Lead tolerance of Betula and Salix in the mining area of Mechernich/Germany

Natural populations of woody perennials on lead-mining sites in the Mechernich area of the Eifel Mountains were investigated with respect to soil factors determining the degree and type of heavy metal tolerance. Salix caprea L. (Goat Willow) grew on soils with up to 17000 mg kg^–1 total lead (ca. 4000 mg kg^–1 ammonium acetate-exchangeable Pb). Betula pendula Roth (Silver Birch) was found on soils containing as much as 29000 mg kg^–1 total lead (7000 mg kg^–1 ammonium acetate-exchangeable Pb). Other woody perennials, with the exception of the dwarf shrub Calluna vulgaris, were not found in the contaminated area even though they did occur in the immediate vicinity. The two lead-tolerant tree species did not form mixed populations.Because of a significantly lower Pb/Ca ratio in Salix soils (2.2) compared with Betula soils (7.4), a calcium-dependent mechanism of lead tolerance is suggested for Salix, but not for Betula.The Betula population could be divided into two groups, each showing a highly significant correlation between root-lead content and exchangeable lead amounts in the soil, but with different levels of lead uptake. The only soil factor distinguishing the two groups was found to be the level of soluble phosphate. A distinctly low level of soil phosphate correlated with a high lead concentration in roots of the one group (30000 mg Pb kg^–1 DW), whereas high phosphate amounts corresponded with a much lower lead concentration in roots of the other (12000 mg Pb kg^–1 DW^–1). Since the correlation between lead in the soil and in plants was similar for the two groups, it is concluded that the type of lead tolerance in Betula is determined by the status of plant phosphate nutrition, rather than by simple phosphate precipitation in the soil.A comparison of growth between different populations of Betula seedlings on homogenized soils from the mining area revealed the Mechaernich population to be a distinct ecotype with respect to lead tolerance. The control population obtained from a non-contaminated area exhibited a lower degree of lead tolerance coupled with a two-step strategy of adaptation to lead.     

Tolerance ofFestuca rubra L. to zinc in relation to mycorrhizal infection

Summary Plant yield of mycorrhizal and non-mycorrhizalFestuca rubra L. was linearly decreased with increasing zinc concentrations in nutrient solution. In all cases, non-mycorrhizal plant growth was significantly greater than that of mycorrhizal plants. Zinc and phosphorus concentrations of root and shoot of mycorrhizal plants were greater in all zinc treatments while mycorrhizal plants showed equal or lower tolerance indices to zinc than non-mycorrhizal plants. Yield depressions of mycorrhizal plants may be the result of enhanced zinc and phosphorus concentrations combined with the cost for growth and maintenance of the mycorrhizal fungi.     

Novelties in the Festuca valesiaca group (Poaceae) from the central Alps

Due to their ploidy levels, their morphological and anatomical characters and their distribution the central alpine Festuca stricta subsp. bauzanina (2n = 8x = 56) and Festuca ovina var. guinochetii (2n = 10x = 70), both belonging to the Festuca valesiaca group, are raised to the specific level. As a new subspecies Festuca bauzanina subsp. rhaetica is described here. The taxa of the polyploid Festuca valesiaca group are morphologically, anatomically and chorologically characterised and an identification key for these taxa in the central Alps is presented.     

The adaptation of Dianthus carthusianorum L. (Caryophyllaceae) to growth on a zinc–lead heap in southern Poland

Plants of Dianthus carthusianorum from a calamine (zinc–lead) waste heap and from unpolluted stands were compared in the field and under controlled laboratory conditions. It was found that the waste-heap plants differed significantly from those in the normal population in respect to the following morphological traits: lower weight of aerial parts, shorter and narrower leaves, smaller number of leaves per plant. In combination with shorter, less numerous shoots, these features reduce the transpiration area of the plants. The general habit (predominance of forms with short and trailing shoots) of the waste-heap population points to adaptation to a xeric environment. Under controlled conditions all of the above traits were maintained through three successive generations. They are thus genetically controlled. Root tolerance tests showed that the waste-heap plants had higher tolerance to zinc and lead. These results indicate that strong selection processes have generated a clear ecotypical differentiation for heavy metal tolerance and drought tolerance in a zinc–lead-rich waste-heap population of D. carthusianorum (variety).     

