Psychro-tolerant nematophagous fungi from the maritime Antarctic

Summary The present investigation examines the comparative growth rates, at various temperatures between 4 and 30°C, of two nematophagous fungiMonacrosporium ellipsosporum (Preuss), (Grove), Cooke and Dickinson andM. cionapagum (Drechsler), (Subramanian), Cooke and Dickinson, both isolated from the Antarctic and from Britain. No psychrophilic species were found although the results clearly show that both the Antarctic isolates were psychro-tolerant, displaying lower minimum, optimum and maximum temperatures for growth than the British isolates. A modified form ofM. ellipsosporum isolated from the Antarctic grew only between 4 and 15°C, indicating it to be much better adapted to such cold habitats than the other isolates examined.     

The living strategy of nematophagous fungi

The infection structures, trophism, and ecological character of nematophagous fungi are reviewed in this article on the basis of data extracted from the literature and the most recent experiments conducted in this area. Traditionally, nematophagous fungi are classified into four groups according to their modes of attacking nematodes: nematode-trapping fungi using adhesive or mechanical hyphal traps, endoparasitic fungi using their spores, eggparasitic fungi invading nematode eggs or females with their hyphal tips, and toxin-producing fungi immobilizing nematodes before invasion. In the present review, we focus on the first two groups. The living strategies of these nematophagous fungi depend on the diversity of their infection structures, such as different traps and spore types, which determine the modes of infecting nematodes. The diversity of trophic modes of nematophagous fungi is an important prerequisite for fungal survival and activity in soil. The abundance and activity of Hirsutella rhossiliensis and H. minnesotensis, representatives of endoparasites and potential biocontrol agents against nematodes, are highly dependent on environmental factors. Comprehensive understanding of the survival and activity of nematophagous fungi in soil is fundamental for the exploitation of these fungi as successful biocontrol agents.     

Ecology of nematophagous fungi: vertical distribution in a deciduous woodland

Summary The distribution of nematophagous fungi in soil collected from a deciduous woodland is compared to various biotic and abiotic soil factors. The microfungi are isolated at all depths down to a maximum of 35 cm. Predators forming constricting rings, adhesive branches and adhesive knobs are restricted to the upper litter and humus layers. The net forming predators and endoparasites are isolated at all depths, although they are significantly more abundant in the lower mineral rich soils. A much greater species diversity of nematophagous fungi is recorded in the upper organic zones.Preliminary soil analysis indicates thatCephalosporium balanoides is independent of all soil variables, while predators able to form traps spontaneously are restricted to the organic soils which are rich in nematodes. Non-spontaneous trap forming predators, which are excellent saprophytes, are isolated from the deeper soils which are low in nutrients. The ecological significance of these results is discussed.     

The distribution,abundance and seasonality of pelagic marine invertebrate larvae in the maritime Antarctic

A survey of pelagic larvae was undertaken between November 1992 and February 1995 at Signy Island, Antarctica (60° 43 minutes S, 45° 36 minutes W). A diver-towed net and hand-held plankton pump were used at five sites of varying depths (6 to 30 m) and benthic substrata, in a combination of monthly and fortnightly samples. Overall larval ecological diversity was much higher than expected, with 131 morphologically distinct larval forms collected, apparently representing most of the benthic phyla present. The species richness observed is comparable with levels recorded at temperate latitudes, and higher than Arctic data and the implications for Thorson''s rule (the inferred cline of reduced pelagic larval diversity towards high latitudes) is discussed. Larval abundances were low (mean 2.6 individuals per cubic metre) which were two to six orders of magnitude lower than peaks in comparable data from temperate and tropical zones. We suggest that the low abundances recorded are a reflection of both slow developmental rates and a high dilution of larvae, reducing synchrony and spreading larvae over larger distances. Three seasonal periods, during which different larval types occur, have been identified. Summer, late summer and winter spawning strategies were discernable, and in some groups larvae occurred throughout the year.     

