Food choice and reproductive success of Folsomia candida feeding on copper-contaminated mycelium of the soil fungus Alternaria alternata

We examined collembolan food preference for fungal mycelium grown on copper-contaminated medium, and the relationship between copper content, food selectivity and collembolan fitness when fed contaminated mycelium.To clarify whether collembolan food selectivity is related to fitness parameters, Folsomia candida were fed mycelium of the dark-pigmented fungus Alternaria alternata grown on medium with different copper concentrations. Copper-contaminated food (fungus grown on 50, 125, 250 and 500 μg Cu g^?1 medium, fresh wt.) was offered together with untreated food for 4 weeks. F. candida fed selectively on the provided mycelium and discriminated clearly between mycelium grown on high and low levels of contamination, distinctly preferring fungus grown on medium with a total copper concentration of 50 and 125 μg g^?1. In contrast, fungus grown on highly contaminated medium (250 and 500 μg g^?1) was avoided. Collembolan food preference generally matched fitness parameters. Reproduction was significantly affected by the total copper concentration of the fungal growth medium. When fed their preferred mycelium, collembolan reproduction was enhanced, whereas a diet of highly contaminated mycelium (250 or 500 μg g^?1) resulted in a strong decrease in reproduction. Adult survival was affected only marginally. Even though heavy metal contamination is a potential stress factor for many soil microarthropods, F. candida is able to discriminate between high and low quality food sources, and even benefits from moderately elevated copper concentrations.     

Recovery of reproduction after drought in the soil living Folsomia candida (Collembola)

Soil dwelling (euedaphic) Collembola have evolved to live in soil pores where the atmosphere is saturated with water vapour. Here we show that reproduction in the euedaphic Folsomia candida ceases during drought where soil water potentials are lower than −7 bar (∼99.4% RH). Recovery of the moulting cycle after cessation of drought showed increasing delay with increasing previous drought pressure. However, reproduction was rapidly resumed. Only in F. candida recovering from the most severe drought (−55 bar, 96% RH) was the time lapse between drought and resumed reproduction significantly different from the control (99.4% RH). Furthermore, the final numbers of eggs were unaffected by the degree of drought exposure. Hence, these results suggest that while even very mild drought will stop the reproductive cycle, subsequent recovery is rapid and complete.     

Defensive strategies of soil fungi to prevent grazing by Folsomia candida (Collembola)

Fungi represent a major part of the living biomass in the upper soil horizon and serve as an important food source for many soil organisms. We hypothesized that certain mycelial characteristics may serve to protect fungi from grazing. Specifically, this study focused on the influence of poisonous or other repellent metabolites and crystalline structures at the hyphal surface on the feeding preference of the soil microarthropod Folsomia candida Willem. The formation of crystalline structures was studied microscopically and the content of certain metabolites such as amanitin and muscarin was investigated using analytical methods. The feeding preference of F. candida was studied in different in-vitro food choice experiments. Additionally, the palatability of the fungal isolates was estimated by the amount of egg clusters laid by F. candida and by analysing the carbon and nitrogen content of the mycelia. F. candida was repelled by fungal species with toxic metabolites or crystals on their hyphal surface, which indicates that these traits serve as feeding protection. F. candida preferred dark-pigmented fungi. Total number of egg clusters and feeding preference were not correlated. However, insects that fed on fungi without repellent characteristics laid the most eggs. The amount of carbon and nitrogen in the mycelium had no influence on feeding behaviour. We conclude that the content of repellent metabolites and crystalline structures at the hyphal surface are defensive strategies of soil fungi and strongly influence feeding preference of F. candida. Other traits such as palatability were less important. Our results help to explain collembolan feeding behaviour and interactions between soil fungi and Collembola.     

The effect of pH and calcium on copper availability to the springtail Folsomia candida in simplified soil solutions

The effect of pH and calcium on copper bioavailability to the springtail Folsomia candida was determined by assessing uptake kinetics upon copper exposure for 15 days in simplified soil solutions. A slight bioaccumulation of copper was observed in all treatments and controls. The effect of exposure concentration and calcium level on copper accumulation was not significant. Although pH and time slightly but significantly affected copper concentrations in the body, copper uptake rates were not significantly different from zero. This suggests that copper uptake in F. candida exposed to sub-lethal copper concentrations in soil pore water is not affected by pH and calcium.     

