A trophic cascade induced by predatory ants in a fig-fig wasp mutualism

Summary 1. A trophic cascade occurs when predators directly decrease the densities, or change the behaviour, of herbivores and thus indirectly increase plant productivity. The predator–herbivore– plant context is well known, but some predators attack species beneficial to plants (e.g. pollinators) and/or enemies of herbivores (e.g. parasites), and their role in the dynamics of mutualisms remains largely unexplored. 2. We surveyed the predatory ant species and studied predation by the dominant ant species, the weaver ant Oecophylla smaragdina, associated with the fig tree Ficus racemosa in southwest China. We then tested the effects of weaver ants on the oviposition behaviour of pollinating and non-pollinating fig wasps in an ant-exclusion experiment. The effects of weaver ants on fig wasp community structure and fig seed production were then compared between trees with and without O. smaragdina. 3. Oecophylla smaragdina captured more non-pollinating wasps (Platyneura mayri) than pollinators as the insects arrived to lay eggs. When ants were excluded, more non-pollinators laid eggs into figs and fewer pollinators entered figs. Furthermore, trees with O. smaragdina produced more pollinator offspring and fewer non-pollinator offspring, shifting the community structure significantly. In addition, F. racemosa produced significantly more seeds on trees inhabited by weaver ants. 4. Oecophylla smaragdina predation reverses the dominance of the two commonest wasp species at the egg-laying stage and favours the pollinators. This behavioural pattern is mirrored by wasp offspring production, with pollinators’ offspring dominating figs produced by trees inhabited by weaver ants, and offspring of the non-pollinator P. mayri most abundant in figs on trees inhabited by other ants. 5. Overall, our results suggest that predation by weaver ants limits the success of the non-pollinating P. mayri and therefore indirectly benefits the mutualism by increasing the reproductive success of both the pollinators and the plant. Predation is thus a key functional factor that can shape the community structure of a pollinator-plant mutualistic system. Key-words: competitive release, fig wasp, mutualism, predation, predator-exclusion experiment, trophic cascade

Biotic carbon feedbacks in a materially-closed soil-vegetation-atmosphere system

The magnitude and direction of the coupled feedbacks between the biotic and abiotic components of the terrestrial carbon cycle is a major source of uncertainty in coupled climate–carbon-cycle models1, 2, 3. Materially closed, energetically open biological systems continuously and simultaneously allow the two-way feedback loop between the biotic and abiotic components to take place4, 5, 6, 7, but so far have not been used to their full potential in ecological research, owing to the challenge of achieving sustainable model systems6, 7. We show that using materially closed soil–vegetation–atmosphere systems with pro rata carbon amounts for the main terrestrial carbon pools enables the establishment of conditions that balance plant carbon assimilation, and autotrophic and heterotrophic respiration fluxes over periods suitable to investigate short-term biotic carbon feedbacks. Using this approach, we tested an alternative way of assessing the impact of increased CO2 and temperature on biotic carbon feedbacks. The results show that without nutrient and water limitations, the short-term biotic responses could potentially buffer a temperature increase of 2.3 °C without significant positive feedbacks to atmospheric CO2. We argue that such closed-system research represents an important test-bed platform for model validation and parameterization of plant and soil biotic responses to environmental changes.

The role of soil microbes in the global carbon cycle: tracking the below-ground microbial processing of plant-derived carbon for manipulating carbon dynamics in agricultural systems.

It is well known that atmospheric concentrations of carbon dioxide (CO2) (and other greenhouse gases) have increased markedly as a result of human activity since the industrial revolution. It is perhaps less appreciated that natural and managed soils are an important source and sink for atmospheric CO2 and that, primarily as a result of the activities of soil microorganisms, there is a soil-derived respiratory flux of CO2 to the atmosphere that overshadows by tenfold the annual CO2 flux from fossil fuel emissions. Therefore small changes in the soil carbon cycle could have large impacts on atmospheric CO2 concentrations. Here we discuss the role of soil microbes in the global carbon cycle and review the main methods that have been used to identify the microorganisms responsible for the processing of plant photosynthetic carbon inputs to soil. We discuss whether application of these techniques can provide the information required to underpin the management of agro-ecosystems for carbon sequestration and increased agricultural sustainability. We conclude that, although crucial in enabling the identification of plant-derived carbon-utilising microbes, current technologies lack the high-throughput ability to quantitatively apportion carbon use by phylogentic groups and its use efficiency and destination within the microbial metabolome. It is this information that is required to inform rational manipulation of the plant–soil system to favour organisms or physiologies most important for promoting soil carbon storage in agricultural soil.

