Reproductive success,spontaneous embryo abortion,and genetic load in flowering plants

Summary Reproductive success is divided into two phases: preemergent (the number of viable seeds that enter the ambient environment) and postemergent (the percentage of progeny that survive to reproduce). We studied preemergent reproductive success (PERS) in flowering plants by measuring the fruit/flower (Fr/Fl) ratio and the seed/ovule (S/O) ratio in a number of species of outcrossing and inbreeding plants, where PERS=the product of (Fr/Fl) and (S/O). In order to determine the influence of the ambient environment (including resource availability) we studied pairs of outcrossing and inbreeding species occurring in the same habitat. Among outcrossing species PERS averaged about 22%, whereas in inbreeding species the average was approximately 90%. The progeny/zygote (P/Z) ratio was studied in hand-pollinated populations in Epilobium angustifolium (a strongly outcrossing species) from populations in Oregon and Utah, by direct observation of embryogenesis at twoday intervals throughout the course of seed development. The P/Z ratio in both populations averaged near 30%, and the developing embryos showed a surprising array of abnormalities that resulted in embryo death. During early development >95% of the ovules had normally developing globular embryos, but beginning with differentiation (cotyledon formation) about 70% of the original globular embryos aborted during the course of embryogenesis and seed development. The clustering of developmental lethals during peroids of major differentiation events parallels the animal model of development. We found little evidence that PERS was limited by the ambient environment (including resource availability), pollination, or factors associated with the inbreeding habit. Instead, PERS was found to be inextricably linked to outcrossing plants, whose breeding systems promote genetic variability. The high incidence of developmental lethals in E. angustifolium and the resulting low P/Z ratio (ca. 30%) is attributed to genetic load (any lethal mutation or allelic combination) possibly working in combination with developmental selection (interovarian competition among genetically diverse embryos). Examples of maternally controlled, fixed patterns of ovule abortion with respect to position or number are discussed. However, we found no need to employ female choice as a hypothesis to explain our results for the extensive, seemingly random patterns of embryo abortion in E. angustifolium and other outcrossing species. A more parsimonious, mechanistic explanation based on genetic load-developmental selection is sufficient to account for the differential survivorship of embryos. Likewise, the traditional concept of a positive growth regulator feedback system based on the number of surviving ovules in an ovary can account for subsequent fruit survivorship.     

Impact of the field slug Deroceras reticulatum on establishment of ryegrass and white clover in mixed swards

The relationship between numbers of Deroceras reticulatum and the establishment of ryegrass cv. Melle and white clover cv. Milkanova was studied on a mini-plot experiment. Assessments 12 and 33 days after sowing showed that Deroceras reticulatum had a proportionally greater impact on survival and growth of clover than on ryegrass. Slugs had no apparent effect on numbers or growth of three species of broad-leaved weeds (Senecio vulgaris, Cardamine hirsuta and Epilobium sp.) which occurred in the field experiment. In pot experiments where seedlings of ryegrass together with one of three clover cultivars, with or without chickweed (Stellaria media), were exposed to Deroceras reticulatum, seedlings of white clover cv. Aran were less affected by slug grazing than cvs S184 or Milkanova. The presence of different clover cultivars did not significantly influence the impact of slugs on growth of ryegrass or chickweed, nor did the presence or absence of chickweed influence the effect of slugs on ryegrass and clover seedlings. Slugs had less impact on the growth of chickweed than on ryegrass or clover. Thus slug grazing would make clover less able to compete with chickweed as well as the other weeds found in the mini-plot experiment.     

Pollen morphology and structure of Tropical and Subtropical American genera of the Piptadenia -group (Leguminosae: Mimosoideae)

A survey of the pollen morphology of the American taxa of the Piptadenia -group (Mimoseae) was done including all together 384 species of the following genera: Piptadenia , Parapiptadenia , Pseudopiptadenia , Piptadeniopsis , Microlobius (= Goldmania ), Stryphnodendron , Adenopodia , Anadenanthera and Mimosa . Because of a high morphological and structural variation the definition and characterization of pollen types turned out to be very difficult. Nevertheless the presence and durability of some pollen characters allow a provisional grouping: (1) always tetrads or polyads, (2) small size 6 (20) 40 mum, (3) unstable shape but currently ovoidal and, if biconvex, ellipsoidal in the outline, (4) number of pollen grains of the polyads very variable and unstable; the most frequent numbers are 8-12-16, (5) pollen grains irregularly united in polyads, with homomorphous exine and, (6) pores always located in the angles of the pollen grains in distal-subdistal position, rarely subdistal, (7) without or with pseudocolpi or subpseudocolpi (never colpi) in the distal face, (8) endexine forming an annulus around the endoaperture with ends of free lamellae with a central white line, (9) the tectal stratum above the pores forming a small vestibular space (except in Anadenanthera ). These palynological characters or most of them confirm the affinity among genera in the Piptadenia -group and its distinction from other taxa of the subfamily Misosoideae with tetrads or polyads.     

