N capture by Plantago lanceolata and Brassica napus from organic material: the influence of spatial dispersion, plant competition and an arbuscular mycorrhizal fungus

This study investigated N capture by Plantago lanceolata L. and Brassica napus L. from complex organic material (dual-labelled with 15N/13C) added either as a thin concentrated layer (discrete patch treatment) or dispersed uniformly with the background sand:soil mix in a 10 cm band (dispersed treatment) when grown in monoculture or in interspecific competition and in the presence or absence of a mycorrhizal inoculum (Glomus mosseae). No 13C enrichments from the organic material were detected in the plant tissues, but 15N enrichments were present. Total plant uptake of N from the organic material on a microcosm basis was not affected by the spatial placement of the organic material, but Plantago monocultures captured less N than the species in interspecific competition (i.e. 23% versus 38% of the N originally added). N capture from Brassica monocultures was no different to either Plantago monocultures or both species in mixture. However, N capture from the organic material by both individual Plantago and Brassica plants was reduced when grown with Brassica plants (by 10-fold and by more than half, respectively). N capture from the organic material was directly related to the estimated root length produced in the sections containing the organic material: the individual that produced the greatest root length captured most N. Strikingly, when the organic material was added as a discrete patch the N captured by Brassica, a non-mycorrhizal species, actually increased when the G. mosseae inoculum was present compared to when G. mosseae was absent (i.e. 35% versus 19% of the N originally added).     

Effect of nitrogen addition,form and clipping on competitive interactions between grassland species

Aims Although the effects of N addition on plant biomass are well understood, we know a lot less about the importance of N form even though some studies have shown different impacts from reduced and oxidized forms of N. Furthermore, responses to grazing are likely to interact with the response to N addition. This experiment investigates the interactive effects of N addition and form with clipping on competition between three grassland species.Methods The three species (Anthoxanthum odoratum L., Plantago lanceolata L. and Prunella vulgaris L.) were grown alone and in combination with factorial additions of deionized water, sodium nitrate and ammonium chloride, and a clipping treatment. Above- and belowground biomass was harvested after 4 months.Important findings In monocultures, the results show increases in biomass with N addition, but clipping resulted in fewer changes with species displaying varying degrees of growth compensation. A. odoratum was the strongest competitor when grown with other species. In monocultures without clipping, N form was not important, but in the presence of clipping and in different species combinations, N form became important. Significant two- and three-way interactive effects were observed showing that complex interactions exist between N addition, clipping and species identity. The results have important implications when considering the effects of N deposition.     

Nutrient availability and atmospheric CO2 partial pressure modulate the effects of nutrient heterogeneity on the size structure of populations in grassland species

BACKGROUND AND AIMS: Size-asymmetric competition occurs when larger plants have a disproportionate advantage in competition with smaller plants. It has been hypothesized that nutrient heterogeneity may promote it. Experiments testing this hypothesis are inconclusive, and in most cases have evaluated the effects of nutrient heterogeneity separately from other environmental factors. The aim of this study was to test, using populations of Lolium perenne, Plantago lanceolata and Holcus lanatus, two hypotheses: (a) nutrient heterogeneity promotes size-asymmetric competition; and (b) nutrient heterogeneity interacts with both atmospheric CO2 partial pressure (P(CO2)) and nutrient availability to determine the magnitude of this response. METHODS: Microcosms consisting of monocultures of the three species were grown for 90 d in a factorial experiment with the following treatments: P(CO2) (37.5 and 70 Pa) and nutrient availability (NA; 40 and 120 mg of N added as organic material) combined with different spatial distribution of the organic material (NH; homogeneous and heterogeneous). Differences in the size of individual plants within populations (size inequality) were quantified using the coefficient of variation of individual above-ground biomass and the combined biomass of the two largest individuals in each microcosm. Increases in size inequality were associated with size-asymmetric competition. KEY RESULTS: Size inequality increased when the nutrients were heterogeneously supplied in the three species. The effects of NH on this response were more pronounced under high nutrient supply in both Plantago and Holcus (significant NA x NH interactions) and under elevated P(CO2) in Plantago (significant P(CO2) x NA x NH interaction). No significant two- and three-way interactions were found for Lolium. CONCLUSIONS: Our first hypothesis was supported by our results, as nutrient heterogeneity promoted size-asymmetric competition in the three species evaluated. Nutrient supply and P(CO2) modified the magnitude of this effect in Plantago and Holcus, but not in Lolium. Thus, our second hypothesis was partially supported.     

