Germination behaviour in populations ofCapsella bursa-pastoris (Cruciferae)

Germination behaviour of variousCapsella bursa-pastoris populations collected from Scandinavia, Middle Europe and the Alps, was tested in unheated, non-illuminated greenhouses (46 populations) and in growth chambers using 5–7 alternating temperature regimes (16 populations). For all populations, the influence of temperature on germination rate is straightforward: the higher the temperature, the greater the germination. Germination capacity, however, may depend on the geographical region. There is also a strong seed age effect on both, rate and capacity of germination. Once dormancy was broken, seeds from all populations were able to germinate over the entire range of temperatures. Some populations revealed a more or less pronounced temperature optimum for germination capacity, others germinated equally well over the entire temperature range. This indicates genetic heterogeneity between populations. However, no correlation between germinability and any environmental pattern was detected. The data indicate thatCapsella bursa-pastoris has adopted a germination strategy which includes a broad temperature tolerance. Germination of wildCapsella plants seems to be regulated by the factors contributing to the inception and breaking of dormancy which depend on pre- and postharvest conditions. Adaptation in germination behaviour inCapsella bursa-pastoris is different from that in other life history traits (flowering behaviour, growth form parameters).     

Variation of growth form parameters inCapsella (Cruciferae)

Growth form parameters ofCapsella bursa-pastoris populations, including a wide range of different environments, have been analyzed from random block field and growth chamber experiments. Changes in one character are often correlated with changes in another. Of special interest are correlations detected with the onset of first flowering. Variation in each of the characters is clearly influenced by both phenotypic and genotypic components. However, genotype — environment interactions are also subject to variation. Therefore, the adaptive significance of a given parameter is not found to be constant over the entire geographical range of the genus. Alpine populations tend to shift from annual to biannual life cycles.Part of a series Adaptation in life history traits of colonizing plant species. Part of a doctoral thesis by the first author. Dedicated to Prof. Dr.Karl Mägdefrau on the occasion of his 80th birthday.     

Biosystematic studies onCapsella bursa-pastoris (Brassicaceae): Enzyme polymorphism in natural populations

The genetic variability of natural populations ofCapsella bursapastoris in North- and Middle-Europe has been estimated by means of enzyme assays. Zymograms of 81 populations have been developed. 17 loci could be identified, and 8 of them can be heterozygous. Genetic variability is greater between populations than within. No correlation between actual population sizes and genetic heterogeneity could be detected. Some electromorphs shift their frequencies proportionally to increasing adversity of climatic conditions, some appear to be constant over the whole area, and others are characterized by an apparently irregular variation pattern. Marginal populations comprise a significantly higher proportion of heterozygous plants than central ones. Apart from this clinal variation pattern, a mosaic pattern, strongly related to habitat conditions, was observed: genetic heterogeneity is greater in more intensively disturbed sites. The pattern of genetic variation in natural populations ofCapsella bursa-pastoris is rather highly influenced by the breeding system.     

Somatic embryogenesis from cell suspension and protoplast cultures ofCapsella bursa-pastoris (L.) Medic

Summary Rapidly growing cell suspension cultures of shepherd’s purse (Capsella bursa-pastoris L. Medic.) were established from leaf-derived calli. These suspensions remained unorganized in the presence of 2,4-D, but underwent extensive root organogenesis in a growth regulator-free liquid medium. Attempts to induce direct embryogenesis in liquid cultures were unsuccessful, but numerous embryos were obtained from cells plated onto growth-regulator-free solid medium. These embryos were frequently abnormal, and secondary embryogenesis was problematic for plant recovery but fertile plants were recovered. Viable protoplasts could readily be isolated from these cell suspensions. After 1 wk of culture, protoplast viability was 62%, and 7% of the cells had divided. Embryogenesis was observed from protoplast-derived microcolonies, plated on growth-regulator-free medium. Although these somatic embryos were difficult to root, plants were recovered. New cell suspensions were more recently established, which were only 4 to 6 mo. old when plant regeneration was attempted. Numerous shoots were obtained when these cells were plated onto growth-regulator-free solid media. However, these shoots differed from the embryos previously obtained in that they readily rooted and rapidly developed into plantlets. This system may allow the use of shepherd’s purse as a gene source for introgression of agronomically interesting traits intoBrassica crop species through protoplast manipulation and somatic hybridization.     

