The introgression of a functional nuclear gene from Poa to Festuca ovina

In sheep's fescue, Festuca ovina, genes coding for the cytosolic enzyme phosphoglucose isomerase, PGIC, are not only found at the standard locus, PgiC1, but also at a segregating second locus, PgiC2. We have used PCR-based sequencing to characterize the molecular structure and evolution of five PgiC1 and three PgiC2 alleles in F. ovina. The three PgiC2 alleles were complex in that they carried two gene copies: either two active genes or one active and one pseudogene. All the PgiC2 sequences were very similar to each other but highly diverged from the five PgiC1 sequences. We also sequenced PgiC genes from several other grass species. Phylogenetic analysis of these sequences indicates that PgiC2 has introgressed into F. ovina from the distant genus Poa. Such an introgression may, for example, follow from a non-standard fertilization with more than one pollen grain, or a direct horizontal gene transfer mediated by a plant virus.     

Alleviation of salt-induced stress on seed emergence using soil additives in a greenhouse

Use of sodium chloride (NaCl) as a deicing salt results in high concentrations of ions in roadside soils, which decreases seedling emergence in these areas. Greenhouse experiments performed in soil culture tested the efficacy of three soil fertilizers, gypsum (CaSO₄), potash (potassium chloride, KCl) and potassium nitrate (KNO₃), in alleviating NaCl stress on seedling emergence of three grass species exhibiting a range of salt tolerance, Poa pratensis (Kentucky bluegrass), Bouteloua gracilis (blue grama), and Puccinellia distans (alkali grass). Two-factor factorial designs were utilized for each species-fertilizer combination. Treatments of 5000 mg/L (0.086 M) NaCl with or without fertilizer, in concentrations that were equal to 0.5, 1, and 2 times the molar equivalent of 5000 mg/L NaCl were applied biweekly. Salt stress on Poa pratensis emergence was alleviated by all fertilizers with CaSO₄ having the greatest effect in alleviating NaCl stress and potash and potassium nitrate alleviating stress at lower treatment levels. Emergence of Bouteloua gracilis and Puccinellia distans was in most cases negatively effected by soil amendments.     

Is <Emphasis Type="Italic">Campanula glomerata</Emphasis> Threatened by Competition of Expanding Grasses?

In East Germany, a high percentage of species-rich, semi-natural dry grasslands has been converted into species-poor communities dominated either by Poa angustifolia or Festuca rupicola. The disappearance of low-intensity types of agricultural land-use such as mowing and grazing has been the cause for the decline of many grassland species, as lack of biomass extraction leads to a new situation in competition. Here we evaluate the performance of the rare forb Campanula glomerata in competition with P. angustifolia and F. rupicola using a replacement design experiment. C. glomerata responded to the presence of these grass species in contrasting ways: whereas the presence of P. angustifolia had a negative effect on C. glomerata biomass and fitness, the presence of F. rupicola affected the forb in a positive way. The results of an additional sowing experiment revealed that the germination and establishment rates of C. glomerata decrease in accordance with increasing density of P. angustifolia. Thus the response of this rare forb varies among neighbour species.     

Frankia in the rhizosphere of nonhost plants: A comparison with root-associated N2-fixing Enterobacter,Klebsiella and Pseudomonas

Bacterial growth in the rhizosphere and resulting changes in plant growth parameters were studied in small aseptic seedlings of birch (Betula pendula and B. pubescens) and grasses (Poa pratensis and Festuca rubra). The seedlings were inoculated with three Frankia strains (Ai1a and Ag5b isolated from native Alnus root nodules and Ai17 from a root nodule induced by soil originating from a Betula pendula stand), and three associative N₂-fixing bacteria (Enterobacter agglomerans, Klebsiella pneumoniae and Pseudomonas sp., isolated from grass roots). Microscopic observations showed that all the Frankia strains were able to colonize and grow on the root surface of the plants tested without addition of an exogenous carbon source. No net growth of the associative N₂-fixers was observed in the rhizosphere, although inoculum viable counts were maintained over the experimental period. Changes in both the biomass and morphology of plant seedlings in response to bacterial inoculation were recorded, which were more dependent on the plant species than on the bacterial strain.     

