The restoration of phytophagous beetles in species-rich chalk grasslands

This study focuses on the restoration of chalk grasslands over a 6-year period and tests the efficacy of two management practices, hay spreading and soil disturbance, in promoting this process for phytophagous beetles. Restoration success for the beetles, measured as similarity to target species-rich chalk grassland, was not found to be influenced by either management practice. In contrast, restoration success for the plants did increase in response to hay spreading management. Although the presence of suitable host plants was considered to dictate the earliest point at which phytophagous beetles could successfully colonized, few beetle species colonized as soon as their host plants became established. Morphological characteristics and feeding habits of 27 phytophagous beetle species were therefore tested to identify factors that limited their colonization and persistence. The lag time between host plant establishment and colonization was greatest for flightless beetles. Beetles with foliage-feeding larvae both colonized at slower rates than seed-, stem-, or root-feeding species and persisted within the swards for shorter periods. Although the use of hay spreading may benefit plant communities during chalk grassland restoration, it did not directly benefit phytophagous beetles. Without techniques for overcoming colonization limitation for invertebrate taxa, short-term success of restoration may be limited to the plants only.

Ecosystem services and food production

By 2030, the world’s human population could rise to 8 billion people and world food demand may increase by 50%. Although food production outpaced population growth in the 20th century, it is clear that the environmental costs of these increases cannot be sustained into the future. This challenges us to re-think the way we produce food. We argue that viewing food production systems within an ecosystems context provides the basis for 21st century food production. An ecosystems view recognises that food production systems depend on ecosystem services but also have ecosystem impacts. These dependencies and impacts are often poorly understood by many people and frequently overlooked. We provide an overview of the key ecosystem services involved in different food production systems, including crop and livestock production, aquaculture and the harvesting of wild nature. We highlight the important ecosystem impacts of food production systems, including habitat loss and degradation, changes to water and nutrient cycles across a range of scales, and biodiversity loss. These impacts often undermine the very ecosystem services on which food production systems depend, as well as other ecosystem services unrelated to food. We argue that addressing these impacts requires us to re-design food production systems to recognise and manage the limitations on production imposed by the ecosystems within which they are embedded, and increasingly embrace a more multifunctional view of food production systems and associated ecosystems. In this way, we should be able to produce food more sustainably whilst inflicting less damage on other important ecosystem services.

Individual biomass factors for beech, oak and pine in Slovakia: a comparative study in young naturally regenerated stands

Biomass conversion and expansion factors (BCEF) which convert tree stem volume to whole tree biomass and biomass allocation patterns in young trees were studied in order to estimate tree and stand biomass in naturally regenerated forests. European beech (Fagus sylvatica L.), Sessile oak (Quercus petraea (Mattuschka) Liebl.) and Scots pine (Pinus sylvestris L.) stands were compared. Seven forest stands of each species were chosen to cover their natural distribution in Slovakia. Species specific BCEF are presented, generally showing a steep decrease in all species in the smallest trees, with the only exception in the case of branch BCEF in beech which grows with increasing tree size. The values of BCEF for all tree compartments stabilise in all species once trees reach about 60-70mm diameter at base. As they grow larger, all species increase their allocation to stem and branches, while decreasing the relative growth of roots and foliage. There are, however, clear differences between species and also between broadleaves and conifers in biomass allocation. This research shows that species specific coefficients must be used if we are to reduce uncertainties in estimates of carbon stock changes by afforestation and reforestation activities.

Biomass partitioning and growth efficiency in four naturally regenerated forest tree species

Current forest growth models and yield tables are almost exclusively based on data from mature trees, reducing their applicability to young and developing stands. To address this gap, young European beech, sessile oak, Scots pine and Norway spruce trees approximately 0 to 10 years old were destructively sampled in a range of naturally regenerated forest stands in Central Europe. Diameter at base and height were first measured in situ for up to 175 individuals per species. Subsequently, the trees were excavated and dry biomass of foliage, branches, stems and roots was measured. Allometric relations were then used to calculate biomass allocation coefficients (BAC) and growth efficiency (GE) patterns in young trees. We found large differences in BAC and GE between broadleaves and conifers, but also between species within these categories. Both BAC and GE are strongly age-specific in young trees, their rapidly changing values reflecting different growth strategies in the earliest stages of growth. We show that linear relationships describing biomass allocation in older trees are not applicable in young trees. To accurately predict forest biomass and carbon stocks, forest growth models need to include species and age specific parameters of biomass allocation patterns.