Flexible responses of insects to changing environmental temperature – early season development of Craspedolepta species on fireweed

Developmental response to temperature during the critical early season growing period was investigated in two congeneric species of Craspedolepta feeding on Epilobium angustifolium at three sites at different altitudes in Norway and the UK. The larval development reaction norm to temperature, measured as accumulated day degrees, was not significantly different between Craspedolepta nebulosa and Craspedolepta subpunctata at sites where they co‐occurred but C. nebulosa development was consistently more advanced at any site. For individual species the reaction norms at the lowest site (Ainsdale, UK) were similar to the intermediate site (Geilo, Norway): and there were no differences between years. Insect size remained relatively constant. However, at the highest site (Haugastøl, Norway), where C. subpunctata is unable to complete its development, the reaction norm for C. nebulosa was significantly higher than at Geilo and the individual insects produced were smaller. These adaptations allow life‐history completion under limiting temperature conditions. An experiment at Ainsdale, to raise the mean temperature by around 2.5°C during the early growing season, resulted in accelerated development in both C. nebulosa and C. subpunctata but development in C. nebulosa was accelerated proportionately more. C. nebulosa thus displays the greater plasticity in developmental response to environmental temperature that allows it to occupy a greater altitudinal and latitudinal range than C. subpunctata, in which the response is less plastic and more canalized. The likely individualistic responses of the two species to climate warming are considered.     

The influence of tetrad shape and intersporal callose wall formation on pollen aperture pattern ontogeny in two eudicot species

Background and Aims

In flowering plants, microsporogenesis is accompanied by various types of cytoplasmic partitioning (cytokinesis). Patterns of male cytokinesis are suspected to play a role in the diversity of aperture patterns found in pollen grains of angiosperms. The relationships between intersporal wall formation, tetrad shape and pollen aperture pattern ontogeny are studied.

Methods

A comparative analysis of meiosis and aperture distribution was performed within tetrads in two triporate eudicot species with contrasting aperture arrangements within their tetrads [Epilobium roseum (Onagraceae) and Paranomus reflexus (Proteaceae)].

Key Results and Conclusions

Intersporal wall formation is a two-step process in both species. Cytokinesis is first achieved by the formation of naked centripetal cell plates. These naked cell plates are then covered by additional thick, localized callose deposits that differ in location between the two species. Apertures are finally formed in areas in which additional callose is deposited on the cell plates. The recorded variation in tetrad shape is correlated with variations in aperture pattern, demonstrating the role of cell partitioning in aperture pattern ontogeny.     

Oenothein B's contribution to the anti-inflammatory and antioxidant activity of Epilobium sp

Willow herb tea or preparation are available and relatively popular in the European market, and claimed to be effective inter alia because of their anti-inflammatory activity. The present study is therefore aimed at comparing the anti-inflammatory and antioxidant activity of extracts of the three most popular Epilobium species (E. angustifolium, E. hirsutum and E. parviflorum) and at juxtaposing this activity against the dominating compounds from the following extracts: oenothein B (OeB), quercetin-3-O-glucuronide and myricetin-3-O-rhamnoside. The phytochemical analysis of the extracts has shown that OeB quantities vary between 20% and 35%, while flavonoids content does not exceed 2%. All extracts have inhibited the activity of hyaluronidase and lipoxygenase with IC₅₀ around 5 μg/ml and 25 μg/ml. The inhibition of hyaluronidase is related with the presence of OeB, a strong inhibitor of this enzyme (IC₅₀ 1.1 μM). Additionally, the extracts inhibited myeloperoxidase (MPO) release from stimulated neutrophils. OeB inhibited MPO release similarly to the anti-inflammatory drug indomethacin with IC₅₀ 7.7 μM and 15.4 μM, respectively. Tested extracts significantly reduced the production of reactive oxygen species (ROS) from f-MLP and PMA induced neutrophils with IC₅₀ 5 μg/ml and 25 μg/ml, respectively. The flavonoids content seems to exert little influence on extracts’ activity, contrary to OeB, whose high concentration explains the activity of extract obtained from Epilobium. Tested currently marketed Epilobium preparations are often wrongly assigned, but we should stress that the level of OeB in all tested herbs was high and always exceeded 2% in raw material.     