Influence of agricultural soils on the growth and reproduction of the bio-indicator Folsomia candida

Soil organisms have been recommended as bio-indicators of soil quality due to their sensitivity to anthropogenic influences and their high degree of site-specificity. The objectives of this study were to determine if the relationship between the soil Collembola, Folsomia candida, and forest soil could be extended to other soils and, if not, to determine the relationship between its life parameters and soils from the different phases of an agricultural rotation sequence. A comparison of growth, reproduction and survival of 1 day old neonates and 10 day old juveniles subjected to a series of different soils (composted manure, pasture soil, forest soil, sand alone, and sand supplemented with yeast) established that the association between F. candida and forest soil quality cannot be automatically transferred to soils from other ecosystems. The test also showed that neonate (1 d) individuals were more sensitive to the different soil treatments than 10 d old juveniles and should be used in future tests. On this basis, neonate F. candida individuals were used to determine if the species could be employed to characterize the agricultural soils from the different phases of an organic or conventional rotation sequence. Results demonstrated that changes in body growth and reproduction constituted suitable criteria to characterize the soil quality of these different phases. The research required to develop these results as a standard bio-indicator test of agricultural soil quality is discussed.     

Assessment of movement patterns in Folsomia candida (Hexapoda: Collembola) in the presence of food

We showed that Folsomia candida (a blind soil-dwelling Collembola) was able to shift from non-directional (random or search strategy) to directional (target-oriented) movements at short distance of food. We measured departure from linearity and access (or not) to food by the springtail according to distance to the target position. Video-records and image analysis were used to obtain numerical data at 0.2 s interval. The probability of food capture within 10 min (maximum duration of the experiment) was negatively related to distance. Two patterns can be observed along successful trajectories in our experimental conditions (22 °C, ambient light, still air), non-directional movement being followed by directional movement when the animals approach food at 25 mm.     

Sensitivity to cadmium of the standard test species Folsomia candida compared to two other species, Onychiurus yodai and Sinella umesaoi (Collembola)

Knowing whether test species used for single-species toxicity tests are representative of other species provides valuable information because contaminant effects are often extrapolated to the community level based on these tests. To evaluate how representative the sensitivity of the standard test collembolan species Folsomia candida is for Collembola, we devised toxicity tests using the collembolan species Onychiurus yodai and Sinella umesaoi, which exhibit life-forms different from F. candida. Sensitivity to cadmium (Cd) was compared between F. candida and the novel test species. The 50% effective concentrations for reproduction were 154.7, 72.2 and 40.9 mg Cd/kg dry soil for O. yodai, F. candida and S. umesaoi, respectively. Thus, the reproductive sensitivity of F. candida to Cd was between that of the other two species. Increasing reproductive sensitivity to Cd corresponded to increasing activity levels and a more surface-dwelling life-form. Our data may facilitate the generalisation of effect data for F. candida to other species.     

Differences in the reproductive success and survival of the rufous treecreeper (Climacteris rufa) between a fragmented and unfragmented landscape

The rufous treecreeper (Climacteris rufa) has declined in abundance in the agricultural regions of southwestern Australia. The patterns of decline are well documented, but the processes that threaten population persistence are poorly understood. I compared the reproductive success and survival of the treecreeper between three sites in an unfragmented landscape and four remnant categories (large, small, grazed and ungrazed) in a fragmented, agricultural landscape. Nest success and annual productivity were significantly higher in the unfragmented landscape, but varied between sites and remnant categories within landscapes. Nest success was lowest in grazed remnants and annual productivity was positively associated with territory size in the fragmented landscape. Fledgling survival rates did not differ between landscapes, but there was a trend for juvenile survival rates to be higher in the unfragmented landscape. I used artificial nests to compare relative predation rates between landscapes, and provisioning rates and prey biomass brought to nestlings to assess differences in food availability. There were no landscape differences in predation rates, but provisioning rates to nestlings and total prey biomass were significantly lower in the fragmented landscape. Mean habitat quality was also lower in the fragmented landscape, although it differed between remnant categories. Reduced reproductive success, juvenile survival, food availability and habitat quality may threaten the viability of the rufous treecreeper population living in the fragmented landscape. Limiting the modification of remaining habitat (e.g. removing stock grazing) and improving habitat quality are required to assist in the conservation of this species.     

Kinetics of Zn and Cd accumulation in the isopod Porcellio scaber exposed to contaminated soil and/or food

To derive accumulation kinetics from different exposure matrices and account for the contribution of different exposure routes, the isopod Porcellio scaber was exposed to Cd and Zn, either in soil or in food, and a combination of both. Accurate uptake and elimination kinetics of Zn and Cd were determined using radioisotopes, allowing non-destructive time measurements. To describe the data, a simple kinetic model was used accounting for the internal distribution of the metals. A strong influence of the elimination kinetics, together with the metal fraction that is compartmentalized in a storage fraction, was found explaining the Cd steady-state level reached in the isopods. Zn turnover kinetics were relatively fast within the first days of exposure and elimination, followed by slow Zn kinetics from the storage compartment. The storage fraction appeared to be inert for elimination of both Zn and Cd within the experimental duration of 32 days and the fraction of metal taken up that was stored in this fraction was influenced by the uptake source. The quantity of metals taken up from soil or food depended on the concentration. For both Zn and Cd, the uptake rate constant from soil equalled the uptake rate constant from food, moreover, the uptake from both routes was shown to be additive. Therefore, it is concluded that the relative contribution of both routes (via soil or food) is explained by the partitioning of metals between soil and food.     

