Urea hydrolysis and transformations in coastal dune sands and soil

Summary Urea hydrolysis was studied in samples taken from a coastal sand dune succession, from uncolonized sand; the rhizosphere ofAmmophila arenaria and soil from the mature dune. Comparisons were made with urea hydrolysis in a fertile loam soil. Urea was hydrolyzed in all sand and soil samples, with complete hydrolysis occurring after 6 and 3 weeks in the rhizosphere sand and dune soil compared with only 4 days in the fertile loam. A third of the added urea, however, was still present in the uncolonized sand samples 6 weeks after the beginning of the incubation period. Urea hydrolysis broadly correlated with urease activity.The liberated NH ₄ ⁺ was oxidized to NO ₃ ^– –N in all samples. Urea stimulated the release of N from native organic matter in the two soils, but not sands, due presumably to the low organic matter content of the latter. Nitrite accumulated in the dune sands and soil, but not in the fertile loam.Although N-Serve (Nitrapyrin) had no effect on urea hydrolysis in any of the treated samples, it inhibited the nitrification of released NH ₄ ⁺ –N. The relevance of these findings to the use of urea as a fertilizer to improve plant growth and dune stabilization is commented upon.     

Assay and properties of some sulphur enzymes in coastal sands

Summary Arylsulphohydrolase; cysteine desulphohydrase and thiosulphate cyanide sulphurtransferase were assayed in coastal sands and their properties determined. Low enzyme activity was detected in sands lacking vegetation, but much higher activities were detected in the rhizospheres of climax vegetation;Hippopha? rhamnoides andAmmophila sp. Properties of the enzymes were generally similar to those quoted for the enzymes in soils.     

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.     

Occurrence and infective potential of the endophyte ofHippophaë rhamnoides L. ssp.rhamnoides in coastal sand-dune areas

Summary The infective potential (IP) of nodule homogenates from field-grownHippophaë rhamnoides L. ssp.rhamnoides was determined by counting the number of nodules formed on test plants after inoculation with various dilutions of the homogenates. The IP was almost constant,i.e. 10⁵ to 10⁶ per gram of fresh nodule material. Methods to store nodule material without loss of IP were tested. The IP of air-dried nodule powders stored at 6°C hardly decreased during a period of more than a year.Data are presented on the IP of soil samples from sites representing various stages of dune formation. BeforeH. rhamnoides appeared, the IP was low: 1 to 36 nodules were formed on test plants per kg of soil. This low IP was due to low numbers of infective endophyte particles in these soils. During the succession of theH. rhamnoides scrub, the IP of the soil increased, due to the increase in the number of endophyte particles in the soil. Gradually, however, nodulation was limited by other environmental factors. The nature of these factors is discussed.     

Evaluation of powdered pine bark for control ofMeloidogyne arenaria andHeterodera glycines on soybean

Commercially avaiable pine bark nuggets (composted) and fresh pine bark were ground into powders and added at rates of 0 to 50 g kg^–1 to field soil infested withMeloidogyne arenaria andHeterodera glycines. The treated soil was maintained moist in the greenhouse for 2 weeks, sampled, and planted with Davis soybean (Glycine max.). Eight weeks after planting, numbers ofM. arenaria andH. glycines in soil decreased with increasing amounts of composted or fresh pine bark. No juveniles were present in soil treated with 5% pine bark. The number of galls and cysts g^–1 root decreased in proportion to the amount of pine bark added to soil. Gall and cyst formation was completely eliminated at the 5% rate. Numbers of saprophagous nematodes were highest in soil with 4–5% pine bark. The activity of several soil enzymes was correlated with the addition of both composted and fresh pine bark. Fresh pine bark powder caused an increase in soil enzyme activity compared to composted pine bark, but did not provide consistent control of gall and cyst formation.Journal Series No. 18-933579 Alabama Agricultural Experiment Station     

Leaching of nitrate and ammonium in heathland and forest ecosystems in Northwest Germany under the influence of enhanced nitrogen deposition

