C. Van Dijk

Plant-specific soil-borne diseases contribute to succession in foredune vegetation

ECOLOGICAL study of the role of soil microorganisms in vegetation succession has focused mainly on organisms affecting plant nutrition, such as mycorrhiza and nitrogen-fixing bacteria. But, soil-borne diseases are involved in the degeneration of Ammophila arenaria (Marram grass) and Hippophaë rhamnoides (Sea buckthorn), two plant species that dominate the coastal foredunes of Europe and are widely planted for sand stabilization. We have used reciprocal transplantation and report here that soil-borne diseases may contribute to the succession of foredune plant species. In pot experiments, plant species that succeed A. arenaria were tolerant of the soil-borne diseases of this species. Plant species that were grown in soils from both previous and later succession stages were reduced most in soils from the later stages. During foredune succession, therefore, plants disappear from sites where the soil has become colonized with specific growth-depressing microorganisms. The soil-borne diseases must have considerable importance for the outcome of interspecific competition and may be involved in patterns of clonal growth. The different sensitivities of plant species for the soil-borne pathogens could be an evolutionary response to selection pressures of the succession stage to which a species is confined by the combined effect of local abiotic and biotic environmental factors.

Plant species diversity as a driver of early succession in abandoned fields: a multi-site approach

Abstract Succession is one of the most studied processes in ecology and succession theory provides strong predictability. However, few attempts have been made to influence the course of succession thereby testing the hypothesis that passing through one stage is essential before entering the next one. At each stage of succession ecosystem processes may be affected by the diversity of species present, but there is little empirical evidence showing that plant species diversity may affect succession. On ex-arable land, a major constraint of vegetation succession is the dominance of perennial early-successional (arable weed) species. Our aim was to change the initial vegetation succession by the direct sowing of later-successional plant species. The hypothesis was tested that a diverse plant species mixture would be more successful in weed suppression than species-poor mixtures. In order to provide a robust test including a wide range of environmental conditions and plant species, experiments were carried out at five sites across Europe. At each site, an identical experiment was set up, albeit that the plant species composition of the sown mixtures differed from site to site. Results of the 2-year study showed that diverse plant species mixtures were more effective at reducing the number of natural colonisers (mainly weeds from the seed bank) than the average low-diversity treatment. However, the effect of the low-diversity treatment depended on the composition of the species mixture. Thus, the effect of enhanced species diversity strongly depended on the species composition of the low-diversity treatments used for comparison. The effects of high-diversity plant species mixtures on weed suppression differed between sites. Low-productivity sites gave the weakest response to the diversity treatments. These differences among sites did not change the general pattern. The present results have implications for understanding biological invasions. It has been hypothesised that alien species are more likely to invade species-poor communities than communities with high diversity. However, our results show that the identity of the local species matters. This may explain, at least partly, controversial results of studies on the relation between local diversity and the probability of being invaded by aliens.

Biotic soil factors affecting the growth and development of Ammophila arenaria

SummaryTo study the origin of replant disease of Ammophila arenaria (L.) Link the growth and development in sand originating from the rhizosphere of a natural Ammophila vegetation was compared with the growth in sand from the sea-floor. In a greenhouse experiment, the growth of Ammophila seedlings in rhizosphere sand, when compared with that in sea sand, was significantly reduced. As sterilization by means of gamma-irradiation increased the biomass production of Ammophila seedlings significantly, it was concluded that the rhizosphere sand contained biotic factors that were harmful to Ammophila. In rhizosphere sand the roots of Ammophila were brown and poorly developed, and the specific uptake rates of N, P and K were reduced. The shoot weight proportion of the total plant dry matter was hardly influenced. In an outdoor experiment with Ammophila seedlings and cuttings, using both sands, the mortality was high and the plants were feeble in rhizosphere sand whereas plants in sea sand grew vigorously. It seems plausible that the plants in rhizophere sand were dessicated because the root system was shallow and badly developed. In the greenhouse experiments, Ammophila cuttings were less sensitive to the inhibiting factors in the rhizosphere than seedlings. This was confirmed in the outdoor experiment. Calammophila baltica (Fluegge ex Schrader) Brand, however, was hardly affected by the harmful biotic factors in the greenhouse. These results are discussed with reference to the ecology of Ammophila. It is assumed that the catching of fresh windblown sand provides Ammophila with a way to escape from harmful biotic soil factors, and it was concluded that degeneration of Ammophila is caused mainly by self-intolerance due to these biotic soil factors.

