Mette Vestergård

Fifty thousand years of Arctic vegetation and megafaunal diet

Although it is generally agreed that the Arctic flora is among the youngest and least diverse on Earth, the processes that shaped it are poorly understood. Here we present 50 thousand years (kyr) of Arctic vegetation history, derived from the first large-scale ancient DNA metabarcoding study of circumpolar plant diversity. For this interval we also explore nematode diversity as a proxy for modelling vegetation cover and soil quality, and diets of herbivorous megafaunal mammals, many of which became extinct around 10 kyr bp (before present). For much of the period investigated, Arctic vegetation consisted of dry steppe-tundra dominated by forbs (non-graminoid herbaceous vascular plants). During the Last Glacial Maximum (25–15 kyr bp), diversity declined markedly, although forbs remained dominant. Much changed after 10 kyr bp, with the appearance of moist tundra dominated by woody plants and graminoids. Our analyses indicate that both graminoids and forbs would have featured in megafaunal diets. As such, our findings question the predominance of a Late Quaternary graminoid-dominated Arctic mammoth steppe.

Interactions Between Bacteria, Protozoa and Nematodes in Soil

Bacteria, protozoa and nematodes interact closely in soil ecosystems. Protozoa and nematodes eat bacteria (and occasionally each other), while bacteria defend themselves using chemical substances, resistant cell walls, irregular shapes and motility. Protozoa and nematodes are very different types of organisms, and hence apply very different feeding mechanisms; thus many protozoa can pick and choose individual bacterial cells, whereas nematodes ingest bacterial patches more uncritically. Protozoa and nematode are both aquatic organisms whose activity depends on available soil water, but differences in size, motility, resting stages and reproductive strategies mean that the soil physico-chemical environment influences the activity of protozoa and nematodes differently. For example, the relative importance of protozoa compared to nematodes may shift towards protozoa in very clay-rich soils. The interactions between the three organism groups have major ecological consequences such as modification of the bacterial communities and increased nitrogen mineralisation, both of which affect plant growth. Increased nitrogen mineralisation will usually be beneficial for plant growth, whereas the grazing induced changes in the bacterial communities can be both beneficial and detrimental to plants. Selective protozoan grazing can favour plant inhibiting bacteria. This may be a problem in clay rich soils where protozoa have better life conditions than nematodes.

Rhizosphere bacterial community composition responds to arbuscular mycorrhiza, but not to reductions in microbial activity induced by foliar cutting

Differences in bacterial community composition (BCC) between bulk and rhizosphere soil and between rhizospheres of different plant species are assumed to be strongly governed by quantitative and qualitative rhizodeposit differences. However, data on the relationship between rhizodeposit amounts and BCC are lacking. Other soil microorganisms, e.g. arbuscular mycorrhizal fungi (AMF), may also influence BCC. We simulated foliar herbivory (cutting) to reduce belowground carbon allocation and rhizodeposition of pea plants grown either with or without AMF. This reduced soil respiration, rhizosphere microbial biomass and bacteriovorous protozoan abundance, whereas none of these were affected by AMF. After labelling plants with (13)CO(2), root and rhizosphere soil (13)C enrichment of cut plants were reduced to a higher extent (24-46%) than shoot (13)C enrichment (10-24%). AMF did not affect (13)C enrichment. Despite these clear indications of reduced rhizosphere carbon-input, denaturing gradient gel electrophoresis (DGGE) of 16S rRNA genes PCR-amplified targeting DNA and RNA from rhizosphere soil did not reveal any effects of cutting on banding patterns. In contrast, AMF induced consistent differences in both DNA- and RNA-based DGGE profiles. These results show that a reduction in rhizosphere microbial activity is not necessarily accompanied by changes in BCC, whereas AMF presence inhibits proliferation of some bacterial taxa while stimulating others.

Trophic interactions between rhizosphere bacteria and bacterial feeders influenced by phosphate and aphids in barley

The aim was to study the effects of P fertilization and leaf aphid attack on the trophic interactions of bacteria and bacterial feeders in the rhizospheres of barley plants. The density of protozoa peaked in the rhizospheres of plants fertilized with N and P, whereas nematodes peaked in the rhizospheres of plants to which only N had been added. Fingerprinting of bacterial communities by length heterogeneity polymerase chain reaction revealed differences in community structure between NP rhizospheres and N rhizospheres as well as aphid-related differences within N rhizospheres. Specifically, α-proteobacteria increased with P addition. To evaluate if differences in bacteria in terms of their quality as food could partly explain the observed differences in protozoan and nematode abundances, growth of the flagellate Cercomonas sp. was assessed with 935 bacteria isolated from the different treatments. This assay indicated that bacterial isolates were of higher food quality to Cercomonas sp. in NP than in N rhizospheres when plants were subjected to aphid attack. Bacteria of high and low food quality for Cercomonas sp., respectively, were fed to the nematode Caenorhabditis elegans and larval production examined. α-Proteobacteria supported the growth of Cercomonas sp. well, whereas Actinobacteria did not. In contrast, C. elegans reproduced poorly on most α-proteobacteria but were able to reproduce well on some Actinobacteria. These results suggest that the different response of protozoa and nematodes to P addition could be mediated through a food quality-related change in community composition of bacteria and that leaf aphid attack may interfere with nutrient effects on bacterial assemblages of rhizospheres.