Trends in the start of the wet season over Africa

A quarter of a century of daily rainfall data from the Global Telecommunications System are used to define the temporal and spatial variability of the start of the wet season over Africa and surrounding extreme south of Europe and parts of the Middle East. From 1978 to 2002, the start of the wet season arrived later in the year for the majority of the region, as time progressed. In some parts of the continent, there was an annual increase in the start date of up to 4 days per year. On average, the start of the wet season arrived 9–21 days later from 1978 to 2002, depending on the threshold used to define the start of the rains (varying from 10–30 mm over 2 days, with no dry period in the following 10 days). It is noted that the inter-annual variability of the start of the wet season is high with the range of start dates varying on average from 116 to 142 days dependent on the threshold used to determine the start date. These results may have important implications for agriculturists on all levels (from the individual farmer to those responsible for regional food supply), as knowledge of potential future climate changes starts to play an increasingly important role in the agricultural decision-making process, such as sowing and harvesting times.

Technologies for biological containment of GM and Non-GM crops

International Perspective The development of GM technology continues to expand into increasing numbers of crops and conferred traits. Inevitably, the focus remains on the major field crops of soybean, maize, cotton, oilseed rape and potato with introduced genes conferring herbicide tolerance and/or pest resistance. Although there are comparatively few GM crops that have been commercialised to date, GM versions of 172 plant species have been grown in field trials in 31 countries. European Crops with Containment Issues Of the 20 main crops in the EU there are four for which GM varieties are commercially available (cotton, maize for animal feed and forage, and oilseed rape). Fourteen have GM varieties in field trials (bread wheat, barley, durum wheat, sunflower, oats, potatoes, sugar beet, grapes, alfalfa, olives, field peas, clover, apples, rice) and two have GM varieties still in development (rye, triticale). Many of these crops have hybridisation potential with wild and weedy relatives in the European flora (bread wheat, barley, oilseed rape, durum wheat, oats, sugar beet and grapes), with escapes (sunflower); and all have potential to cross-pollinate fields non-GM crops. Several fodder crops, forestry trees, grasses and ornamentals have varieties in field trials and these too may hybridise with wild relatives in the European flora (alfalfa, clover, lupin, silver birch, sweet chestnut, Norway spruce, Scots pine, poplar, elm, Agrostis canina, A. stolonifera, Festuca arundinacea, Lolium perenne, L. multiflorum, statice and rose). All these crops will require containment strategies to be in place if it is deemed necessary to prevent transgene movement to wild relatives and non-GM crops. Current Containment Strategies A wide variety of GM containment strategies are currently under development, with a particular focus on crops expressing pharmaceutical products. Physical containment in greenhouses and growth rooms is suitable for some crops (tomatoes, lettuce) and for research purposes. Aquatic bioreactors of some non-crop species (algae, moss, and duckweed) expressing pharmaceutical products have been adopted by some biotechnology companies. There are obvious limitations of the scale of physical containment strategies, addressed in part by the development of large underground facilities in the US and Canada. The additional resources required to grow plants underground incurs high costs that in the long term may negate any advantage of GM for commercial productioNatural genetic containment has been adopted by some companies through the selection of either non-food/feed crops (algae, moss, duckweed) as bio-pharming platforms or organisms with no wild relatives present in the local flora (safflower in the Americas). The expression of pharmaceutical products in leafy crops (tobacco, alfalfa, lettuce, spinach) enables growth and harvesting prior to and in the absence of flowering. Transgenically controlled containment strategies range in their approach and degree of development. Plastid transformation is relatively well developed but is not suited to all traits or crops and does not offer complete containment. Male sterility is well developed across a range of plants but has limitations in its application for fruit/seed bearing crops. It has been adopted in some commercial lines of oilseed rape despite not preventing escape via seed. Conditional lethality can be used to prevent flowering or seed development following the application of a chemical inducer, but requires 100% induction of the trait and sufficient application of the inducer to all plants. Equally, inducible expression of the GM trait requires equally stringent application conditions. Such a method will contain the trait but will allow the escape of a non-functioning transgene. Seed lethality (‘terminator’ technology) is the only strategy at present that prevents transgene movement via seed, but due to public opinion against the concept it has never been trialled in the field and is no longer under commercial development. Methods to control flowering and fruit development such as apomixis and cleistogamy will prevent crop-to-wild and wild-to-crop pollination, but in nature both of these strategies are complex and leaky. None of the genes controlling these traits have as yet been identified or characterised and therefore have not been transgenically introduced into crop species. Neither of these strategies will prevent transgene escape via seed and any feral apomicts that form are arguably more likely to become invasives. Transgene mitigation reduces the fitness of initial hybrids and so prevents stable introgression of transgenes into wild populations. However, it does not prevent initial formation of hybrids or spread to non-GM crops. Such strategies could be detrimental to wild populations and have not yet been demonstrated in the field. Similarly, auxotrophy prevents persistence of escapes and hybrids containing the transgene in an uncontrolled environment, but does not prevent transgene movement from the crop. Recoverable block of function, intein trans-splicing and transgene excision all use recombinases to modify the transgene in planta either to induce expression or to prevent it. All require optimal conditions and 100% accuracy to function and none have been tested under field conditions as yet. All will contain the GM trait but all will allow some non-native DNA to escape to wild populations or to non-GM crops. There are particular issues with GM trees and grasses as both are largely undomesticated, wind pollinated and perennial, thus providing many opportunities for hybridisation. Some species of both trees and grass are also capable of vegetative propagation without sexual reproduction. There are additional concerns regarding the weedy nature of many grass species and the long-term stability of GM traits across the life span of trees. Transgene stability and conferred sterility are difficult to trial in trees as most field trials are only conducted during the juvenile phase of tree growth. Bio-pharming of pharmaceutical and industrial compounds in plants Bio-pharming of pharmaceutical and industrial compounds in plants offers an attractive alternative to mammalian-based pharmaceutical and vaccine production. Several plantbased products are already on the market (Prodigene’s avidin, β-glucuronidase, trypsin generated in GM maize; Ventria’s lactoferrin generated in GM rice). Numerous products are in clinical trials (collagen, antibodies against tooth decay and non-Hodgkin’s lymphoma from tobacco; human gastric lipase, therapeutic enzymes, dietary supplements from maize; Hepatitis B and Norwalk virus vaccines from potato; rabies vaccines from spinach; dietary supplements from Arabidopsis). The initial production platforms for plant-based pharmaceuticals were selected from conventional crops, largely because an established knowledge base already existed. Tobacco and other leafy crops such as alfalfa, lettuce and spinach are widely used as leaves can be harvested and no flowering is required. Many of these crops can be grown in contained greenhouses. Potato is also widely used and can also be grown in contained conditions. The introduction of morphological markers may aid in the recognition and traceability of crops expressing pharmaceutical products. Plant cells or plant parts may be transformed and maintained in culture to produce recombinant products in a contained environment. Plant cells in suspension or in vitro, roots, root cells and guttation fluid from leaves may be engineered to secrete proteins that may be harvested in a continuous, non-destructive manner. Most strategies in this category remain developmental and have not been commercially adopted at present. Transient expression produces GM compounds from non-GM plants via the utilisation of bacterial or viral vectors. These vectors introduce the trait into specific tissues of whole plants or plant parts, but do not insert them into the heritable genome. There are some limitations of scale and the field release of such crops will require the regulation of the vector. However, several companies have several transiently expressed products in clinical and pre-clinical trials from crops raised in physical containment.