To study the impact of high atmospheric nitrogen deposition on the leaching of NO₃^– and NH₄⁺ beneath forest and heathland vegetation, investigations were carried out in adjacent forest and heathland ecosystems in Northwest Germany. The study area is subjected to high deposition of nitrogen ranging from 15.9 kg ha^–1 yr^–1 in bulk precipitation to 65.3 kg ha^–1 yr^–1 beneath a stand of Pinus sylvestris L. with NH₄–N accounting for 70–80% of the nitrogen deposited. Considerable leaching of nitrogen compounds from the upper horizons of the soil, mostly as nitrate, occurred at most of the forest sites and below a mixed stand of Calluna vulgaris (L.) Hull. and Erica tetralix, but was low in a Betula pubescens Ehrh. swamp forest as well as beneath Erica tetralix L. wet heath and heath dominated by Molinia caerulea(L.) Moench. Ground water concentrations of both NO₃–N and NH₄–N did not exceed 1 mg L^–1 at most of the sites investigated.     

Does salt stress constrain spatial distribution of dune building grasses Ammophila arenaria and Elytrichia juncea on the beach?

Rising sea levels threaten coastal safety by increasing the risk of flooding. Coastal dunes provide a natural form of coastal protection. Understanding drivers that constrain early development of dunes is necessary to assess whether dune development may keep pace with sea‐level rise. In this study, we explored to what extent salt stress experienced by dune building plant species constrains their spatial distribution at the Dutch sandy coast. We conducted a field transplantation experiment and a glasshouse experiment with two dune building grasses Ammophila arenaria and Elytrigia juncea. In the field, we measured salinity and monitored growth of transplanted grasses in four vegetation zones: (I) nonvegetated beach, (II) E. juncea occurring, (III) both species co‐occurring, and (IV) A. arenaria dominant. In the glasshouse, we subjected the two species to six soil salinity treatments, with and without salt spray. We monitored biomass, photosynthesis, leaf sodium, and nutrient concentrations over a growing season. The vegetation zones were weakly associated with summer soil salinity; zone I and II were significantly more saline than zones III and IV. Ammophila arenaria performed equally (zone II) or better (zones III, IV) than E. juncea, suggesting soil salinity did not limit species performance. Both species showed severe winter mortality. In the glasshouse, A. arenaria biomass decreased linearly with soil salinity, presumably as a result of osmotic stress. Elytrigia juncea showed a nonlinear response to soil salinity with an optimum at 0.75% soil salinity. Our findings suggest that soil salinity stress either takes place in winter, or that development of vegetated dunes is less sensitive to soil salinity than hitherto expected.     

Growth,ionic balance,proton excretion,and nitrate reductase activity in Alnus and Hippophaë supplied with different sources of nitrogen

Summary Pre-cultivated, nodulated and non-nodulated plants of black alder (Alnus glutinosa) and sea buckthorn (Hippophaë rhamnoides ssp.rhamnoides) were grown on different N sources, with and without acidity control. Dry matter yields were lowest when plants were supplied with only NO ₃ ^– and were much greater when NH ₄ ⁺ was supplied either alone or in combination with NO ₃ ^– as long as the external pH was controlled; the final yields of the N₂-fixing plants were relatively low, especially withH. rhamnoides. Without acidity control, yields were greatly reduced in the presence of NH ₄ ⁺ .Proton or hydroxyl-ion effluxes, calculated on the basis of plant analyses, agreed well with measured excretion values. Without pH adjustment, the total proton efflux into the external solution was greater inA. glutinosa than inH. rhamnoides.Both species, but particularlyA. glutinosa, displayed the highest nitrate reductase activity in the roots.     

Effects of plant species on phosphorus availability in a range of grassland soils

