The performance of plants in any one generation can be influenced not just by the prevailing biotic and abiotic factors, but also by those factors experienced by the parental generation. These maternal effects have been recorded in an array of plant species, but most studies tend to focus on abiotic factors over two generations. Here we show that maternal effects in the annual forb Senecio vulgaris may be influenced by beneficial arbuscular mycorrhizal fungi and insect herbivory over four successive generations. These effects were very much determined by seed provisioning, wherein C:N:P ratios were altered by both fungi and aphids. There was little evidence of epigenetic changes induced by the fungi or insects, instead the driving forces seemed to be allocation of N and P to the seeds. However, changes in seed chemistry were not cumulative over generations, often decreases in seed nutrient content were followed by recovery in subsequent generations. The changes in seed stoichiometry can have important consequences for viability, germination and subsequent seedling growth rates. We conclude that studies of maternal effects need to be conducted over multiple generations, and also need to be multifactorial, involving variation in abiotic factors such as water and nutrients, combined with biotic factors.

This study investigated the effects of light and soil fertility, on arbuscular mycorrhizal fungi (AMF) colonization, and the growth responses (height and dry mass) of Syzygium seedlings. Seedlings of four Syzygium spp. were grown for 2 y in six different light treatments at the research station of the Sinharaja Forest, Sri Lanka. The light treatments exposed seedlings to: (1) 3%; (2) 16%; (3) 50%; (4) 100% of full sun (control); (5) short periods (2 h d−1) of direct sunlight; and (6) long periods (6 h d−1) of direct sunlight. In the 16% of full sun treatment five sets of fertilizer applications supplied: (1) magnesium; (2) potassium; (3) phosphorus; (4) all three nutrients combined; and (5) no fertilizer (control). The Syzygium species had the greatest mycorrhizal colonization in brighter treatments that provided direct light. Comparison across species revealed S. firmum to have moderate mycorrhizal colonization but high total dry mass. Syzygium operculatum had high percentages of mycorrhizal colonization while S. rubicundum had low percentages of mycorrhizal colonization especially in deep shade. Syzygium makul showed moderate levels of mycorrhizal colonization and dry mass, but low height growth. Among fertilizer applications, phosphorus enhanced seedling growth and mycorrhizal colonization for all species. However, species showed decreased growth with high amounts of potassium and combined fertilizer applications. Results suggest that AMF colonization will be highest, and Syzygium spp. growth greatest, beneath canopy openings large enough to receive direct sun in phosphorus-rich soils.

Interspecific variation in seed size can affect seedling establishment in the light-limited understorey of rain forests. Large seeds with extensive seed reserves may allow seedlings to (1) persist in the deep shade, thereby increasing the likelihood of exposure to a canopy gap (Leishman & Westoby 1994, Saverimuttu & Westoby 1996); (2) promote growth, thereby allowing seedlings to reach higher light strata or to develop sufficient photosynthetic tissue to meet their energy needs in low light (Foster 1986); and (3) allow compensation for tissue losses to herbivores and pathogens (Armstrong & Westoby 1993, Foster 1986, Harms & Dalling 1997). However, comparisons of shade tolerance among species that vary in seed size have produced conflicting results; short-term seedling survival was strongly positively associated with seed size in several studies (Grime & Jeffrey 1965, Leishman & Westoby 1994), but only weakly in others (Augspurger 1984, Grubb & Metcalfe 1996).

The ecology and forest management of tropical montane cloud forests of the Neotropics have attracted little scientific attention so far, and understanding of the ecosystem is still fragmentary (Churchill et al. 1995). The montane forests of southern Ecuador are an outstanding biodiversity hotspot for vascular plants (Barthlott et al. 1996), including a multitude of tree species (Madsen & Øllgard 1994). However, identification of species, studies of phenology, productivity, seed production and growth of seedlings have only just begun (Homeier & Breckle 2002). None of the trees has been investigated with respect to their mycorrhizal status, although it is well established that mycorrhizal symbioses are not only important for survival of trees in nutrient-poor habitats (Read 1991) and play a key role for nutrient cycling and nutrient retention in the humus layers (Medina & Cuevas 1993, Rilling et al. 2001), but also have an important impact on the composition of plant communities (Allen et al. 1995, Kottke 2002, van der Heijden et al. 1998).