Above-belowground interactions govern the course and impact of biological invasions

We often consider invasive organisms as one of the most serious threats to global biodiversity, and we want to meet this threat. This, however, requires that we understand invasion in an evolutionary and ecological context; in particular, we must understand interactions between plants and their soil environment. Further, we must understand the temporal context of the invasion, which typically include an initial acute phase and a later chronic phase where equilibrium is reestablished. We claim that invasions fall into two broad categories. Some have irreversible effects, others, however, are reversible, where the exotic organism disappears or its negative impact decreases.

Nematode communities of natural and managed beech forests – a pilot survey

Summary The soil nematode communities of natural beech forests and managed beech forests were surveyed in order to examine which community parameters, if any, would be suited to differentiate between the two management regimes. Nematodes were collected from mineral soil at three sites, each including managed and adjacent natural beech forests. Following enumeration and identification of nematodes to family level, relative abundance of trophic groups, adult/juvenile ratio, Shannon-index, Plant Parasitic Index (PPI) and Maturity Index (MI) were determined. A clear separation of samples according to site was found in a Canonical Community Ordination (CANOCO) that related nematode data and various soil parameters in each sample. Beech forests of Mid-Zealand (Suserup) had significantly lower sand content, higher pH, higher PPI and higher nematode diversity according to the Shannon-index than the forests of North Zealand (Farum and Rankeskov). A distinct difference in the distribution of families was observed between sites, which could be governed by differences in texture and pH. The MI of the two old natural forest sites (Farum and Suserup) was significantly higher than the comparable managed sites, while this was not the case for Rankeskov, which is in a less mature state. There was a significantly higher adult/juvenile ratio and higher relative abundance of bacterial feeders in the natural forests compared to the managed forests. The apparent relation between pH and Maturity Index in beech forests is discussed. We suggest dead wood input to be the driving variable leading to the observed differences in the nematode community between managed and natural forests of Zealand, Denmark. The marked site differences found in this study emphasizes the need to carefully choose reference areas where soil conditions etc. are very similar to the managed forest in question when reference schemes for nature-based forestry are being developed.

Long-term and realistic global change manipulations had low impact on diversity of soil biota in temperate heathland

In a dry heathland ecosystem we manipulated temperature (warming), precipitation (drought) and atmospheric concentration of CO2 in a full-factorial experiment in order to investigate changes in below-ground biodiversity as a result of future climate change. We investigated the responses in community diversity of nematodes, enchytraeids, collembolans and oribatid mites at two and eight years of manipulations. We used a structural equation modelling (SEM) approach analyzing the three manipulations, soil moisture and temperature, and seven soil biological and chemical variables. The analysis revealed a persistent and positive effect of elevated CO2 on litter C:N ratio. After two years of treatment, the fungi to bacteria ratio was increased by warming, and the diversities within oribatid mites, collembolans and nematode groups were all affected by elevated CO2 mediated through increased litter C:N ratio. After eight years of treatment, however, the CO2-increased litter C:N ratio did not influence the diversity in any of the four fauna groups. The number of significant correlations between treatments, food source quality, and soil biota diversities was reduced from six to three after two and eight years, respectively. These results suggest a remarkable resilience within the soil biota against global climate change treatments in the long term.

Bioaccumulation of cadmium in soil organisms – With focus on wood ash application

Harvesting whole-tree biomass for biofuel combustion intensifies removal of nutrients from the ecosystem. This can be partly amended by applying ash from the combustion back to the system and thus recycle the nutrients. However, besides being rich in inorganic nutrients, ash also contains trace amounts of heavy metals. Due to the risk of toxic effects and trophic transfer of heavy metals, especially cadmium, legislation usually restricts the use of ash as a soil amendment. In order to provide researchers and governmental agencies with a tool to assess the risk of cadmium bioaccumulation in specific soil systems after ash application, we review: 1) the properties of ash; 2) the chemical and toxic properties of cadmium; 3) the key factors affecting cadmium bioavailability, cadmium uptake-, storage- and elimination-abilities in soil organisms and the risk of cadmium accumulation and biomagnification in the soil food web; 4) how ash impact on soil can change the risk of cadmium bioaccumulation. We conclude that for assessing the risk of cadmium bioaccumulation for specific sites, it is necessary to consider both the type and composition of ash, the soil conditions and organism composition on the site. On a general basis, we conclude that granulated ashes low in cadmium content, applied to low pH soils with high organic matter content, in systems with low abundances of earthworms, isopods and gastropods, will have a low risk of cadmium accumulation.