Vegetative conversion from grass to forest may influence soil nutrient dynamics and availability. A short-term (40 weeks) glasshouse experiment was carried out to investigate the impacts of ryegrass (Lolium perenne) and radiata pine (Pinus radiata) on soil phosphorus (P) availability in 15 grassland soils collected across New Zealand using ³³P isotopic exchange kinetics (IEK) and chemical extraction methods. Results from this study showed that radiata pine took up more P (4.5–33.5 mg P pot^–1) than ryegrass (1.1–15.6 mg pot^–1) from the soil except in the Temuka soil in which the level of available P (e.g., E _1min P_i, bicarbonate extractable P_i) was very high. Radiata pine tended to be better able to access different forms of soil P, compared with ryegrass. There were no significant differences in the level of water soluble P (Cp, intensity factor) between soils under ryegrass and radiata pine, but the levels of Cp were generally lower compared with original soils due to plant uptake. The growth of both ryegrass and radiata pine resulted in the redistribution of soil P from the slowly exchangeable P_i pool (E _{> 10m} P_i, reduced by 31.8% on the average) to the rapidly exchangeable P_i (E _1min-1d P_i, E _1d-10m P_i) pools in most soils. The values of R/r ₁ (the capacity factor) were also generally greater in most soils under radiata pine compared with ryegrass. Specific P mineralisation rates were significantly greater for soils under radiata pine (8.4–21.9%) compared with ryegrass (0.5–10.8%), indicating that the growth of radiata pine enhanced mineralisation of soil organic P. This may partly be ascribed to greater root phosphatase activity for radiata pine than for ryegrass. Plant species × soil type interactions for most soil variables measured indicate that the impacts of plant species on soil P dynamics was strongly influenced by soil properties.     

Soil H2-uptake in relation to soil properties and rhizobial H2-production

Summary The biological nature of soil H₂-consumption has been investigated. Soil microorganisms were capable to remove H₂ present in the gas phase at concentrations in the range of 200 ppm at rates varying between 0.2 and 1.0 l.min^–1. 100 g^–1. Free soil enzymes did not contribute significantly at the H₂ concentrations tested. Oxygen seemed to be the predominant electron acceptor. The influence of microbiological and physical soil properties on the H₂-uptake activity was examined for 38 soils.A highly significant correlation between biomass-C and H₂-uptake rate of the soil was noted, suggesting that the latter parameter might be useful as an indirect estimation of soil microbial biomass. The correlation was however not applicable for soils recently grown with legumes. Indeed, soya plants nodulated with aRhizobium strain with a weak hydrogen uptake capability, strongly increased the hydrogen oxidizing capability of the surrounding soil.     

Invasive plants and their escape from root herbivory: a worldwide comparison of the root-feeding nematode communities of the dune grass <Emphasis Type="Italic">Ammophila arenaria</Emphasis> in natural and introduced ranges

Invasive plants generally have fewer aboveground pathogens and viruses in their introduced range than in their natural range, and they also have fewer pathogens than do similar plant species native to the introduced range. However, although plant abundance is strongly controlled by root herbivores and soil pathogens, there is very little knowledge on how invasive plants escape from belowground enemies. We therefore investigated if the general pattern for aboveground pathogens also applies to root-feeding nematodes and used the natural foredune grass Ammophila arenariaas a model. In the late 1800s, the European A. arenariawas introduced into southeast Australia (Tasmania), New Zealand, South Africa, and the west coast of the USA to be used for sand stabilization. In most of these regions, it has become a threat to native vegetation, because its excessive capacity to stabilize wind-blown sand has changed the geomorphology of coastal dunes. In stable dunes of most introduced regions, A. arenaria is more abundant and persists longer than in stabilized dunes of the natural range. We collected soil and root samples and used additional literature data to quantify the taxon richness of root-feeding nematodes on A.␣arenaria in its natural range and collected samples from the four major regions where it has been introduced. In most introduced regions A. arenaria did not have fewer root-feeding nematode taxa than the average number in its natural range, and native plant species did not have more nematode taxa than the introduced species. However, in the introduced range native plants had more feeding-specialist nematode taxa than A. arenaria and major feeding specialists (the sedentary endoparasitic cyst and root knot nematodes) were not found on A. arenaria in the southern hemisphere. We conclude that invasiveness of A. arenaria correlates with escape from feeding specialist nematodes, so that the pattern of escape from root-feeding nematodes is more alike escape from aboveground insect herbivores than escape from aboveground pathogens and viruses. In the natural range of A. arenaria, the number of specialist-feeding nematode taxa declines towards the margins. Growth experiments are needed to determine the relationship between nematode taxon diversity, abundance, and invasiveness of A. arenaria.     

Etudein situ de l'influence de l'humidité et de la teneur en nitrate d'un sol dunaire sur l'accumulation et la réduction du nitrate chez l'oyat (Ammophila arenaria L.)