In normal populations of the common grass Holcus lanatus there is a polymorphism for arsenate resistance, manifested as suppressed phosphate uptake (SPU), and controlled by a major gene with dominant expression. A natural population of SPU plants had greater arbuscular-mycorrhizal colonization than wild type, nonSPU plants. It was hypothesized that, in order to survive alongside plants with a normal rate of phosphate (P) uptake, SPU plants would be more dependent on mycorrhizal associations. We performed an experiment using plants with SPU phenotypes from both arsenate mine spoils and uncontaminated soils, as well as plants with a nonSPU phenotype. They were grown with and without a mycorrhizal inoculum and added N, which altered plant P requirements. We showed that grasses with SPU phenotypes accumulated more shoot P than nonSPU plants, the opposite of the expected result. SPU plants also produced considerably more flower panicles, and had greater shoot and root biomass. The persistence of SPU phenotypes in normal populations is not necessarily related to mycorrhizal colonization as there were no differences in percentage AM colonization between the phenotypes. Being mycorrhizal reduced flower biomass production, as mycorrhizal SPU plants had lower shoot P concentrations and produced fewer flower panicles than non-mycorrhizal, nonSPU plants. We now hypothesize that the SPU phenotype is brought about by a genotype that results in increased accumulation of P in shoots, and that suppression of the rate of uptake is a consequence of this high shoot P concentration, operating by means of a homeostatic feedback mechanism. We also postulate that increased flower production is linked to a high shoot P concentration. SPU plants thus allocate more resources into seed production, leading to a higher frequency of SPU genes. Increased reproductive allocation reduces vegetative allocation and may affect competitive ability and hence survival, explaining the maintenance of the polymorphism. As mycorrhizal SPU plants behave more like nonSPU plants, AM colonization itself could play a major part in the maintenance of the SPU polymorphism.

Under defined laboratory conditions it was shown that two glucosinolate-containing plant species, Tropaeolum majus and Carica papaya, were colonized by arbuscular mycorrhizal (AM) fungi, whereas it was not possible to detect AM fungal structures in other glucosinolate-containing plants (including several Brassicaceae). Benzylglucosinolate was present in all of the T. majus cultivars and in C. papaya it was the major glucosinolate. 2-Phenylethylglucosinolate was found in most of the non-host plants tested. Its absence in the AM host plants indicates a possible role for the isothiocyanate produced from its myrosinase-catalysed hydrolysis as a general AM inhibitory factor in non-host plants. The results suggest that some of the indole glucosinolates might also be involved in preventing AM formation in some of the species. In all plants tested, both AM hosts and non-hosts, the glucosinolate pattern was altered after inoculation with one of three different AM fungi (Glomus mosseae, Glomus intraradices and Gigaspora rosea), indicating signals between AM fungi and plants even before root colonization. The glucosinolate induction was not specifically dependent on the AM fungus. A time-course study in T. majus showed that glucosinolate induction was present during all stages of mycorrhizal colonization.

Afrothismia winkleri develops fleshy rhizomes, densely covered with small root tubercles, narrowing to filiform roots with age. The exclusively intracellular mycorrhizal fungus has distinct morphologies in different tissues of the plant. In the filiform root the hyphae grow straight and vesicles are borne on short hyphal stalks. The straight hyphae are present in the epidermis of the root tubercles, but change to loosely coiled and swollen hyphae in the rhizome tissue. No penetration from epidermis to root cortex was found. From the rhizome, a separating cell layer permits only one or rarely two hyphal penetrations into the cortex of each root tubercle. The hyphae proceed apically within the root hypodermis in a spiral row of distinctively coiled hyphae, branches of which colonize the inner root cortex. In the inner root cortex the hyphal coils degenerate to amorphous clumps. In older roots the cortex itself also deteriorates, but epidermis, hypodermis, endodermis and central cylinder persist. The mycorrhizal pattern in A. winkleri is interpreted as an elaborate exploitation system whereby the fungus provides carbon and nutrients to the plant and, simultaneously but spatially distinct, its hyphae are used to translocate and store the matter within the plant. Several features indicate that the endophyte is an arbuscular mycorrhizal fungus.

