This work explored whether the natural abundances of carbon and nitrogen isotopes could be used to describe the movement of C and N within wheat plants; we also considered whether isotopic analyses of aphids or their honeydew would substitute for direct analysis of phloem exudate.

The δ13C of ears and roots (sinks) most closely matched those of the sugars+organic acids fraction (sources) in both growth stages; phloem δ13C matched that of leaf blade sugars. Xylem exudate δ13C matched no other putative (and measured) source in the ear-forming stage and matched that of whole roots and ears in the grain-filling stage. The δ15N of grain and roots (sinks) resembled that of leaf amino acids (sources) in the ear-forming stage. In the grain-filling stage, ear δ15N continued to resemble that of leaf amino acids, and δ15N of roots most closely resembled that of whole leaves. In the grain-filling stage, phloem δ15N fell between that of leaf blade amino acids and that of whole leaves and was 15N-depleted relative to internal and external NO3−-N. In both growth stages, xylem exudate δ15N was less than that of soil NO3−-N and and more than that of residual soil N after mineral N extraction. The isotopic values are generally in agreement with data from other approaches, such as isotope labelling; they show NO3−-N reduction in both shoots and roots of wheat and significant N recycling (root-shoot-phloem-root) and C movement.

Aphids might serve as a substitute for isotopic analysis of phloem δ15N, having the same value as their food source. Their excreta was 15N-enriched relative to phloem.

Phytoplankton infections by fungal parasites in the upper, mixed layer of a mesotrophic northern temperate lake were analysed according to the following parameters: host and parasite species, host population density and prevalence of infection, resting spore formation by the parasite, and the lowest host density at which parasites appeared. The phytoplankton taxa recorded included the Cyanobacteria, Dinomastigota, Chrysophyceae, Bacillariophyceae, Chlorophyceae, Cryptophyceae and Haptophyceae, but infection was never found in the last two classes. The parasites belonged almost exclusively to the monocentric Chytridiomycetes. Fungal epidemics occurred at all times of the year. Parasites appeared at population densities as low as about 1 cell ml−1 in some host species, with infection prevalence sometimes exceeding 80%. The proportion of the total phytoplankton biovolume infected by fungi was usually much <1%, but occasionally reached 10%. Parasitism proved to be highly species-specific, with one parasite species usually infecting only one host species. In the case of Zygorhizidium planktonicum, which infected both Asterionella formosa and Synedra acus, there is evidence that two species-specific formae speciales, each infecting only one of these two host species, are present in the lake.

To study the response of non-mycorrhizal and mycorrhizal maize plants to drought, the changes in the pools of non-structural carbohydrates and amino acids were analysed in leaves and roots of two maize cvs. Plants well colonized by the arbuscular mycorrhizal fungus Glomus mosseae (Nicol. & Gerd.) (60% of root length infected) and comparable non-mycorrhizal plants were subjected to moderate drought stress by reducing the water supply. This stress induced a conspicuous increase in the trehalose pool in the mycorrhizal roots, probably because it was accumulated by the fungal symbiont. Furthermore, glucose and fructose were accumulated in leaves and roots of non-mycorrhizal plants but not in the mycorrhizal ones. Starch disappeared completely from the leaves of both mycorrhizal and non-mycorrhizal plants in response to drought. Activities of soluble acid invertase and trehalase were also measured. Acid invertase activity increased during drought in the leaves of both non-mycorrhizal and mycorrhizal plants whilst in the roots it was unaffected in non-mycorrhizal plants and decreased in the mycorrhizal ones. Without drought stress, trehalase activity was considerably higher in the leaves and roots of mycorrhizal plants than in those of non-mycorrhizal plants. It increased conspicuously during drought, primarily in the leaves of non-mycorrhizal plants. A drought-induced accumulation of amino acids as well as imino acids was found in roots and leaves of both mycorrhizal and non-mycorrhizal plants; leaves of mycorrhizal plants accumulated more imino acids than those of non-mycorrhizal ones. Our results show that drought stress and the presence of a mycorrhizal fungus have a considerable effect on carbon partitioning, imino acid and amino acid accumulation in maize plants.

