Internal cephalodia of Lobaria pulmonaria were examined in two British locations, one in western Scotland (Isle of Skye) and the other in England (Cumbria, Lake District), sites which were predicted to have contrasting levels of fixed nitrogen deposition. Cephalodia were found to occupy a small proportion of the total biomass averaging (<1%) in both areas. Modelled nitrogen deposition in the sites in Cumbria was 3–6 times higher than in the Skye sites but the cephalodium relative biomass was not significantly different. It is suggested that local climatic and topographic effects might have reduced the estimated rate of nitrogen deposition at the former sites.

The aim of this investigation was to determine the NH4Cl concentration threshold, above which negative physiological effects would occur in the nitrophytic lichen Xanthoria parietina. Over a 10 month period, X. parietina thalli growing on roof tiles were exposed weekly to NH4Cl concentrations of 0·04, 0·17, 0·34 or 0·69 M. Nitrogen (N) uptake from ammonium and the concentrations of total thallus N and biont markers (chlorophyll a, ergosterol and chitin) were measured on four occasions, over the experimental period. Xanthoria parietina was able to assimilate a significant quantity of the applied ammonium. However, lichens exposed to the two higher concentrations suffered damage to both the photobiont and the mycobiont, as evidenced by reduced chlorophyll a and ergosterol concentrations, while lichens exposed to the two lower concentrations showed no significant changes in either chlorophyll a or ergosterol that could be related to the ammonium inputs. Xanthoria parietina tolerated a weekly irrigation of at least 0·17 M NH4Cl, corresponding to an N deposition of c. 1000 kg ha−1 yr−1, suggesting that this species has a very high tolerance to N pollution.

Temperate forests are recipients of anthropogenic nitrogen (N) deposition. Because growth in these ecosystems is often limited by N availability, elevated N inputs from the atmosphere can influence above- and belowground production in forests. Although fine-root production is the largest component of belowground production in forests, it is unclear whether or how increases in Navailability to forest trees accompanying increased N deposition might influence fine-root growth. Uncertainties as to how fine-root dynamics (i.e. production and turnover) vary in relation to soil N availability contribute to this problem. Although fine-root biomass typically decreases along soil N availability gradients in forests, it is unclear whether fine-root production and turnover also decrease along these gradients. Here, four possible relationships between fine-root turnover, fine-root production, and forest soil N availability are evaluated to develop a general hypothesis about changes in rooting dynamics that might accompany increases in N deposition. The four possible relationships are as follows. (1) Fine-root turnover rates do not systematically change with N availability in forest soils. If this is true, then fine-root production rates decrease with fine-root biomass in relation to soil N availability, and increased N deposition could lead to decreased fine-root production in forests. (2) Decreases in photosynthate allocation belowground along N availability gradients will function to slow fine-root turnover (or increase life span) as N availability increases with N deposition, thereby dramatically decreasing fine-root production. (3) Fine-root production might increase with N availability even though fine-root biomass typically decreases with N availability. This could occur if fine-root metabolism and turnover increase (life span decreases) with soil N supply. Increases in fine-root production accompanying increases in N availability, if large enough, could result in constant proportions of forest production being allocated to fine roots as soil N availability increases with N deposition. (4) Although fine-root turnover and production might both increase as N becomes more available to tree roots, the proportional allocation of total primary production to fine roots could decrease. Identifying the most likely of these four possibilities requires intersite comparisons of forest root dynamics along gradients of soil N availability and N deposition. Collective results of studies that use sequential sampling of fine-root biomass to estimate production suggest that fine-root turnover and production either; do not vary systematically, or that they decrease as N availability increases. By contrast, studies using ecosystem C or N budgets suggest that fine-root turnover and production both increase with N availability and that similar increases might be expected with elevated N deposition. It is argued here that assumptions underlying most biomass-based estimates of fine-root production are more suspect than are assumptions underlying element budget-based estimates. If so, it is likely that N deposition will function to decrease forest fine-root biomass but to stimulate fine-root turnover and production. However, increases in fine- root turnover and production could eventually decrease if chronically elevated N deposition leads to forest stand mortality.

