in the British Isles. The readiness with which hybridisation occurs between taxa of differing base number and ploidy level, and similarities revealed in chromosome morphology, meiotic pairing and in DNA C-value, suggest to the authors that these two genera are best amalgamated underFallopia.     

Gene variation and gene flow inOrchis morio (Orchidaceae) from Italy

Data are presented on genetic variation at 27 enzyme loci of the Green-Winged orchid,Orchis morio, in 18 population samples from Italy. The existence in Italy of two subspecies, i.e. subspp.morio andpicta, is not supported by allozyme data. No genetic heterogeneity was found betweenmorio-like andpicta-like samples and specimens. Moreover, morphological transition between the two forms was observed in different Italian populations. The parameters of genetic variability estimated forO. morio populations are consistent with those found among monocotyledon plants, and among those outcrossing, animal-pollinated and with wind-dispersed seeds. Genetic diversity of ItalianO. morio is mostly within populations. Correspondingly, low values of interpopulational genetic distance were found. This appears to be due to high levels of gene flow, which were estimated with different methods. The lack ofO. longicornu from Italian samples, as well as of any hybrid withO. morio (F₁, backcrossed or recombinant individuals) is demonstrated on the basis of genetic data. It is concluded that recurrent reports ofO. longicornu from Italy are due to confusion withO. morio or with otherOrchis species.     

Taxonomical investigations onHelleborus viridis s. l. (Ranunculaceae) in Northern Italy

The taxon commonly namedHelleborus viridis in Lombardy (NW. Italy) differs from both, subsp.viridis and subsp.occidentalis, and is shown to fall within the variation range ofH. odorus subsp.laxus which has been reported so far for NE. Italy only. TrueH. viridis within Italy grows only in the Maritime Alps.     

Stress-induced male sterility and mixed mating in the island plantCedronella canariensis (Lamiaceae)

Reaction norms were determined in two natural populations of the perennial plantIris pumila from contrasting light conditions at two levels of photosynthetically active radiation (105 and 35 µmol m^–2s^–1). Seedlings developed from free-pollinated seed families sampled individually from 19 and 12 randomly selected clonal genotypes in an exposed and a shaded population respectively, were grown in a growth room using a mixed model nested factorial experimental design and scored for a number of phenological and morphological traits. Ambient light conditions significantly affected all of the juvenile traits measured. In both populations, relatively convergent norms of reactions were revealed for leaf appearance dates, as well as for leaf number, suggesting the importance of these traits for functional adjustment to prevailing light conditions. Reaction norms for morphological traits, however, were largely nonparallel, but although often crossing each other, a significant family-by-treatment interaction in the ANCOVAs performed was not detected for any of the traits measured, indicating a small amount of genetic variation for plasticity in these populations. The extent of plastic response to light availability was determined and expressed as an index that estimates the percentage change in a trait value from a high to a low light level. Evolutionary and ecological implications of the observed plastic variations are also discussed.     

Microgeographic edaphic differentiation in allozyme polymorphisms of wild barley (Hordeum spontaneum,Poaceae)

Allozymic variation in proteins encoded by 22 loci was analyzed electrophoretically in 278 individual plants of wild barley,Hordeum spontaneum, the progenitor of cultivated barley, in four 100 meter transects, in Israel, each equally subdivided into basalt and terra rossa soil types. Significant differentiation according to soil was found in 9 alleles. Our results suggest that allozyme polymorphisms in wild barley are at least partly adaptive, and differentiate by edaphic natural selection rather than by stochastic processes, and/or neutrality of allozymic variants.     

Genetic differentiation among populations ofArabis serrata (Brassicaceae) along its geographic distribution