Résumé L'étude du fonctionnement du cycle de l'azote dans les milieux dunaires des c?tes méditerranéennes fran?aises a conduit à analyser le comportement de l'oyat,Ammophila arenaria, au cours de son développement en réponse aux variations de l'humidité et de la teneur en nitrate du sol. Les mesures d'activité nitrate réductase et de la teneur en nitrate dans les feuilles ont été effectuées simultanément avec les mesures des teneurs en eau et en nitrate du sol. Cette plante a des activités nitrate réductase ne dépassant pas 0,27 μmoles h⁻¹ g⁻¹ matière fra?che, qui n'ont pu être mesuréesin vivo qu'en présence de nitrate exogène dans le milieu d'incubation. Il appara?t que la plante réduit ou accumule préférentiellement le nitrate selon que l'on se situe en période humide ou sèche. Pour des valeurs voisines ou inférieures à 2% d'humidité dans les vingt premiers centimètres de sol, l'activité nitrate réductase décro?t et des quantités importantes de nitrate s'accumulent (169 μg N.NO₃.g⁻¹ matière sèche). Dans ces conditions, le nitrate dans la solution du sol peut atteindre une concentration élevée (6 meq.l⁻¹). Inversement, au-delà de 2% d'humidité le nitrate endogène s'épuise et l'activité nitrate réductase augmente. La régulation de l'assimilation et de l'accumulation du nitrate par l'oyat dépend de son alimentation hydrique et done des quantités de nitrate absorbées à partir de la solution du sol et véhiculées dans la sève brute.

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In situ study of the influence of dune soil humidity and nitrate contents on the nitrate accumulation and reduction in marram (Ammophila arenaria L.)

Summary The search for a better understanding of the nitrogen cycle in the sand dunes of french mediterranean beaches have driven us to study the habits ofAmmophila arenaria. In this way, we have followed its responses to different soil water potentials and nitrate contents of the soil during its development. Nitrate reductase activities and nitrate contents of leaves have been measured simultaneously with water and nitrate contents of soil. This plant has a slight nitrate reduction activity, not more than 0,27 μmoles h⁻¹ g⁻¹ fresh matter, which could be measuredin vivo only by the addition of nitrate to the incubation media. It seems that the plant reduces or else it stores, in a preferential way the nitrate whether it is in a humid or dry period. If humidity values are near or below 2%, the marram diminishes its assimilation activity and stores important amounts of nitrate (169 μg N−NO₃ ⁻.g⁻¹ dry matter). In these conditions the nitrate can be highly concentrated in the soil solutions (6 meq.l⁻¹). Opposite, above 2% of humidity, the nitrate reductase activity decreases and the endogenous nitrate is consumed. The regulation of the assimilation and storage of nitrate by the marram is mainly a function of the soil humidity and so, of the amount of nitrate taken from the soil solution and carried into the xylem.

     

Utilisation of soil organic P by agroforestry and crop species in the field,western Kenya

A field experiment in western Kenya assessed whether the agroforestry species Tithonia diversifolia (Hemsley) A. Gray, Tephrosia vogelii Hook f., Crotalaria grahamiana Wight & Arn. and Sesbania sesban (L) Merill. had access to forms of soil P unavailable to maize, and the consequences of this for sustainable management of biomass transfer. The species were grown in rows at high planting density to ensure the soil under rows was thoroughly permeated by roots. Soil samples taken from beneath rows were compared to controls, which included a bulk soil monolith enclosed by iron sheets within the tithonia plot, continuous maize, and bare fallow plots. Three separate plant biomass samples and soil samples were taken at 6-month intervals, over a period of 18 months. The agroforestry species produced mainly leaf biomass in the first 6 months but stem growth dominated thereafter. Consequently, litterfall was greatest early in the experiment (0–6 months) and declined with continued growth. Soil pH increased by up to 1 unit (from pH 4.85) and available P increased by up to 38% (1 g P g^–1) in agroforestry plots where biomass was conserved on the field. In contrast, in plots where biomass was removed, P availability decreased by up to 15%. Coincident with the declines in litterfall, pH decreased by up to 0.26 pH units, plant available P decreased by between 0.27 and 0.72 g g^–1 and P_o concentration decreased by between 8 and 35 g g^–1 in the agroforestry plots. Declines in P_o were related to phosphatase activity (R²=0.65, P<0.05), which was greater under agroforestry species (0.40–0.50 nmol MUB s^–1 g^–1) than maize (0.28 nmol MUB s^–1 g^–1) or the bare fallow (0.25 nmol MUB s^–1 g^–1). Management of tithonia for biomass transfer, decreased available soil P by 0.70 g g^–1 and P_o by 22.82 g g^–1. In this study, tithonia acquired P_o that was unavailable to maize. However, it is apparent that continuous cutting and removal of biomass would lead to rapid depletion of P stored in organic forms.     