Co-adaptation of seed dormancy and flowering time in the arable weed Capsella bursa-pastoris (shepherd's purse)

Background and Aims

The duration of the plant life cycle is an important attribute that determines fitness and coexistence of weeds in arable fields. It depends on the timing of two key life-history traits: time from seed dispersal to germination and time from germination to flowering. These traits are components of the time to reproduction. Dormancy results in reduced and delayed germination, thus increasing time to reproduction. Genotypes in the arable seedbank predominantly have short time to flowering. Synergy between reduced seed dormancy and reduced flowering time would create stronger contrasts between genotypes, offering greater adaptation in-field. Therefore, we studied differences in seed dormancy between in-field flowering time genotypes of shepherd''s purse.

Methods

Genotypes with early, intermediate or late flowering time were grown in a glasshouse to provide seed stock for germination tests. Secondary dormancy was assessed by comparing germination before and after dark-incubation. Dormancy was characterized separately for seed myxospermy heteromorphs, observed in each genotype. Seed carbon and nitrogen content and seed mass were determined as indicators of seed filling and resource partitioning associated with dormancy.

Key Results

Although no differences were observed in primary dormancy, secondary dormancy was weaker among the seeds of early-flowering genotypes. On average, myxospermous seeds showed stronger secondary dormancy than non-myxospermous seeds in all genotypes. Seed filling was similar between the genotypes, but nitrogen partitioning was higher in early-flowering genotypes and in non-myxospermous seeds.

Conclusions

In shepherd''s purse, early flowering and reduced seed dormancy coincide and appear to be linked. The seed heteromorphism contributes to variation in dormancy. Three functional groups of seed dormancy were identified, varying in dormancy depth and nitrate response. One of these groups (FG-III) was distinct for early-flowering genotypes. The weaker secondary dormancy of early-flowering genotypes confers a selective advantage in arable fields.     

Spatial distribution of mRNA during pre-fertilization ovule development in Capsella bursa-pastoris

Summary In situ hybridization of sections of the ovary and ovule of Capsella bursa-pastoris with ³H-polyuridylic acid [³H-poly(U)] showed the presence of polyadenylic acid-containing RNA [poly(A) + RNA] in the cells of the placenta and nucellus. During megasporogenesis there was a decrease in ³H-poly(U) binding activity of the nucellar cells concomitant with the appearance of poly(A) + RNA in the integuments. As the typical eight-nucleate embryo sac was formed, ³H-poly(U) binding was not apparent around the quartet of nuclei at the chalazal end, while it persisted at the micropylar end. Both the egg and synergids as well as the chalazal proliferating tissue showed high concentrations of poly(A) + RNA in their cytoplasm. The results suggest a role for transient localizations of poly(A) + RNA during female sporogenesis and gametogenesis in C. bursa-pastoris.     

Multivariate and cladistic analyses of the purple-flowered species ofErysimum (Cruciferae) from the Iberian Peninsula

A taxonomically difficult purple-flowered group within the genusErysimum, restricted to the Iberian Peninsula, is analyzed by multivariate and cladistic analyses. 51 specimens have been scored for 14 characters. Both principal components and discriminant analyses provide support to the recognition of the five species considered by the author, namelyE. linifolium, E. lagascae, E. baeticum, E. popovii, andE. cazorlense. Cladistic analysis, using 7 characters resulted in a single most parsimonious cladogram containing no homoplasies. The pattern of morphologic divergence follows a clear NW.-SE. trend, which is congruent with the topology of the cladogram. This trend significantly affects growth-form as well as fruit characters, both providing the main grounds for species recognition. The different behavior and significance of several characters in both kinds of analysis is discussed. The co-occurrence of morphologically similar individuals differing in the flower color is discussed, too. Possible explanations for this phenomenon involve hybridization in a wide sense or, alternatively, rejecting the assumption of monophyly for the group.     