Colonization of roots by arbuscular mycorrhizal (AM) fungi in several annual crops in two consecutive seasons was compared with, in the second season, the density of fungal propagules in the soil with the use of a bioassay. Root density decreased down the soil profile in both years in all crops, and a high proportion of roots were mycorrhizal throughout the profile. AM colonization decreased down the profile in cotton and lablab in the second season only. The bioassay indicated that most propagules of AM fungi in soils under cotton were located near the surface, with virtually no propagules at 1 m. The absence of propagules at depth indicates a lack of mycelium deep in the soil, and suggests that mycorrhizas are primarily initiated in the surface soil and that the fungi colonize the root system mostly through secondary spread down the profile. The use of AM colonization in the field as an indicator of propagule density and symbiotic function should be qualified by an understanding of the depth in the soil from which roots were extracted.

Interactive effects of elevated atmospheric CO2 and arbuscular mycorrhizal (AM) fungi on biomass production and N2 fixation were investigated using black locust (Robinia pseudoacacia). Seedlings were grown in growth chambers maintained at either 350 μmol mol−1 or 710 μmol mol−1 CO2. Seedlings were inoculated with Rhizobium spp. and were grown with or without AM fungi. The 15N isotope dilution method was used to determine N source partitioning between N2 fixation and inorganic fertilizer uptake. Elevated atmospheric CO2 significantly increased the percentage of fine roots that were colonized by AM fungi. Mycorrhizal seedlings grown under elevated CO2 had the greatest overall plant biomass production, nodulation, N and P content, and root N absorption. Additionally, elevated CO2 levels enhanced nodule and root mass production, as well as N2 fixation rates, of non- mycorrhizal seedlings. However, the relative response of biomass production to CO2 enrichment was greater in non-mycorrhizal seedlings than in mycorrhizal seedlings. This study provides strong evidence that arbuscular mycorrhizal fungi play an important role in the extent to which plant nutrition of symbiotic N2-fixing tree species is affected by enriched atmospheric CO2.

The extent of arbuscular mycorrhizal colonization was assessed in 10 field-collected plant species, representing three annual forbs, three perennial forbs, three perennial grasses and one annual grass. Each root system of each plant was split into four portions, and for each portion, mycorrhizal structures were revealed with epifluorescence microscopy (under which only arbuscules are generally visible) and three commonly used stains (Chlorazol Black E, Acid Fuchsin and Trypan Blue). The aim of the study was not to evaluate the efficacy of each method, but to compare results obtained by each under standard laboratory conditions. The recorded colonization levels of arbuscules, total arbuscular mycorrhizal fungal material and total fungal (arbuscular mycorrhizal+non-arbuscular mycorrhizal) material differed significantly between visualization methods in a number of species. However, there were also interactions between stain and plant species, indicating that the performance of a stain is dependent on the plant species being examined. In some cases (e.g. Plantago lanceolata), each visualization method produced the same colonization level, while in others (e.g. Dactylis glomerata), each method gave a different result. These data therefore suggest that the level of mycorrhizal colonization recorded in any particular plant species at a particular time is dependent on the technique employed.

The outcome of dual infection of the grass Lolium perenne L. by arbuscular mycorrhizal (AM) fungi and the parasitic angiosperm Rhinanthus minor L. was investigated in a glasshouse study. Colonization of L. perenne roots by AM fungi was significantly reduced by the presence of R. minor, as was host growth which fell by 44–51%. It was concluded that these two responses were linked, with AM colonization declining in response to the reduction in availability of host carbon. Parasite growth and reproductive output rose by 58% and 47% respectively when the hosts were mycorrhizal. These trends were unrelated to the attachment success of the parasite, but were accompanied by a significant increase in the formation of secondary haustoria. The benefits afforded the parasite when the hosts were mycorrhizal were attributed to increased carbon and nutrient flux resulting from alternations in sink strength. Host responses to parasitism and mycorrhizal colonization were not affected by the interaction between the two symbionts. However, the suggestion is made that the interaction between the AM fungi and parasite might have long-term ecological implications for the host species via its impact on parasite fecundity.