Levels and distribution of genetic variation were investigated in ten populations of the perennialArabis serrata distributed along a latitudinal gradient throughout Japan. Populations of this endemic species occupy predominantly three types of habitats: limestone and derived soils, volcanic and disturbed sites. Previous studies showed that plants ofA. serrata are differentiated in morphological and ecological traits under both natural conditions and common garden experiments suggesting genetic differentiation among populations. To assess the degree of genetic differentiation under different habitats ofA. serrata populations, we analyzed the isozyme genetic structure. Ten populations, located in mountains of central and northern Honshu and Hokkaido, were analyzed by starch gel enzyme electrophoresis. Fourteen loci of eight enzymes were resolved and six loci were monomorphic for all the populations. Populations sampled maintain low levels of genetic variation (P=0.16;H=0.05;A=1.16) compared to that maintained by other outcrossing seed plants. In some cases, few or no heterozygous individuals were detected, and consequently, mean observed heterozygosity was zero or near zero. Six (29%) of the fixation indices,F, estimated deviated significantly from Hardy Weinberg genotypic expectations indicating a deficiency of heterozygotes in most of the cases. The mean genetic identity (Nei'sI) between population pairs was 0.852 and indicates a moderate level of genetic differentiation among populations.Arabis serrata has most of its genetic variation distributed among rather than within populations. The among-population component of the total genetic diversity (G _ST mean value) was 0.416, indicating genetic differentiation between populations. There groups of populations were recognized in an unrooted tree generated by the Neighbor-Joining method. These results suggest groups of populations differentiated regionally. Estimates of interpopulational gene flow (Nm) were very variable (range 0.049–3.718) with a meanNm=1.203 for all populations.     

Cytogenetical changes of offsprings from the induced tetraploid hybrid betweenRanunculus silerifolius (2n = 16) andR. chinensis (2n = 16) (Ranunculaceae)

Cytogenetical studies were carried out on the successive generations of offsprings from the induced tetraploid hybrid (2n = 32) betweenRanunculus silerifolius (2n = 16) andR. chinensis (2n = 16). Aneuploids, 2n = 30 to 35, frequently occurred. In latter subsequent generations the deviation of aneuploids increased, but the proportion of euploids decreased, accompanied by the reduction of fertility of pollen grains and seed sets. F₂ and F₄ PMCs constantly exhibited meiotic abnormality, i.e. formation of quadrivalents and univalents. The speciation process ofR. cantoniensis (2n = 32), which was presumed to arise from tetraploid hybrids between the above two species, is discussed on the basis of the above evidences.Former contributions of this series areOkada & Tamura (1977) andOkada (1984).     

A new species ofThottea (Aristolochiaceae) from India

Thottea ponmudiana sp. nova from Kerala (India) can readily be distinguished from its closest allyTh. siliquosa and all the other known species of the genus by its yellow flowers with purple eyes, deeply lobed perianth with strongly reflexed margins, uniseriate stamens united in three bundles and strongly 4-angled, green, glabrescent fruits.     

Vergleichende cytotaxonomische Untersuchungen anRanunculus seguieri und der Artengruppe desR. alpestris (Ranunculaceae)

Morphological and cytological investigations as well as crossing experiments were carried out withRanunculus seguieri Vill. and 4 species of theRanunculus alpestris L. group (R. alpestris L.,R. traunfellneri Hoppe,R. bilobus Bertol.,R. crenatus Waldst. & Kit.). ForR. seguieri andR. alpestris, localities and distribution are given in addition to extensive diagnoses. A key to the species includes morphological characteristics and distribution data forR. traunfellneri, R. bilobus, andR. crenatus. New diagnostic characters are described. Crossing experiments betweenR. seguieri and the species of theR. alpestris group were unsuccessful. All 5 species have a chromosome number of 2n = 16, the record forR. bilobus is new. There is no statistically significant difference between the karyotypes ofR. seguieri andR. alpestris s. str. Nevertheless, according to morphological evidence and crossing experiments,R. seguieri is not closely related to theR. alpestris group.

     

Self-incompatibility inCrocus vernus subsp.vernus (Iridaceae)

Outcross, self- and mixed pollinations were performed inCrocus vernus subsp.vernus, a species with bicellular pollen, dry stigmas and hollow style. No differences were noted among the above pollinations concerning the germination of pollen and the growth of pollen tubes until the top of ovary. Within 45 min after pollinations 62% of pollen grains germinated. Pollen tubes penetrated the papilla cuticle extending along the papilla wall; on entry into stigmatic lobes they continued growth in the stylar secretion to ovarian locules. Here, however, self-pollen tubes failed to reach or to enter the ovule micropyle; while pollen tubes from either outcross- or mixed pollinations grew until fertilizing ovules. These observations gave evidence of a self-incompatibility system inCrocus, which appeared to be neutralized by mentor effect. The ovary as site of incompatibility response is discussed.