Nematophagous fungi from the maritime antarctic: factors affecting distribution

Soil collected from throughout the maritime Antarctic was analyzed to determine the effects of a range of abiotic and biotic soil parameters on the distribution of nematophagous fungi. Endoparasites were far more abundant than predatory fungi due to the greater efficiency of endoparasites in attracting and infecting nematodes, which resulted in a rapid completion of the infection cycle. This allowed endoparasites to colonise even the most exposed soil habitats where conditions favourable for nematode activity were restricted to only a few hours daily. 76% of all endoparasites isolated formed adhesive conidia. The success of this group of endoparasites was due to the ability of the conidia to attract nematodes.Spontaneous trap forming predators were far more abundant than nonspontaneous trap formers. The former have a clear competitive advantage over the latter, as they are able to attract nematodes to the conidia which subsequently germinate and form traps only when induced by direct chemical stimulation of nematodes. These predators are able to use internal reserves only and so rapidly respond to short periods when the soil is unfrozen and the nematodes are active.Although nematodes were utilized by all the species isolated, many were also capable of using Protozoa and Rotifera as a food source. While endoparasites were capable of infecting Rotifera, they were never observed infecting, or were isolated from soil containing, Protozoa. The distribution of endoparasites and spontaneous trap forming predators was largely independent of abiotic soil parameters. They were however significantly associated with the presence and abundance of suitable prey. These species survive solely on nematodes and therefore do not require prolonged periods of suitable soil conditions in order to produce vegetative mycelium to adsorb nutrients saprophytically from the soil, which is a significant ecological advantage over the other groups of microfungi found in the Antarctic.     

Diversity of endophytic fungi associated with bryophyte in the maritime Antarctic (King George Island)

Bryophytes comprise one of the richest microfungal microhabitats in the Antarctic environment. The maritime Antarctic is very vulnerable to rapid environmental change caused by global warming. The aim of this study was to investigate the importance of bryophytes as a microhabitat for fungal species in the maritime Antarctic by surveying endophytic fungal diversity from several bryophytes including Andreaea sp., Barbilophozia hatcheri, Chorisodontium aciphyllum, Polytrichum alpinum, Polytrichum strictum, Sanionia uncinata, and Warnstorfia sarmentosa. We collected 13 bryophyte samples at four localities on Barton Peninsula, King George Island. In total, 31 endophytic fungi morphotypes were isolated from bryophyte tissues by a thorough surface sterilization method. Using internal transcribed spacer sequence analysis, 16 endophytic fungal strains belonging to Ascomycota (12), Basidiomycota (1), Oomycota (1), and Zygomycota (2) phyla were obtained. Our results suggest the presence of a diverse range of fungal species even in a very limited area, and those bryophytes play an important role in conserving fungal diversity in this harsh environment. Growth rate measurements at a wide range of temperatures confirmed that most of the fungal strains were both mesophilic and psychrotolerant. This is the first report of endophytic fungi in Antarctic moss tissue by fluorescence in situ hybridization.     

Extracellular enzymes and the pathogenesis of nematophagous fungi

Nematophagous fungi are an important group of soil microorganisms that can suppress the populations of plant-parasitic nematodes. The pathogenic mechanisms of nematophagous fungi are diverse: They can be parasitical–mechanical through producing specialized capturing devices, or toxin-dependent. During infections, a variety of virulence factors may be involved against nematodes by nematophagous fungi. In this review, we present up-to-date information on the modes of infection by nematophagous fungi. The roles of extracellular hydrolytic enzymes and other virulence factors involved in infection against nematodes were summarized. The biochemical properties and peptide sequences of a special group of enzymes, the serine proteases, were compared, and their implications in infections were discussed. We also discussed the impact of emerging new techniques on our understanding of this unique group of fungi.     