Nodulation potential of soils from red alder stands covering a wide age range

Red alder (Alnus rubra Bong.) stands in the Pacific Northwest are the common first stage in succession following disturbance. These stands are highly productive and contribute a large amount of N to the soils as a result of their N₂-fixing symbiosis with Frankia. As these alder stands age, the soils not only increase in total N, but concentrations of NO^– ₃ increase and pH decreases as a result of nitrification. The objective of this study was to determine how the nodulation capacity of Frankia varies as red alder stands age and if differences in nodulation capacity are related to changes in soil properties. Nodulation capacity was determined by a red alder seedling bioassay for soils from red alder stands in the Oregon coast range covering a wide range of ages. Six chronosequences were sampled, each containing a young, an intermediate, and an older alder stand. Soil total N, total C, NO^– ₃, NH⁺ ₄, and pH were measured on the same soil samples. These factors as well as alder stand characteristics were compared with nodulation capacity in an attempt to identify soil characteristics typical in developing alder stands that most strongly affect nodulation capacity. Soil pH and NO^– ₃ concentration were highly correlated with nodulation capacity and with each other. Cluster analysis of the sites using these two variables identified two groups with distinctly different nodulation capacities. The cluster with the higher nodulation capacity was lower in NO^– ₃ and higher in pH than the other cluster, which included the majority of sites. There was substantial overlap in the age ranges for the two clusters and there was no significant correlation between age and nodulation capacity. Thus nodulation capacity appears to be most closely related to soil properties than to stand age.     

Biomass production by alders on four abandoned agricultural soils in Québec

The production of aboveground tissue of three alder species (Alnus crispa (Ait.) Pursh,A. rugosa (Du Roi) Spreng. andA. glutinosa (L) Gaertn.) on four sites ranged from 0.4 t ha^–1 yr^–1 to 4.0 t ha^–1 yr^–1 after four growing seasons. Large differences were observed among the four sites studied and among species. Soil nutrient levels affected the biomass production and foliar symptoms of P and Mg deficiency occurred withA. crispa andA. rugosa. Because of their poor aboveground biomass production (0.4–1.4 t ha^–1 yr^–1),A. crispa andA. rugosa should be used mainly as nurse trees. For its higher potential for biomass production (up to 4.0 t ha^–1 yr^–1), and its apparent higher ability to use P and Mg on deficient sites,A. glutinosa should be used preferably toA. crispa andA. rugosa for the production of biomass.     

Persistence ofBacillus thuringiensis andBacillus cereus in soil supplemented with grass or manure

Summary Persistance of inocula ofBacillus thuringiensis spores, parasporal crystals, andBacillus cereus spores in soil supplemented with dried-grass or partly composted, dried-chicken manure (100 mg supplement per 900 mg soil,^–0.01 MPa water availability, 25°C) were monitored over a period of up to 64 days by dilution plating and bioassay with larvae ofPieris brassicae. The inoculantB. thuringiensis population increased 22 x in level in grass-supplemented soil, but declined in manure-supplemented soil to 0.22 x the original level. TheB. cereus inocula declined in both soil treatments to approximately 0.1 x the original level. Insecticidal activity of theB. thuringiensis parasporal crystal decreased exponentially in grass and manuresupplemented soils, with half-lives of approximately 9.5 and 8.5 days respectively.     