Modeling plant transpiration under limited soil water: comparison of different plant and soil hydraulic parameterizations and preliminary implications for their use in land surface models

Accurate estimates of how soil water stress affects plant transpiration are crucial for reliable land surface model (LSM) predictions. Current LSMs generally use a water stress factor, β, dependent on soil moisture content, θ, that ranges linearly between β = 1 for unstressed vegetation and β = 0 when wilting point is reached. This paper explores the feasibility of replacing the current approach with equations that use soil water potential as their independent variable, or with a set of equations that involve hydraulic and chemical signaling, thereby ensuring feedbacks between the entire soil–root–xylem–leaf system. A comparison with the original linear θ-based water stress parameterization, and with its improved curvi-linear version, was conducted. Assessment of model suitability was focused on their ability to simulate the correct (as derived from experimental data) curve shape of relative transpiration versus fraction of transpirable soil water. We used model sensitivity analyses under progressive soil drying conditions, employing two commonly used approaches to calculate water retention and hydraulic conductivity curves. Furthermore, for each of these hydraulic parameterizations we used two different parameter sets, for 3 soil texture types; a total of 12 soil hydraulic permutations. Results showed that the resulting transpiration reduction functions (TRFs) varied considerably among the models. The fact that soil hydraulic conductivity played a major role in the model that involved hydraulic and chemical signaling led to unrealistic values of β, and hence TRF, for many soil hydraulic parameter sets. However, this model is much better equipped to simulate the behavior of different plant species. Based on these findings, we only recommend implementation of this approach into LSMs if great care with choice of soil hydraulic parameters is taken

Predicted soil organic carbon stocks and changes in Jordan between 2000 and 2030 made using the GEFSOC modelling system

Estimates of soil organic carbon (SOC) stocks and changes under different land use systems can help determine vulnerability to land degradation. Such information is important for countries in and areas with high susceptibility to desertification. SOC stocks, and predicted changes between 2000 and 2030, were determined at the national scale for Jordan using The Global Environment Facility Soil Organic Carbon (GEFSOC) Modelling System. For the purpose of this study, Jordan was divided into three natural regions (The Jordan Valley, the Uplands and the Badia) and three developmental regions (North, Middle and South). Based on this division, Jordan was divided into five zones (based on the dominant land use): the Jordan Valley, the North Uplands, the Middle Uplands, the South Uplands and the Badia. This information was merged using GIS, along with a map of rainfall isohyets, to produce a map with 498 polygons. Each of these was given a unique ID, a land management unit identifier and was characterized in terms of its dominant soil type. Historical land use data, current land use and future land use change scenarios were also assembled, forming major inputs of the modelling system. The GEFSOC Modelling System was then run to produce C stocks in Jordan for the years 1990, 2000 and 2030. The results were compared with conventional methods of estimating carbon stocks, such as the mapping based SOTER method. The results of these comparisons showed that the model runs are acceptable, taking into consideration the limited availability of long-term experimental soil data that can be used to validate them. The main findings of this research show that between 2000 and 2030, SOC may increase in heavily used areas under irrigation and will likely decrease in grazed rangelands that cover most of Jordan giving an overall decrease in total SOC over time if the land is indeed used under the estimated forms of land use. (C) 2007 Elsevier B.V. All rights reserved.

Complexation of copper by sewage sludge-derived dissolved organic matter: Effects on soil sorption behaviour and plant uptake

The complexation of Cu by sewage sludge-derived dissolved organic matter (SSDOM) is a process by which the environmental significance of the element may become enhanced due to reduced soil sorption and, hence, increased mobility. The work described in this paper used an ion selective electrode procedure to show that SSDOM complexation of Cu was greatest at intermediate pH values because competition between hydrogen ions and Cu for SSDOM binding sites, and between hydroxyl ions and SSDOM as Cu ligands, was lowest at such values. Batch sorption experiments further showed that the process of Cu complexation by SSDOM provided an explanation for enhanced desorption of Cu from the solid phase of a contaminated, organic matter-rich, clay loam soil, and reduced adsorption of Cu onto the solid phase of a sandy loam soil. Complexation of Cu by SSDOM did not affect uptake of Cu by spring barley plants, when compared to free ionic Cu, in a sand-culture pot experiment. However, it did appear to lead to greater biomass yields of the plant; perhaps indicating that the Cu-SSDOM complex had a lower toxicity towards the plant than the free Cu ion.