Cereal-based wholecrop silages: a potential conservation measure for farmland birds in pastoral landscapes

Declines of farmland birds have been pronounced in landscapes dominated by lowland livestock production and densities of seed-eating birds are particularly low in such areas. Modern livestock production often entails a simple cropping system dominated by ley grassland and maize grown for animal feed. These crops often lack invertebrate and seed resources for foraging birds and can be hostile nesting environments. Cereal-based wholecrop silages (CBWCS) offer potential benefits for farmland birds because they can be grown with minimal herbicide applications and can be spring-sown with following winter stubbles. We compared the biodiversity benefits and agronomic yields of winter-sown wheat and spring-sown barley as alternatives to grass and maize silage in intensive dairy livestock systems. Seed-eating birds foraged mainly in CBWCS fields during summer, and mainly on barley stubbles during winter and this reflected the higher densities of seed-bearing plants therein. Maize and grass fields lacked seed-bearing vegetation and were strongly avoided by most seed-eating birds. Production costs of CBWCS are similar to those of maize and lower than those of grass silage. Selective (rather than broad-spectrum) herbicide application on spring barley crops increased forb cover, reduced yields (by 11%) but caused only a small (<4%) increase in production costs. CBWCS grown with selective herbicide and with following winter stubbles offer a practical conservation measure for seed-eating farmland birds in landscapes dominated by intensively-managed grassland and maize. However, the relatively early harvesting of CBWCS could destroy a significant proportion of breeding attempts of late-nesting species like corn bunting (Emberiza calandra) or yellow wagtail (Motocilla flava). Where late-breeding species are likely to nest in CBWCS fields, harvesting should be delayed until most nesting attempts have been completed (e.g. until after 1st August in southern Britain). (C) 2010 Elsevier Ltd. All rights reserved.