Seed weight increases with altitude in the Swiss Alps between related species but not among populations of individual species

Seed weight is a crucial plant life history trait, determining establishment success and dispersal ability. Especially in stressful environments, larger seeds may be selected at the expense of seed number, because larger seeds have a better chance of giving rise to an established offspring. We tested the hypotheses that between related species-pairs and among populations of single species a similar trend for increasing seed weight with increasing altitude should be present. Firstly, we measured seed weights from 29 species-pairs, with one species occurring in lowland areas and a congeneric species from high altitudes. Seeds of the alpine species were 28±8% larger than seeds from lowland species (P<0.01). Compared to the related lowland species, 55% of the alpine species had heavier seeds, 3% (one species) had lighter, and 41% had seeds of approximately equal weight. Secondly, we compared seed weights among populations of four species from different habitats and with different life histories. Seeds from between 11 and 34 populations per species were sampled along altitudinal gradients of 800–1,500 m (ca. 800 m in Scabiosa lucida, ca. 1,000 m in Saxifraga oppositifolia, ca. 1,000 m in Epilobium fleischeri, and ca. 1,500 m in Carex flacca). In all the four species, we found no indication for heavier seeds at higher altitudes. Our results indicate a selection pressure for species with heavier seeds at higher altitude, but the trend does not seem to operate across all cases. Phylogenetic constraints may limit the correlation among altitude and seed weight, operating particularly against selection for larger seed size, the closer populations and species are related to each other.     

THE EFFECT OF INBREEDING IN DIPLOID AND TETRAPLOID POPULATIONS OF EPILOBIUM ANGUSTIFOLIUM (ONAGRACEAE): IMPLICATIONS FOR THE GENETIC BASIS OF INBREEDING DEPRESSION

The partial dominance model for the evolution of inbreeding depression predicts that tetraploids should exhibit less inbreeding depression than their diploid progenitors. We tested this prediction by comparing the magnitude of inbreeding depression in tetraploid and diploid populations of the herbaceous perennial Epilobium angustifolium (Onagraceae). Inbreeding depression was estimated in the greenhouse for three tetraploid and two diploid populations at four life stages. The mating system of a tetraploid population was estimated and compared to a previous estimate for diploids. Tetraploids showed less inbreeding depression than diploids at all life history stages, and these differences were significant for seed-set and cumulative fitness, but not for germination, survival, or plant dry mass at nine weeks. This result suggests that the genetic basis of inbreeding depression may differ among life stages. The primary selfing rate of the tetraploid population was r = 0.43, which is nearly identical to that of a diploid population (r = 0.45), indicating that differences in inbreeding depression between diploids and tetraploids are probably not due to differences in the mating system. Cumulative inbreeding depression, calculated from the four life history stages, was significantly higher for diploids () than for tetraploids (), supporting the partial dominance model of inbreeding depression.     

Phytometric estimation of fertility of waterlogged rich-fen peats usingEpilobium hirsutum L.

Summary The above-ground net production of herbaceous vegetation in 16 stands of waterlogged rich fen in Broadland, Norfolk UK, has been compared with the growth ofEpilobium hirsutum seedlings (in controlled environment) on replicate peat samples from each site and with the chemical composition of extracts from sub-samples of each peat (Ca, Mg, Na, K, Fe, Mn, inorganic P, inorganic N). There were no consistent or significant relationships between measured concentrations of major nutrients (K, N, P) in peat samples and netin situ production of the vegetation or with growth ofE. hirsutum seedlings on each peat. There was a strong, significant (p<0.001) relationship between above ground productionin situ and growth ofE. hirsutum seedlings. Seedling growth thus provided a simple, sensitive phytometric measure of fertility of the waterlogged peats (pH range 6.0–7.0). One possible limitation is the apparent sensitivity ofE. hirsutum seedlings to high manganese concentrations.     

Vapour Phase Activation,Intracellular Motility and Germination in the Tripororate Pollen of Epilobium angustifolium

The germinability of newly released pollen of Epilobium angustifolium in liquid medium is low, but it is enhanced by a short period in an atmosphere of 85–95% RH without access to liquid water or solutes. During this period the thickened pectic layers of the intine at the germination apertures of the grains are eroded and prospective tube tips emerge. Concomitantly polarised movement of organelles begins in the vegetative cells, and wall precursor vesicles (P-particles) accumulate at the apertures. In pollen held continuously in the humid environment transition to cylindrical growth follows, with the establishment of the inverse-fountain pattern of circulation in the apical part of the tubes.