Changes in the production and turnover of roots in forests and grasslands in response to rising atmospheric CO2 concentrations, elevated temperatures, altered precipitation, or nitrogen deposition could be a key link between plant responses and longer-term changes in soil organic matter and ecosystem carbon balance. Here we summarize the experimental observations, ideas, and new hypotheses developed in this area in the rest of this volume. Three central questions are posed. Do elevated atmospheric CO2, nitrogen deposition, and climatic change alter the dynamics of root production and mortality? What are the consequences of root responses to plant physiological processes? What are the implications of root dynamics to soil microbial communities and the fate of carbon in soil? Ecosystem-level observations of root production and mortality in response to global change parameters are just starting to emerge. The challenge to root biologists is to overcome the profound methodological and analytical problems and assemble a more comprehensive data set with sufficient ancillary data that differences between ecosystems can be explained. The assemblage of information reported herein on global patterns of root turnover, basic root biology that controls responses to environmental variables, and new observations of root and associated microbial responses to atmospheric and climatic change helps to sharpen our questions and stimulate new research approaches. New hypotheses have been developed to explain why responses of root turnover might differ in contrasting systems, how carbon allocation to roots is controlled, and how species differences in root chemistry might explain the ultimate fate of carbon in soil. These hypotheses and the enthusiasm for pursuing them are based on the firm belief that a deeper understanding of root dynamics is critical to describing the integrated response of ecosystems to global change.

Regular (monthly) additions of NH4NO3 (4–12 g N m−2 yr−1) were made over a period of 8 yr (1989–98) to areas of moorland in North Wales dominated by the ericaceous shrub Calluna vulgaris. Results from the early stages of the experiment (1990–94) have shown marked and dose-related increases in shoot extension and canopy height in response to the nitrogen treatments, with significantly higher shoot nitrogen contents. The nitrogen-related stimulation in the growth of the C. vulgaris canopy over this period has resulted in large accumulations of litter on the high-nitrogen-treated plots (6.6 kg m−2 in plots treated with 12 g N m−2 yr−1, compared with 3.8 kg m−2 for the water controls). Litter nitrogen concentrations were also significantly increased at the higher rates of nitrogen addition, leading to a doubling of the total return of nitrogen to the litter layer over the experimental period. These changes in vegetation structure were associated with large reductions in the abundance of the bryophyte and lichen species normally present under the untreated canopy. Results since 1994, however, show little increase in shoot extension in response to the nitrogen treatments, with no clear dose response to increasing levels of addition. These findings are associated with a dose-related increase in the susceptibility of the nitrogen-treated areas of the C. vulgaris canopy to late winter injury, characterized as browning of the shoot tips in early to late spring. These results indicate that deleterious effects are now accumulating as a result of the long-term addition of nitrogen to these moorland plots.

A field and laboratory based bioassay has been developed to investigate the effects of the quantity and duration of simulated pollutant nitrogen (N) deposition on root-surface phosphomonoesterase (PME) activities in calcareous and acid grasslands. Seedlings of Plantago lanceolata were transplanted to a calcareous grassland and Agrostis capillaris seedlings were grown in microcosms containing soil from an acid grassland that had received either 7 yr (long-term) N additions or 18 months (short-term) N and phosphorus (P) additions. The bioassay revealed that short-term N treatments had little effect on the enzyme activity, whereas long-term N additions significantly increased PME activity within 7 d of transplanting into the field plots. Root-surface PME activity of A. capillaris was significantly reduced in soil that received additions of P. In the plots receiving long-term additions of N, a strong relationship was observed between extractable soil ammonium and root-surface PME activity. Soil ammonium concentrations accounted for 67% of the variation in PME activity of P. lanceolata in the calcareous grassland, and 86% of the variation in PME activity of A. capillaris in the acid grassland. These results provide evidence that N deposition may have considerable effects on the demand and turnover of P in ecosystems that are approaching or have reached N saturation.