Production and genetic analysis of partial hybrids in intertribal crosses between Brassica species (B. rapa, B. napus) and Capsella bursa-pastoris

Capsella bursa-pastoris (L.) Medic (2n = 4x = 32) is a natural double-low (erucic acid < 1%, glucosinolates < 30 micromol/g) germplasm and shows high degree of resistance to Sclerotinia sclerotiorum. Hybridizations were carried out between two Brassica species viz. B. rapa (2n = 20) and B. napus (2n = 38) as female and C. bursa-pastoris as male parent to introduce these desirable traits into cultivated Brassica species. Majority of F(1) plants resembled female parents in morphology and only a few expressed some characters of male parent, including the white petals. Based on cytological observation of somatic cells, the F(1) plants were classified into five types: two types from the cross with B. rapa, type I had 2n = 27-29; type II had 2n = 20; three types from the crosses with B. napus, type III was haploids with 2n = 19; type IV had 2n = 29; type V had 2n = 38. One to two chromosomes of C. bursa-pastoris were detected in pollen mother cells (PMCs) of type I plant by genomic in situ hybridization (GISH), together with chromosomal segments in ovary cells and PMCs of some F1 plants. Amplified fragment length polymorphism (AFLP) bands specific for the male parent, novel for two parents and absent bands in Brassica parents were generated in majority of F1 plants, even in Brassica-types and haploids, indicating the introgressions at various levels from C. bursa-pastoris and genomic alterations following hybridization. Some Brassica-type progeny plants had reduced contents of erucic acid and glucosinolates associated with improved resistance to S. sclerotiorum. The cytological and molecular mechanisms behind these results are discussed.     

Cytochemical heterochromatin differentiation inSinapis alba (Cruciferae) using a simple air-drying technique for producing chromosome spreads

The fluorochrome and Giemsa chromosome banding patterns and the Ag-NOR histochemical staining ofSinapis alba are described. Two major types of heterochromatin can be distinguished, one of which contains GC-rich DNA. The observations are discussed as they relate to the known satellite DNAs ofS. alba. — A simple air-drying technique for producing spreads of plant mitotic chromosomes is presented. Different materials and staining techniques were tested showing that the method has wide applications.Dedicated in gratitude to Prof. DrElisabeth Tschermak-Woess on the occasion of her 70th birthday.     

Cytotaxonomy and breeding system of the genusBiscutella (Cruciferae)

Chromosome counts were determined for 46 populations ofBiscutella representing 28 taxa. The genus was found to contain diploid taxa with 2n = 12, 16 and 18, tetraploid taxa with 2n = 36 and hexaploid taxa having 2n = 54.B. laevigata L. s. l. consists of diploid and tetraploid populations which are poorly differentiated morphologically. TetraploidB. laevigata s. l. and hexaploidB. variegata Boiss. & Reuter (s. l.) are characterized by chromosomal instability. The variation in chromosome numbers and the occurrence of polyploidy is discussed in relation to the taxonomy of the genus. An investigation of the breeding system showed that most of the annual species were self-compatible and partly inbreeding and most of the perennial species self-incompatible and, therefore, outbreeding, while one annual species,B. cichoriifolia Loisel., showed both systems.     

Rainfall and flowering synchrony in a tropical shrub: Variable selection on the flowering time ofErythroxylum havanense

Summary We tested the adaptive significance of flowering synchrony by means of a quantitative analysis of selection and by flowering induction experiments with the deciduous shrubErythroxylum havanense. Temporal schedules of flower and fruit production were determined for a local population (in three sites) in a Mexican seasonal forest for 2 years (1987–1988). The consequences of natural variation in flowering time (flowering initiation day) on maternal reproductive success (fecundity) were evaluated. We observed high levels of inter- and intraindividual flowering synchrony in 1987, but not in 1988 and this contrast was related to differences in rainfall patterns between the two years. A significant proportion (15.4%) of the phenotypic variation in flowering initiation day was accounted for by environmental variance. The expression of phenotypic variance of flowering time and, consequently, the opportunity for selection to act, are controlled by annual variation in rainfall. Despite the between-year difference in flowering synchrony, we detected a relatively intense directional selection on flowering initiation day in both years, but selection coefficients were of opposite sign (standardized directional gradients were –0.326 and 0.333 for 1987 and 1988, respectively). For both years there was a significant relationship between individual relative fitness and the number of neighbouring flowering plants in a given day, suggesting positive frequency-dependent selection.     