Variation in mycorrhizal growth response influences competitive interactions and mechanisms of plant species coexistence

Plant species vary in their growth response to arbuscular mycorrhizal (AM) fungi, with responses ranging from negative to positive. Differences in response to AM fungi may affect competition between plant species, influencing their ability to coexist. We hypothesized that positively responding species, whose growth is stimulated by AM fungi, will experience stronger intraspecific competition and weaker interspecific competition in soil containing AM fungi, while neutrally or negatively responding species should experience weaker intraspecific and stronger interspecific competition. We grew Plantago lanceolata, which responds positively to AM fungi, and Bromus inermis, which responds negatively to AM fungi, in an additive response surface competition experiment that varied the total density and relative frequency of each species. Plants were grown in sterilized background soil that had been inoculated with whole soil biota, which includes AM fungi, or a microbial wash, that contained other soil microbes but no AM fungi, or in sterilized soil that contained no biota. The positively responding P. lanceolata was more strongly limited by intraspecific than interspecific competition when AM fungi were present. By contrast, the presence of AM fungi decreased the strength of intraspecific competition experienced by the negatively responding B. inermis. Because AM fungi are almost always present in soil, strong intraspecific competition in positively responding species would prevent them from outcompeting species that respond neutrally or negatively to AM fungi. The potential for increased intraspecific competition to offset growth benefits of AM fungi could, therefore, be a stabilizing mechanism that promotes coexistence among plant species.     

The Dynamic Process of Interspecific Interactions of Competitive Nitrogen Capture between Intercropped Wheat (Triticum aestivum L.) and Faba Bean (Vicia faba L.)

Wheat (Triticum aestivum L.)/faba bean (Vicia faba L.) intercropping shows significant overyielding and high nitrogen (N)-use efficiency, but the dynamics of plant interactions have rarely been estimated. The objective of the present study was to investigate the temporal dynamics of competitive N acquisition between intercropped wheat and faba bean with the logistic model. Wheat and faba bean were grown together or alone with limited N supply in pots. Data of shoot and root biomass and N content measured from 14 samplings were fitted to logistic models to determine instantaneous rates of growth and N uptake. The superiority of instantaneous biomass production and N uptake shifted from faba bean to wheat with their growth. Moreover, the shift of superiority on N uptake occurred 7–12 days earlier than that of biomass production. Interspecific competition stimulated intercropped wheat to have a much earlier and stronger superiority on instantaneous N uptake compared with isolated wheat. The modeling methodology characterized the temporal dynamics of biomass production and N uptake of intercropped wheat and faba bean in different planting systems, which helps to understand the underlying process of plant interaction for intercropping plants.     

Condition-specific competition allows coexistence of competitively superior exotic oysters with native oysters

1. Trade-offs between competitive ability and tolerance of abiotic stress are widespread in the literature. Thus, condition-specific competition may explain spatial variability in the success of some biological invaders and why, in environments where there is small-scale environmental variability, competitively inferior and superior species can coexist. 2. We tested the hypothesis that differences in abiotic stress alter the outcome of competitive interactions between the native Sydney rock oysters Saccostrea glomerata and exotic Pacific oysters Crassostrea gigas by experimentally testing patterns of intra- and interspecific competition across a tidal elevation gradient of abiotic stress at three sites on the east coast of Australia. 3. At low and mid-intertidal heights, exotic C. gigas were able to rapidly overgrow and smother native S. glomerata, which grew at c. 60% of the exotic's rate. In high intertidal areas, where C. gigas displayed about 80% mortality but similar growth rates to S. glomerata, the native oyster was not affected by the presence of the exotic species. 4. Asymmetrical effects of the exotic species on the native could not be replicated by manipulating densities of conspecifics, confirming that effects at low and mid-intertidal heights were due to interspecific competition. 5. Our results suggest that the more rapid growth of C. gigas than S. glomerata comes at the cost of higher mortality under conditions of abiotic stress. Thus, although C. gigas may rapidly overgrow S. glomerata at low and mid tidal heights, the native oyster will not be competitively excluded by the exotic due to release from competition at high intertidal elevations. 6. The success of trade-offs in explaining spatial variation in the outcome of competitive interactions between C. gigas and S. glomerata strengthen the claim that these may be a useful tool in the quest to produce general predictive models of invasion success.     