Cumulative germination curves were recorded for carrot (Daucus carota L.) seeds at a range of constant temperatures (T) and water potentials (Ψ) in the laboratory and under variable soil conditions in 15 seed-bed environments in the field. A single base temperature (Tb), a distribution of base water potentials (Ψb(G)) for percentiles (G) of the population and the hydrothermal time constant (Hg) were determined from laboratory data. Although less effective at low Ψ, it was possible, using these germination parameters, to satisfactorily describe the effect of T and Ψ on germination rates under constant conditions according to the threshold models of thermal and hydrothermal time. These models were applied to field data with the condition that the germination process ceased if T<Tb for thermal time and additionally Ψ<Ψb(G) for hydrothermal time.

Neither model accurately predicted germination patterns in the field. However, the pattern of germination was adequately described in most situations by a modified threshold model in which the predicted progress towards germination was unaffected by soil Ψ, provided it remained above Ψb(G), and was therefore more rapid under variable seed bed conditions than hydrothermal time. In this modified threshold model, the condition Ψ<Ψb(G) had to be fulfilled at the initiation of radicle extension before germination occurred. This result implies that the initiation of radicle growth operates as a moisture-sensitive step that can determine germination and seedling emergence timing under variable soil-moisture conditions.

Seedling emergence was also recorded in the field and used to determine, separately, the impact of germination and post-germination growth on the variation in seedling emergence patterns. The analysis suggests that delays in seedling emergence occur largely in the germination phase, but that seedling losses and variation in the spread of seedling emergence times within the population occur largely during the post-germination growth phase.

Patterns of photobiont diversity were examined in some Nostoc-containing lichens using the nucleotide sequence of the cyanobacterial tRNALeu (UAA) intron. Lichen specimens collected in northwestern USA were analysed and the sequence data were compared with tRNALeu(UAA) intron sequences previously obtained from lichens in northern Europe. Generally, it is the species identity of a lichen rather than the geographical origin of the specimen that determines the identity of the cyanobiont. Identical intron sequences were found in Peltigera membranacea specimens collected in Oregon (USA) and in Sweden, and very similar sequences were also found in Nephroma resupinatum thalli collected in Oregon and Finland. Furthermore, in mixed assemblages where two Peltigera species grew in physical contact with each other, the different lichen species housed different photobiont strains. There is however not a one-to-one relation between mycobiont and photobiont as some intron sequences were found in more than one lichen species, and different intron sequences were found in different samples of some lichen taxa. Peltigera venosa exhibited a higher level of photobiont diversity than any other lichen species studied, and several intron sequences could for the first time be obtained from a single thallus. It is not clear whether this is evidence of lower cyanobiont specificity, or reflects an ability to exhibit different degrees of lichenization with different Nostoc strains. In one specimen of P. venosa, which contained bipartite cyanosymbiodemes and tripartite, cephalodiate thalli, both thallus types contained the same intron sequence.

Ectomycorrhizal seedlings of Scots pine (Pinus sylvestris) inoculated with the nitrotolerant Laccaria bicolor and the nitrophobic Suillus bovinus were exposed to ambient (350 μl l−1) and elevated (700 μl l−1) [CO2]. After 79 d the seedlings were labelled for 28 d with 14CO2, after which they were harvested. 14C was determined in shoots, roots plus mycorrhizas, soil, and below-ground respiration; nitrogen was determined in shoots and roots. Total net 14C uptake increased under elevated [CO2]. The extra carbon did not increase the shoot mass but was translocated to the roots and resulted in a decreased shoot-to-root ratio in the Suillus-inoculated seedlings. Laccaria-inoculated seedlings did not incorporate the additional carbon in root or fungal tissue but only increased below-ground respiration. S. bovinus acquired or transferred nitrogen better than L. bicolor and enabled the seedlings to perform better with regard to net carbon uptake under elevated [CO2]. This resulted in nitrogen concentrations in shoots of Suillus-inoculated seedlings that were twice as high as in Laccaria-inoculated seedlings, irrespective of [CO2]. The higher nitrogen concentration in the shoots resulted in a doubling of the 14C uptake per unit shoot mass. Our results suggest that the ability of ectomycorrhizal Scots pine seedlings to respond positively to elevated atmospheric [CO2] is strongly fungal-species specific.

The host range of ectomycorrhizal (ECM) fungi in Britain was examined by compilation of a data matrix from published literature sources, based primarily on accounts of sporocarp associations with particular host genera. Information was gathered for 577 species of ECM fungi belonging to 51 genera, and 25 genera of host trees, representing the majority of ECM fungal species and host genera recorded in Britain.