Effect of wet and dry cycles in calcareous soil on mineral nutrient uptake of two grasses,Agrostis stolonifera L. and Festuca ovina L.

Shoot uptake of mineral nutrients (Ca, Cu, Fe, K, mg, Mn, P, S, Zn) by Agrostis stolonifera L. was compared with Festuca ovina L. under wet and dry cycles. Such conditions are typical for A. stolonifera sites, whereas F. ovina is growing mostly on consistently drier and better-drained soils. Plants were grown in a glasshouse, at controlled temperature and light conditions, using two moisture regimes, one constant at 60% WHC (water holding capacity), one wet/dry fluctuating between 35 and 100% WHC. Above ground and total biomass production was lower under wet/dry treatment than at constant water regime in F. ovina, but did not differ between regimes in A. stolonifera. Shoot uptake of most elements was severely reduced in F. ovina at the wet/dry regime. Shoot uptake and concentrations of most elements studied (Cu, K, Mn, P, S, Zn) were lower (p<0.05) under wet/dry treatment than at constant regime in A. stolonifera and tended to be lower also of Fe and Mg. Differences in biomass production observed are consistent with field evidence that A. stolonifera grows in sites which are periodically flooded but may become quite dry during other periods, and that F. ovina is limited to sites which are consistently drier and better drained. Evidence from the present study, however, does not support any view that alternating wet and dry cycles, as typical of A. stolonifera field sites, would be beneficial to nutrient acquisition of this species but that biomass production may develop normally at the lower uptake of most mineral nutrients measured under the wet/dry regime. Such regimes are decidedly unfavourable to both growth and nutrient acquisition of F. ovina.     

Moisture conditions and the presence of bryophytes determine fescue species abundance in a dry calcareous grassland

Festuca ovina is the abundant matrix-forming species and F. rubra a subordinate species in shallow-soil calcareous grasslands. F. pratensis is a transient species, occurring sparsely in this community. We hypothesised that the different abundances of these three species are primarily due to the differential effect of moisture conditions on their germination and early establishment, and that the effect of the pattern of rainfall intensity depends on the presence or absence of a bryophyte layer. We studied the dependence of the germination and establishment of the three fescue species on the moisture conditions both in the laboratory and in the patches of intact grassland community (microcosms). In a laboratory germination experiment, F. pratensis showed the highest, F. rubra , the intermediate and F. ovina, the lowest drought tolerance. In microcosms, the establishment of F. ovina was the highest. At the same time, the annual mortality of seedlings of F. ovina was the lowest. All three species responded positively to an increasing irrigation level. Differently from F. ovina, F. rubra showed a positive response only in plots from which the bryophyte layer had been removed, while F. pratensis responded positively to both irrigation and bryophyte removal. We conclude that moisture conditions have a differential effect on the three fescue species mainly in the seedling establishment, not in the germination phase. For the successful establishment of F. rubra and F. pratensis, the coincidence of high rainfall and local disturbance, removing bryophytes, is required. The presence or absence of bryophytes had no effect on establishment in dry years, while in rainy years the removal of bryophytes has a clear positive effect.     