Effects of yeast on the growth and reproduction of the saprophagous millipedePolydesmus angustus (Diplopoda, Polydesmidae)

The millipedePolydesmus angustus was reared in the laboratory from hatching to maturity, reproduction and death. Two food types were used: dead leaves alone or dead leaves supplemented monthly with dry food yeast (Saccharomyces cerevisiae) at a rate not exceeding 5% of leaf dry weight. Growth, survival, adult live weight and fertility were compared between females reared on the two diets. Although the species was able to complete its life cycle on dead leaves alone, several parameters were strongly affected by the addition of yeast: growth was significantly faster, adult females became significantly larger and there was a 4.3-fold increase in fertility. Only survival was unaltered by the addition of yeast. The comparison between these laboratory results and field data on female fertility and live weight suggests that the natural diet of millipedes includes foods of higher quality than dead leaves. Possible sources of high-quality food in natural conditions are discussed.     

The role of floral biology and breeding system on the reproductive success of the narrow endemic Petrocoptis viscosa rothm. (Caryophyllaceae)

Morphological and functional characteristics of flowers may have major effects on their reproductive success. Here, we report a study on the characteristics of flowers of Petrocoptis viscosa, a herb species endemic to the northwest Iberian Peninsula, restricted to crevices in limestone outcrops, and currently occurring in only three populations within an area of less than 30 km². We also investigated the effects of pollen source on indicators of reproductive success. The results show that Petrocoptis viscosa flowers are well adapted to autogamy. Inbreeding depression (as determined by comparison of results obtained after selfing and outcrossing) was negligible for fruit production (δ=−0.05) and mean seed number (δ=−0.11), low for seed germination percentage (δ=0.08), but relatively high for mean seed weight (δ=0.23). The spatial structure and small size of population of Petrocoptis viscosa may mean that inbreeding is frequent in natural habitats. Adaptation of plants to autogamy may therefore be energetically beneficial (i.e. less wasted expenditure on rewards and flowers).     

Effects of Medicago sativa, Taraxacum officinale and Bromus sterilis on the density and diversity of Collembola in grassy arable fallows of different ages

The density and diversity of Collembola of nine grassy arable fallows of different ages were investigated in a factorial design with the factors ‘plant species’ (legume: Medicago sativa, herb: Taraxacum officinale, grass: Bromus sterilis) and ‘age class’ (2-3, 6-8 and 12-15 years) including the random effect ‘site’ (1-9). In May 2008, four plots were selected randomly at each fallow. Within each plot five M. sativa, T. officinale and B. sterilis plants were extracted with their associated soil using steel cylinders. The material from each plant species was used for extraction of Collembola and for determination of environmental parameters. Thus, the new aspect of the present study compared to other field studies investigating the relationships between plant and Collembolan communities is the focus on the “micro-scale”, investigating the Collembolan communities of the soil associated with single plants.We found that species richness and density of total and euedaphic Collembola were significantly higher in B. sterilis than in T. officinale samples with the M. sativa samples being intermediate. Fine-root feeding euedaphic Collembola particularly benefited from the higher amount of fine roots in B. sterilis samples. We also discovered that the age of the fallows had no significant influence on the number of Collembolan species and the density of the Collembolan groups. Notably, however, species of the epedaphic genera Lepidocyrtus and Sminthurinus were associated with the 12-15 year-old fallows and presumably benefited from the high number of plant species in the old fallows. Finally, canonical correspondence analysis (CCA) indicated that the soil organic matter content and the microbial biomass, both potential food sources for many Collembolan species, were important structuring forces for the Collembolan communities.     