Pronounced variation was recorded in the number of ECM fungal species associated with different host genera, with over 200 species recorded with Betula, Fagus, Pinus and Quercus. There was a positive linear relationship (r2=0·47, P=0·007) between the number of species of ECM fungi associated with different host genera and the total area occupied by each tree genus in Britain (both values log-transformed). There was also variation in the number of species of ECM fungi which were apparently specific to particular host genera, values ranging from zero (in 15 genera) to >40 in the case of Betula and Fagus. In total, 233 fungal species appeared to be specific to a single host genus (i.e. 40% of those surveyed). Comparison of the ECM mycota associated with different host genera by PCA accounted for 17% of the total variation, with genera belonging to the Fagaceae (Quercus, Fagus and Castanea) tending to cluster together, indicating a degree of overlap in their ECM associates. Exotic conifer species, which displayed a lower ECM diversity than would be expected from their distributional areas, were characterized by a high degree of overlap with the ECM associates of Pinus and Betula.

These results indicate that the abundance of different genera of host trees and variation in host specificity could provide a basis for understanding patterns of diversity in ECM fungi within Britain.

We assessed the ability of several populations of the metal-hyperaccumulator species, Thlaspi caerulescens, to mobilize non-labile cadmium in soils historically contaminated by Pb/Zn mine spoil or sewage sludge. Radio- labile Cd was determined chemically as an ‘E-value’, [CdE], and biologically as an ‘L-value’, [CdL]. For comparison, chloride-extractable Cd, [Cdchlor], was also determined using 1 M CaCl2 as a single-step soil extractant. Values of [CdL] were measured for six populations of T. caerulescens that varied substantially in their ability to assimilate soil Cd, and a non-accumulator species with a similar growth habit, Lepidium heterophyllum. Seeds were sown in soil spiked with 109Cd and grown for 9–12 wk in a controlled environment room. Values of [CdL] were determined from the specific activity of 109Cd and concentration of Cd in the plant leaves. For the six soils studied, [CdE] ranged from 4.9 to 49% of total soil Cd [CdT]. Values of [CdL] were, in general, in close agreement with both [CdE] and [Cdchlor] and substantially less than [CdT]. However, [CdL] showed no correlation with the concentration of Cd in plant tissue, [Cdshoot]. This suggests that, in the soils studied, T. caerulescens did not mobilize non-labile soil Cd by producing root exudates or altering rhizosphere pH. The results imply that there may be significant restrictions to metal bioavailability, even to hyperaccumulator species, in heavily contaminated soils in which a large proportion of the metal may be present in ‘non-labile’ forms.

Salt-affected soil is a major problem and has led to large areas of the earth's surface being unable to support agriculture. Wheat is not tolerant to high levels of salt; however, a number of its wild relatives, in particular Thinopyrum bessarabicum Savul. and Rayss, which grows on the Black Sea shore in the Crimea, are tolerant to high levels of NaCl. Thus the introgression of genes conferring salt tolerance from Th. bessarabicum into Triticum aestivum is one way in which salt-tolerant wheat varieties can be produced. This paper reviews the progress made in developing salt-tolerant wheat carrying Th. bessarabicum genes and salt-tolerant wheat/Th. bessarabicum amphiploids.

Elevated [CO2] has been shown to protect photosynthesis and growth of wheat against moderately elevated [O3]. To investigate the role of ozone exclusion and detoxification in this protection, spring wheat (Triticum aestivum L. cv. Wembley) was grown from seed, in controlled-environment chambers, under reciprocal combinations of [CO2] at 350 or 700 μmol mol−1 and [O3] peaking at <5 or 60 nmol mol−1, respectively. Cumulative ozone dose to the mesophyll and antioxidant status were determined throughout flag leaf development. Catalase activity correlated with rates of photorespiration and declined in response to elevated [CO2] and/or [O3]. Superoxide dismutase activity was not significantly affected by either condition. Neither ascorbate nor glutathione content was enhanced by elevated [CO2]. In wheat, at moderately elevated [O3], our results show that stomatal exclusion plays a major role in the protective effect of elevated [CO2] against O3 damage.

