Atmospheric deposition of nitrogen is responsible for widespread changes in the structure and function of sensitive seminatural ecosystems. The proposed reduction in emissions of nitrogenous pollutants in Europe under the Gothenburg Protocol raises the question of whether affected ecosystems have the potential to recover to their previous condition and, if so, over what timescale. Since 1998, we have monitored the response of a lowland heathland in southern England following the cessation of a long-term nitrogen addition experiment, and subsequent management, assessing changes in vegetation growth and chemistry, soil chemistry and the soil microbial community. Persistent effects of earlier nutrient loading on Calluna growth and phenology, and on the abundance of lichens, were apparent up to 8 years after nitrogen additions ceased, indicating the potential for long-term effects of modest nutrient loading (up to 15.4 kg N ha À1 yr À1 , over 7 years) on heathland ecosystems. The size and activity of the soil microbial community was elevated in former N-treated plots, 6-8 years after additions ceased, suggesting a prolonged effect on the rate of nutrient cycling. Although habitat management in 1998 reduced nitrogen stores in plant biomass, effects on belowground nitrogen stores were small. Although some parameters (e.g. soil pH) recover pretreatment levels relatively rapidly, others (e.g. vegetation cover and microbial activity) respond much more slowly, indicating that the ecological effects of even small increases in nitrogen deposition will persist for many years after deposition inputs are reduced. Indeed, calculations suggest that the additional soil nitrogen storage associated with 7 years of experimental nitrogen inputs could sustain the observed effects on plant growth and phenology for several decades. Carry over effects on plant phenology and sensitivity to drought suggest that the persistence of vegetation responses to nitrogen deposition should be integrated into long-term assessments of the impact of global climate change on sensitive ecosystems.

Increases in the emissions and associated atmospheric deposition of nitrogen (N) have the potential to cause significant changes to the structure and function of N-limited ecosystems. Here, we present the results of a long-term (13 year) experiment assessing the impacts of N addition (30 kg ha(-1)  yr(-1) ) on a UK lowland heathland under a wide range of environmental conditions, including the occurrence of prolonged natural drought episodes and a severe summer fire. Our findings indicate that elevated N deposition results in large, persistent effects on Calluna growth, phenology and chemistry, severe suppression of understorey lichen flora and changes in soil biogeochemistry. Growing season rainfall was found to be a strong driver of inter-annual variation in Calluna growth and, although interactions between N and rainfall for shoot growth were not significant until the later phase of the experiment, N addition exacerbated the extent of drought injury to Calluna shoots following naturally occurring droughts in 2003 and 2009. Following a severe wildfire at the experimental site in 2006, heathland regeneration dynamics were significantly affected by N, with a greater abundance of pioneering moss species and suppression of the lichen flora in plots receiving N additions. Significant interactions between climate and N were also apparent post fire, with the characteristic stimulation in Calluna growth in +N plots suppressed during dry years. Carbon (C) and N budgets demonstrate large increases in both above- and below-ground stocks of these elements in N-treated plots prior to the fire, despite higher levels of soil microbial activity and organic matter turnover. Although much of the organic material was removed during the fire, pre-existing treatment differences were still evident following the burn. Post fire accumulation of below-ground C and N stocks was increased rapidly in N-treated plots, highlighting the role of N deposition in ecosystem C sequestration.

Indirect competition is often mediated by plant responses to herbivore feeding damage and is common among phytophagous insect species. Plant-mediated responses may be altered by abiotic conditions such as nutrient supply, which can affect plant growth, morphology, and the concentration of primary and secondary metabolites. Nutrient supply can be manipulated by the type and amount of fertilizer applied to a plant. Brassica oleracea plants were grown in several types of fertilizer, including those commonly used in sustainable and conventional agricultural systems. The occurrence of indirect competition between two phytophagous species from different feeding guilds (a phloem-feeder and leaf-chewer) was assessed. The leaf-chewer reduced aphid populations on plants growing in most fertilizer treatments, but not on those in the ammonium nitrate fertilizer treatment, which caused the highest concentration of foliar nitrogen. The potential consequences of our findings are discussed for phytophagous species in conventional and sustainable agricultural systems.

The results from three long-term field manipulation studies of the impacts of increased nitrogen deposition (0-120 kg N ha −1 yr −1 ) on lowland and upland heathlands in the UK were compared, to test if common responses are observed. Consistent increases in Calluna foliar N content and decreases in litter C:N ratios were found across all sites, while increases in N leaching were not observed at any site over the range 0-80 kg ha −1 yr −1 . However, the response of Calluna biomass did vary between sites, possibly reflecting site differences in nutrient status and management histories. Five versions of a simulation model of heathland responses to N were developed, each reflecting different assumptions about the fate and turnover of soil N. Model outputs supported the deduction from mass balance calculations at two of the field sites that N additions have resulted in an increase in immobilisation; the latter was needed to prevent the model overestimating measured N leaching. However, this version of the model significantly underestimated Calluna biomass. Model versions, which included uptake of organic N by Calluna and re-mobilisation of N from the soil organic store provided some improvement in the fit between modelled and field biomass data, but re-mobilisation also led to an overestimation of N leaching. Quantification of these processes and their response to increased N deposition are therefore critical to interpreting experimental data and predicting the long-term impacts of atmospheric deposition on heathlands and moorlands.

Plant and animal community composition changes at higher elevations on mountains. Plant and animal species richness generally declines with elevation, but the shape of the relationship differs between taxa. There are several proposed mechanisms, including the productivity hypotheses; that declines in available plant biomass confers fewer resources to consumers, thus supporting fewer species. We investigated resource availability as we ascended three aspects of Helvellyn mountain, UK, measuring several plant nutritive metrics, plant species richness and biomass. We observed a linear decline in plant species richness as we ascended the mountain but there was a unimodal relationship between plant biomass and elevation. Generally, the highest biomass values at mid-elevations were associated with the lowest nutritive values, except mineral contents which declined with elevation. Intra-specific and inter-specific increases in nutritive values nearer the top and bottom of the mountain indicated that physiological, phenological and compositional mechanisms may have played a role. The shape of the relationship between resource availability and elevation was different depending on the metric. Many consumers actively select or avoid plants based on their nutritive values and the abundances of consumer taxa vary in their relationships with elevation. Consideration of multiple nutritive metrics and of the nutritional requirements of the consumer may provide a greater understanding of changes to plant and animal communities at higher elevations. We propose a novel hypothesis for explaining elevational diversity gradients, which warrants further study; the ‘nutritional complexity hypothesis’, where consumer species coexist due to greater variation in the nutritional chemistry of plants.