The relationship between precipitation chemistry and the concentrations of nitrogen ([N]) and phosphorus ([P]) in the cushion-forming lichen Cladonia portentosa (Dufour) Coem. (=C. impexa (Harm)) was investigated. Samples of C. portentosa were collected from heathlands and upland moorlands close to 31 rural sampling stations in the UK Acid Deposition Monitoring Network, which provides data on wet deposition and NO2 concentrations in air. The [N] and [P] were measured in the top 5 mm of lichen thalli (thallus apices) and also in a horizontal stratum between 40–50 mm from the apices (thallus base). The [N] (per unit dry mass) was 0·08–1·82% and [P] was 0·04–0·17%, depending on collection site and lichen fraction analysed. Concentrations of both elements were c. 2–5 times greater in the apices than in the basal strata, and [N] and [P] values in each stratum were strongly positively correlated. Lichen [N] was positively correlated with N deposition: this relationship was stronger when using [N] values for thallus bases than for the apices. By contrast, thallus [N] was poorly correlated with [N] values in precipitation. When [NO2] in air was included together with NO3− deposition in a linear regression model explaining thallus base N, the model fit was significantly improved, whereas modelled values of NH3 deposition rate for the heathland sites did not correlate with lichen [N]. It is suggested that the proportionately greater enrichment of [N] in the thallus base might reflect a perturbation of internal recycling of thallus N at polluted sites. Thallus [P] was generally weakly linked to wet N deposition but positively correlated with [NO2] in air. It is not known whether the trend for increasing thallus [P] values indicates decreasing lichen growth rate and reduced growth dilution of P in polluted areas, or is due to regional variation in P deposition rate.

Adventitious rooting might be induced in recalcitrant woody genotypes by infection with Agrobacterium rhizogenes, and, in some cases, might also require exogenous auxin. The objective of the present study was to determine how agrobacteria trigger root formation in the stem of a recalcitrant woody microcutting, which cytological events result from the combined presence of infection and exogenous auxin, and which types of roots are induced by infection. Microcuttings of a recalcitrant walnut (Juglans regia), infected or not with A. rhizogenes strain 1855, were cultured with either indolebutyric acid (IBA), IAA, or without exogenous hormones, to induce rhizogenesis. They were cytohistologically and ultrastructurally investigated at various times in culture. Southern blot and PCR analyses were performed to verify the frequency of transgenic, chimeric and bacterium-containing roots. The infection was sufficient per se to stimulate rhizogenesis. Rooting on the infected cuttings was enhanced by exogenous IBA, which accelerated and increased root meristemoid formation, in comparison with without-hormone treatment. Meristemoids were organized both directly by the cambial cells and indirectly by the callus, and showed a pluricellular origin. Inter and intracellular bacteria were observed in the stem throughout the culture period (30 d). They were preferentially present in the vessels, and mainly in those showing polyphenol deposition. In the infected IAA-treated cultures, a high level of secondary xylem formation occurred instead of rhizogenesis. Nontransformed roots were preferentially produced by the infected cuttings treated with the auxins. Bacterium-containing and chimeric roots were produced by infected cuttings independently of the treatment. Thus, in a recalcitrant walnut, nontransformed root meristemoids are stimulated by combining infection and exogenous indolebutyric acid. Furthermore, the persistence of bacteria in the stem during the culture and the pluricellular origin of the meristemoids explain the presence of the bacterium-containing and chimeric roots.

