Arbuscular mycorrhizas are present on Spitsbergen.

A previous study of 76 plant species on Spitsbergen in the High Arctic concluded that structures resembling arbuscular mycorrhizas were absent from roots. Here, we report a survey examining the roots of 13 grass and forb species collected from 12 sites on the island for arbuscular mycorrhizal (AM) colonisation. Of the 102 individuals collected, we recorded AM endophytes in the roots of 41 plants of 11 species (Alopecurus ovatus, Deschampsia alpina, Festuca rubra ssp. richardsonii, putative viviparous hybrids of Poa arctica and Poa pratensis, Poa arctica ssp. arctica, Trisetum spicatum, Coptidium spitsbergense, Ranunculus nivalis, Ranunculus pygmaeus, Ranunculus sulphureus and Taraxacum arcticum) sampled from 10 sites. Both coarse AM endophyte, with hyphae of 5-10 µm width, vesicles and occasional arbuscules, and fine endophyte, consisting of hyphae of 1-3 µm width and sparse arbuscules, were recorded in roots. Coarse AM hyphae, vesicles, arbuscules and fine endophyte hyphae occupied 1.0-30.7, 0.8-18.3, 0.7-11.9 and 0.7-12.8% of the root lengths of colonised plants, respectively. Principal component analysis indicated no associations between the abundances of AM structures in roots and edaphic factors. We conclude that the AM symbiosis is present in grass and forb roots on Spitsbergen.

A meta-analysis of plant responses to dark septate root endophytes.

• Dark septate endophytes (DSE) frequently colonize roots in the natural environment, but the effects of these fungi on plants are obscure, with previous studies indicating negative, neutral or positive effects on plant performance. • In order to reach a consensus for how DSE influence plant performance, meta-analyses were performed on data from 18 research articles, in which plants had been inoculated with DSE in sterile substrates. • Negative effects of DSE on plant performance were not recorded. Positive effects were identified on total, shoot and root biomass, and on shoot nitrogen (N) and phosphorus contents, with increases of 26-103% in these parameters for plants inoculated with DSE, relative to uninoculated controls. Inoculation increased total, shoot and root biomass by 52-138% when plants had not been supplied with additional inorganic N, or when all, or the majority, of N was supplied in organic form. Inoculation with the DSE Phialocephala fortinii was found to increase shoot and root biomass, shoot P concentration and shoot N content by 44-116%, relative to uninoculated controls. • The analyses here suggest that DSE enhance plant performance under controlled conditions, particularly when all, or the majority, of N is available in organic form.

Widespread association between the ericoid mycorrhizal fungus Rhizoscyphus ericae and a leafy liverwort in the maritime and sub-Antarctic.

A recent study identified a fungal isolate from the Antarctic leafy liverwort Cephaloziella varians as the ericoid mycorrhizal associate Rhizoscyphus ericae. However, nothing is known about the wider Antarctic distribution of R. ericae in C. varians, and inoculation experiments confirming the ability of the fungus to form coils in the liverwort are lacking. Using direct isolation and baiting with Vaccinium macrocarpon seedlings, fungi were isolated from C. varians sampled from eight sites across a 1875-km transect through sub- and maritime Antarctica. The ability of an isolate to form coils in aseptically grown C. varians was also tested. Fungi with 98-99% sequence identity to R. ericae internal transcribed spacer (ITS) region and partial large subunit ribosomal (r)DNA sequences were frequently isolated from C. varians at all sites sampled. The EF4/Fung5 primer set did not amplify small subunit rDNA from three of five R. ericae isolates, probably accounting for the reported absence of the fungus from C. varians in a previous study. Rhizoscyphus ericae was found to colonize aseptically-grown C. varians intracellularly, forming hyphal coils. This study shows that the association between R. ericae and C. varians is apparently widespread in Antarctica, and confirms that R. ericae is at least in part responsible for the formation of the coils observed in rhizoids of field-collected C. varians.

Survival of antarctic soil metazoans at -80 degree C for six years.