A novel method for assessing risks to pollinators from plant protection products using honeybees as a model species

Pollination is one of the most important ecosystem services in agroecosystems and supports food production. Pollinators are potentially at risk being exposed to pesticides and the main route of exposure is direct contact, in some cases ingestion, of contaminated materials such as pollen, nectar, flowers and foliage. To date there are no suitable methods for predicting pesticide exposure for pollinators, therefore official procedures to assess pesticide risk are based on a Hazard Quotient. Here we develop a procedure to assess exposure and risk for pollinators based on the foraging behaviour of honeybees (Apis mellifera) and using this species as indicator representative of pollinating insects. The method was applied in 13 European field sites with different climatic, landscape and land use characteristics. The level of risk during the crop growing season was evaluated as a function of the active ingredients used and application regime. Risk levels were primarily determined by the agronomic practices employed (i.e. crop type, pest control method, pesticide use), and there was a clear temporal partitioning of risks through time. Generally the risk was higher in sites cultivated with permanent crops, such as vineyard and olive, than in annual crops, such as cereals and oil seed rape. The greatest level of risk is generally found at the beginning of the growing season for annual crops and later in June–July for permanent crops.

Simulating the effects of grassland management and grass ensiling on methane emission from lactating cows

A dynamic, mechanistic model of enteric fermentation was used to investigate the effect of type and quality of grass forage, dry matter intake (DMI) and proportion of concentrates in dietary dry matter (DM) on variation in methane (CH(4)) emission from enteric fermentation in dairy cows. The model represents substrate degradation and microbial fermentation processes in rumen and hindgut and, in particular, the effects of type of substrate fermented and of pH oil the production of individual volatile fatty acids and CH, as end-products of fermentation. Effects of type and quality of fresh and ensiled grass were evaluated by distinguishing two N fertilization rates of grassland and two stages of grass maturity. Simulation results indicated a strong impact of the amount and type of grass consumed oil CH(4) emission, with a maximum difference (across all forage types and all levels of DM 1) of 49 and 77% in g CH(4)/kg fat and protein corrected milk (FCM) for diets with a proportion of concentrates in dietary DM of 0.1 and 0.4, respectively (values ranging from 10.2 to 19.5 g CH(4)/kg FCM). The lowest emission was established for early Cut, high fertilized grass silage (GS) and high fertilized grass herbage (GH). The highest emission was found for late cut, low-fertilized GS. The N fertilization rate had the largest impact, followed by stage of grass maturity at harvesting and by the distinction between GH and GS. Emission expressed in g CH(4)/kg FCM declined oil average 14% with an increase of DMI from 14 to 18 kg/day for grass forage diets with a proportion of concentrates of 0.1, and on average 29% with an increase of DMI from 14 to 23 kg/day for diets with a proportion of concentrates of 0.4. Simulation results indicated that a high proportion of concentrates in dietary DM may lead to a further reduction of CH, emission per kg FCM mainly as a result of a higher DM I and milk yield, in comparison to low concentrate diets. Simulation results were evaluated against independent data obtained at three different laboratories in indirect calorimetry trials with COWS consuming GH mainly. The model predicted the average of observed values reasonably, but systematic deviations remained between individual laboratories and root mean squared prediction error was a proportion of 0.12 of the observed mean. Both observed and predicted emission expressed in g CH(4)/kg DM intake decreased upon an increase in dietary N:organic matter (OM) ratio. The model reproduced reasonably well the variation in measured CH, emission in cattle sheds oil Dutch dairy farms and indicated that oil average a fraction of 0.28 of the total emissions must have originated from manure under these circumstances.