The inter- and intracellular distribution of phosphate in non-mycorrhizal and ectomycorrhizal roots of Pinus sylvestris was analysed by energy-dispersive X-ray spectroscopy after cryofixation, freeze-drying, pressure infiltration with resin and dry cutting of the material. The results show (1) that in non-mycorrhizal roots the distribution of phosphate across the root cross section is relatively homogenous, (2) that the intracellular distribution of phosphate depends on the external supply, and (3) that the phosphate absorption can be increased by a supply of NH4. Under phosphate starvation, the phosphate is mainly localized in the cell cytoplasm, whereas under high external phosphate this element accumulates in the vacuoles of cortical cells. It can be inferred that at low external conditions phosphate is mainly located in the cytoplasm of cells to protect metabolism from deficiency, that phosphate is translocated to the shoot via symplastic and apoplastic pathways, and that especially the vacuolar pool size is regulated by the external phosphate supply. The nutrition of the host plant under phosphate starvation was improved by a mycorrhizal infection with Suillus bovinus. However, this effect might not be the same for all ectomycorrhizal infections and supply conditions. With high external phosphate an ectomycorrhizal infection with S. bovinus had no effect on intracellular phosphate within the roots, the shoot contents of mycorrhizal plants were less than those of non-mycorrhizal seedlings and translocation rate across the mycorrhizal interface appeared to be independent of the availability of phosphate. Intracellular phosphate in fungal cells of mycorrhizal roots was independent of the external supply and seemed to be regulated by the formation of polyphosphates. A supply of (NH4)2HPO4, which led to an increase in cytoplasmic phosphate levels in the hyphae of the Hartig net of the Paxillus involutus mycorrhiza also led to higher phosphate in the plant cell compartments. Therefore, it can be inferred that the translocation of phosphate from fungus to host plant across the mycorrhizal interface is regulated by the phosphate concentration in the cytoplasm of the Hartig net and by the efflux rate into the interfacial apoplast.

Ascospore germination, thallus initiation, and areole and prothallus development in the lichen Rhizocarpon lecanorinum Anders were examined using light, fluorescence and scanning electron microscopy. The ascospore germ hyphae remain very short and do not form a prothallus-like mycelium. Instead, a compact soredium-like granule develops directly from sporeling contact with a compatible species of Trebouxia. Diffuse initial stages involving non-trebouxioid algae are lacking. The onset of thallus differentiation is marked by the deposition of rhizocarpic acid in an incipient cortical layer within the apical part of the granule. As pigmentation and cortex-formation transform this structure into a typical areole, radiating prothallus hyphae are simultaneously initiated from its basal margin. Most areoles formed subsequently in the marginal prothallus lack subtending melanized hyphae and apparently stem from overgrowth by the prothallus of photobiont cells on, or in, the substratum. Apothecia reach maturity in thalli as small as 2 mm in diameter. It is proposed that the lack of diffuse hyphal growth in sporelings and telescoped morphogenesis of R. lecanorinum are part of a life history strategy geared to precocious, heavy investment in ascospore production. The R. lecanorinum–Trebouxia symbiosis has a number of features which make it well-suited for further studies of the life history and development of prothallus-forming crustose lichens with sexually reproducing mycobionts.

To investigate nitrogen assimilation in Lolium perenne L. colonized by the arbuscular mycorrhizal (AM) fungus Glomus fasciculatum (Thax. sensu Gerd.), nitrate uptake, key enzyme activities, and 15N incorporation into free amino acids were measured. After a 4-h labelling period with [15N]nitrate, 15N content was higher in roots and shoots of AM-plants than in those of control plants. Glutamine synthetase (GS) and nitrate reductase (NR) activities were increased in shoots of AM-plants, but not in roots. More label was incorporated into amino acids in shoots of AM plants. Glutamine, glutamate, alanine and γ-aminobutyric acid were the major sinks for 15N in roots and shoots of control and AM plants. Interactions between mycorrhizal colonization, phosphate and nitrate nutrition and NR activity were investigated in plants which received different amounts of phosphate or nitrate. In shoots of control plants, NR activity was not stimulated by high levels of phosphate nutrition but was stimulated by high levels of nitrate. At 4 mM nitrate in the nutrient solution, NR activity was similar in control and AM plants. We concluded that mycorrhizal effects on nitrate assimilation are not mediated via improved phosphate nutrition, but could be due to improved nitrogen uptake and translocation.