Invasive Grass Alters Litter Decomposition by Influencing Macrodetritivores

The results of nitrogen (N) fertilization experiments have shown inconsistent rates of plant litter decomposition, a phenomenon that may be explained by dispropotionate influence of animal detritivores (macro-detritivores) on litter mass loss versus that of microbial decomposers, whose activity may be dependent on inorganic N. In turn, macrodetritivores may be influenced by plant species composition via their selection of optimal food resources and habitats. In our experiment, fertilizer had no apparent effect on litter decomposition, suggesting that microbial decomposers did not use the additional inorganic N and/or that macrodetritivores had a greater influence on decomposition. Manipulation of macrodetritivores suggested that plant species composition—dominated in this study by Festuca arundinacea, an exotic, invasive grass, and Aster ericoides, a native forb—caused shifts in detrivore communities and/or feeding patterns that tended to increase litter mass loss. Canopy cover of F. arundinacea and A. ericoides ranged from 0% to 11%, suggesting that low-intensity invasion may produce significant changes in ecosystem function, such as decomposition.     

Food Preference of Wireworms Analyzed with Multinomial Logit Models

Many species of wireworms (larvae of click beetles, Elateridae) are poly- phagous root herbivores. In grasslands under restoration succession with various grass species, we aim to determine the role of wireworms in aboveground vegetation succession. Therefore, it is crucial to know whether wireworms prefer some food plants to others. We have investigated the root preference to different grass species for Agriotes obscurus and Athous haemorrhoidalis and whether these preferences can be explained by covariates. In Experiment 1, individual wireworms could choose between four different plants, one of each species (Anthoxanthum odoratum, Festuca rubra, Holcus lanatus, and Lolium perenne). In Experiment 2, groups of wireworms were released into the soil in the center of 16 plants (4 from each species). We used multinomial logit models (MLMs) to analze the data. In the appendix the use of multinomial response models is clarified with a fictitious example, using the SAS statistic software package. No preference was found in Experiment 1. In Experiment 2 we found differences in attractiveness of plant species depending on wireworm species: A. obscurus preferred grass species from nutrient-rich grasslands (L. perenne and H. lanatus). Both wireworm species disliked F. rubra. The distance from the release point influenced the probability of being found at a certain place at the end of the experiment: wireworms tended to stay in the proximity of the release point. A. haemorrhoidalis was more often found farther from the point of release than A. obscurus. Dispersal was farther from the release point in experiments with young plants (6 weeks) compared to older ones (9 weeks). Results are discussed in a broad ecological context.     

Taxonomic studies of the genusPoa of Japan

With the aim of making clear the boundaries between species in thePoa acroleucahisauchii-nipponica aggregate, chromosomes and morphological features of 746 collections gathered from 125 localities in Japan were examined. For morphological observations, the voucher specimens of 95 collections whose chromosome numbers were reported previously (Tateoka, 1985) were also used. Tetraploids (2n=28) and hexaploids (2n=42), as well as a few pentaploids which were the hybrids between 4× and 6×, were found. By examining morphological features of these collections, two groups were recognized in tetraploids and one in hexaploids. The two tetraploid groups corresponded toPoa acroleuca Steud. andP. hisauchii Honda, and the hexaploid group toP. nipponica Koidz. It was confirmed that the hairiness on the internerve surface of lemma, ligule hairiness and the length ratio of anther/ lemma are the most important features for discriminating between these species; panicle shape, leaf shape and anther length are also helpful for the identification. The ambiguity of the boundaries between species which was hitherto present in the taxonomy of this species aggregate was unrelated to the creation of nature itself but was attributable to the insufficiency of our research work.     