Net nitrogen mineralization in soils under six indigenous tree species,an abandoned pasture and a secondary forest in the atlantic lowlands of costa rica

Nitrogen mineralization, nitrification potentials, pH, total N, C, extractable P and cations were measured in soils under 4-year-old, mono-specific stands of six fast-growing, native tree species, an abandoned pasture, and a 20-year-old secondary forest, as part of a study on the use of indigenous tree species for rehabilitation of soil fertility on degraded pastures at the La Selva Biological Station in the Atlantic humid lowlands of Costa Rica. Soil net nitrification potential rates were higher under two N-fixing, leguminous species,Stryphnodendron microstachyum Poepp. et Endl. (1.1–1.9 mg kg^–1 day^–1) andDalbergia tucurensis Donn. Smith (0.7–1.5 mg kg^–1 day^–1), than under the non-N-fixing trees in the plantation,Vochysia guatemalesis Don. Sm.,Vochysia ferruginea Mart,Dipteryx panamensis (Pittier) Record and Mell andHyeronima alchorneoides Fr. Allemao (0.2–0.8 mg kg^–1 day^–1). Values under the N-fixing trees were comparable to those found in secondary forest. There were no statistically significant differences in soil total N or in other nurtients between the species. Results of pH measurements done before and after incubation did not show any clear evidence of a pH drop attributable to nitrification.     

Biotic soil factors affecting the growth and development of Ammophila arenaria

Summary To 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.     

Effect of soil compaction on root growth and uptake of phosphorus

Summary Zea mays L. andLolium rigidum Gaud. were grown for 18 and 33 days respectively in pots containing three layers of soil each weighing 1 kg. The top and bottom layers were 100 mm deep and they had a bulk density of 1200 kg m^–3, while the central layer of soil was compacted to one of 12 bulk densities between 1200 and 1750 kg m^–3. The soil was labelled with³²P and³³P so that the contribution of the different layers of soil to the phosphorus content of the plant tops could be determined. Soil water potential was maintained between –20 and –100 kPa.Total dry weight of the plant tops and total root length were slightly affected by compaction of the soil, but root distribution was greatly altered. Compaction decreased root length in the compacted soil but increased root length in the overlying soil. Where bulk density was 1550 kg m^–3, root length in the compacted soil was about 0.5 of the maximum. At that density, the penetrometer resistance of the soil was 1.25 and 5.0 MPa and air porosity was 0.05 and 0.14 at water potentials of –20 and –100 kPa respectively, and daytime oxygen concentrations in the soil atmosphere at time of harvest were about 0.1 m³m^–3. Roots failed to grow completely through the compacted layer of soil at bulk densities 1550 kg m^–3. No differences were detected in the abilities of the two species to penetrate compacted soil.Ryegrass absorbed about twice as much phosphorus from uncompacted soil per unit length of root as did maize. Uptake of phosphorus from each layer of soil was related to the length of root in that layer, but differences in uptake between layers existed. Phosphorus uptake per unit length of root was higher from compacted than from uncompacted soil, particularly in the case of ryegrass at bulk densities of 1300–1500 kg m^–3.     

Soil microorganisms control plant ectoparasitic nematodes in natural coastal foredunes

Belowground herbivores can exert important controls on the composition of natural plant communities. Until now, relatively few studies have investigated which factors may control the abundance of belowground herbivores. In Dutch coastal foredunes, the root-feeding nematode Tylenchorhynchus ventralis is capable of reducing the performance of the dominant grass Ammophila arenaria (Marram grass). However, field surveys show that populations of this nematode usually are controlled to nondamaging densities, but the control mechanism is unknown. In the present study, we first established that T. ventralis populations are top-down controlled by soil biota. Then, selective removal of soil fauna suggested that soil microorganisms play an important role in controlling T. ventralis. This result was confirmed by an experiment where selective inoculation of microarthropods, nematodes and microbes together with T. ventralis into sterilized dune soil resulted in nematode control when microbes were present. Adding nematodes had some effect, whereas microarthropods did not have a significant effect on T. ventralis. Our results have important implications for the appreciation of herbivore controls in natural soils. Soil food web models assume that herbivorous nematodes are controlled by predaceous invertebrates, whereas many biological control studies focus on managing nematode abundance by soil microorganisms. We propose that soil microorganisms play a more important role than do carnivorous soil invertebrates in the top-down control of herbivorous ectoparasitic nematodes in natural ecosystems. This is opposite to many studies on factors controlling root-feeding insects, which are supposed to be controlled by carnivorous invertebrates, parasitoids, or entomopathogenic nematodes. Our conclusion is that the ectoparasitic nematode T. ventralis is potentially able to limit productivity of the dune grass A. arenaria but that soil organisms, mostly microorganisms, usually prevent the development of growth-reducing population densities.