Analysis of nematodes and soil-borne fungi fromAmmophila arenaria (Marram grass) in Dutch coastal foredunes by multivariate techniques

A survey was carried out at nine locations in the Dutch coastal foredunes to identify the species of soil borne fungi and nematodes associated withAmmophila arenaria (Marram grass).Ammophila arenaria is a sand binding grass that is very important for the stabilization of coastal foredunes. Degeneration of the plants occurs at stabilized sites and is supposed to be caused by a combination of soil-borne fungi and nematodes. Canonical correspondence analysis (CCA) and two-way indicator species analysis (TWINSPAN) were used to examine which fungal and nematode species usually coexist in the rhizosphere of vigorous and early declining stands ofA. arenaria. In total, 47 species of fungi and 10 genera of plant-parasitic nematodes were found. According to CCA, the community of soil organisms of stands that were more than 10 years old was significantly different from recently established stands of 3 years old. Also, the community of soil organisms isolated from calcareous locations differed significantly from that of lime-poor locations. No relationship between the vigour of the plants (vigorous vs. early declining) and the soil borne species composition was found, although in roots of vigorous stands, the number of nematodes was higher than that of early declining stands. A relatively large group of soil organisms occurred generally. This group possibly contains an ubiquitous pathocomplex that cause the growth reducing effects of biotic origin which generally occur inA. arenaria. Analysis of this group of nematodes and fungi by TWINSPAN resulted in 9 different combinations of concurring soil organisms of which 5 combinations were present at all investigated locations. Two of the latter combinations contained both nematodes and fungi. The first contained three endoparasitic nematodes (Meloidogyne maritima, Heterodera spp. andPratylenchus sp.) that concurred with the fungusMucor hiemalis. The second group containedHeterodera spp.,Telotylenchus ventralis, Filenchus sp. together with the potentially plant-pathogenic fungiMicrodochium bolleyi and Fusarium culmorum, as well as the fungiMortierella sp. andTrichoderma harzianum, all in relatively high numbers.It is concluded that both CCA and TWINSPAN are valuable exploratory techniques, especially when used in combination, to detect possible combinations of soil organisms which may be involved in the degeneration ofA. arenaria. Further identifications of harmful organisms should be obtained from experiments.

Effect of vegetation manipulation of abandoned arable land on soil microbial properties

Abstract The effect of vegetation composition on various soil microbial properties in abandoned arable land was investigated 2 years after agricultural practice had terminated. Microbial numbers and processes were determined in five replicate plots of each of the following treatments: continued agricultural practice (monoculture of buckwheat in 1997), natural colonization by the pioneer community (arable weeds), and manipulated colonization from low (four species, three functional groups: grasses, forbs and legumes) or high diversity (15 species, three functional groups) seed mixtures from plant species that are characteristic of abandoned fields in later successional stages. The results indicated that differences in above-ground plant biomass, plant species composition and plant species diversity had no significant effect on soil microbial processes (net N mineralization, short-term nitrification, respiration and Arg ammonification), microbial biomass C and N (fumigation-incubation) or colony-forming units of the major microbial groups. Hence, there were no indications that soil microbial processes responded differently within 2 years of colonization of abandoned arable land by later successional plants as compared to that by plants from the natural pioneer weed community. Therefore, it seems that during the first few years after arable field abandonment, plants are more dependent on the prevailing soil microbiological conditions than vice versa.

Colonization of the root zone ofAmmophila arenaria by harmful soil organisms

The role of harmful soil organisms in the degeneration ofAmmophila arenaria at coastal foredunes was examined by the growing of seedlings ofA. arenaria in soil samples collected from its root zone. Three sites, each representing a successive stage in foredune succession were examined: (1) a highly mobile dune (sand accretion of 80 cm year−1) with vigorousA. arenaria, colonizing only the upper 30-cm of the annually deposited layer of sand, (2) a mobile dune with vigorousA. arenaria (sand accretion of 22 cm year−1) and a 1-metre soil profile completely colonized by roots and (3) a stable dune (no sand accretion) with degeneratedA. arenaria and young roots mainly present in the upper 0–10 cm.In the upper part of the highly mobile site, the presence of harmful soil organisms was confined to the root layers and at the mobile site for all depth layers a significant growth reduction ofA. arenaria was observed due to the activity of harmful soil organisms. At the stable site, however, growth had only been reduced in some of the depth layers. At all sites newly formed roots ofA. arenaria had been colonized by harmful soil organisms within one year.If present in sand prior to root growth harmful soil organisms reduced root length and root hair formation severely and they enhanced branching of the roots. It is concluded that harmful soil organisms initiate degeneration ofA. arenaria in stable dunes by attack of the root system, which makes the plants suffer from abiotic stress.