Do endogeic earthworms change plant competition? A microcosm study

Plants compete for limited resources. Although nutrient availability for plants is affected by resource distribution and soil organisms, surprisingly few studies investigate their combined effects on plant growth and competition. Effects of endogeic earthworms (Aporrectodea jassyensis), root-knot nematodes (Meloidogyne incognita) and the spatial distribution of ¹⁵N labelled grass litter on the competition between a grass (Lolium perenne), a forb (Plantago lanceolata) and a legume (Trifolium repens) were investigated in the greenhouse. Earthworms promoted N uptake and growth of L. perenne. Contrastingly, shoot biomass and N uptake of T. repens decreased in the presence of earthworms. P. lanceolata was not affected by the earthworms. We suggest that earthworms enhanced the competitive ability of L. perenne against T. repens. Nematodes increased the proportion of litter N in each of the plant species. Litter distribution (homogeneous vs. patch) did not affect the biomass of any plant species. However, P. lanceolata took up more ¹⁵N, when the litter was homogeneously mixed into the soil. The results suggest that endogeic earthworms may affect plant competition by promoting individual plant species. More studies including decomposers are necessary to understand their role in determining plant community structure.     

Arbuscular mycorrhizal fungi can transfer substantial amounts of nitrogen to their host plant from organic material

Nitrogen (N) capture by arbuscular mycorrhizal (AM) fungi from organic material is a recently discovered phenomenon. This study investigated the ability of two Glomus species to transfer N from organic material to host plants and examined whether the ability to capture N is related to fungal hyphal growth. Experimental microcosms had two compartments; these contained either a single plant of Plantago lanceolata inoculated with Glomus hoi or Glomus intraradices, or a patch of dried shoot material labelled with (15)N and (13)carbon (C). In one treatment, hyphae, but not roots, were allowed access to the patch; in the other treatment, access by both hyphae and roots was prevented. When allowed, fungi proliferated in the patch and captured N but not C, although G. intraradices transferred more N than G. hoi to the plant. Plants colonized with G. intraradices had a higher concentration of N than controls. Up to one-third of the patch N was captured by the AM fungi and transferred to the plant, while c. 20% of plant N may have been patch derived. These findings indicate that uptake from organic N could be important in AM symbiosis for both plant and fungal partners and that some AM fungi may acquire inorganic N from organic sources.     

施肥对苗期紫茎泽兰和黑麦草相对竞争力的影响

为种植黑麦草替代控制紫茎泽兰提供依据,运用取代实验法,研究了施肥(氮、磷)对苗期紫茎泽兰和黑麦草的相对竞争力以及生长表现的影响。结果表明:增施氮肥和磷肥均能够提高紫茎泽兰的相对竞争力,而仅磷肥对黑麦草的竞争力略有促进作用;但在各种施肥水平下,黑麦草的竞争力仍然明显强于紫茎泽兰。增施氮肥可以显著提高紫茎泽兰的株高、分枝数和干质量,而磷肥仅在较高时才显著提高其干质量;增施氮肥虽然可以提高黑麦草的分蘖数量,但对其干质量没有影响,而增施磷肥(2次)可以显著提高其干质量;在竞争中紫茎泽兰植株生长的能量分配(用根茎比表示)对磷肥不敏感。建议适当增施磷肥、不施或少施氮肥,以提高黑麦草的替代控制效果。     

Growth rates of rhizosphere microorganisms depend on competitive abilities of plants and N supply

Abstract

Plant‐microbial interactions under N‐limiting conditions are governed by competitive abilities of plants for N. Our study aimed to examine how two plant species of strawberry, Fragaria vesca L. (native species) and Duchesnea indica (Andrews) Focke (an invasive plant in central Europe), growing in intra‐specific and inter‐specific competition alter the functions of rhizosphere microorganisms in dependence on N availability. By intra‐specific competition at low N level, a 2.4‐fold slower microbial‐specific growth rate was observed under D. indica characterized by smaller root biomass and lower N content in roots compared with F. vesca. By inter‐specific competition of both plants at low N level, microbial growth rates were similar to those for D. indica indicating that plants with stronger competitive abilities for N controls microbial community in the rhizosphere. Since a high N level smoothed the differences between plant species in root and microbial biomass as well as in microbial growth rates under both intra‐specific and inter‐specific competition, we conclude that competitive abilities of plant species were crucial for microbial growth in the rhizosphere only under N imitation.     