On the basis of the evidence that insect fungivory has the potential to affect fungal reproductive fitness, we investigated the effects of two specialist ciid beetles (Octotemnus glabriculus and Cis boleti) on the reproductive potential of their host fungus, Coriolus versicolor. We found, from field data, a negative correlation between the number of individuals of O. glabriculus inhabiting C. versicolor fruit bodies and the percentage of the fungal spore-producing surface (hymenium) that was functional. By contrast, the number of C. boleti inhabiting C. versicolor fruit bodies did not correlate with the percentage of functional hymenium. Experimentally, O. glabriculus and C. boleti reduced the reproductive potential of C. versicolor by 58% and 30%, respectively, whereas the combined trophic activity of both beetles caused a reduction of 64%. This latter effect was not significantly different from that caused by O. glabriculus alone. These findings disagree with previous assertions that insect fungivory on fruit bodies has only neutral effects on fungal fitness. We conclude that in the short-term, fungivory by ciids significantly decreases the area of functional hymenium of C. versicolor and is likely to reduce fungal reproductive fitness. Within this perspective the evolution of certain fungal characteristics (i.e. chemical composition, consistency and phenology) can be interpreted as being driven by fungivory.

Ozone effects on plant species mixtures could depend on the characteristics of the species involved, their mixing ratio, or on environmental conditions. Predicting long-term effects on the dynamics of plant communities requires an understanding of the interactions involved. The present experiment was designed to determine the effects of ozone on grassland species in relation to mixing ratio and soil water content (irrigation) using binary mixtures. The grass Trisetum flavescens was grown in potted replacement-series mixtures with Centaurea jacea (Experiment A) or Trifolium pratense (Experiment B). The plants were exposed to three concentrations of ozone in open-top chambers in two irrigation treatments. Total above-ground dry weight over three growth periods was measured. The competitive ability of T. flavescens was expressed as the competitive ratio (CRT). In Experiment B, total above-ground dry weight was reduced by elevated ozone and by reduced soil moisture, and significant interactions were found for ozone × irrigation and ozone × ratio. In Experiment A these effects were not significant. Under well watered conditions, CRT tended to be reduced by elevated ozone in Experiment A, but increased significantly in Experiment B, indicating the importance of the competing species in modifying the ozone effect on T. flavescens. In both experiments reduced irrigation decreased the magnitude of ozone effects on biomass production, which could be related to observed reductions in specific leaf conductance. The results suggest that under well watered conditions the effect of elevated ozone on the competitive balance between species depends on the species mixture, but that the mixing ratio is less important.

In a study on the mechanism of stimulated petiole elongation in submerged plants, oxygen concentrations in petioles of the flood-tolerant plant Rumex palustris were measured with micro-electrodes. Short-term submergence lowered petiole partial oxygen pressure to c. 19 kPa whereas prolonged submergence under continuous illumination depressed oxygen levels to c. 8–12 kPa after 24 h. Oxygen levels in petioles depended on the presence of the lamina, even in submerged conditions, and on available light. In darkness, petiole oxygen levels in submerged plants dropped quickly to values as low as 0.5–4 kPa. It is hypothesized that prolonged submergence in the light is accompanied by a decrease in carbon dioxide in the petiole. Submergence-enhanced petiolar elongation rate was compared with emergent plants. Peak daily elongation rates occurred at the end of the dark period in emergent plants, but in the middle of the light period in submerged plants. We suggest that this shift in daily elongation pattern is induced by dependence of growth on photosynthetically derived oxygen in submerged plants. Implications of reduced oxygen for ethylene production are raised. Levels of 1- aminocyclopropane-1-carboxylic acid synthase and 1-aminocyclopropane-1-carboxylic acid oxidase and ethylene sensitivity are cited as potential factors in hypoxia-induced ethylene release.

This study was conducted to determine whether xylem abscisic acid (ABA) concentration is altered by mycorrhizal symbiosis of cowpea plants grown in drying soil, and to determine whether stomatal sensitivity to xylem ABA is altered by the symbiosis. We allowed the entire root zone to dry and found that at high soil water contents (θ), mycorrhizal plants had higher stomatal conductance (gs) and lower xylem [ABA] than did non-mycorrhizal plants, but the difference disappeared at low θ, probably because of stomatal closure. The altered gs and xylem [ABA] were apparently not related to plant water status since shoot water potential, xylem sap osmotic potential and shoot water content were similar for mycorrhizal and non-mycorrhizal plants across the range of soil moisture. These differences were also not related to P nutrition or plant size. The relationship of gs to xylem [ABA] was not affected by the symbiosis, indicating that either stomatal sensitivity to xylem ABA was not affected by other xylem constituents, or that more than one xylem constituent was altered by the symbiosis but was offsetting in its effect on gs. We conclude that the symbiosis altered gs non-hydraulically, and that the factor might be xylem ABA.