Responses of leaf gas exchange and above-ground growth of beech (Fagus sylvatica L.) and Norway spruce (Picea abies Karst.) to atmospheric CO2 enrichment (374 μl l−1 vs. 590 μl l−1) and increased wet deposition of N (5 vs. 50 kg N ha−1 a−1) in combination with two natural forest soil types (‘acidic’ and ‘calcareous’) were studied in large open-top chambers. Eight juvenile beech and spruce trees from different provenances, together with a ground cover composed of five understorey species, were established in each of 32 model ecosystems. Both beech and spruce showed sustained enhancement of photosynthesis in response to atmospheric CO2 enrichment during the first 2 yr of treatment. Nevertheless, switching measurement CO2 concentrations revealed partial downward adjustment of photosynthesis in trees grown in elevated CO2, beech generally showing more pronounced downward adjustment than spruce. The responsiveness of photosynthesis to CO2 enrichment did not vary significantly among trees from different provenances. Stomatal conductance was reduced under elevated CO2 in both tree species. In spruce, the radial growth of the main stem and the annual production of wood (shoot-wood dry mass of current-year lateral shoots), needle dry mass, and assimilation area per tree were stimulated both by CO2 enrichment and increased N deposition, but were not significantly affected by soil type by year 2. In contrast, in beech, the radial growth of the stem and the total leaf number, foliage dry mass, and assimilation area per tree were all not significantly affected by elevated CO2 and increased N deposition when responses of the two soil types were pooled, but were greater on calcareous than on acidic soil by year 2. However, CO2 interacted with soil type in beech: irrespective of the N deposition rate, saplings showed growth stimulation on the calcareous soil but responded negatively to CO2 enrichment on the acidic soil (where growth was slower). Our results suggest that complex interactions between CO2, species and soil quality need to be accounted for when attempting to predict forest development in a future CO2-rich world.

Evidence that enhanced reactive nitrogen deposition is affecting semi-natural terrestrial ecosystems comes from historic increases in plant tissue N concentrations, correlations between tissue N concentrations and present-day total atmospheric N deposition, changes in plant amino-acid composition and effects on N assimilation. The ecological significance of such changes in biomarkers is uncertain. This paper explores the ecological significance of reactive atmospheric N deposition through a review of previous experimental findings and new experimental evidence from an acidic and a calcareous grassland, both showing phosphorus limitation, and a N-limited Calluna vulgaris (L.) Hull heathland in upland Britain. Nitrogen addition in the range 0–20 g N m−2 yr−1 initially (years 0–4) increased the growth of Calluna and a decline in some subordinate species. In subsequent years, shoot extension was not stimulated, but winter injury was observed from 1993 onwards, suggesting a strong interaction between N supply and climatic conditions. By contrast, the grasslands showed a small decrease in the cover of higher plants in later years (6–7) of the experimental treatments (0–14 g N m−2 yr−1) and no growth stimulation. All N treatments reduced the bryophyte cover in the acidic grassland. There were marked effects on below-ground processes, including a sustained stimulation of N mineralization in the grassland soils, and an increase in the bacterial utilization of organic substrates in the heathland, as measured in BIOLOG plates. The results strongly suggest the importance of atmospheric N deposition on microbially driven processes in soils, and are discussed in relation to the scale of potential ecosystem changes and their reversibility by pollution abatement.

Because nitrogen is the mineral nutrient needed in largest amounts by plants, it is usually also the limiting factor for plant growth in terrestrial ecosystems (Vitousek & Howarth, 1991). Consequently, the deposition of oxidized and reduced N compounds will almost invariably have large effects in these systems, and because N availability not only regulates plant growth but also that of organisms at other trophic levels, disturbances of several ecosystem processes might occur. The alternations introduced by deposition of atmospheric N compounds are both of a quantitative and of a qualitative nature. Moreover, N deposition can have phytotoxic as well as growth-stimulating effects.

This short commentary gives a personal view of some of the possible consequences of N deposition on plants. It refers particularly to the oral presentations given by Professor Heinz Rennenberg and Dr Marta Peréz-Soba, and to the discussions held after their talks where appropriate. Separate attention is given to four different consequence of anthropogenic N-deposition: N-availability; N-form, N-uptake by the shoot, and the period of N-uptake. Finally, I have tried to adopt an ecosystem perspective and discuss briefly the concept of critical loads of N.

As a result of increasing anthropogenic nitrogen deposition, N availability in many forest ecosystems, which are normally N-limited, has been enhanced. We discuss the impacts of this increased N availability on the ectomycorrhizal (ECM) symbiosis which is generally regarded as an adaptation to nutrient limited conditions. Nitrogen deposition can influence fruit-body formation by ECM fungi, the production and distribution of the extraradical mycelium in the soil and the formation of ectomycorrhizas.