Plantago lanceolata L. was grown for 104 d with (M) or without (NM) arbuscular mycorrhizal colonization under conditions of ambient (CAMB=350 μl l−1) and elevated (CELEV=540 μl l−1) CO2. Sequential harvests (H) were taken at 41 (H1), 76 (H2) and 104 d (H3) to determine the time-course of mycorrhizal influence on the response of the plant to CO2 enrichment. Total yields of M plants were greater than those of NM from H2 onwards. Plants in the MELEV treatment were significantly larger than those in the MAMB at 104 d. There were significant but much smaller differences in yield between NMELEV and NMAMB. The differences in total yield arose through impact of CELEV on both shoots and roots. Total root length was greater in MELEV than in MAMB only at H3, but total length of mycorrhizal root was greater at H2 and H3. The percentages of root length colonized and that occupied by arbuscules and vesicles were greater in MELEV than in MAMB at the last two harvests, indicating increased sequestration of carbon in internal fungal structures. Though extraradical hyphal lengths were greater in MELEV than in MAMB at H2 and H3, the differences were not significant. Phosphorus inflow and P content of M plants were higher than those of NM plants at H2 and H3, and were higher in MELEV than in MAMB at H3. anova revealed no significant interactions between CO2 and mycorrhizal treatment.

The results are discussed in relation to carbon sequestration in mycorrhizal systems and likely impacts of CO2 enrichment on P. lanceolata grown under field conditions. The importance of sequential harvesting for realistic determination of responses to CO2 is stressed.

Roots of Lablab purpureus (L.) Sweet were treated with tri-iodobenzoic acid (TIBA), kinetin or with nodulation factors (Nod factors) purified from Rhizobium sp. NGR234 and grown in the presence of a mycorrhizal inoculum (Glomus mosseae (Nicol. & Gerd.) Gerdemann & Trappe. Colonization by the mycorrhizal fungus was increased from <30% to c. 65% of root length when roots were treated with these growth regulators. Moreover, treatment of mycorrhizal L. purpureus roots with Nod factors or TIBA strongly induced sporocarp formation of Glomus mosseae. In parallel, the pool size of the fungal disaccharide trehalose was significantly affected in roots treated with TIBA and Nod factors alone, and with TIBA combined with all effectors, and increased from 0·06 mg g−1 d. wt in control roots to up to 1·7 mg g−1 d. wt (TIBA+kinetin). Conversely, the sucrose pool decreased from 5% d. wt to less than a half in roots treated with Nod factors. Activities of trehalase were significantly enhanced in mycorrhizal roots by the treatment with Nod factors or TIBA. When Nod factors and TIBA were added in combination, these activities were strongly enhanced suggesting synergism between these growth regulators.

Arbuscular mycorrhizal (AM) fungi form mutualistic symbioses with the root systems of most plant species. These mutualisms regulate nutrient exchange in the plant–soil interface and might influence the way in which plants respond to increasing atmospheric CO2. In other experiments, mycorrhizal responses to elevated CO2 have been variable, so in this study we test the hypothesis that different genera of AM fungi differ in their response, and in turn alter the plant's response, to elevated CO2. Four species from three genera of AM fungi were tested. Artemisia tridentata Nutt. seedlings were inoculated with either Glomus intraradices Schenck & Smith, Glomus etunicatum Becker & Gerdemann, Acaulospora sp. or Scutellospora calospora (Nicol. & Gerd.) Walker & Sanders and grown at either ambient CO2 (350 ppm) or elevated CO2 (700 ppm). Several significant inter-specific responses were detected. Elevated CO2 caused percent arbuscular and hyphal colonization to increase for the two Glomus species, but not for Acaulospora sp. or S. calospora. Vesicular colonization was not affected by elevated CO2 for any fungal species. In the extra-radical phase, the two Glomus species produced a significantly higher number of spores in response to elevated CO2, whereas Acaulospora sp. and S. calospora developed significantly higher hyphal lengths. These data show that AM fungal taxa differ in their growth allocation strategies and in their responses to elevated CO2, and that mycorrhizal diversity should not be overlooked in global change research.

Plant roots enter symbiotic as well as pathogenic interactions with fungi in the rhizosphere. We studied the response of bean (Phaseolus vulgaris L. cv. Saxa) roots to infection by the arbuscular mycorrhizal fungus Glomus mosseae (Nicol. & Gerd.) Gerdemann & Trappe and the pathogen Fusarium solani (Mart.) Sacc. f. sp. phaseoli (Burkholder) Snyder & Hansen. In a time-course study of the symbiotic interaction between bean roots and G. mosseae, covering all stages of mycorrhiza development, we detected little change in the expression of the defence-related genes chitinase, β-1,3-glucanase and phenylalanine ammonia-lyase compared with non-mycorrhizal control roots. The only difference observed was a transient increase in chalcone synthase transcripts at later stages of mycorrhizal root colonization. In interactions with the pathogen, a marked induction of chitinase and phenylalanine ammonia-lyase expression was observed at the level of both the transcripts and enzyme activities. Class I β-1,3-glucanase levels strongly increased at the transcript level, but there was little change in the overall β-1,3-glucanase enzyme activity. In the non-host interaction between common bean and Fusarium solani (Mart.) Sacc. f. sp. pisi (Linford) Snyder & Hansen defence responses increased only slightly and transiently, if at all.