The reaction of some sand-dune plant species to experimentally imposed environmental change: a reductionist approach to stability

Coastal dunes in western Europe have long been under grazing and recreational pressure, but the vegetative cover appears to be remarkably resilient. In a series of experiments on Ynyslas Dunes, Cardigan Bay (Wales, U.K.), trampling, protection, fertiliser, and herbicide treatments were monitored using non-destructive and destructive sampling methods. This paper discusses the behaviour of 12 important species after these treatments on three major dune habitats: Agrostis tenuis, Ammophila arenaria, Bellis perennis, Festuca rubra, Galium verum, Leontodon taraxacoides, Ononis repens, Plantago lanceolata, Poa subcaerulea, Thymus praecox, Homalothecium lutescens, Tortula ruraliformis.Although stability is usually thought of as a feature of the community, a reductionist approach suggests that grasses react quickly but temporarily, dicotyledonous forbs often show slower reaction times, each species reacting in its own way to changes in the environment. A tentative classification of our study species is possible.Nomenclature follows Smith (1978), for bryophytes and Clapam, Tutin & Warburg (1962), for angiosperms.     

Structure of the natural transgene PgiC2 in the common grass Festuca ovina

Background

A horizontal gene transfer has brought an active nuclear gene, PgiC2, from a polyploid Poa species (P. palustris or a close relative) into the common grass sheep''s fescue (Festuca ovina). The donor and the receptor species are strictly reproductively separated, and PgiC2 occurs in a polymorphic state within F. ovina. The active gene copy is normally closely linked to a very similar pseudogene.

Methodology/Principal Findings

By genome walking we have obtained the up- and downstream sequences of PgiC2 and of corresponding genes in the donor and recipient species. Comparisons of these sequences show that the complete upstream region necessary for the gene''s expression is included in the transferred segment. About 1 kb upstream of PgiC2 a fragment with transposition associated properties has been found (TAF). It is present in P. palustris and its polyploid relatives, though not at the homologous position, and is absent from many other grasses, including non-transgenic F. ovina plants. It is possible that it is a part of a transposing element involved in getting the gene into a transferring agent and/or into the recipient chromosome.

Conclusions/Significance

The close similarity of the up- and downstream regions with the corresponding regions in P. palustris excludes all suggestions that PgiC2 is not a HGT but the result of a duplication within the F. ovina lineage. The small size of the genetic material transferred, the complex nature of the PgiC2 locus, and the associated fragment with transposition associated properties suggest that the horizontal transfer occurred via a vector and not via illegitimate pollination.     

Relationship between Contents of Chlorophyll (a+b) (SPAD values) and Nitrogen of Some Temperate Grasses

In a field experiment the chlorophyll (a+b) (SPAD readings) and nitrogen contents of three grass species (Festuca arundinacea Schreb., Lolium perenne L., and Lolium multiflorum Lam.) and three intergeneric hybrids of Festuca pratensis Huds. × Lolium multiflorum Lam. and Festuca arundinacea Schreb. × Lolium multiflorum Lam. were measured. Close relationships were found between SPAD readings and nitrogen leaf content (r ² = 0.873^** 0.491^** and 0.938^**) for the 1^st, 2^nd, and 3^rd cut, respectively. SPAD readings and N contents were closely correlated (r = 0.836^**) confirming that SPAD measurements could be used in grass selection and/or breeding for high N concentration in herbage. This revised version was published online in August 2006 with corrections to the Cover Date.     