A sample of the liverwort Cephaloziella varians was collected on 1 January 1999 at Rothera Point on the Wright Peninsula, Adelaide Island, western Antarctic Peninsula and was partially dried and then frozen at -80 degree C. The sample was rapidly defrosted to c. 10 degree C after six years and two months of storage at this temperature. Nematodes, tardigrades and a bdelloid rotifer present in the sample were found to have survived. Of the 159 nematodes recovered from the sample, 49 (31%) were alive: of the tardigrades and rotifers, two of 15 (13%) and one of 48 (2%) had survived, respectively. A Chi-square test showed that there was a significant association between nematode taxon and survival: a greater proportion of Coomansus gerlachei individuals were alive than of Rhyssocolpus paradoxus. A Chi-square test also showed that there was a significant association between phylum and survival: a significantly greater proportion of nematodes or tardigrades were alive than of bdelloid rotifers. We conclude that Antarctic soil metazoans are capable of surviving long-term exposure to low sub-zero temperatures and that there may be taxon-specific effects of freezing on survival.

UV-B radiation arising from stratospheric ozone depletion influences the pigmentation of the Antarctic moss Andreaea regularis.

Changes to the radiative environment arising from stratospheric ozone (O(3)) depletion and subsequent associations between these changes and the pigmentation of the moss Andreaea regularis were measured in late austral spring and early summer 1998 at Rothera Point on the western Antarctic Peninsula (67 degrees S, 68 degrees W). A strong relationship between O(3) column depth and the ratio of UV-B to PAR irradiance ( F(uv-b)/ F(par)) was recorded at ground level ( r(2)=92%, P<0.001). Weaker, but significant, associations between O(3) column depth and ground level unweighted and biologically effective UV-B radiation (UV-B(be)) were also found. Regression analyses indicated that F(uv-b)/ F(par) was the best predictor for concentrations of UV-B screening pigments and total carotenoids extracted from plant tissues. Concentrations of these pigments were loosely ( r(2)= ca. 30%) but significantly ( P<0.01) positively associated with F(uv-b)/ F(par). Concentrations of UV-B screening pigments were also positively associated with irradiances and daily doses of unweighted UV-B and UV-B(be) radiation. The concentrations of chlorophylls a and b were apparently unaffected by O(3) depletion. The data derived from this study suggest that changes to the radiative environment associated with stratospheric O(3) depletion influence the pigmentation of A. regularis. As a corollary, flavonoids are shown to be present in tissues of A. regularis.

Effects of elevated ultraviolet radiation and endophytic fungi on plant growth and insect feeding in Lolium perenne, Festuca rubra, F. arundinacea and F. pratensis.

Plants of perennial ryegrass (Lolium perenne L.), red fescue (Festuca rubra L.), tall fescue (F. arundinacea Schreb.) and meadow fescue (F. pratensis Huds) were exposed at an outdoor facility located in Edinburgh, UK to modulated levels of UV-B radiation (280-315 nm) using banks of cellulose diacetate filtered UV-B fluorescent lamps that also produce UV-A radiation (315-400 nm). The plants were derived from a single clone of each species and were grown both with and without colonization by naturally-occurring fungal endophytes. The UV-B treatment was a 30% elevation above the ambient erythemally-weighted level of UV-B during July to October. Growth of treated plants was compared with plants grown under elevated UV-A radiation alone produced by banks of polyester filtered lamps and with plants grown at ambient levels of solar radiation under banks of unenergized lamps. At the end of the treatment period, sample leaves were collected for feeding trials with the desert locust Schistocerca gregaria (Forsk). The UV-B treatment produced no effects on the aboveground biomass of any of the four grasses. The UV-B treatment and the UV-A control exposure both increased plant height and the number of daughter plants formed by rhizome growth in F. rubra. There were significant effects of endophyte presence on the total fresh and dry weights of F. arundinacea and F. rubra, on fresh weight only in F. pratensis, and on the fresh and dry weights of inflorescence in F. arundinacea and L. perenne. There were no effects of UV treatments on the absolute amounts of leaf consumed or on the feeding preferences of locusts for leaves with or without endophyte in three species: F. rubra, F. arundinacea and L. perenne. In F. pratensis there was no effect of UV treatment on the weight of leaves consumed but a significant UV x endophyte interaction caused by a marked change in feeding preference between leaves with and without endophyte that differed between the UV-B treatment and UV-A control exposures. The alkaloid compounds known as lolines were analysed in leaves of F. pratensis and were only found in plants grown with endophyte. However, there was no significant relationship between total loline content and insect feeding preference. These effects illustrate the potential complexities of species interactions under increasing levels of UV-B. The experiment also demonstrates the importance of appropriate controls in UV lamp supplementation experiments for interpretation of both plant growth and insect feeding effects.