Genetic diversity of red-fleshed apples (Malus)

Anthocyanins are flavonoid pigments imparting red, blue, or purple pigmentation to fruits, flowers and foliage. These compounds are powerful antioxidants in vitro, and are widely believed to contribute to human health. The fruit of the domestic apple (Malus x domestica) is a popular and important source of nutrients, and is considered one of the top ‘functional foods’—those foods that have inherent health-promoting benefits beyond basic nutritional value. The pigmentation of typical red apple fruits results from accumulation of anthocyanin in the skin. However, numerous genotypes of Malus are known that synthesize anthocyanin in additional fruit tissues including the core and cortex (flesh). Red-fleshed apple genotypes are an attractive starting point for development of novel varieties for consumption and nutraceutical use through traditional breeding and biotechnology. However, cultivar development is limited by lack of characterization of the diversity of genetic backgrounds showing this trait. We identified and cataloged red-fleshed apple genotypes from four Malus diversity collections representing over 3,000 accessions including domestic cultivars, wild species, and named hybrids. We found a striking range of flesh color intensity and pattern among accessions, including those carrying the MYB10 R 6 allele conferring ectopic expression of a key transcriptional regulator of anthocyanin biosynthesis. Although MYB10 R 6 was strongly associated with red-fleshed fruit among genotypes, this allele was neither sufficient nor required for this trait in all genotypes. Nearly all red-fleshed accessions tested could be traced back to ‘Niedzwetzkyana’, a presumed natural form of M. sieversii native to central Asia.

Determining the cost of in-field mitigation options to reduce sediment and phosphorus loss

The Mitigation Options for Phosphorus and Sediment (MOPS) project investigated the effectiveness of within-field control measures (tramline management, straw residue management, type of cultivation and direction, and vegetative buffers) in terms of mitigating sediment and phosphorus loss from winter-sown combinable cereal crops using three case study sites. To determine the cost of the approaches, simple financial spreadsheet models were constructed at both farm and regional levels. Taking into account crop areas, crop rotation margins per hectare were calculated to reflect the costs of crop establishment, fertiliser and agro-chemical applications, harvesting, and the associated labour and machinery costs. Variable and operating costs associated with each mitigation option were then incorporated to demonstrate the impact on the relevant crop enterprise and crop rotation margins. These costs were then compared to runoff, sediment and phosphorus loss data obtained from monitoring hillslope-length scale field plots. Each of the mitigation options explored in this study had potential for reducing sediment and phosphorus losses from arable land under cereal crops. Sediment losses were reduced from between 9 kg ha−1 to as much as 4780 kg ha−1 with a corresponding reduction in phosphorus loss from 0.03 kg ha−1 to 2.89 kg ha−1. In percentage terms reductions of phosphorus were between 9% and 99%. Impacts on crop rotation margins also varied. Minimum tillage resulted in cost savings (up to £50 ha−1) whilst other options showed increased costs (up to £19 ha−1 for straw residue incorporation). Overall, the results indicate that each of the options has potential for on-farm implementation. However, tramline management appeared to have the greatest potential for reducing runoff, sediment, and phosphorus losses from arable land (between 69% and 99%) and is likely to be considered cost-effective with only a small additional cost of £2–4 ha−1, although further work is needed to evaluate alternative tramline management methods. Tramline management is also the only option not incorporated within current policy mechanisms associated with reducing soil erosion and phosphorus loss and in light of its potential is an approach that should be encouraged once further evidence is available.

Effects of seed addition on beetle assemblages during the re-creation of species-rich lowland hay meadows

Species-rich lowland hay meadows are of conservation importance for both plants and invertebrates; however, they have declined in area across Europe as a result of conversion to other land uses and management intensification. The re-creation of these grasslands on ex-arable land provides a valuable approach to increasing the extent and conservation value of this threatened habitat. Over a 3-year period a replicated block design was used to test whether introducing seeds promoted the re-creation of both plant and phytophagous beetle assemblages typical of a target hay meadow. Seeds were harvested from local hay meadows, and applied to experimental plots in the form of either green hay or brush harvesting seeds. Green hay spreading achieved the greatest success in re-creating plant and phytophagous beetle assemblages. While re-creation success increased over time for both taxa, for the phytophagous beetles the greatest increase in re-creation success relative to the establishment year also occurred where green hay was applied. We also considered the phytophagous beetles in terms of functional traits that describe host plant specificity, larval feeding location and dispersal. Phytophagous beetle functional trait composition was most similar to the target hay meadow assemblage where some form of seed addition was used, i.e. hay spreading or brush harvested seeds. This study identified the importance of introducing target plant species as a mechanism to promote the re-creation of phytophagous beetle communities. Seed addition methods (e.g. green hay spreading) are crucial to successful hay meadow re-creation.