Les pelouses à Festuca paniculata du Tessin (Suisse) dans un contexte Alpin

Vittoz P., Selldorf P., Eggenberg S. and Maire S. 2005. Festuca paniculata meadows in Ticino (Switzerland) and their Alpine environment. Bot. Helv. 115: 33–48.Festuca paniculata (L.) Schinz & Thellung locally dominates montane and subalpine meadows of the Alps and other mountains of southern Europe. Vegetation relevés were carried out in Switzerland and northern Italy to study the site conditions under which Festuca paniculata meadows occur in this part of the Alps, their species composition and phytosociological status, and their relationship to Festuca paniculata meadows described previously from the French Alps (Centaureo-Festucetum spadiceae) and Austrian Alps (Hypochaerido uniflorae-Festucetum paniculatae). The Swiss meadows were found to have a similar ecology to those in France and Austria. They occur mostly between 1600 and 2100 m a.s.l on steep slopes with southern aspect, generally on crystalline rocks, but sometimes on calcareous rocks if soils have been decalcified. The species composition of the Swiss meadows is closer to the Austrian than to the French communities, and we attribute them to the association Hypochaerido uniflorae-Festucetum paniculatae with the new subassociation polygaletosum chamaebuxi. Climate is probably the main factor separating vegetation units in the Alps: the Centaureo-Festucetum spadiceae occurs where summers are dry, whereas the Hypochaerido uniflorae-Festucetum paniculatae occurs where rainfall is not a limiting factor in summer.Manuscrit accepté le 10 février 2005     

Phytoremediation of Metal-Contaminated Soil in Temperate Humid Regions of British Columbia,Canada

The suitability of five plant species was studied for phytoextraction and phytostabilisation in a region with temperate maritime climate of coastal British Columbia, Canada. Pot experiments were conducted using Lolium perenne L (perennial rye grass), Festuca rubra L (creeping red fescue), Helianthus annuus L (sunflower), Poa pratensis L (Kentucky bluegrass) and Brassica napus L (rape) in soils treated with three different metal (Cu, Pb, Mn, and Zn) concentrations. The bio-metric characters of plants in soils with multiple-metal contaminations, their metal accumulation characteristics, translocation properties and metal removal were assessed at different stages of plant growth, 90 and 120 DAS (days after sowing). Lolium was found to be suitable for the phytostabilisation of Cu and Pb, Festuca for Mn and Poa for Zn. Metal removal was higher at 120 than at 90 days after sowing, and metals concentrated more in the underground tissues with less translocation to the aboveground parts. Bioconcentration factors indicate that Festuca had the highest accumulation for Cu, Helianthus for Pb and Zn and Poa for Mn.     

Variable effects of endophytic fungus on seedling establishment of fine fescues

Seedborne systemic endophytic fungi of grasses are thought to be plant mutualists, because they have been shown to improve their host’s resistance against biotic and abiotic stresses. The interactions in plant–endophyte associations vary from mutualistic to parasitic with environmental conditions and the genotypes of interacting species. The possible pros and cons of endophytic fungi are expected to be most evident during the seedling establishment, where host fitness is most directly affected. If this holds true, endophytes may play a focal role in local adaptation of hosts to different environments. We examined if endophyte-infected and uninfected seeds and seedlings of two native grass species, Festuca rubra and F. ovina, differ in seed germination and seedling growth rates under greenhouse conditions. The germination of F. rubra seeds was also studied in the field. This is the first time that the effects of Epichloë endophyte on seedling establishment of fine fescues from natural populations have been experimentally evaluated. Mother plant (seed family) had a marked effect on many response variables in both grass species. Length and mean biomass of tillers of endophyte-infected (E+) F. ovina seedlings were lower, but root:shoot ratios were higher than in endophyte-free (E?) seedlings. In F. rubra, the effects of the endophyte were dependent on the habitat where the seeds were collected. The E+ seeds from river banks germinated faster than E+ seeds from meadows, and E+ seedlings from the river banks produced fewer but taller and heavier tillers than the other seedlings. Our data suggest that the effects of the endophyte infection on the seedling stage of fine fescues are dependent the species of grass, host genetic background and mother plant habitat. The germination strategy and growth form of E+ red fescue seedlings from river banks may be beneficial to surviving in the harsh conditions of that habitat.     