Methane in maritime Antarctic freshwater lakes

Summary Methane was found to occur in all freshwater lakes, irrespective of trophic status, sampled during this preliminary investigation at Signy Island, South Orkney Islands, Antarctica. Methane accumulated in the water column of these lakes during the winter period when ice cover prevented wind-induced mixing. Maritime Antarctic lakes are usually subject to wind-induced complete mixing during the summer open-water period but two major exceptions to the rule were found during this study. Methanogenesis occurred in both littoral and profundal regions of oligotrophic Sombre Lake. The presence of a substantial algal mat stabilized the Eh status of underlying sediments at the littoral site. Methane production was confined to the sediments in both littoral and profundal sediments during the study period (December–March) but in winter probably migrated to the sediment surface at the profundal site. All Signy Island lakes sampled were sulphate-poor and addition of sulphate markedly inhibited methanogenesis. Radio-isotope studies indicated that the H₂/CO₂ pathway was probably the predominant route for methanogenesis in these sediments through the acetate pathway appeared equally important at the sediment surface. In the absence of sulphate, sulphate reducers probably acted as net hydrogen donors to the methanogens. The process rate was permanently limited by the consistent low temperature (annual range 1–3°C). Rates increased with increasing temperature over the range 4–32°C, but no evidence was found to suggest cold sensitivity or psychrophily. The optimum temperature for methanogenesis was in excess of 30°C, temperatures never experienced at Signy Island. Rates of methanogenesis during the study period (Dec–Mar) ranged from 0.29 to 0.45 mg of carbon m^-2 and on an annual basis methanogenesis was calculated equivalent to 13% of the organic carbon deposition rate.     

The comparison of isolation techniques for nematophagous fungi from soil

Using a single soil, a comparison was made of six well-known techniques for the isolation of nematophagous fungi. Each technique was tested for its ability to isolate the fungi present with regard to the number of species recovered, the number of plates needed to be 95% sure of isolating all the species possible and the amount of soil required. The advantages and disadvantages of each method are discussed. The results show that specific techniques are required for the isolation of endoparasites and predators respectively. The soil sprinkling technique and the Baermann funnel technique are shown to be the most efficient methods overall, with six and seven replicates required for 95% probability of isolating all the predator and endoparasite species respectively.     

Quantification of predatory and endoparasitic nematophagous fungi in soil

Methods were developed to quantify predatory and endoparasitic fungi in soil. The methods were based on previously developed detection techniques and combined with a most probable number estimation. The methods were applied to an agricultural soil fertilized with farmyard manure. Large amounts of farmyard manure resulted in increased amounts of organic matter, numbers of propagules of predatory and endoparasitic fungi, and numbers of bacteria and nematodes.     

An insect introduction to the maritime Antarctic

Although several invertebrates have been introduced by Man into the Antarctic, no holometabolous insects have survived to colonize terrestrial habitats successfully. Data are presented on the survival of populations of a chironomid midge, together with an enchytraeid worm, for 17 years in a maritime Antarctic site at Signy Island, South Orkney Islands. Both species are thought to have been introduced on plant material transplanted from either South Georgia or the Falkland Islands or both in 1967. Population densities average 25718m^-2 for the dipteran larvae and 3243 m^-2 for the worms. Successful completion of the midge's life cycle was indicated by emergence of brachypterous adults and oviposition (the population is parthenogenetic with only females present). Although both taxa are capable of supercooling to between -13 and -26°C, this capacity may not be sufficient in a severe winter to avoid lethal freezing. Four potential cryoprotectants were found in insect extracts, but in concentrations (< 1 % fresh weight) unlikely to influence cold hardiness. Both invertebrates appear to be pre-adapted for survival in much harsher conditions than they normally experience, by the extension of existing physiological mechanisms. It is concluded that the main limitations to invertebrate colonization of suitable Antarctic land areas by soil-dwelling species are geographical.     

The effect of replication on the probability of isolating nematophagous fungi from soil

The exact number of replicates required for a given probability of isolating all the species of nematophagous fungi present in a single sample of soil was measured. Using the soil sprinkling technique twenty replicate sub-samples of a sample of farm soil were examined for species. Eight replicates were required under these circumstances to be 95% sure of isolating all the species present. An equation to calculate the number of replicates to achieve a given probability of 100% species isolation is also given.     