An AM fungus and a PGPR intensify the adverse effects of salinity on the stability of rhizosphere soil aggregates of Lactuca sativa

A mesocosm experiment was conducted to examine the effect of an arbuscular mycorrhizal (AM) fungus (Glomus mosseae (Nicol & Gerd.) Gerd. & Trappe) and a plant growth-promoting rhizobacterium (PGPR) (Pseudomonas mendocina Palleroni), alone or in combination, on the structural stability of the rhizosphere soil of Lactuca sativa L. grown under two levels of salinity. The plants inoculated with P. mendocina had significantly greater shoot biomass than the control plants at both salinity levels, whereas the mycorrhizal inoculation was only effective in increasing shoot biomass at the moderate salinity level. The aggregate stability of soils inoculated with the PGPR and/or G. mosseae significantly decreased with increasing saline stress (about 29% lower than those of soils under non-saline conditions). Only the inoculated soils showed higher concentrations of sodium (Na) under severe saline stress. The severe salinity stress decreased the glomalin-related soil protein (GRSP) concentration, but the highest values of GRSP were recorded in the inoculated soils. Our findings suggest that the use of AM fungi and/or a PGPR for alleviating salinity stress in lettuce plants could be limited by their detrimental effect on soil structural stability.     

Soil inoculation with the biocontrol agent Clonostachys rosea and the mycorrhizal fungus Glomus intraradices results in mutual inhibition, plant growth promotion and alteration of soil microbial communities

Interactions between the biocontrol fungus Clonostachys rosea IK 726 and a tomato/Glomus intraradices BEG87 symbiosis were examined with and without wheat bran, which served as a food base for C. rosea. In soil without wheat bran amendment, inoculation with C. rosea increased plant growth and altered shoot nutrient content resulting in an increase and decrease in P and N content, respectively. Inoculation with G. intraradices had no effect on plant growth, but increased the shoot P content. Dual inoculation with G. intraradices and C. rosea followed the pattern of C. rosea in terms of plant growth and nutrient content. Wheat bran amendment resulted in marked plant growth depressions, which were counteracted by both inoculants and dual inoculation increased plant growth synergistically. Amendment with wheat bran increased the population density of C. rosea and reduced mycorrhizal fungus colonisation of roots. The inoculants were mutually inhibitory, which was shown by a reduction in root colonisation with G. intraradices in treatments with C. rosea and a reduction in colony-forming units (cfu) of C. rosea in treatments with G. intraradices, irrespective of wheat bran amendment. Moreover, both inoculants markedly influenced soil microbial communities examined with biomarker fatty acids. Inoculation with G. intraradices increased most groups of microorganisms irrespective of wheat bran amendment, whereas the influence of C. rosea on other soil microorganisms was affected by wheat bran amendment. Overall, inoculation with C. rosea increased and decreased most groups of microorganisms without and with wheat bran amendment, respectively. In conclusion, despite mutual inhibition between the two inoculants this interaction did not impair their observed plant growth promotion. Both inoculants also markedly influenced other soil microorganisms, which should be further studied in relation to their plant growth-promoting features.     

Effect of soil bacteria on the ability of polycyclic aromatic hydrocarbons (PAHs) removal by Trametes versicolor and Irpex lacteus from contaminated soil

The effect of bacteria represented by indigenous soil microflora or a mixture of soil bacteria Pseudomonas aeruginosa and Rhodococcus erythropolis on fungal growth, extracellular enzyme production and polycyclic aromatic hydrocarbons (PAHs) biodegradation efficiency in soil of white-rot fungi Trametes versicolor and Irpex lacteus was investigated. Both fungi were able to colonize soil. The growth yields measured by ergosterol were about two-fold in I. lacteus after 10 weeks. Laccase was produced in T. versicolor cultures in the presence or absence of bacteria but live bacteria reduced the laccase levels in soil about 5 times. Manganese-dependent peroxidase (MnP) was not detected in T. versicolor cultures. The amounts of MnP and laccase in I. lacteus cultures were not affected by the presence of bacteria. T. versicolor was more efficient in PAH removal for all PAHs tested although its capacity to colonize soil was lower. The removal rates of PAHs by T. versicolor in sterile soil were 1.5-fold, 5.8-fold and 1.8-fold for 2-3-ring, 4-ring and 5-6-ring PAHs, compared to I. lacteus, respectively. I. lacteus showed a low efficiency of removal of pyrene, benzo[a]anthracene and benzo[k]fluoranthene, compared to T. versicolor, whereas chrysene and benzo[b]fluoranthene were degraded by neither fungus. The main effect of the presence of the indigenous microflora or R. erythropolis and P. aeruginosa was a significant decrease of degradation of total PAHs by both T. versicolor and I. lacteus. Weak fungal/bacterial synergistic effects were observed in the case of removal of acenapthylene, benzo[a]pyrene, dibenzo[a,h]anthracene and benzo[g,h,i]perylene by I. lacteus and acenapthylene by T. versicolor. However, the bacterial effects were different in the two fungi. PAH abiotic losses represented 15 and 21% of the total PAHs after 5 and 10 weeks, respectively; naphthalene and acenaphthene were removed from the soil due to volatilization.