The study reported was conducted to establish the impacts of photochemical oxidants (primarily ambient ozone, O3) on the yield of watermelon (Citrullus lanatus) at a site on the east coast of Spain. Fruit yield and quality were monitored in plots established in a commercial watermelon field exposed, in open-top chambers (OTCs), to non-filtered air (NFA; near-ambient levels of ozone) or charcoal-filtered air (decreased levels of photochemical oxidants including O3; CFA), or to ambient air (AA), during the 1988 and 1989 growing seasons. Ambient levels of O3 were found to exceed present UN-ECE (United Nations Economic Commission for Europe; Convention on Long-Range Transboundary Air Pollution) critical level guidelines for the protection of crop yield by approx. twofold in 1988 and by approx. fivefold in 1989. Plants exposed to NFA and AA developed visible O3 injury on the upper surface of sun-exposed older leaves, and fruit yield (annual marketable fruit weight and number) was found to be depressed in OTCs ventilated with NFA in comparison with those receiving CFA. Consistent with inter-annual variations in O3 exposure, greater yield losses were experienced in 1989 (39%) than in 1988 (19%), an effect mediated predominantly by a decline in fruit number rather than average fruit weight. Exposure to ambient levels of ozone also slightly decreased fruit quality (4–8% decline in soluble solids content). Leaf gas exchange measurements made in the field in 1988 revealed effects of O3 on fruit yield and quality to be associated with a decline in the net CO2 assimilation rate per unit leaf area under light saturation (Asat) and stomatal conductance to water vapour (gs), and enhanced rates of dark respiration. A/ci curves (where A is the net CO2 assimilation rate per unit leaf area and ci is the mole fraction of CO2 in the leaf intercellular air space) constructed for plants grown in laboratory-based closed chambers, and exposed to an accumulated O3 exposure similar to that experienced by plants in the field, suggested that the likely cause of the decline in photosynthetic capacity was (1) a decrease in the amount and/or activity of Rubisco and (2) an impaired capacity for regeneration of ribulose 1,5-bisphosphate, which was not mediated through changes in the photochemical efficiency of photosystem II (Fv/Fm, where Fv is variable chlorophyll a fluorescence and Fm is maximum chlorophyll a fluorescence). No shift in the relative stomatal limitation to photosynthesis was observed under the influence of O3, suggesting that the decline in gs induced by the pollutant in both field and laboratory was the result, and not the cause, of the decrease in Asat. Ozone exposure also caused a decrease in C isotope discrimination (approx. 0.5‰), a shift that revealed a departure from predicted theory based on supporting leaf gas exchange measurements. The study demonstrates that ambient levels of photochemical oxidants on the Spanish Mediterranean coast are high enough to adversely influence the yield and physiology of an economically important crop grown in the region, and the magnitude of the effects was greater than would be predicted from exposure–yield-loss relationships for grain and fodder crops in central and northern Europe. Implications for the derivation of UN-ECE ozone standards are discussed, along with the mechanistic basis of the observed yield decreases.

The hypaphorine concentration in Pisolithus tinctorius Coker & Couch hyphae colonizing Eucalyptus roots was 3 to 5 times higher than in adjacent parts of the fungal colony. This phenomenon, observed 24 h after inoculation, was also recorded in several-month-old, well-established ectomycorrhizas. Accumulation was controlled by specific root-derived diffusible molecules: it can be induced through a membrane, but not by non-host plants. In pure culture, high hypaphorine concentration was found only in the youngest mycelium, i.e. the outer 2 mm of the colony. Fungal hypaphorine had no IAA-like activity on Eucalyptus root development and therefore could not be considered as an auxin analogue; instead, a strong reduction of root hair elongation was recorded.

Relationships between nitrogen (N) content and growth are routinely measured in plants. This study determined the effects of N on the separate morphological and physiological components of plant growth, to assess how N-limited growth is effected through these components. Lettuce (Lactuca sativa) plants were grown hydroponically under contrasting N-supply regimes, with the external N supply either maintained continuously throughout the period of study, or withdrawn for up to 14 d. Richards' growth functions, selected using an objective curve-fitting technique, accounted for 99.0 and 99.1% of the variation in plant dry weight for control and N-limited plants respectively. Sublinear relationships occurred between N and relative growth rates under restricted N-supply conditions, consistent with previous observations. There were effects of treatment on morphological and physiological components of growth. Leaf weight ratio increased over time in control plants and decreased in N- limited plants. Shoot:root ratio followed a similar pattern. On a whole-plant basis, assimilation of carbon decreased in N-limited plants, a response paralleled by differences in stomatal conductance between treatments. Changes in C assimilation, expressed as a function of stomatal conductance to water vapour, suggest that the effects of N limitation on growth did not result directly from a lack of photosynthetic enzymes. Relationships between plant N content and components of growth will depend on the availability of different N pools for remobilization and use within the plant.