Seedlings of Betula pendula Roth. were grown in the presence of Paxillus involutus (Batsch) Fr., and metabolic changes during mycorrhiza formation were examined by measuring organic acid and amino acid pools and related enzyme activities, following sequential harvests. Glutamine, aspartate and asparagine pools were always lower in infected roots than in non-infected roots, especially during Hartig net initiation and formation. Glutamate concentration was similar in both tissues. Citrate and malate were the two major organic acids detected and their concentrations were equal in infected and non-infected roots. Aspartate aminotransferase, glutamine synthetase, NAD-dependent malate dehydrogenase and glucose-6-phosphate dehydrogenase activities were higher in infected roots than non-infected roots. For all enzymes revealed on polyacrylamide gels, both root and fungal isoforms were present in infected roots. Quantitative changes in enzyme capacities and metabolite pools indicated that mycorrhiza formation caused a re-arrangement of the main metabolic pathways during the very early stages following contact, which might be related to the structural changes.

The photosynthetic and respiratory activities of the leaves of a tropical tree, cacao, Theobroma cacao L., grown under shading, were determined in relation to leaf age and light conditions within the canopy, in order to gain a deeper insight into characteristics of the leaf population composing a canopy.

The specific leaf weight and leaf water content varied with the height of leaves from the ground and/or the irradiance. The net photosynthetic rate also varied markedly depending on leaf age and irradiance. The leaves in full or nearly full irradiance (<70% full daylight) attained the maximum rate c. 60 d after the leaves emerged, and the rate decreased to nearly zero at a leaf age of c. 270 d.

The nocturnal leaf respiratory activity was proportional to the photosynthetic activity during the day, but the proportional ratio was not constant throughout the survival period of leaves. Within the closed canopy, the upper leaves, which much higher ratios of respiratory to photosynthetic activity, under full or nearly full irradiance, tended to have much shorter mean longevities (c. 160 d) than the lower leaves under a lower irradiance (c. 310 d).

Two parameters were applied to estimate the degree of stress induced by exposure of the lichen Ramalina duriaei (De Not.) Bagl. to air pollution in urban-industrial sites in Israel. Changes in the spectral reflectance response of the thallus and in the production of stress-ethylene, were compared with the concentration of sulphate-S, V, Ni, K, Pb, P, Cu, Cr, Fe, Mn and Mg in either in situ or in transplanted thalli of the lichen.

The concentrations of Cu, Mg, Fe, Ni, Pb, V, Cr, Mn and sulphate-S in in situ thalli collected in a nature reserve in an industrial town (Ashdod, SW Israel) were higher than those detected in in situ thalli collected in a forest (HaZorea Forest) in the north-east of Israel. The high concentrations of these mineral elements in R. duriaei from the nature reserve coincided with a high rate of production of stress-ethylene and with a low normalized difference vegetation index (NDVI) indicating that changes in the spectral reflectance response relate to damage to the chlorophyll.

Lichens collected in the unpolluted forest in February 1994, either resuspended in the same site or transplanted in nine sites in the Ashdod region, and retrieved in November of the same year, contained a higher concentration of sulphate-S, Pb, Cu, Ni, V and Cr in the urban-industrial sites than in most of the rural localities. Statistical analysis revealed a positive correlation between the amount of ethylene produced by R. duriaei and the concentration of V and Ni in the same thalli.

To test the integrity of the ethylene-producing system, lichens were soaked in either 5 mM or 10 mM FeCl2 at pH 3·7. Thalli exposed to a very polluted environment produced less ethylene after this treatment. The NDVI correlated positively with the concentration of K in the thallus but negatively with the concentration of Mn, Ni, Pb and sulphate-S in the same material.