The effect of living plants on root decomposition of four grass species

We tested whether living plant roots of Holcus lanatus and Festuca ovina can affect the decomposition rate of dead roots of Holcus lanatus, Festuca rubra, Anthoxanthum odoratum and Festuca ovina. Moreover, we investigated whether this effect is dependent on the decomposing root species or the nitrogen supply during the growth of the roots. The selected perennial grass species are typical of grassland habitats in a range from high to low nitrogen availability: H. lanatus, F. rubra, A. odoratum and F. ovina. Seedlings of these species were homogeneously labelled with ¹⁴CO₂ for eight weeks. Plants were grown on soil at two nitrogen levels: one without additional nitrogen and one with nitrogen addition (14 g N m⁻²).
At the start of the decomposition experiment ¹⁴C labelled roots were separated from soil and incubated in litterbags (mesh width 1 mm) in fresh soil. These ¹⁴C labelled roots were left to decompose for 19 weeks in an open greenhouse in soil planted with H. lanatus or F. ovina and in unplanted soil. After the incubation period, the decomposition of the ¹⁴C labelled roots of the four species was measured. The mass and ¹⁴C losses from the dead roots were calculated and the living plant biomass and C, N and P contents of the living plants were measured.
Living plant roots of F. ovina had positive effects on the decomposition rate of F. ovina root litter, but dead A. odoratum roots from the N fertilized treatment decomposed slower in the presence of living F. ovina plants. It seems likely that living plants like F. ovina exude carbon compounds that stimulate the growth of soil microbes and thereby increase dead root decomposition and mineralization. Root decomposition rates differed among the species. We found no evidence to support our hypothesis that dead roots of high fertility species (i.e. H. lanatus and F. rubra) decompose faster than dead roots of low fertility species (i.e. A. odoratum and F. ovina). In unplanted soil, the mass loss and total ¹⁴C loss from A. odoratum dead roots were higher than those from H. lanatus, F. rubra and F. ovina dead roots. Dead roots of F. ovina lost less mass and total ¹⁴C than dead roots of H. lanatus.     

Responses to Mg/Ca balance in an Iranian serpentine endemic plant, Cleome heratensis (Capparaceae) and a related non-serpentine species, C. foliolosa

A soil Ca/Mg quotient greater than unity is generally considered necessary for normal plant growth but some serpentine plants are adapted to much lower Ca/Mg quotients, resulting from a major cation imbalance in their substrata. In order to investigate the growth and tolerance responses of serpentine and non-serpentine species to varied Ca/Mg quotients, controlled nutrient solution experiments were performed using an a newly reported Iranian endemic serpentine plant, Cleome heratensis Bunge et Bien. Ex Boiss. and a related non-serpentine species Cleome foliolosa DC. and a Eurasian Ni-hyperaccumulating species Alyssum murale Waldst. and Kit. Seedlings were grown in modified Hoagland’s solutions with varying Ca and Mg concentrations (0.2–2.5 and 0.5–10 mM, respectively) in a fully factorial randomised block design. The yields of the two serpentine plants increased significantly as Mg concentrations in the nutrient solution were increased from 0.5 to 4 mM but decreased in the 10 mM Mg treatment. For C. foliolosa yields decreased significantly from 0.5 to 10 mM Mg, indicating the sensitivity of this non-serpentine plant, and the relative tolerance of the serpentine plants to extremely high levels of Mg. Shoot and root Mg and Ca concentrations in C. heratensis and A. murale were higher than those in C. foliolosa in the low and moderate Mg treatments, supporting the view that many serpentine plants have a relatively high requirement for Mg. Maximum Mg concentrations were found in the roots of C. heratensis. Yields of C. heratensis and A. murale did not change significantly as Ca levels in nutrient solution increased from 0.2 to 2.5 mM Ca, However the yield of C. foliolosa increased significantly from 0.2 to 1.5 mM Ca, indicating sensitivity in this non-serpentine plant and tolerance of the two serpentine plants to low levels of Ca correlated with tissue Ca concentrations, probably because of a greater ability for Ca uptake at low-Ca availability. Calcium deficiency in the low-Ca treatments could be a reason for reduced yield in the non-serpentine plants.     