TIP1 is a gene defined by an X-ray induced allele tip1–2 and a previously described EMS-induced allele tip1−1. TIP1 is involved in plant cell growth. tip1–2 plants display growth defects throughout the plant and exhibit defects in both root-hair and pollen-tube growth. tip1–2 plants are partly male sterile resulting from a combination of pollen germination and pollen-tube defects; their root-hairs are short, exhibit a tendency to branch and 2–4 hairs can initiate from each hair cell. They are also slightly dwarf in stature as a result of a general decrease in cell growth indicating that TIP1 activity is required for general cell growth. We propose a role for TIP in both the initiation and maintenance of growth in tip-growing cells. In addition TIP1 activity is required for normal cell expansion (non-tip cell growth) indicating that TIP1 is not exclusively involved in tip-growth.

The effects of mycorrhiza formation in combination with elevated CO2 concentrations on carbon metabolism of Norway spruce (Picea abies) seedlings and aspen (Populus tremula × Populus tremuloides) plantlets were analysed. Plants were inoculated for 6 wk with the ectomycorrhizal fungi Amanita muscaria and Paxillus involutus (aspen only) in an axenic Petri-dish culture at 350 and 700 μl l−1 CO2 partial pressure. After mycorrhiza formation, a stimulation of net assimilation rate was accompanied by decreased activities of sucrose synthase, an increased activation state of sucrose-phosphate synthase, decreased fructose-2,6-bisphosphate and starch, and slightly elevated glucose-6-phosphate contents in source leaves of both host species, independent of CO2 concentration. Exposure to elevated CO2 generally resulted in higher net assimilation rates, increased starch as well as decreased fructose-2,6-bisphosphate (aspen only) content in source leaves of both mycorrhizal and nonmycorrhizal plants. Our data indicate only slightly improved carbon utilization by mycorrhizal plants at elevated CO2. They demonstrate however, that both factors which modulate the sink-source properties of plants increase the capacity for sucrose synthesis in source leaves mainly by allosteric enzyme regulation.

The photosynthate costs of processes (amino acid and protein synthesis and turnover, and pH regulation) associated with the utilization of nitrate (NO3−), ammonium (NH4+) or glutamine (Gln) for plant growth were estimated. Based on these estimates, the effects of these forms of nitrogen (N) on the carbon balance of plants and on shoot–root biomass allocation were evaluated. The results indicated that NO3− as an N source for plant growth is not substantially more expensive to utilize than either NH4+ or Gln, particularly in the long term when costs due to protein turnover dominate the total costs of N utilization. It is also suggested that the photosynthate use in processes associated with N assimilation has little impact on the carbon balance of plants, and hence on shoot–root biomass allocation.

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.

Seedling mortality of Alyssum serpyllifolium ssp. lusitanicum and A. murale, both nickel hyperaccumulators, was reduced by increasing concentrations of metal within plant tissues when inoculated with the fungi Pythium mamillatum or P. ultimum, both of which cause damping-off disease of seedlings. Pythium mamillatum, isolated from nickel-rich serpentine soil, was more tolerant of nickel than P. ultimum, isolated from low-metal control soil, and was more pathogenic than P. ultimum towards seedlings containing high concentrations of metal. These results support the hypothesis that metal hyperaccumulation by plants is closely linked to increased protection against disease.

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 effects of low levels of nitrogen addition (7·7 and 15·4 kg N ha−1 yr−1) on plant sensitivity to biotic and abiotic stress were studied at a lowland heath in the south of England that has received N treatments since 1989. Larval growth rates and adult weights of heather beetles were found to be significantly higher when insects were reared on plants that had received additional N, with implications for insect survival and reproductive success. Electrolyte leakage measurements failed to reveal any significant impact of N addition on plant sensitivity to frost episodes in early winter. In April, however, there was some evidence of slightly decreased frost hardiness in plants receiving additional N. Accelerated spring bud burst also suggested earlier physiological activity in N-treated plots. The rate of water loss from excised shoots of Calluna vulgaris (L.) Hull was significantly faster in plants receiving additional N, although no difference in plant water potential was measured in the field after a prolonged dry spell. Whilst experimental N addition had only a small effect on plant sensitivity to abiotic stress, the relationship between enhanced deposition and increased insect performance was clear, with the potential for substantially increased insect damage at deposition rates around the critical load contributing to the formation of gaps in the Calluna canopy.