It may be concluded that the in situ thalli of R. duriaei in the nature reserve are severely stressed and possibly endangered by the pollution produced by the combustion of heavy fuel oil in the local power plant and oil refineries and by acid rain.

After cooling the roots and shoot apex of barley, there is a reduction in sucrose export from, and an increased total accumulation of carbohydrate and induction of fructan synthesis in, source leaves. We investigated carbohydrate accumulation in individual leaf cells. Using the microsampling technique and microfluorometric enzymatic assay, the amounts of various carbohydrates were measured in mesophyll, bundle sheath and epidermal cells. Epidermal cells take no part in carbohydrate partitioning and have very low concentrations of sugars even when leaf export is reduced. Even without cooling, however, there are remarkable changes in sucrose concentration (up to 200 mM) in mesophyll and bundle sheath cells during the photoperiod. The activity of acid invertase in samples from individual cells was very low, c. 3 μmol sucrose (g f. wt)−1 h−1. Interrelationships between these cell types and the control of fructan metabolism are discussed.

Hordeum spontaneum shows a large genetic variation and occupies a wide range of different habitats. The aim of this study was to quantify variation in growth characteristics of H. spontaneum from different sites in Israel and to relate this variation to different environmental conditions. To this end, 84 accessions of 21 populations were grown in a growth chamber in near-optimal conditions and a range of physiological, morphological, allocation- related and chemical characteristics were measured. These parameters included rates of photosynthesis, shoot and root respiration, specific leaf area, biomass allocation and seed mass. Averaged over all traits variation explained by differences between populations was 26%, between accessions 21%, whereas that within accessions was 53%. By contrast with most genetic studies, we found variation between populations larger than between accessions. The largest between-population variation (46%) was for morphological traits. In particular, seed mass, leaf thickness and leaf width differed strongly between populations. Variation in growth characteristics between populations was poorly related to mean annual rainfall, mean humidity or January temperature at the sites of origin. We expect that differences between populations to be larger and correlation with environmental parameters stronger in plants grown in stressful conditions. According to our study, seed mass is more important than relative growth rate in determining variation in early plant biomass in H. spontaneum.

To predict changes in the range and density of plant species as a consequence of elevated atmospheric CO2, it is essential to characterize the effect of elevated CO2 on key components of plant life history stages such as rate of establishment and maturation of individuals, reproductive output and offspring fitness. We measured vegetative response, phenology, reproductive output and seed quality of four annual C3 plant species, grown from seed to senescence under ambient and elevated (ambient+200 ppm) CO2 and ambient and reduced (33% shade) light. Few previous studies have included all stages of a plant's life as well as characteristics of the next generation. Elevated CO2 had no effect on the vegetative attributes of Cardamine hirsuta, Spergula arvensis or Poa annua whereas Senecio vulgaris produced longer leaves and greater biomass. Both Senecio and Poa had faster maturation times. The vegetative response of Senecio was not translated into increased seed output, although seed mass and carbon[ratio ]nitrogen ratios were significantly increased. By contrast, Poa showed no vegetative response to elevated CO2, but had significantly increased seed production. Thus we found no evidence for a simple translation from vegetative response to elevated CO2 into reproductive response. There was also no consistent light-mediated response to elevated CO2 among the four species. However, the effect of reduced light (33% shade) on vegetative and reproductive output was consistent across the four species and significantly stronger than the effect of elevated CO2. On the basis of this glasshouse study, we predict that Poa would be most likely of the four species to show significant increases in population size and migration potential, as a result of increased reproductive output, under elevated atmospheric CO2. However, this response may be relatively small compared with variation in growth and reproduction as a result of environmental heterogeneity in resources such as light.