Formation of Root Epidermal Transfer Cells in Plantago

The root ultrastructure and transmembrane electron transport activities of two Plantago species have been examined with respect to alterations in response to Fe deficiency, exogenously supplied auxin, and the presence of chromium in the external medium. Both species showed increased ferric reductase activity upon Fe starvation, but they differed in the maximum rates. The addition of chromium to the nutrient solution led to a further enhancement in Fe-ethylenediaminetetraacetate reduction by Fe-deficient plants. In roots of Plantago lanceolata, the enhanced redox activity is associated with the formation of transfer cells in the epidermis. Similar characteristics of rhizodermal cells were observed in Fe-sufficient roots 3 d after application of the auxin analog 2,4-dichlorophenoxy-acetic acid. No structural adaptations occurred in roots of Plantago maritima. A quantitative estimation of the frequencies of transfer cells in root segments of Fe-deficient plants that differ in reduction activity revealed no correlation between the two phenomena. It is concluded that the area of plasmalemma infoldings is not specialized for the enhanced reduction of extracytoplasmatic Fe in response to Fe deficiency. The role of transfer cells in the adaptation to suboptimal Fe availability and the mechanisms triggering their formation are discussed.     

Competition, traits and resource depletion in plant communities

Although of primary importance to explain plant community structure, general relationships between plant traits, resource depletion and competitive outcomes remain to be quantified across species. Here, we used a comparative approach to test whether instantaneous measurements of plant traits can capture both the amount of resources depleted under plant cover over time (competitive effect) and the way competitors perceived this resource depletion (competitive response). We performed a large competition experiment in which phytometers from a single grass species were transplanted within 18 different monocultures grown in a common-garden experiment, with a time-integrative quantification of light and water depletion over the phytometers’ growing season. Resource-capturing traits were measured on both phytometers (competitive response traits) and monocultures (competitive effect traits). The total amounts of depleted light and water availabilities over the season strongly differed among monocultures; they were best estimated by instantaneous measurements of height and rooting depth, respectively, performed when either light or water became limiting. Specific leaf area and leaf water potential, two competitive response traits measured at the leaf level, were good predictors of changes in phytometer performance under competition, and reflected the amount of light and water, respectively, perceived by plants throughout their lifespan. Our results demonstrated the relevance of instantaneous measures of plant traits as indicators of resource depletion over time, validating the trait-based approach for competition ecology. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

The effect of solar UV radiation on four plant species occurring in a coastal grassland vegetation in The Netherlands

During the summer of 1992, growth and some physiological parameters of four native plant species occurring in a coastal grassland in The Netherlands, were studied after reduction of solar UV irradiance using different cut-off filters. Biomass production, morphology and photosynthesis of all species tested were unaffected by the different treatments. Litter production of Plantago lanceolata was increased in the absence of the total UV waveband, indicating a possible role for this waveband in plant senescence. Depletion of the total UV waveband from sunlight resulted in alterations in biomass allocation in Calamagrostis epigeios and Urtica dioica while no changes were observed in P. Ianceolata and Verbascum thapsus. In C. epigeios an increase in the specific leaf area was observed, whereas in U. dioica root weight per total plant weight was decreased resulting in an increase in the shoot/root ratio. Both photosynthetic and UV-absorbing pigment concentrations were altered by the different filter applications. When compared to control plants receiving full sunlight, depletion of UV-B resulted in a significant increase in chlorophyll concentration in U. dioica leaves, this however did not affect photosynthetic rate. The presence of UV-B radiation enhanced the UV-absorbance of leaf extracts of all species except P. lanceolata. Optical characteristics of the leaves were also changed. Both the quantity ( P. lanceolata and U. dioica ) and the quality (all species) of radiation transmitted by the leaves was affected by the different treatments.     

Soil nutrient heterogeneity interacts with elevated CO2 and nutrient availability to determine species and assemblage responses in a model grassland community

Interactive effects of atmospheric CO(2) concentration ([CO(2)]), soil nutrient availability and soil nutrient spatial distribution on the structure and function of plant assemblages remain largely unexplored. Here we conducted a microcosm experiment to evaluate these interactions using a grassland assemblage formed by Lolium perenne, Plantago lanceolata, Trifolium repens, Anthoxanthum odoratum and Holcus lanatus. Assemblages exhibited precise root foraging patterns, had higher total and below-ground biomass, and captured more nitrogen when nutrients were supplied heterogeneously. Root foraging responses were modified by nutrient availability, and the patterns of N capture by interactions between nutrient distribution, availability and [CO(2)]. Greater above-ground biomass was observed under elevated CO(2) only under homogeneous conditions of nutrient supply and at the highest availability level. CO(2) interacted with nutrient distribution and availability to determine foliar percentage N and below : above-ground biomass ratios, respectively. Interactions between nutrient distribution and CO(2) determined the relative contribution to above-ground biomass of four of the species. The responses of dominant and subordinate species to [CO(2)] were dependent on the availability and distribution of nutrients. Our results suggest that soil nutrient distribution has the potential to influence the response of plant species and assemblages to changes in [CO(2)] and nutrient availability.     