The relationship between mycorrhizal colonisation and phosphorus acquired by seedlings of the arbuscular mycorrhizal tree Oubanguia alata Bak f. (Scytopetalaceae) and the ectomycorrhizal tree Tetraberlinia moreliana Aubr. (Caesalpiniodeae) was evaluated at low and high inorganic phosphorus availability. AM colonisation was positively correlated with phosphorus uptake by O. alata at low, but not at high phosphorus availability. Seedlings growth was positively related to arbuscular mycorrhizal colonisation at both low and high phosphorus availability, suggesting that growth promotion by arbuscular mycorrhizas is not simply related to an increase of phosphorus uptake. In contrast, phosphorus uptake by T. moreliana was correlated with EM colonisation at both low and high phosphorus availability, but there was no relationship between growth and ectomycorrhizal colonisation. Promotion of phosphorus uptake by arbuscular mycorrhizas and ectomycorrhizas at low phosphorus availability is consistent with the co-occurrence of the two types of mycorrhiza in tropical rain forests where available soil phosphorus is low. However, ectomycorrhizal colonisation may also be of advantage where inputs of phosphorus rich litter raise the phosphorus status of the soil, as seen in the groves of ectomycorrhizal trees in Korup National Park, and may be one of the factors reinforcing local dominance by these trees.

Mean δ15N of whole-plants of lettuce and barley varied by [les ]3‰ when given a chemically and isotopically uniform N source. This variation was related to the presence, absence and species of arbuscular mycorrhizal (AM) fungi and to external N concentration. A highly AM-susceptible plant (lettuce) responded to treatments differently than a less susceptible one (barley). The largest change in whole plant δ15N was related to the experimental combination most likely to be found in field conditions: species of fungus interacting with varying external concentrations of N.

The mechanisms underlying observed variations of plant δ15N are not understood, nor can they be approached directly using δ15N. However, descriptive data, such as presented here, are important to the development of a mechanistic model, because they suggest relationships for future research, using inter alia15N-enriched tracers. They also confirm that plant N sources, cannot be identified using plant δ15N, even when the type of mycorrhizal association (endo vs. ecto) is known.

The arbuscular mycorrhizas of bluebell (Hyacinthoides non-scripta (L.) Chouard ex Rothm.) involve several symbiotic fungi of the order Glomales. We have previously simplified the system by ignoring the taxonomic diversity of the fungi, but it is unlikely that all fungal species contribute in the same manner or to the same extent to the functioning of the symbiosis. To discover how many and which fungi take part in the bluebell mycorrhiza we sought to identify the range of arbuscular mycorrhizal (AM) fungi found in bluebell roots sampled during a complete growing season, September to June.

Although the taxonomy of the Glomales by their spores is not yet fully understood, identification is, to a large extent, possible. Arbuscular mycorrhizal communities are usually characterized by their spores but, since spores can rarely be directly associated with individual plants or plant species, a more satisfactory approach would be to identify fungal symbionts where they interact with the host plant, in the roots. Unfortunately, the intraradical mycelia of the fungi are less easily distinguished than the spores and, as yet, classification is possible only to the family level.

We have developed a method whereby different AM fungal taxa in the roots of bluebell can be distinguished by objective assessment. A large suite of morphological characteristics of the fungi in roots was recorded in samples taken at monthly intervals. Hierarchical cluster analysis of the resulting data separated six distinct AM fungal morphotypes (Scutellospora type, Acaulospora type, three Glomus types and fine endophytes) and a classification system created by which identification by eye was possible.

We compared the fungi in roots with glomalean spores in soil from the root zone of the same bluebell plants. Two species occurred in most samples, Scutellospora dipurpurescens Morton & Koske (emend. Walker, 1993) and Acaulospora koskei Blaskowski. A further six occurred sporadically, five Acaulospora spp. and Glomus rubiforme Gerdemann & Trappe) Almeida & Schenck (=Sclerocystis rubiformis). The presence of a single species of Scutellospora was consistent with a Scutellospora root morphotype which varied little. By contrast, the diversity of Acaulospora in the spore assemblage was reflected by variation within the Acaulospora morphotype. Glomus spores were very rarely found in field collections, yet Glomus morphotypes were found to be an important component of the bluebell mycorrhiza.