Formation of abandoned penguin rookery ecosystems in the maritime Antarctic

Due to regional glacio-isostatic uplift of land during the Holocene period, a large number of penguin populations from several rookeries began to nest on the newly formed beaches and emerged rocks, leaving previously colonized higher cliffs. Chronology and extent of raised beaches, lichenometric age estimation of Usnea antarctica and radiocarbon-dating of guano and subfossil penguin bones in the abandoned nesting sites suggest the progress of the downward movement of many rookeries about 2500 years ago, accelerated during the last millennium, when wide bands of 4 to 6 m raised beaches were formed. Patterns of relic ornithogenic soil formation in the abandoned rookeries (focussed on leaching, erosion processes and humus horizon creation), as well as the succession of vegetation in the age sequence of abandoned nesting sites, were investigated in detail at Stranger Point, King George Island. The chemical composition of Deschampsia antarctica growing in the area of active, abandoned rookeries and in control sites provides direct and indirect evidence of current and past ornithogenic manuring on nutrition of vegetation. Received: 10 January 1996 / Accepted: 8 September 1996     

Epiphytic diatoms in two freshwater maritime Antarctic lakes

SUMMARY.

• 1 An ecological study of two small maritime Antarctic lakes on Signy Island, South Orkney Islands, was undertaken from January 1986 to March 1987. Analysis of diatom counts from the lakes provided examples of oligotrophic and mesotrophic ecosystems.
• 2 A diverse community of 104 epiphytic taxa was identified. Twenty-eight taxa had a percentage abundance greater than 1% in both lakes. Distinctive dominant taxa were identified from each lake.
• 3 A variety of ordination techniques was performed on the abundance data and a principal components analysis demonstrated differences in the diatom assemblages between the two lakes. Clear separations of sites and species were evident between the lakes, and accounted for the greatest percentage variance.
• 4 Species composition varied with depth within each lake and was also important in influencing changes in assemblage composition between sites.
• 5 A redundancy analysis indicated that species composition was correlated to concentrations of nitrogen and phosphorus but the total variance accounted for by the four physical and chemical factors measured was low (24%).

     

Modelling reproductive effort in sub-and maritime Antarctic mosses

A comparison is made of the reproductive effort (RE), considered as the investment in sporophyte relative to gametophyte biomass, of eight species of moss occurring at sub-and maritime Antarctic sites. Six of the species showed smaller sporophytes and game-tophytes at the climatically more extreme maritime Antaretic sites and one species showed no size difference between regions. The remaining species, although showing no regional difference, showed some evidence of a reverse pattern, with higher altitude samples having greater biomass than lower altitude samples. Spore counts indicated a measure of compensation in maritime Antarctic samples, with no significant decrease in spore output in several species despite smaller sporophyte biomass. The relationship between sporophyte (S) and gametophyte (G) biomass within samples was described by an allometric curve (S=aG ^b ) which gave a better fit than a straight line for six species. This form of model allows comparisons of patterns of RE to be made between samples with non-or partially overlapping size distributions, even when the relationship involves size-dependence. An allometric curve was not appropriate for describing samples of one species (Andreaea regularis), and insufficient data were available to identify any relationship in Polytrichum alpinum. The exponent (b) differed between species, but there were no statistically significant differences between exponents from samples of the same species. Samples of two species could further be described by the same coefficient (a), indicating that they lie on the same curve. However, samples of three species from sub-Antarctic South Georgia gave significantly higher coefficients, indicating increased RE relative to maritime Antarctic populations.     

Ovicidal effect of nematophagous fungi on Taenia taeniaeformis eggs

This work evaluated the ovicidal effect of the nematophagous fungi Monacrosporium sinense (SF53), Monacrosporium thaumasium (NF34) and Pochonia chlamydosporia (VC1) on Taenia taeniaeformis eggs in laboratory conditions. T. taeniaeformis eggs were plated on 2% water-agar with the grown isolates and control without fungus and examined at seven and fourteen days post-inoculation. At the end of the experiment, P. chlamydosporia showed ovicidal activity (P < 0.01) on T. taeniaeformis eggs unlike the other two species, mainly for internal egg colonization with percentage results of 32.2–54.0% at 7th and 14th day, respectively. The other fungi only showed lytic effect without morphological damage to eggshell. Results demonstrated that P. chlamydosporia was in vitro effective against Taenia taeniaeformis eggs unlike the other fungi. In this way, the use of P. chlamydosporia is suggested as a potential biological control agent for eggs of this cestode.