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The role of soil fauna in crucial ecosystem services such as nutrient cycling remains poorly quantified, mainly because of the overly reductionistic approach adopted in most experimental studies. Given that increasing nitrogen inputs in various ecosystems influence the structure and functioning of soil microbes and the activity of fauna, we aimed to quantify the role of the entire soil nematode community in nutrient mineralization in an experimental set-up emulating nutrient-rich field conditions and accounting for crucial interactions amongst the soil microbial communities and plants. To this end, we reconstructed a complex soil foodweb in mesocosms that comprised largely undisturbed native microflora and the entire nematode community added into defaunated soil, planted with Lolium perenne as a model plant, and amended with fresh grass-clover residues. We determined N and P availability and plant uptake, plant biomass and abundance and structure of the microbial and nematode communities during a three-month incubation. The presence of nematodes significantly increased plant biomass production (+9%), net N (+25%) and net P (+23%) availability compared to their absence, demonstrating that nematodes link below- and above-ground processes, primarily through increasing nutrient availability. The experimental set-up presented allows to realistically quantify the crucial ecosystem services provided by the soil biota.

Root-feeding nematodes are major soil-borne pests in agriculture. In natural ecosystems, their abundance can be strongly controlled by natural enemies. In coastal foredune soil, the abundance of the ectoparasitic nematode Tylenchorhynchus ventralis is controlled by local interactions with soil microorganisms. If not controlled, T. ventralis reduces growth and performance of the host plant Ammophila arenaria. In the present study, we examine if the nematodes may sense the presence of soil microorganisms and, if so, they are able to actively avoid their enemies. First, using Petri dishes with agar medium we examined if T. ventralis can choose between A. arenaria seedlings inoculated with or without soil microorganisms. We observed that there was a trend (although non-significant) in nematode migration towards the non-inoculated plants. If the seedlings were not present, the nematodes did not make any choice and stayed in the centre of the Petri dish. Then, using Y-tubes filled with sterilized dune soil, we examined if T. ventralis could choose between A. arenaria roots with or without microorganisms. We also included treatments of microbial suspensions without plants and a microbe-free filtrate. We observed that the nematodes preferred roots without microorganisms. Microorganisms alone or roots with microbial filtrate did not influence nematode choice significantly. We conclude that the nematode T. ventralis is able to choose roots without soil microorganisms when having roots with them as alternative. Such avoidance could explain why biological control of nematodes in field is not always effective, especially when microbial antagonists accumulate in specific parts of the rhizosphere.

Root-feeding nematodes may play an important role in generating spatial and temporal variation in natural plant communities, but little is known about the performance of the nematodes in the plant root zone. We studied the emergence, survival and reproductive success of the cyst nematode Heterodera arenaria , a root feeding-specialist that occurs in mobile dunes. The host plants of H. arenaria are buried regularly by windblown sand to which the plants respond by upward clonal expansion. As a consequence, the nematodes have to migrate upwards in the soil profile to find new roots for feeding and reproduction, however, not all juveniles migrate. We tested the hypothesis that the offspring from migrated individuals would perform better than from individuals that remained behind and discuss the advantage of this dualistic behavior. The individual performance of the cyst nematodes was better when their juveniles migrated to the new root layer. In the field, in the new root layer the cysts had more eggs and juveniles than cysts collected from the 1-year-old root layer. Under controlled conditions, cysts from the new root layer released their first juveniles faster than cysts from the 1-year-old root layer. However, the juveniles that do not migrate might be crucial for the persistence of the population. In the past decade in two extremely dry years, we could not find cysts in the new root layer. Apparently, summer drought constrains cyst formation, so that the cysts produced in the older root layers serve as a reservoir for the population. Such reservoirs could become more important when climate change will result in increased incidence of summer droughts.