Because some important species are not represented in spore assemblages in the field and those that are found can only be associated with vegetational groups, not individual plants or single species, glomalean spore populations provide only a partial account of the fungi which contribute to arbuscular mycorrhizas. Although it is still not possible to identify AM fungi in roots with the same precision as their spores, the method reported here permits assessment of diversity in the roots of individual plant species, which may be applied to the investigation of mycorrhizal function and demography in natural ecosystems.

Roots of bluebell (Hyacinthoides non-scripta (L.) Chouard ex Rothm.) are colonized by a range of fungal symbionts from several genera of the order Glomales. Using the identification scheme described in Merryweather & Fitter (1998), arbuscular mycorrhizal (AM) fungi in bluebell roots were quantified throughout the single growing season of 1994–5 and compared with populations of spores found in the soil around the roots.

In the early part of the growing season, when its activity is entirely subterranean (autumn and winter), bluebell habitually associates with a Scutellospora morphotype which is almost certainly S. dipurpurescens Morton & Koske (emend. Walker, 1993) whose spores occur in the root zone. This is the time of maximum phosphorus inflow and bulb-stored carbohydrate utilization by this mycorrhiza. A diverse flora of other AM fungal morphotypes (Acaulospora and Glomus), which might also form mycorrhizas with other plants in the vicinity of bluebells, invade the roots later in the season (spring), when P inflow is reduced and carbohydrate is available as fresh photosynthate. Their contribution to the mycorrhiza might be less than that of Scutellospora, particularly in terms of P assimilation.

Both AM fungi in roots and glomalean spores recovered from soil around bluebell roots showed a significant degree of correlation with the vegetation within which the test plants grew. In the case of AM fungi in roots, Scutellospora showed no special preference for either, but Glomus correlated with a canopy of sycamore (Acer pseudoplatanus L.) and Acaulospora with oak (Quercus petraea (Mattuschka) Liebl.). Spores which most closely resembled S. dipurpurescens and Acaulospora gerdemannii Schenck & Nicolson were significantly more numerous under sycamore, but a spore like Acaulospora koskei Blaskowski, the most numerous and frequently encountered glomalean spore in the system, showed no preference for areas dominated by either tree. There was no significant relationship between AM fungal populations in bluebell roots and glomalean spores recovered from associated soil.

The two spore taxa most frequently found in the vicinity of bluebell roots (A. koskei and S. dipurpurescens) were also found in lower numbers in soil from a region of the field site in which bluebell was absent, indicating that the main bluebell AM fungi do not exclusively associate with that host.

Many hypotheses have been developed to explain the adaptive nature of insect galls. One of these, the nutrition hypothesis, states that gall formers have advantages over other insects because gall tissue provides a better (higher quality) food source than unmodified tissue. However, this has rarely been experimentally tested. In a test of this hypothesis, we grew plants of Cirsium arvense (L.) Scop. in a factorial design with two main treatments: the addition of nitrogen (to enhance foliar N levels) and of fungicide (to reduce colonization of roots by arbuscular mycorrhizal fungi). Mycorrhizal fungi have been shown previously to reduce the N concentration of host plants. Plants were exposed to adult gall flies, Urophora cardui L., and maintained through one season to allow maturation of galls.

Reduction of the percentage mycorrhizal colonization by fungicide resulted in an elevation of total stem N comparable to that achieved by N addition, but gall N concentration remained unchanged in all treatments. Nitrogen application elevated stem N levels when mycorrhizal fungi were present, but application of both compounds together did not result in any increase over either single treatment. Fungicide application resulted in larger galls, which contained more larval chambers, with more live, and heavier, larvae. However, the main effects of N were not significant, as N addition only increased fly performance on plants where mycorrhizas were not reduced.

It is suggested that U. cardui gall inhabitants can manipulate N at an optimal level and thus might conform to a modified version of the nutrition hypothesis. Mycorrhizal colonization might reduce gall fly performance by delaying the appearance, or impairing the quality, of secondary nutritive tissue in the gall. Future tests of the nutrition hypothesis should include a consideration of the plant's mycorrhizal status.