Competition between two grass species with and without grazing over a productivity gradient

Soil nutrient-level and herbivory are predicted to have opposing effects on the allocation pattern of the competitive dominant plant species. Lower stem and higher leaf allocation are favoured when plants are grazed, whereas a higher stem allocation is favoured at high nutrient levels. Grazing by hares and geese can prevent invasion of the tall Elymus athericus, into short vegetation of Festuca rubra, at unproductive stages of salt-marsh succession but not at more productive stages. We hypothesise that the negative effect of herbivory on Elymus decreases due to increasing soil nitrogen levels and shifts the competitive balance towards this species. We tested how simulated grazing and nitrogen availability affected the competitive balance between adult plants of both grass species in a greenhouse experiment. Elymus had a higher above-ground biomass production, invested relatively more in stem and root tissue and had a larger shoot length than Festuca. The above-ground relative yield of Elymus in mixtures of both species increased with increasing nitrogen levels. This indicates that Elymus was the superior competitor at high soil fertility. Although clipping removed relatively more biomass from Elymus than from Festuca and exceeded the observed biomass removal in field conditions, it did not change the competitive balance between both species. Decreasing effects of herbivory due to increasing nitrogen levels are not a likely explanation for the invasion of Elymus in productive marshes. The results suggest that once Elymus has established it can easily invade vegetation dominated by Festuca irrespective of grazing by herbivores such as hares and geese. Herbivory by small herbivores may mainly retard the invasion of this plant by influencing establishment itself.     

Competitive responses of the rare Viola elatior and the common Viola mirabilis

Viola mirabilis is abundant in Estonia; Viola elatior is rare. We tested whether these species differ in their competitive responses to a common grass. We used a pot experiment in which individuals of each violet species grew with 0, 2, 4, or 8 individuals of Festuca ovina in natural soil. The response patterns of shoot and root mass and leaf numbers of the two species were similar: the biomass and leaf number of both violet species decreased with increasing grass density. The decrease of root mass of the rare V. elatior was significantly greater, and the decrease of leaf number tended to be greater, than that of the common V.␣mirabilis. We conclude that the stronger competitive response of V. elatior compared to V. mirabilis might be one reason behind its lower regional and local abundance.     

Role of topography and soils in grassland structuring at the landscape and community scales

The relative importance of abiotic factors in community assembly is debated and thought to be dependent on the scale. I investigated the relative role of topography and soils as structuring agents at the landscape and the community scales in 126 subalpine calcareous grasslands in the Pyrenees, in terms of species composition and abundance. I wished to know: (1) the role of abiotic factors in the organization of plant communities across the landscape; (2) how much of the variation in community distribution was accounted for by abiotic factors; and (3) how well their role applied to the distribution of dominant species at the landscape and the community scales. The hypothesis was: abiotic factors play an important role in community distribution in the landscape, but species interactions are more important within communities. Multivariate methods generated four communities, organized in two contrasting groups along the main vegetation axis, which explained 13% of the variation: mesic grasslands (Nardus stricta and Festuca nigrescens communities) and xeric grasslands (Carex humilis and Festuca gautieri communities). Mesic communities were more acidic and fertile than xeric communities. Changes in the abiotic environment, accounting for up to 80% of the variation in the vegetation, were smooth, while the transition between xeric and mesic grasslands was sharp in terms of species composition. The distribution in the landscape of the first main species from each community was closely related to abiotic factors, which modeled poorly the abundance of the main species at smaller scales. At the within-community scale, the explanatory power of biotic relationships was community dependent, producing the most significant models for plants highly dominant within their communities, such as N. stricta and F. gautieri. Contrary to current hypothesis, there was a shift from mainly positive relationships among dominant species in fertile mesic communities to mainly negative in infertile xeric ones.     