An ineffective strain type of Frankia in the soil of natural stands of Alnus glutinosa (L.) Gaertner

An ineffective strain type of Frankia of unknown strain composition, coded AgI-WD1 was discovered in the soil of wet dune slacks where A. glutinosa was the dominant tree species.Strain type AgI-WD1 was recognized by the development of slow growing root nodules on A. glutinosa testplants inoculated with soil suspensions. Microscopical examination of these nodules showed extremely reduced development of vesicles, normal development of intracellular clusters of hyphae and absence of sporangia. The stability of characteristics of this strain type such as the expression of root nodule symbiosis and ineffectivity of symbiontic N-fixation was demonstrated through ‘subculture’ of ineffective root nodules in successive hydrocultures of A. glutinosa. The nodulation process also differed from normal effective root nodules by the occurrence of resistance to strain type AgI-WD1 among part of the half-siblings of A. glutinosa used in the nodulation tests. Strain type AgI-WD1 was detected in the soil of different dune slacks which are inundated for a large part of the year and in a nearby peatbog covered with alder. The contribution of this strain type to soil populations of Frankia was demonstrated by nodulation potentials that were up to 500 times higher than that of the concurrent effective strain type AgSp-. The distribution of strain type AgI-WD1 appeared to be restricted to sites with water-logged soil conditions. Nodulation experiments pointed to potentials for competitive interactions between effective and ineffective strain thpes, especially to a density dependent reduction of nodule type AgI-WD1 by strain type AgSp-. The impact of competitive interactions is also affected by host trees that are resistant to AgI-WD1. The occurrence of resistance in the study areas was suggested by resistance among seedlings of a local seedbatch (±70% of the half-siblings) and by the absence of ineffective root nodules at site VD7-1, despite a high nodulation potential of the soil population of strain type AgI-WD1.

Effects of sand movement by wind on nematodes and soil-borne fungi in coastal foredunes

In stabilized dunesAmmophila arenaria (marram grass) degenerates due to a process involving soil-borne pathogens and parasites. This leads to exposure of the sand surface so that wind erosion may create blowouts.Ammophila rejuvenates on the edges of the blowouts, where the sand has accumulated. We tested the hypothesis that such rejuvenation of plants may be related to a reduction of the plant-parasitic nematodes and fungal propagules during the process of wind-driven transport.Field measurements in blowouts during storm events indicated that the drifted sand contained relatively low numbers of plant pathogenic fungi and plant-parasitic nematodes. A wind tunnel experiment showed that drifting sand may indeed reduce the numbers of fungi and nematodes. Although most fungi were attached to sand particles, they were also affected by the wind-borne sand movement. Sand that had been deposited by wind was made up of a larger proportion of large-sized particles. In our experiment the relatively small particles were lost during transport.Stirring the soil (part of the forces of natural winds) by mixing for 15 min. with a propeller mixer at 1500 rpm significantly reduced the number of nematodes and fungi. Both sand movement in the wind tunnel and intensive stirring of the sand enhanced the growth ofAmmophila test plants in a bioassay. It was concluded that in wind-blown sand the pathogen inoculum is reduced. Therefore, serious consideration should be given to allowing controlled reactivation of blow-outs to rejuvenate decliningAmmophila in stabilized foredunes.Implications for dune management are briefly discussed.

Effects of rhizosphere soil, vesicular-arbuscular mycorrhizal fungi and phosphate onPlantago major L. ssp.pleiosperma Pilger

Biotic factors in the rhizosphere and their effect on the growth ofPlantago major L. ssp.pleiosperma Pilger (Great plantain) were studied. In a pot experiment the effect on shoot growth of the addition of 2.5% rhizosphere soil at four levels of phosphate was highly dependent on the availability of phosphate: a promoting effect at low phosphate levels was observed while a reducing effect occurred at higher phosphate levels. As the roots were infected with vesicular-arbuscular mycorrhizal (VAM) fungi in the treatment with rhizosphere soil, two other experiments were set up to separate effects of the indigenous VAM fungi from effects of the total rhizosphere population. The uptake of phosphate and shoot growth was not decreased at higher phosphate availability when VAM inoculum was added alone or in combination with rhizosphere soil. The growth reducing effect of the rhizosphere soil could therefore not be ascribed only to mycorrhizal infection. The results suggest that biotic factors in the rhizosphere soil affect the phosphate uptake ofPlantago major ssp.pleiosperma. This may, under conditions of phosphate limitation, lead to an increase of phosphate stress and, subsequently, a growth reduction. Futhermore, it is concluded that VAM fungi, as part of the rhizosphere population, may compensate this phosphate stress by enhancing the phosphate uptake.