Soil nutrient heterogeneity and competitive ability of three grass species (<Emphasis Type="Italic">Festuca ovina,Arrhenatherum elatius</Emphasis> and <Emphasis Type="Italic">Calamagrostis epigejos</Emphasis>) in experimental conditions

We studied the effects of differences in root growth and nutrient pool on the competitive ability of Festuca ovina (short grass), Arrhenatherum elatius and Calamagrostis epigejos (tall grasses) grown in monocultures and in mixtures of homogeneous and heterogeneous environments during two growing seasons. Analysis of variance revealed a significant effect of plant species on nutrient concentrations in above-ground biomass and of substrate type on contents of N, K, Ca, Mg in biomass. The ANOVA also confirmed the significant effect of competitive environment on the concentration of N, K in above-ground biomass. In heterogeneous environments, both tall grasses (in competition with F. ovina) were able to produce more roots in the nutrient-rich patches and to accumulate more nitrogen in plant tissues, which was associated with higher yield of their above-ground biomass. Thus, the relative competitive ability for nutrients of both tall grasses was higher than that of F. ovina. This competitive ability of A. elatius to C. epigejos increased in heterogeneous treatments.     

Drei neue Arten desFestuca ovina-Formenkreises (Poaceae) aus dem Osten Österreichs

WithinFestuca ovina agg. two hexaploid taxa:F. brevipila andF. carnuntina, and one tetraploid:F. eggleri, are described as new species.F. pseudovina Hack. exWiesb. is better treated asF. valesiaca Schleich. exGaudin subsp.parviflora (Hack.), comb. nova.

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Drei neue Arten desFestuca ovina-Formenkreises (Poaceae) aus dem Osten Österreichs

     

Contrasting effects of tropospheric ozone on five native herbs which coexist in calcareous grassland

The aim of this study was to examine the effects of increased tropospheric ozone concentrations on the growth and morphology of five native herbs commonly found to coexist in calcareous grassland in areas of Britain and continental Europe: Anthyllis vulneraria L., Cirsium acaule (L.) Scop., Festuca ovina L., Pilosella offtcinarum F. Shultz & Shultz-Bip and Lotus comiculatus L. In a chronic fumigation (mean O₃ concentration of 71 ppb (71 nl 1^?1) for 7 h d^?1 AOT40 4585 ppb-h) which lasted for 21 d, the effects of ozone were assessed using classical growth analysis. Large reductions in mean relative growth rates for shoot and root weight and root length were observed for the two legumes (Fabaceae) Lotus corniculatus and Anthyllis vulneraria, although these were only statistically significant for Lotus corniculatus. Significant reductions in specific root length (length per unit dry weight) were found for Cirsium acaule and Pilosella officinarum (Asteraceae), while for Festuca ovina (Poaceae) the allometric coefficient was reduced significantly following exposure to ozone. An acute fumigation (mean O₃ concentration of 196 ppb, 7 h) resulted in a range of visible injury, from no injury (Festuca ovina and Pilosella officinarum) through moderate levels of injury (Cirsium acaule and Lotus corniculatus) to extensive and widespread injury (Anthyllis vulneraria). Scoring of visible damage showed that this was only statistically significant for the two legumes, Lotus corniculatus and Anthyllis vulneraria. These results suggest that native herbs may differ in their sensitivity to tropospheric ozone. Both chronic and acute exposures revealed that members of the Fabaceae may be most sensitive to ozone pollution, but the study also suggests that subtle changes in root morphology occurred for members of the Asteraceae. These findings are discussed in relation to the critical levels of ozone set recently for plants and the implications of increasing tropospheric ozone for the conservation of native plant communities.     

DasFestuca vivipara-Problem in den Alpen

The differences betweenFestuca vivipara (tetraploid) andF. ovina subsp.supina (diploid), two often confused taxa, are demonstrated in regard to morphology, leaf anatomy, ecology and distribution. New maps illustrate the distribution ofF. vivipara in the Alps and the Northern Hemisphere. The development of different polyploidy levels withinF. vivipara as a consequence of occasional sexual processes is discussed. (English Summary on p. 39.)