Indicators of nitrate in wetland surface and soil-waters: interactions of vegetation and environmental factors

This paper describes a new bio-indicator method for assessing wetland ecosystem health: as such, the study is particularly relevant to current legislation such as the EU Water Framework Directive, which provides a baseline of the current status Of Surface waters. Seven wetland sites were monitored across northern Britain, with model construction data for predicting, eco-hydroloplical relationships collected from five sites during 1999, Two new sites and one repeat site were monitored during 2000 to provide model test data. The main growing season for the vegetation, and hence the sampling period, was May-August during both years. Seasonal mean concentrations of nitrate (NO3-) in surface and soil water samples during 1999 ranged from 0.01 to 14.07 mg N 1(-1), with a mean value of 1.01 mg N 1(-1). During 2000, concentrations ranged from trace level (<0.01 m- N 1(-1)) to 9.43 mg N 1(-1), with a mean of 2.73 mg N 1(.)(-1) Surface and soil-water nitrate concentrations did not influence plant species composition significantly across representative tall herb fen and mire communities. Predictive relationships were found between nitrate concentrations and structural characteristics of the wetland vegetation, and a model was developed which predicted nitrate concentrations from measures of plant diversity, canopy structure and density of reproductive structures. Two further models, which predicted stem density and density of reproductive structures respectively, utilised nitrate concentration as one of the independent predictor variables. Where appropriate, the models were tested using data collected during 2000. This approach is complementary to species-based monitoring, representing a useful and simple too] to assess ecological status in target wetland systems and has potential for bio-indication purposes.

Competition in tree row agroforestry systems. 3. Soil water distribution and dynamics

The purpose of this study was to test the hypothesis that soil water content would vary spatially with distance from a tree row and that the effect would differ according to tree species. A field study was conducted on a kaolinitic Oxisol in the sub-humid highlands of western Kenya to compare soil water distribution and dynamics in a maize monoculture with that under maize (Zea mays L.) intercropped with a 3-year-old tree row of Grevillea robusta A. Cunn. Ex R. Br. (grevillea) and hedgerow of Senna spectabilis DC. (senna). Soil water content was measured at weekly intervals during one cropping season using a neutron probe. Measurements were made from 20 cm to a depth of 225 cm at distances of 75, 150, 300 and 525 cm from the tree rows. The amount of water stored was greater under the sole maize crop than the agroforestry systems, especially the grevillea-maize system. Stored soil water in the grevillea-maize system increased with increasing distance from the tree row but in the senna-maize system, it decreased between 75 and 300 cm from the hedgerow. Soil water content increased least and more slowly early in the season in the grevillea-maize system, and drying was also evident as the frequency of rain declined. Soil water content at the end of the cropping season was similar to that at the start of the season in the grevillea-maize system, but about 50 and 80 mm greater in the senna-maize and sole maize systems, respectively. The seasonal water balance showed there was 140 mm, of drainage from the sole maize system. A similar amount was lost from the agroforestry systems (about 160 mm in the grevillea-maize system and 145 mm in the senna-maize system) through drainage or tree uptake. The possible benefits of reduced soil evaporation and crop transpiration close to a tree row were not evident in the grevillea-maize system, but appeared to greatly compensate for water uptake losses in the senna-maize system. Grevillea, managed as a tree row, reduced stored soil water to a greater extent than senna, managed as a hedgerow.

National and sub-national assessments of soil organic carbon stocks and changes: The GEFSOC modelling system