In coastal foredunes, the cyst nematode Heterodera arenaria has been supposed to play a role in degeneration of the pioneer grass Ammophila arenaria (marram grass). However, recent field surveys and field inoculation experiments suggested that the abundance of this cyst nematode is controlled by the host plant. Here, we test pathogenicity of H. arenaria by inoculating a range of densities of second-stage juveniles (J2) onto A. arenaria in sterilised soil. Plant root biomass was reduced after 6 weeks of growth, but not after 13 weeks, when cyst formation was completed. Shoot biomass was not influenced. In the field, H. arenaria occurred in sand and on roots of Elymus farctus and A. arenaria in mobile dunes, but not on A. arenaria in stable dunes. Although we observed Heterodera cysts and juveniles in later succession stages, this cyst nematode appeared to be H. hordecalis. In a host suitability test, H. arenaria produced cysts on E. farctus and A. arenaria, while there were almost no cysts produced when J2 had been added to the later succession species Festuca rubra ssp. arenaria, Carex arenaria, Elymus athericus and Calamagrostis epigejos. Heterodera arenaria did not negatively affect its natural host plants. However, H. arenaria reduced root growth of one later succession plant species, E. athericus, which was a poor host for H. arenaria. The effects of H. arenaria inoculation on plant biomass production did not differ when J2 were added to plants growing in non-sterilised soil. We conclude that H. arenaria is specialised on the pioneer grasses E. farctus and A. arenaria, and also on mobile dunes as a habitat. Our results confirm recent suggestions that H. arenaria may cause little direct growth reduction to its natural host plants. [KEYWORDS: AMMOPHILA ARENARIA ; BIOTROPHIC PARASITE ; COASTAL FOREDUNE ; HETERODERA HORDECALIS ; HOST SPECIFICITY]

peer-reviewed Plants rely on microorganisms to mobilize organically and inorganically bound sulfur (S) and phosphorus (P) in which the plant can then readily utilize. The aim of this study was to investigate the role of S- and P-mobilizing bacteria in plant growth promotion in biochar-amended soil, which has been rarely investigated so far. Pot experiments of Lolium perenne were established on S and P limited soil with 1% or 2% biochar (Miscanthusxgiganteus) or without biochar (control) for a period of 126days. Both biochar amendments resulted in significant plant growth promotion. Rhizobacteria capable of growing with (1) S from aromatic sulfonates, (2) P from phosphate esters, (3) P from phosphonates, and (4) P from tri-calcium phosphates as sole source of S or P, respectively, were significantly more abundant in the biochar treatments. 16S rRNA gene-based rhizobacteria community analysis revealed a significant biochar treatment effect. Abundance of nematodes feeding on bacteria was also significantly increased in the biochar treatments. Diversity analysis of rhizospheric asfA and phnJ genes revealed broad sequence diversities in bacterial sulfonate and phosphonate-mineralizing capabilities. These findings suggest that biochar amendment enhances microbially mediated nutrient mobilization of S and P resulting in improved plant growth. ACCEPTED peer-reviewed

Several studies have suggested that plants are able to control the development of specialist herbivorous invertebrates, but not that of generalists. Plants are alleged to have evolved tolerance against specialists in order to suppress the development of more damaging generalists through competition. Here, we tested whether specialist plant parasitic nematodes in the root zone of the natural dune grass Ammophila arenaria are able to suppress the development of a generalist plant parasitic nematode and therewith protect the plant. We added a generalist ectoparasite (Tylenchorhynchus ventralis) and specialist endoparasites (Heterodera arenaria, Pratylenchus penetrans, Meloidogyne maritima) in different densities to A. arenaria. We also tested whether sequential inoculation of the specialists had an additional competitive effect on T. ventralis. Our results show that the specialist endoparasitic nematodes indeed suppressed the development of the generalist T. ventralis, but only when the specialists were added to the plant in relatively high densities that exceeded the field density of the specialist endoparasitic nematodes. Therefore, we conclude that competition by specialist nematodes is not a likely mechanism for the regulation of the generalist plant parasite T. ventralis. Sequential inoculation of endoparasites did not influence the development of T. ventralis more than inoculation at the same time. Despite their inhibiting effect on the development of T. ventralis, the endoparasites did not counteract the negative effect of T. ventralis on plant biomass. On the contrary, they themselves had a negative effect on shoot biomass of A. arenaria, although no effect was found on root biomass. We discuss our results in relation to other mechanisms that may regulate the population density of T. ventralis. [KEYWORDS: Competition; Generalist; Plant parasitic nematode; Population regulation; Specialist; Temporal nematode dynamics]