Growth responses of two provenances of European beech (Fagus sylvatica) were studied. The seedlings were grown in closed-top chambers at four temperature regimes (−2.9 °C below ambient, ambient, +2.3 °C and +4.8 °C above ambient) in combination with two CO2 partial pressures (40 Pa and 74 Pa). Growth was followed by making destructive harvests c. every 25 d from germination in early June to senescence in late September. Allocation patterns were significantly affected by the temperature regimes. However, changes in dry matter allocation and morphology associated with the different treatments at a given time were mostly a result of differences in tree size. Temperature regimes only had a significant effect on the relative growth rate, RGR, at the beginning of the experiment. In contrast to temperature, high [CO2] increased RGR throughout the experiment when compared with plants of equal size. As the trees increased in size net assimilation rate, NAR, decreased but the effect of [CO2] on both NAR and RGR had a tendency to increase. Increases in NAR caused by elevated [CO2] were partly counteracted by reductions in the leaf area ratio, LAR. Reductions in LAR were caused by concomitant reductions in specific leaf area, SLA, whereas the level of [CO2] did not significantly affect leaf weight ratio, LWR, nor other dry weight ratios. The interactions between temperature and [CO2] are highly dependent on whether they are expressed as instantaneous values for plants at a common age or instantaneous values at a common size (and thereby extracting the effects of ontogenetic drift). When comparing instantaneous values at common sizes, the positive effect of [CO2] on RGR increased with plant size in every temperature regime. This also occurred in every temperature regime when comparing plants of equal age but the response to [CO2] was less. The effect of [CO2] on RGR was dependent on growth temperature. The positive effects of elevated [CO2] on RGR were less than the positive effect on photosynthesis. The two provenances did not differ significantly in the response of RGR to [CO2] which is in agreement with measurements of photosynthesis.

Patterns produced by foraging mycelial systems of decomposer fungi might be the visual evidence of strategies for obtaining differently distributed carbon sources. Mycelial systems might also be responsive to unevenly distributed mineral nutrients in the soil. The nature of the mycelial response might differ between species inhabiting eutrophicated disturbed sites, such as Stropharia caerulea (Kriesel), and inhabitants of more typical woodland soils, such as Phanerochaete velutina DC.: (Pers.) Parmasto. We examined the effect of enrichment of soil by addition of different concentrations of N, P and potassium on inoculum decay rate and on extension rate, biomass production and fractal dimension (determined by image analysis) of extra-resource mycelia of S. caerulea and P. velutina extending from wood blocks across compacted non-sterile soil in trays.

S. caerulea was more responsive than P. velutina to both type and quantity of soil nutrient enrichment. P. velutina grew more rapidly, produced more biomass and decayed inocula more rapidly than S. caerulea under most conditions, rates of growth and resource decay of the former remaining stable over a range of soil nutrient enrichments. In contrast, morphology of S. caerulea altered on enriched soils compared to controls with greater and more pronounced fanning (divergence of separate hyphae) of the foraging mycelial front. These changes were reflected in fractal values. Extension rate and biomass production of S. caerulea was less than controls on N-amended soil, but greater on P-amended soils, with the greatest biomass produced on 0·13–0·27 mg P g−1 air-dried soil. Decay rate of inocula was often positively related with mycelial production rates. These responses indicate extra-resource mycelium of S. caerulea is able to respond rapidly to a range of soil nutrient enrichments, and this is discussed in terms of the different foraging strategies exhibited by the two fungi.

Recent research in whole-plant stomatal physiology, conducted largely with potted plants in controlled environments, suggests that stomatal conductance (gs) might be more closely linked to plant chemical variables than to hydraulic variables. To test this in a field situation, seasonal gs was examined in relation to a number of plant and environmental variables in 11 temperate, deciduous forest tree species. Stomatal conductance was generally better correlated with environmental variables (air temperature, vapor pressure deficit, PPFD) than with plant variables, and slightly better correlated with plant hydraulic variables (shoot water and osmotic potentials) than with plant chemical variables (xylem sap ABA concentration, xylem sap pH). We examined a model, developed previously for maize, which describes regulation of gs by xylem sap ABA concentration with leaf water status acting to modify stomatal sensitivity to the ABA signal. This model explained slightly more variation in seasonal gs in the forest trees than did single plant variables but not more variation than most single environmental variables. Response surface models, especially those incorporating environmental variables, were more consistently successful at explaining gs across species.