Soil organic carbon (SOC) plays a vital role in ecosystem function, determining soil fertility, water holding capacity and susceptibility to land degradation. In addition, SOC is related to atmospheric CO, levels with soils having the potential for C release or sequestration, depending on land use, land management and climate. The United Nations Convention on Climate Change and its Kyoto Protocol, and other United Nations Conventions to Combat Desertification and on Biodiversity all recognize the importance of SOC and point to the need for quantification of SOC stocks and changes. An understanding of SOC stocks and changes at the national and regional scale is necessary to further our understanding of the global C cycle, to assess the responses of terrestrial ecosystems to climate change and to aid policy makers in making land use/management decisions. Several studies have considered SOC stocks at the plot scale, but these are site specific and of limited value in making inferences about larger areas. Some studies have used empirical methods to estimate SOC stocks and changes at the regional scale, but such studies are limited in their ability to project future changes, and most have been carried out using temperate data sets. The computational method outlined by the Intergovernmental Panel on Climate Change (IPCC) has been used to estimate SOC stock changes at the regional scale in several studies, including a recent study considering five contrasting eco regions. This 'one step' approach fails to account for the dynamic manner in which SOC changes are likely to occur following changes in land use and land management. A dynamic modelling approach allows estimates to be made in a manner that accounts for the underlying processes leading to SOC change. Ecosystem models, designed for site scale applications can be linked to spatial databases, giving spatially explicit results that allow geographic areas of change in SOC stocks to be identified. Some studies have used variations on this approach to estimate SOC stock changes at the sub-national and national scale for areas of the USA and Europe and at the watershed scale for areas of Mexico and Cuba. However, a need remained for a national and regional scale, spatially explicit system that is generically applicable and can be applied to as wide a range of soil types, climates and land uses as possible. The Global Environment Facility Soil Organic Carbon (GEFSOC) Modelling System was developed in response to this need. The GEFSOC system allows estimates of SOC stocks and changes to be made for diverse conditions, providing essential information for countries wishing to take part in an emerging C market, and bringing us closer to an understanding of the future role of soils in the global C cycle. (C) 2007 Elsevier B.V. All rights reserved.

Plant roots release phospholipid surfactants that modify the physical and chemical properties of soil

Plant root mucilages contain powerful surfactants that will alter the interaction of soil solids with water and ions, and the rates of microbial processes. The lipid composition of maize, lupin and wheat root mucilages was analysed by thin layer chromatography and gas chromatography-mass spectrometry. A commercially available phosphatidylcholine (lecithin), chemically similar to the phospholipid surfactants identified in the mucilages, was then used to evaluate its effects on selected soil properties. The lipids found in the mucilages were principally phosphatidylcholines, composed mainly of saturated fatty acids, in contrast to the lipids extracted from root tissues. In soil at low tension, lecithin reduced the water content at any particular tension by as much as 10 and 50% in soil and acid-washed sand, respectively. Lecithin decreased the amount of phosphate adsorption in soil and increased the phosphate concentration in solution by 10%. The surfactant also reduced net rates of ammonium consumption and nitrate production in soil. These experiments provide the first evidence we are aware of that plant-released surfactants will significantly modify the biophysical environment of the rhizosphere.

Acid-base reactions between an acidic soil and plant residues

The elemental composition of residues of maize (Zea mays), sorghum (S. bicolor), groundnuts (Arachis hypogea), soya beans (Glycine max), leucaena (L. leucocephala), gliricidia (G. sepium), and sesbania (S. sesban) was determined as a basis for examining their alkalinity when incorporated into an acidic Zambian Ferralsol. Potential (ash) alkalinity, available alkalinity by titration to pH 4 and soluble alkalinity (16 It water extract titrated to pH 4) were measured. Potential alkalinity ranged from 3 73 (maize) to 1336 (groundnuts) mmol kg(-1) and was equivalent to the excess of their cation charge over inorganic anion charge. Available alkalinity was about half the potential alkalinity. Cations associated with organic anions are the source of alkalinity. About two thirds of the available alkalinity is soluble. Residue buffer curves were determined by titration with H2SO4 to pH 4. Soil buffer capacity measured by addition of NaOH was 12.9 mmol kg(-1) pH(-1). Soil and residue (10 g:0.25 g) were shaken in solution for 24 h and suspension pH values measured. Soil pH increased from 4.3 to between 4.6 (maize) and 5.2 (soyabean) and the amounts of acidity neutralized (calculated from the rise in pH and the soil buffer capacity) were between 3.9 and 11.5 mmol kg(-1), respectively. The apparent base contributions by the residues (calculated from the buffer curves and the fall in pH) ranged between 105 and 350 mmol kg(-1) of residue, equivalent to 2.6 and 8.8 mmol kg(-1) of soil, respectively. Therefore, in contact with soil acidity, more alkalinity becomes available than when in contact with H2SO4 solution. Available alkalinity (to pH 4) would be more than adequate to supply that which reacts with soil but soluble alkalinity would not. It was concluded that soil Al is able to displace cations associated with organic anions in the residues which are not displaced by H+, or that residue decomposition may have begun in the soil suspension releasing some of the non-available alkalinity. Soil and four of the residues were incubated for 100 days and changes in pH, NH4+ and NO3- concentrations measured. An acidity budget equated neutralized soil acidity with residue alkalinity and base or acid produced by N transformations. Most of the potential alkalinity of soyabean and leucaena had reacted after 14 days, but this only occurred after 100 days for gliricidia, and for maize only the available alkalinity reacted. For gliricidia and leucaena, residue alkalinity was primarily used to react with acidity produced by nitrification. Thus, the ability of residues to ameliorate acidity depends not only on their available and potential alkalinity but also on their potential to release mineral N. (C) 2004 Elsevier B.V. All rights reserved.