Phialophora graminicola, a dark septate fungus, is a beneficial associate of the grass Vulpia ciliata ssp. ambigua.

Seedlings of the annual grass Vulpia ciliata ssp. ambigua were grown in sterilized sand with a dark septate root fungus, Phialophora graminicola, which had been isolated from a natural population of the grass. Tiller number and shoot, root and total biomass of seedlings grown with P. graminicola were enhanced relative to uninoculated control plants in a growth room and a glasshouse experiment. Root length of seedlings grown with P. graminicola was significantly increased, but no effects of the fungus on root diameter, number of root hairs or specific root length were recorded. Root nitrogen content and shoot, root and total phosphorus contents of seedlings grown with the fungus were enhanced, but shoot nitrogen concentration of these plants was reduced. Shoot biomass and specific root length of inoculated plants were positively associated with the number of P. graminicola colonies re-isolated from roots. These data indicate that P. graminicola acts as a beneficial associate of V. ciliata ssp. ambigua seedlings under controlled conditions.

Multi-functionality and biodiversity in arbuscular mycorrhizas.

Plant roots in natural ecosystems are typically colonized by a wide range of fungi. Some of these are pathogenic, others appear to be opportunistic and have no apparent impact, while mycorrhizal fungi are generally regarded as mutualistic. Of the various types of mycorrhizal fungi, the arbuscular mycorrhizal (AM) association is by far the most abundant and widespread. While the most widely accepted model of AM function depends upon plants benefiting from the facilitation of phosphorus uptake, recent data from field-based studies in temperate ecosystems indicate that only plant species with poorly branched root systems benefit from AM fungi in this way: species with highly branched root systems may benefit in other ways, such as by being protected against root pathogenic fungi. These two responses apparently represent extremes along a continuum of AM benefit determined by root system architecture.

Rhizosphere and root-infecting fungi and the design of ecological field experiments.

As part of a wider study into the role of soil fungi in the ecology of the winter annual grass, Vulpia ciliata ssp. ambigua (Le Gall) Stace & Auquier, we applied the fungicides benomyl and prochloraz to three natural populations of the grass growing in East anglia, United Kingdom. The rhizosphere and rootinfecting fungi associated with the three populations were analysed each month between February and May 1992 when plants set seed. There were marked differences between the fungal floras associated with each of the three populations of V. ciliata, despite the fact that associated plant species and soil nutrient status were broadly similar between sites. This was attributed to wide differences in soil pH between the three populations. Prochloraz did not affect fungal abundance, but benomyl decreased the isolation frequencies of Fusarium oxysporum from roots and the frequencies of Penicillium and Trichoderma spp. isolated from rhizosphere soil, and increased the frequency of isolation of Mucor hiemalis from the rhizosphere of V. ciliata. There were also significant increases in the isolation frequencies of F. oxysporum from roots and M. hiemalis, Trichoderma spp. and Phoma fimeti from the rhizosphere of V. ciliata as plants matured. The significance of these results for the design of ecological field experiments are discussed in light of a previous study which has shown that asymptomatic root-infecting fungi can affect plant fecundity and hence abundance in natural populations of V. ciliata. We propose that differences in microbial communities between sites, controlled in part by soil chemistry, are a major factor determining plant performance under field conditions.