A mathematical model of mucilage expansion in myxospermous seeds of Capsella bursa-pastoris (shepherd's purse)

Background and Aims

Myxospermy is a term which describes the ability of a seed to produce mucilage upon hydration. The mucilage is mainly comprised of plant cell-wall polysaccharides which are deposited during development of those cells that comprise the seed coat (testa). Myxospermy is more prevalent among those plant species adapted to surviving on arid sandy soils, though its significance in determining the ecological fitness of plants is unclear. In this study, the first mathematical model of myxospermous seed mucilage expansion is presented based on seeds of the model plant species Capsella bursa-pastoris (shepherd''s purse).

Methods

The structures underpinning the expansion process were described using light, electron and time-lapse confocal micrographs. The data and experimental observations were used to create a mathematical model of myxospermous seed mucilage expansion based on diffusion equations.

Key Results

The mucilage expansion was rapid, taking 5 s, during which the cell mucilage volume increased 75-fold. At the level of the seed, this represented a 6-fold increase in seed volume and a 2·5-fold increase in seed surface area. These increases were shown to be a function of water uptake (16 g water g⁻¹ mucilage dry weight), and relaxation of the polymers which comprised the mucilage. In addition, the osmotic pressure of the seed mucilage, estimated by assessing the mucilage expansion of seeds hydrated in solutions of varying osmotic pressure, was –0·54 MPa (equivalent to 0·11 m or 6·6 g L⁻¹ NaCl).

Conclusions

The results showed that the mucilage may be characterized as hydrogel and seed-mucilage expansion may be modelled using the diffusion equation described. The potential of myxospermous seeds to affect the ecological services provided by soil is discussed briefly.     

Variation in flowering behaviour inVulpia (Poaceae)

Some abnormalities in the androecial structure and behaviour withinVulpia sect.Vulpia are described. These features help to trace the evolutionary history of cleistogamous taxa with a single micranthous stamen from chasmogamous taxa with three macranthous stamens.     

Chromosomal variability in natural populations of Chironomus cingulatus meigen (Diptera, Chironomidae)

Chromosomal polymorphism has been studied in seven natural populations of Chironomus cingulatus from Western Europe, Western Siberia, and the Republic of Sakha (Yakutia). The banding sequences pool of the species includes 15 banding sequences. Chromosomal polymorphism was revealed in five out of seven chromosomal arms. Arm B is the most polymorphic with four banding sequences. There are three banding sequences in arm A. Arms D, E, and G have two banding sequences. None of the chromosome rearrangements were revealed in arms C and F. The populations of C. cingulatus differ clearly in their number and frequency of banding sequences, which indicates that different gene sequences are adaptive in different populations.     

Evaluation ofChaoborus predation on natural populations of herbivorous zooplankton in a eutrophic lake

Hydrobiologia - The effect ofChaoborus predation on natural zooplankton communities was tested in enclosure experiments during June, July and August in the eutrophic Frederiksborg Slotssø,...     