     

Interspecific interactions under fertilization among Elymus nutans,Festuca sinensis and Festuca ovina on the eastern Qinghai‐Tibetan plateau

A field replacement experiment was used to study the interspecific interaction among three perennial grasses (Elymus nutans, Festuca sinensis and Festuca ovina) that are distributed widely on the east Qinghai‐Tibetan plateau. The experiment consisted of four different species mixtures at four seeding densities and two fertilizer levels. Above‐ground biomass, relative yield and complementary effect were determined from harvested shoot dry weights. The results showed that above‐ground biomass was greater in all species mixtures than in monocultures. The difference between the observed and expected relative yield was greater than zero in all mixtures for E. nutans and was greater than zero in the F. sinensis/F. ovina mixture, but was below zero in all other mixtures for F. sinensis, and was below zero in all mixtures for F. ovina. The complementary effect was more negative across all seeding densities except at a seeding density of 400 seeds/m², and was negative across all mixtures except the F. sinensis/F. ovina mixture. In addition, fertilization had an insignificant impact on the relative yield, but a significant impact on the complementary effect. Fertilization enhanced negative interspecific interaction among the species.     

Festuca curvula (Poaceae) — neu für Österreich

The octoploidFestuca curvula Gaudin (F. ovina agg.) is new for the Austrian flora. Epidermal structures indicate relationships toF. pallens. Its distribution ranges from the Pyrenees and French Massif Central through the Alps to western Austria.

     

Tolerance to copper and zinc of Acidithiobacillus thiooxidans isolated from sewage sludge

The effect of copper and zinc ions on sulphur oxidation by Acidithiobacillus thiooxidans, strain SFR01, isolated from anaerobic sewage sludge was assessed, resulting in tolerance levels up to 20 and 200 mmol l^–1 for copper and zinc, respectively. The tolerance levels obtained were higher than the concentration of copper and zinc usually found in the collected sewage sludge. The tolerance levels obtained indicate no constraints for sludge bioleaching of those metals due to their toxicities to the indigenous A. thiooxidans.     

Selective adaptation to copper of populations ofAgrostis tenuis andFestuca rubra (Poaceae)

Populations ofAgrostis Tenuis andFestuca rubra, tolerant and non-tolerant towards Zn and Pb, were studied in regard to their adaptability by natural selection, towards a third metal (Cu) and the time required for the appearance of such tolerance. It was found that Zn and Pb tolerant and non-tolerant populations are likely to select the character of Cu-tolerance within the course of only one generation. More specifically, Zn and Pb tolerant populations show a better adaptability compared with non-tolerant ones. The amount of tolerance to copper obtained by artificial selection is comparable with that of populations growing naturally on copper mines.     

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.     

Metal uptake byCaryophyllaceae species from metalliferous soils in northern Greece

Plant samples collected from 17 populations of 9 taxa of the generaDianthus, Minuartia, Scleranthus, andSilene (Caryophyllaceae), growing in 14 metalliferous sites of N Greece, and surface soil samples from their growth area were analysed for Cu, Pb, Zn, Cd, Ni, Cr, Fe, Mn, Ca, Mg. Metal uptake varies considerably between the populations. A resistance mechanism excluding Cu and Pb from the aerial parts of the plants over a wide range of the soil concentrations up to a critical soil level appeared to exist, whereas Zn concentrations in plants were analogously related to those of the soil.Scleranthus perennis subsp.perennis showed the highest Cu concentration (205 µg/g), whereasMinuartia cf.bulgarica hyperaccumulated Pb (1175 µg/g). Ca concentrations in plants were in most cases much higher than those in soil, whereas the contrary was true for Mg. As a result the Ca/Mg ratio, which was in almost all cases lower than 1 in the soil, was much increased in the plants.     