Effects of dry-deposited sulphur dioxide on fungal decomposition of angiosperm tree leaf litter II. Chemical content of leaf litters.

Ash (Fraxinus excelsior L.), birch (Betula spp.), hazel (Corylus avellana L.), sessile oak [Quercus petraea (Mattuschka) Liebl.] and sycamore (Acer pseudoplatanus L.) leaf litters from a non-polluted and a heavily sulphur dioxide (SO2 )-polluted woodland were fumigated with environmentally-realistic concentrations (0.010-0.030 µl l-1 ) of SO2 for 16-68 wk in an open-air field-fumigation experiment. Fumigation markedly increased sulphate and protons in leachates from the litters and decreased calcium and magnesium contents of the leaves. However, there were few differences in the responses between leaf litters from the two woodlands. This was attributed to rapid sulphate wash-out from the litters from the heavily polluted woodland, so that the litters from the two sites quickly reached the same sulphate status.

Effects of dry-deposited sulphur dioxide on fungal decomposition of angiosperm tree leaf litter III. Decomposition rates and fungal respiration.

Ash (Fraxinus excelsior L.), birch (Betula spp.), hazel (Corylus avellana L.), sessile oak [Quercus petraea (Mattuschka) Liebl.] and sycamore (Acer pseudoplatanus L.) leaf litters from a virtually non-polluted and a heavily sulphur dioxide (SO2 )-polluted woodland were fumigated with environmentally-realistic concentrations (0.010-0.030µl l-1 ) of SO2 for 16-68 wk in an open-air field-fumigation experiment. Fumigation inhibited the respiration (CO2 evolution) and decomposition rates of the leaf litters. However, there were few differences in the responses between leaf litters from the two woodlands. In addition, pure cultures of four saprotrophic fungi were grown individually on irradiated hazel litter and exposed to c. 0.030µl l-1 of gaseous SO2 , for 28 d in the laboratory. The gas inhibited the respiration of Phoma exigua Desm. and Phoma macrostoma Mont, but not the respiration of Cladosporium cladosporioides (Fres.) de Vries or Coniothyrium quercinum Sacc. var. glandicola Grove. These results in part substantiated findings of previous experiments examining the effects of SO2 on the structures of saprotrophic fungal communities. The effects of SO2 on fungal decomposition of angiosperm tree leaf litter as possible causes of forest decline are discussed.

Effects of dry-deposited sulphur dioxide on fungal decomposition of angiosperm tree leaf litter I. Changes in communities of fungal saprotrophs.

Comparisons of the saprotrophic fungi isolated from ash (Fraxinus excelsior L.), birch (Betula spp.), hazel (Corylus avellana L.), pedunculate oak (Quercus robur L.), sessile oak [Q. petraea (Mattuschka) Liebl.] and sycamore (Acer pseudoplatanus L.) leaf litters from three woodlands exposed to low, medium and high levels (c. 0.0→ 0.060µl l-1 ) of sulphur dioxide (SO2 ) showed that the composition of the fungal communities differed between sites. Fumigation of angiosperm tree leaf litters from the least and the most polluted site with environmentally-realistic concentrations (0.010-0.030 µl1 ) of SO2 for 16-68 wk in an open-air field-fumigation experiment resulted in marked changes in the composition of the fungal communities in the leaf litters, comparable with differences found between the woodland sites. Cladosporium spp., Epicoccutn nigrum Link, Fitsarium spp. and Phoma exigua Desm. were less commonly isolated from leaf litters exposed to SO2 , whereas Coniothyrium quercinum Sacc. var. glandicola Grove, Cylindrocarpon orthosporum (Sacc.) Wollenw. and Penicillium spp. were more frequently isolated from fumigated litters. However, few differences could be detected in the response to SO2 of the mycofloras of leaf litters originating from different woodland sites. In general, SO2 did not affect the total extent of fungal occupancy of the microsites in the litter, as fungal species which decreased in abundance on exposure to the gas appeared to be replaced in the litter by other species more tolerant to the gas. SO2 therefore appeared to be selectively toxic to saprotrophic fungi isolated from these litters.