Impacts of defoliation on the growth and physiology of sugar maple (Acer saccharum Marsh.) and trembling aspen (Populus tremuloides Michx.) were examined in ambient and CO2-enriched atmospheres. Saplings were grown for 70 d in controlled environments, wherein CO2 mole fractions averaged either 356 μmol mol−1 or 645 μmol mol−1, under a PPF of 500 μmol m−2 s−1. On day 49 of the study, 50% of the leaf area was removed from a subset of each species in both CO2 environments. Relative growth rate (rgr) and its physiological and morphological determinants were monitored before and after defoliation. For non-defoliated saplings of both species, a slight stimulation of rgr (c. 5%) in elevated CO2 led to a modest increase (9–11%) in final sapling weight. In the case of maple, the minimal growth response corresponded with minor CO2 effects on specific leaf area (sla) and leaf weight ratio (lwr), and an apparent CO2-induced down-regulation of photosynthetic metabolism. For aspen, the CO2 stimulation of photosynthesis was largely offset by a decrease in sla. Responses to defoliation differed markedly between species and CO2 environments. Defoliation decreased maple rgr in ambient CO2, whereas the opposite occurred in elevated CO2. The latter led to complete recovery of plant weight (compensation), and was attributed to a defoliation-induced increase in carbon allocation to new leaves, along with a reversal of photosynthetic CO2 acclimation in that foliage. In both environments, aspen rgr increased after defoliation, facilitating almost full compensation. Defoliation increased light penetration into the aspen canopy, and it was estimated that the resultant stimulation of photosynthesis in lower leaves would have more than offset the concomitant decrease in lwr. CO2 enrichment might substantially enhance the ability of certain tree species to recover from herbivory. Moreover, responses to elevated CO2 might be largest in the presence of stresses, such as herbivory, that decrease plant source[ratio ]sink ratios.

The physiological potential for acquisition of atmospheric ammonia (NH3) was investigated in three European meadow grasses (Arrhenatherum elatius, Bromus erectus and Brachypodium pinnatum) competing in chalk grasslands. Experiments were carried out with plants cultivated for about three months on a soil–sand mixture at high root nitrogen supply or in nutrient solutions at low root nitrogen supply. Two different root nitrogen regimes were applied to the solution-grown plants: 130 μmol NO3 plant−1 wk−1 (approx. 50 kg nitrogen ha−1 in three months); or 130 μmol NO3− plus 130 μmol NH4+ plant−1 wk−1. Each regime was combined with two levels of NH3 fumigation (0 and 70 nmol mol−1 air for 24 d). Uptake of gaseous NH3 in the shoots was investigated under controlled environmental conditions including NH3 concentrations ranging from 0 to 30 nmol mol−1 air. Concurrently, photosynthesis, glutamine synthetase activity, nitrogen allocation, biomass allocation and apoplastic cation composition were measured. For A. elatius, the influence of photorespiration on NH3 acquisition was also assessed. Independently of plant nitrogen status, ammonia compensation points in A. elatius and B. erectus plants were <0.5 nmol mol−1. The total leaf conductance to NH3 absorption remained constant at increasing NH3 concentrations, showing that the capacity for assimilation was unaltered. Whereas internal factors in the leaves did not cause differences in the potential for NH3 acquisition between the species, other factors of NH3 acquisition were quite different: B. erectus had higher stomatal conductance and, thus, higher NH3 uptake rates per unit leaf area compared to A. elatius and B. pinnatum; higher stomatal conductances of B. erectus were to a large extent offset by a lower leaf area per plant, resulting from a lower growth rate and thicker leaves than in the two other species. The rate of photorespiration in Arrhenatherum constituted at least 15% of the net photosynthetic rate. Surprisingly, suppression of photorespiration indicated that NH3 uptake was supported by photorespiration. Bromus responded to fumigation with 70 nmol NH3 mol−1 air for 24 d by lowering the root[ratio ]shoot ratio and increasing the nitrogen concentration in the stem dry matter. The total leaf conductance to NH3 uptake decreased in all three species upon exposure to NH3, while the stomatal conductance was unaffected. The NH3 exposure caused lower apoplastic concentrations of H+, Mg2+ and Ca2+ in A. elatius and B. erectus.