Zinc toxicity in ectomycorrhizal Pinus sylvestris

Six strains of ectomycorrhizal fungi were compared for their ability to increase zinc tolerance in Pinus sylvestris L. seedlings. Pioneer and late-stage fungi as well as one strain collected from a Zn-polluted site were included in the study. The accumulation of zinc in the host plants was determined at two different sublethal substrate Zn concentrations. The mycobionts varied considerably in their protection of the autobionts against zinc toxicity. Several fungal species reduced zinc accumulation in the pine seedlings. A Thelephora terrestris (Ehrh.) Fr. Strain, however, increased the Zn concentration in its host plants. Specific zinc-retaining capacity of the mycelium and density of the extramatrical mycelium were important features determining the effectiveness of the zinc retention in the fungal symbiont.The experiments were carried out in a growth chamber where plants were cultivated in root observation chambers. The determination of the extent of substrate colonization showed that an elevated zinc concentration in a substrate might be an important stress factor for the growth of an ectomycorrhizal fungus. The growth of the extramatrical mycelium of some species was inhibited at the highest zinc concentration supplied. A determination of the in vitro zinc tolerance of the fungi could not always predict their tolerance as symbionts, where the latter characteristic did not determine the Zn retention capacity of their mycelium.In this paper we also discuss the possibility that on Zn-polluted soils ectomycorrhizal species and strains are selected that are Zn-tolerant and, in addition, are able to protect their own energy source, the autobiont, from toxicity. These mycorrhizal fungi act as a safety net, that can immobilize large amounts of zinc, thus preventing transport to the host plant.     

Tolerance and bioaccumulation of heavy metals in five populations of Cistus ladanifer L. subsp. ladanifer

The effects of heavy metals on the growth, mineral composition (P, K, Fe and Mn) and metal accumulation of five populations of Cistus ladanifer subsp. ladanifer from NE Portugal were investigated in hydroponic experiments. Plants were exposed to increasing concentrations (0–2000 M) of one of eight heavy metals: Cd, Co, Cr, Cu, Mn, Ni, Pb or Zn. Populations of C. ladanifer, whose origin was ultramafic soils (S and UB) or soils developed on basic rocks (B), showed a higher tolerance to the metals Cd, Co, Cr, and Mn, and a considerable degree of tolerance to Ni. In contrast, populations originating on acid-rock soils (M and SC) showed higher tolerance to the metals Cu and Zn. Populations showed different patterns of metal accumulation and distribution in the plant parts, suggesting different mechanisms of metal tolerance are used. The more Cd-, Co- and Mn-tolerant populations (S, UB, B and SC (Cd)) showed accumulation of these three metals in the shoots (shoot:root metal concentration ratios (S:R) > 1). Shoot concentrations of up to 309 g Cd g^–1, 2667 g Co g^–1 and 6214 g Mn g^–1 were found in these populations. The populations, UB and M, showed considerable tolerance to Ni and Zn, respectively. These populations accumulated up to 4164 g Ni g^–1 and 7695 g Zn g^–1 in their shoot tissues, and these metals were efficiently transported from the roots to aerial parts (S:R > 3 (Ni), S:R > 1 (Zn)). In contrast, the S and SC populations maintained higher growth rates in the presence of Ni and Zn, respectively, but showed exclusion mechanisms of metal tolerance: reduced Ni and Zn transport to shoots (S:R < 1). Cistus ladanifer was not able to efficiently transport Cr, Cu or Pb from its roots to its aerial parts (S:R ranged from 0–0.4). The more Cu-tolerant populations, M and SC, showed a greater restriction of Cu transport to the shoots than the ultramafic- or basic-rock populations. Significant changes in the plant mineral composition were found, however, concentrations were generally above mineral deficiency levels. Based on these preliminary results the possible usefulness of this plant for phytoremediation technologies is discussed. However, further investigations are necessary to evaluate its growth and metal accumulation under soil and field conditions.