Suppression of host photosynthesis by the parasitic plant Rhinanthus minor

BACKGROUND AND AIMS: Parasitism is well understood to have wide-ranging deleterious effects on host performance in species thus far characterized. Photosynthetic performance reductions have been noted in the Striga-Zea mays association; however, no such information exists for facultative hemiparasitic plants and their hosts, nor are the effects of host species understood. METHODS: Chlorophyll fluorimetry was used to study the effects of parasitism by the hemiparasite Rhinanthus minor on the grass Phleum bertolinii and the forb Plantago lanceolata, and the effects of host species on the photosynthetic apparatus of R. minor. KEY RESULTS: Parasitism by Rhinanthus led to a significant decrease in the host, and total (host + parasite) biomass in Phleum; however, in Plantago, no significant repression of growth was noted. Maximum quantum yield (F(v)/F(m)) was reduced in parasitized Plantago, relative to control plants, but not in Phleum. F(v)/F(m) was significantly lower in R. minor parasitizing Phleum than Plantago, suggesting Phleum to be a superior host to Plantago for R. minor. Steady-state quantum yield (Phi(PSII)) was significantly depressed in parasitized Phleum, but only at low irradiances in Plantago. Phi(PSII) was very low for R. minor grown on Plantago, but not Phleum. CONCLUSIONS: Shown here is the first evidence of the suppression of host photosynthesis by a facultative hemiparasitic plant, which has significant effects on total biomass production. Host identity is a significant factor in parasite success, with the forb Plantago lanceolata exhibiting apparent chemical as well as previously identified physical defences to parasitism. It is proposed that the electron transport rate (as denoted by Phi(PSII)) represents the limiting factor for biomass accumulation in this system, and that Plantago is able to suppress the growth of Rhinanthus by suppressing the electron transport rate.     

Phenotypic plasticity in seedling defense strategies: compensatory growth and chemical induction

Phenotypic plasticity in growth (leading to compensation) and secondary chemical production (leading to induction) in response to herbivory are key defense strategies in adult plants, but their role in seedling defense remains unclear. A pair of greenhouse studies was conducted to investigate compensation and induction in seedlings and juvenile plants, using Plantago lanceolata (Plantaginaceae) and the specialist buckeye caterpillar Junonia coenia (Nymphalidae) as a model system. Plants received 50% defoliation at two and four weeks of age, and groups of plants were harvested one week after herbivory and six to eight weeks after herbivory to investigate the duration of the responses. Plants damaged at two weeks showed no chemical induction and fully compensated for the lost leaf tissue by ten weeks of age. Plants damaged at four weeks showed a significant reduction in iridoid glycosides one week after herbivory and achieved full shoot compensation by ten weeks of age at the expense of root biomass. These results indicate that P. lanceolata seedlings use compensation, but not chemical induction, as a defense strategy. This research highlights the importance of considering ontogeny in studies of plant–herbivore interactions and suggests that seedling defense may differ markedly from adult plant defense.     

Growth, respiration and nutrient acquisition by the arbuscular mycorrhizal fungus Glomus mosseae and its host plant Plantago lanceolata in cooled soil

Although plant phosphate uptake is reduced by low soil temperature, arbuscular mycorrhizal (AM) fungi are responsible for P uptake in many plants. We investigated growth and carbon allocation of the AM fungus Glomus mosseae and a host plant (Plantago lanceolata) under reduced soil temperature. Plants were grown in compartmented microcosm units to determine the impact on both fungus and roots of a constant 2.7 °C reduction in soil temperature for 16 d. C allocation was measured using two (13)CO(2) pulse labels. Although root growth was reduced by cooling, AM colonization, growth and respiration of the extraradical mycelium (ERM) and allocation of assimilated (13)C to the ERM were all unaffected; the frequency of arbuscules increased. In contrast, root respiration and (13)C content and plant P and Zn content were all reduced by cooling. Cooling had less effect on N and K, and none on Ca and Mg content. The AM fungus G. mosseae was more able to sustain activity in cooled soil than were the roots of P. lanceolata, and so enhanced plant P content under a realistic degree of soil cooling that reduced plant growth. AM fungi may therefore be an effective means to promote plant nutrition under low soil temperatures.     