Available data from long-term N deposition studies indicate that the most prominent effects might be discernible above-ground (i.e. on the formation of fruit bodies). ‘Generalist’ species, forming a symbiosis with a wide range of tree species, seem to be less affected by increased N availability than ‘specialist’ species, especially those living in symbiosis with conifers. However, the importance of below-ground investigations to determine the impacts of N deposition on the ECM symbiosis must not be underestimated. Culture experiments show an optimum N concentration for the formation of extraradical mycelium and mycorrhizas. Often, negative effects only become visible at comparatively high N concentrations, but the use of a few easily cultivated species of ECM fungi, which are adapted to higher N concentrations, undermines our ability to generalize.

So far, N deposition experiments in the field have only shown minor changes in the below-ground mycorrhizal population, as estimated from the investigation of mycorrhizal root tips. However, effects on the ECM mycelium, which is the main fungal component in terms of nutrient uptake, cannot be excluded and need further consideration.

Because the photoassimilate supply from the plant to the fungal partner is crucial for the maintenance of the ECM symbiosis, we discuss the possible physiological implications of increasing N inputs on the allocation of C to the fungus. Together with ultrastructural changes, physiological effects might precede obvious visible changes and might therefore be useful early indicators of negative impacts of increasing N inputs on the ECM symbiosis.

Man's activities pose a number of threats to the functioning, structure and diversity of natural and semi-natural ecosystems. One of the main threats is the increase in concentrations in air pollutants in this century (Wellburn, 1988; Tamm, 1991). This paper is a commentary on the effects of tropospheric ozone (O3) and airborne nitrogen deposition (both oxidized (NOx) and reduced (NHy) compounds) on natural and semi-natural ecosystems, based upon the oral presentations and the discussions during the Symposium, extended with a personal overview and some suggestions about future challenges for research. The most important effects of these air pollutants on natural and semi-natural vegetation are summarized and evaluated in ecological terms, with respect to the functioning and structure of unaffected systems. Air pollutants are transported over both short and long distances (as far as a few thousand km) before being deposited on surface water, vegetation or soil. In this way, vegetation over a large area or in remote regions can be influenced by airborne pollutants (see Fowler et al. (1998); Asman, Sutton & Schjørring (1998)).

The global cycles of carbon and nitrogen are being perturbed by human activities that increase the transfer from large pools of non-reactive forms of the elements to reactive forms that are essential to the functioning of the terrestrial biosphere. The cycles are closely linked at all scales, and global change analyses must consider C and N cycles together. The increasing amount of N originating from fossil fuel combustion and deposited to terrestrial ecosystems as nitrogen oxides could increase the capacity of ecosystems to sequester C, thereby removing some of the excess carbon dioxide from the atmosphere and slowing the development of greenhouse warming. Several global and ecosystem models have calculated the amount of C sequestration that can be attributed to N deposition, based on assumptions about the allocation of N among ecosystem components with different C[ratio ]N ratios. They support the premise that, since industrialization began, N deposition has been responsible for an increasing terrestrial C sink, but there is great uncertainty whether ecosystems will continue to retain exogenous N. Whether terrestrial ecosystems continue to sequester additional C will depend in part on their response to increasing concentrations of atmospheric carbon dioxide, widely thought to be constrained by limited N availability. Ecosystem models generally support the conclusion that responses to increasing concentrations of carbon dioxide will be greater, and the range of possible responses will be wider, in ecosystems where increased N inputs originate as atmospheric deposition. The interactions between N deposition and increasing carbon dioxide concentrations could be altered considerably, however, by additional factors, including N saturation of ecosystems, changes in community composition, and climate change. Nitrogen deposition is also linked to global change issues through the volatile losses of nitrous oxide, which is a potent greenhouse gas, and the role of nitrogen oxides in the production of tropospheric ozone, which could interact with plant responses to elevated carbon dioxide. Any consideration of the role of N deposition in global change issues must also balance the projected responses against the serious detrimental impact of excess N on the environment.

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.