Evaluation of the potential role of water in spread of conidia of the Neotyphodium endophyte of Poa ampla

Neotyphodium endophytes are asexual, filamentous fungi, mutualistically associated with diverse cool season grasses. Infected seeds and vegetative organs of infected host plants are the only known modes of propagation of the asexual endophytes. In the last decade certain Epichloë and Neotyphodium-infected grass species have been shown to have epiphyllous structures of the endophytes, hyphae, conidiophores, and conidia, growing on leaf blades. The production of epiphyllous conidia suggests the possibility that some of these endophytes may have the ability for plant-to-plant spread. The objective of this study was to determine the possible mechanisms involved in liberation and dispersal of asexual spores of Neotyphodium growing in vitro and epiphyllously on leaves of Poa ampla. Our results indicate that water dispersal is the most likely means of dissemination of conidia of the Neotyphodium sp. Wind generated by dry compressed air does not dislodge the conidia from slide cultures or from P. ampla leaves.     

A recently silenced, duplicate PgiC locus in Clarkia

Previous electrophoretic analysis showed that 17 diploid species of the wildflower Clarkia (Onagraceae) have two cytosolic isozymes of phosphoglucose isomerase (PGIC; EC 5.3.1.9), whereas 15 other diploid species have a single PGIC. Molecular studies revealed that the two isozymes in the former species are encoded by duplicate genes, PgiC1 and PgiC2, whereas the single isozyme in the latter is always encoded by PgiC1. Phylogenetic analysis of the nucleotide sequences implied that PgiC2 was silenced four times independently in the genus. Here we describe a psi PgiC2 from C. mildrediae, a species in which only PgiC1 is expressed. The discovery of the psi PgiC2 is significant because it confirms a formal prediction of the phylogenetic analysis. The psi PgiC2 includes 5,039 nucleotides corresponding to 18 of the 23 exons of PgiC, as well as the intervening introns and 3' nontranslated region. The absence of an increase of nucleotide substitutions in its "exons" suggests that the gene was silenced recently. The present study appears to be the first to establish that a specific duplicate gene locus regularly expressed in a group of related plant species has been silenced in one of them. The multiple independent silencings of PgiC2 suggest that it remained functional but inessential in ancestral lineages. We discuss the possibility that PgiC2 may have been preserved in these lineages by selection against mutants causing defective PGIC1- PGIC2 heterodimers.      

A DNA marker for the duplicated cytosolic PGI genes in sheep's fescue (Festuca ovina L.)

Active duplicate PgiC genes in sheep's fescue, Festuca ovina, are associated with a PCR marker of specific length (about 370 bp, of which 231 are in an intron). Using this marker, the frequency of plants with duplicate genes is estimated to be about 10% in a population from southern Sweden. The close molecular similarity between the electrophoretically different duplicated genes is in accordance with the conclusion reached earlier that they are indeed alleles at the same locus.     

Balance of Competitive and Facilitative Effects of Exotic Trees on a Native Patagonian Grass

The balance between facilitation and competition in plants changes with species characteristics and environmental conditions. Facilitative effects are common in natural ecosystems, particularly in stressful environments or years. Contrarily, in artificial associations of plants, such as agroforestry systems, some authors have suggested that even when facilitative effects may occur, net balance of tree effects on grasses is usually negative, particularly in dry environments. The aim of this study was to determine the net effect of the exotic ponderosa pine on the native grass Festuca pallescens (St. Ives) Parodi in agroforestry systems in Patagonia. Soil water content, plant water status, and relative growth were measured in the grass growing in different treatments (determined by tree cover level) during two growing seasons with contrasting climatic conditions. Facilitative effects of trees over grass water status were recorded only when water availability was high. A net negative effect was detected on dates when soil water content was very low and evaporative demand was high. The strength of these negative effects depended on tree density and climatic conditions, being higher in treatments with lower tree canopy cover. These results indicate that the positive effect of trees could only be expected under relatively low stress conditions. However, relative growth of grasses was always similar in plants growing in forested plots than in open grassland. Differences in biomass allocation for grasses growing in shade and open habitats may reconcile these contrary results. Our results highlight the importance of the physiology of a species (relative drought and shade tolerance) in determining the response of a plant to a particular interacting species.