High nitrogen : phosphorus ratios reduce nutrient retention and second-year growth of wetland sedges

Shifts from nitrogen (N)- to phosphorus (P)-limited growth due to high N deposition may alter the functioning of wetland vegetation. This experiment tested how N vs P deficiency affects the growth and nutrient use of wetland sedges. Five wetland Carex species were grown at nine N : P supply ratios (0.6-405) with two absolute levels of N and P. Biomass and nutrient concentrations were determined after one and two growing seasons. Shoot biomass was maximal at N : P supply ratios of 15-26 after one season but 5-15 after two seasons. Photosynthesis after the first season, second-year growth, leaf longevity, and the fraction of nutrient supply retained by plants over two seasons were all negatively related to N : P supply ratios, with small effects of absolute supply. The five Carex species responded similarly to N : P ratios but differed in nutrient resorption efficiency and biomass allocation. Plants treated with high N : P ratios appeared to lose nutrients below ground. Such losses may reduce plant performance in P-limited wetlands affected by high N deposition.     

Interactions between root and shoot competition vary among species

Understanding how the competition varies with productivity is essential for differentiating among alternative models of plant community organization. Prior attempts to explain shifts in root and shoot competition along gradients have generally assumed an additive interaction between the two competitive forms, using an experimental design which does not fully separate both above‐ and belowground processes. At the most basic level, few field studies have separated root and shoot competition, and we have limited knowledge about both the relative importance of these processes, and how they interact to affect plant growth in the field. Presented here are findings from a field study in which root and shoot competition were experimentally separated by using root exclusion tubes and neighbor tiebacks in an early successional community. Individuals of four species (Abutilon theophrasti, Amaranthus retroflexus, Rumex crispus, and Plantago lanceolata) were grown at two levels of fertilization with full competition, aboveground competition only, belowground competition only, or neither above‐ nor belowground competition. Competition was measured as competitive response, which is the natural log of the relative biomass of a target plant grown with competition compared to growth without competition. In contrast to predictions from current models of productivity‐competition relationships, but in agreement with other experimental studies, there was no change in the strengths or root, shoot, or total competition with a modest increase in productivity. Despite no effect of fertilization on the strength of competition, the form of interaction between root and shoot competition varied both as a function of species identity and fertilization. For both of the rosette forming species, the combined effects of root and shoot competition were less than predicted assuming no interaction (a “negative interaction”), with one species switching from a negative to an additive interaction with fertilization. The fact that fertilization caused a shift in the root‐shoot interaction, but not in the total strength of root and shoot competition, suggests that the root‐shoot interaction is itself a highly labile variable. If root‐shoot interactions are common in natural systems, then simply measuring the strength of one form of competition in no way provides any information about the overall importance of that competitive form to plant growth.     

Functional groups' performances as influenced by nitrogen,phosphorus and nodule inhibition of legumes

Aims Nitrogen (N) and phosphorus (P) constitute essential elements for plant growth and their availability influence species diversity in herbaceous plant communities. Legumes exhibit relatively high abundance in N-limited soils. Moreover, the legumes' N:P ratios are much higher than those of the other plant species grown in the same site, probably because they are able to fix atmospheric N 2. The objective of this study was to determine how the relative proportion in N and P availability and the restriction of legumes to fix atmospheric N 2 affect: (i) the primary productivity of plant species, (ii) species composition and (iii) N and P concentrations of species.Methods In an outdoor experiment, mixtures containing grasses, legumes and non-legume forbs were established in 32 containers under four soil treatments (control, N addition, P addition and disinfected soil), in a completely randomized design with eight replicates. Plant growth was examined when N and P were limited in the control soil:sand mixture, in a pot experiment sown with Plantago lanceolata .Important findings The pot experiment indicated that both N and P were limiting for the growth of P. lanceolata. Soil treatments affected primary productivity and species composition. Legumes had a relatively high abundance in the control and their growth was favoured, especially that of Medicago sativa, by P addition. Grasses' growth was increased by the addition of N. Inhibition of rhizobia resulted in poor growth of legumes and concomitant higher growth of grasses, in comparison to the control. The N:P ratios of non-legume species differed between treatments and were always higher in the legume species, even in the disinfected soil. The latter provides evidence that the high N concentrations found in legumes are a physiological characteristic of this specific group of plants.