The REFLEX project: Comparing different algorithms and implementations for the inversion of a terrestrial ecosystem model against eddy covariance data

We describe a model-data fusion (MDF) inter-comparison project (REFLEX), which compared various algorithms for estimating carbon (C) model parameters consistent with both measured carbon fluxes and states and a simple C model. Participants were provided with the model and with both synthetic net ecosystem exchange (NEE) of CO2 and leaf area index (LAI) data, generated from the model with added noise, and observed NEE and LAI data from two eddy covariance sites. Participants endeavoured to estimate model parameters and states consistent with the model for all cases over the two years for which data were provided, and generate predictions for one additional year without observations. Nine participants contributed results using Metropolis algorithms, Kalman filters and a genetic algorithm. For the synthetic data case, parameter estimates compared well with the true values. The results of the analyses indicated that parameters linked directly to gross primary production (GPP) and ecosystem respiration, such as those related to foliage allocation and turnover, or temperature sensitivity of heterotrophic respiration, were best constrained and characterised. Poorly estimated parameters were those related to the allocation to and turnover of fine root/wood pools. Estimates of confidence intervals varied among algorithms, but several algorithms successfully located the true values of annual fluxes from synthetic experiments within relatively narrow 90% confidence intervals, achieving >80% success rate and mean NEE confidence intervals <110 gC m−2 year−1 for the synthetic case. Annual C flux estimates generated by participants generally agreed with gap-filling approaches using half-hourly data. The estimation of ecosystem respiration and GPP through MDF agreed well with outputs from partitioning studies using half-hourly data. Confidence limits on annual NEE increased by an average of 88% in the prediction year compared to the previous year, when data were available. Confidence intervals on annual NEE increased by 30% when observed data were used instead of synthetic data, reflecting and quantifying the addition of model error. Finally, our analyses indicated that incorporating additional constraints, using data on C pools (wood, soil and fine roots) would help to reduce uncertainties for model parameters poorly served by eddy covariance data.

Efficiency, enforcement and revenue tradeoffs in participatory forest management: an example from Tanzania

Where joint forest management has been introduced into Tanzania, ‘volunteer’ patrollers take responsibility for enforcing restrictions over the harvesting of forest resources, often receiving as an incentive a share of the collected fine revenue. Using an optimal enforcement model, we explore how that share, and whether villagers have alternative sources of forest products, determines the effort patrollers put into enforcement and whether they choose to take a bribe rather than honestly reporting the illegal collection of forest resources. Without funds for paying and monitoring patrollers, policy makers face tradeoffs over illegal extraction, forest protection and revenue generation through fine collection.

The effects of a changing pollution climate on throughfall deposition and cycling in a forested area in southern England

This study compares two sets of measurements of the composition of bulk precipitation and throughfall at a site in southern England with a 20-year gap between them. During this time, SO2 emissions from the UK fell by 82%, NOx emissions by 35% and NH3 emissions by 7%. These reductions were partly reflected in bulk precipitation, with deposition reductions of 56% in SO4,38% in NO3, 32% in NH4, and 73% in H+. In throughfall under Scots pine, the effects were more dramatic, with an 89% reduction in SO4 deposition and a 98% reduction in H+ deposition. The mean pH under these trees increased from 2.85 to 4.30. Nitrate and ammonium deposition in throughfall increased slightly, however. In the earlier period, the Scots pines were unable to neutralise the high flux of acidity associated with sulphur deposition, even though this was not a highly polluted part of the UK, and deciduous trees (oak and birch) were only able to neutralise it in summer when the leaves were present. In the later period, the sulphur flux had reduced to the point where the acidity could be neutralised by all species — the neutralisation mechanism is thus likely to be largely leaching of base cations and buffering substances from the foliage. The high fluxes are partly due to the fact that these are 60–80 year old trees growing in an open forest structure. The increase in NO3 and NH4 in throughfall in spite of decreased deposition seems likely due to a decrease in foliar uptake, perhaps due to the increasing nitrogen saturation of the catchment soils. These changes may increase the rate of soil microbial activity as nitrogen increases and acidity declines, with consequent effects on water quality of the catchment drainage stream.