Pleiotropic effects of flowering time genes in the annual crucifer Arabidopsis thaliana (Brassicaceae)

Variation in flowering time of Arabidopsis thaliana was studied in an experiment with mutant lines. The pleiotropic effects of flowering time genes on morphology and reproductive yield were assessed under three levels of nutrient supply. At all nutrient levels flowering time and number of rosette leaves at flowering varied among mutant lines. The relationship between these two traits depended strongly on nutrient supply. A lower nutrient supply first led to an extension of the vegetative phase, while the mean number of leaves at flowering was hardly affected. A further reduction resulted in no further extension of the vegetative phase and, on average, plants started flowering with a lower leaf number. At low nutrients, early flowering affected the timing of production of siliques rather than the total output, whereas late flowering was favorable at high nutrients. This may explain the fact that many plant species flower at a relatively small size under poor conditions. Flowering time genes had pleiotropic effects on the leaf length, number of rosette and cauline leaves, and number of axillary flowering shoots of the main inflorescence. Silique production was positively correlated with the number of axillary shoots of the main inflorescence; the number of axillary primordia appeared to have a large impact on reproductive yield.     

Variation in amount of enzyme protein in natural populations

Among strains of Drosophila melanogaster each derived from a single fertilized female taken from natural populations, there is variation in both alcohol dehydrogenase (ADH) activity and the amount of ADH protein. The correlation between ADH activity and number of molecules over all strains examined is 0.87 or 0.96 in late third instar larvae depending on whether the substrate is 2-propanol or ethanol. With respect to the two common electrophoretic allozymic forms, F and S, segregating in these populations, the FF strains on the whole have higher ADH activities and numbers of ADH molecules than the SS strains. Over all strains examined, enzyme extracts from FF strains have a mean catalytic efficiency per enzyme molecule higher than that of enzyme extracts from SS strains when ethanol is the substrate, and much higher when 2-propanol is the substrate. One FF strain had an ADH activity/ADH protein ratio characteristic of SS strains.     

Das Blühverhalten vonCapsella bursa-pastoris (Brassicaceae)

Capsella bursa-pastoris (L.)Med. is a partially self-pollinating, autogamous plant. The selfing-rate depends on ecological factors. High atmospheric humidity at temperatures over 15 °C and low light intensity, i.e. cloudy and rainy weather, lead to almost exclusive self-pollination, while dry and sunny weather favours outcrossing. At low temperatures (about 4–10 °C) anthesis is prolongated up to five-fold, but allogamy is reduced. Day-length distinctly influences the beginning of flowering.Capsella bursa-pastoris is not a day-neutral plant (as was supposed up to now) but behaves as a quantitative long-day plant.

Teil einer Publikationsreihe über die Populationsökologie vonCapsella bursa-pastoris (L.)Med.     

Genetic variation in natural and laboratory populations of the marsupial Sminthopsis crassicaudata

Gel electrophoresis of blood proteins has detected allelic variation at five loci (TRF, PGD, SOD, ADA, GPI) in a laboratory colony of the dasyurid marsupial Sminthopsis crassicaudata. Family data show no significant departures from Mendelian expectations. Analysis of blood from wild-caught progenitors of the colony revealed significant differences in gene frequency between groups of animals captured from different parts of southern and central Australia and showed that there are two major population clusters. These interpopulation differences are particularly marked at the TRF locus and indicate that the river Murray is a barrier for this species.This project was supported by a grant from the Australian Research Grants Scheme.     

Herbivory and local genetic differentiation in natural populations of Arabidopsis thaliana (Brassicaceae)

To explore genetic variation in defence against the natural herbivores of Arabidopsis thaliana, we transplanted genotypes between a dune habitat and inland habitat in both of which A. thaliana occurred naturally. In previous years we had observed that the specialist weevils Ceutorhynchus atomus and C. contractus (Curculionidae) fed conspicuously on flowers and fruits of A. thaliana in the dunes, while these weevils were always rare in inland habitats. Taking all plants together, total fruit damage was indeed much higher in our experimental plots in the dune habitat (59.7%) relative to the inland garden habitat (18.9%). Within a habitat, additional differences existed between plants of different origins, pointing to genetic differences in ecologically relevant characters; plants of inland origin flowered a week earlier, grew better and produced more fruits than plants of dune origin. However, plants of inland origin experienced more total fruit damage by the specialist weevils (75.4%) than plants of dune origin (44.0%) when the two types grew side by side in the dune habitat. Escape from herbivory gives dune genotypes an advantage in their native habitat, whereas stronger growth and higher survival gives inland genotypes an edge under garden conditions.