Use of an artificial root to examine the influence of 8-hydroxyquinoline on soil microbial activity and bacterial community structure

Invasive plant species have been shown to alter the microbial community composition of the soils they invade and it is suggested that this below-ground perturbation of potential pathogens, decomposers or symbionts may feedback positively to allow invasive success. Whether these perturbations are mediated through specific components of root exudation are not understood. We focussed on 8-hydroxyquinoline, a putative allelochemical of Centaurea diffusa (diffuse knapweed) and used an artificial root system to differentiate the effects of 8-hydroxyquinoline against a background of total rhizodeposition as mimicked through supply of a synthetic exudate solution. In soil proximal (0-10 cm) to the artificial root, synthetic exudates had a highly significant (P < 0.001) influence on dehydrogenase, fluorescein diacetate hydrolysis and urease activity. in addition, 8-hydroxyquinoline was significant (p = 0.003) as a main effect on dehydrogenase activity and interacted with synthetic exudates to affect urease activity (p = 0.09). Hierarchical cluster analysis of 16S rDNA-based DGGE band patterns also identified a primary affect of synthetic exudates and a secondary affect of 8-hydroxyquinoline on bacterial community structure. Thus, we show that the artificial rhizosphere produced by the synthetic exudates was the predominant effect, but, that the influence of the 8-hydroxyquinoline signal on the activity and structure of soil microbial communities could also be detected. (C) 2009 Elsevier Ltd. All rights reserved.

Pea-barley intercropping for efficient symbiotic N-2-fixation, soil N acquisition and use of other nutrients in European organic cropping systems

Complementarity in acquisition of nitrogen (N) from soil and N-2-fixation within pea and barley intercrops was studied in organic field experiments across Western Europe (Denmark, United Kingdom, France, Germany and Italy). Spring pea and barley were sown either as sole crops, at the recommended plant density (P100 and B100, respectively) or in replacement (P50B50) or additive (P100B50) intercropping designs, in each of three cropping seasons (2003-2005). Irrespective of site and intercrop design, Land Equivalent Ratios (LER) between 1.4 at flowering and 1.3 at maturity showed that total N recovery was greater in the pea-barley intercrops than in the sole Crops Suggesting a high degree of complementarity over a wide range of growing conditions. Complementarity was partly attributed to greater soil mineral N acquisition by barley, forcing pea to rely more on N-2-fixation. At all sites the proportion of total aboveground pea N that was derived from N-2-fixation was greater when intercropped with barley than when grown as a sole crop. No consistent differences were found between the two intercropping designs. Simultaneously, the accumulation Of Phosphorous (P), potassium (K) and sulphur (S) in Danish and German experiments was 20% higher in the intercrop (P50B50) than in the respective sole crops, possibly influencing general crop yields and thereby competitive ability for other resources. Comparing all sites and seasons, the benefits of organic pea-barley intercropping for N acquisition were highly resilient. It is concluded that pea-barley intercropping is a relevant cropping strategy to adopt when trying to optimize N-2-fixation inputs to the cropping system. (C) 2009 Elsevier B.V. All rights reserved.

Plant species diversity, plant biomass and responses of the soil community on abandoned land across Europe: idiosyncracy or above-belowground time lags

The relationship between plant species diversity, productivity and the development of the soil community during early secondary succession on former arable land across Europe is investigated. The enhancement of biomass production due to the increase in initial plant species diversity and the consequent stimulation of soil microbial biomass and abundance of soil invertebrates are examined.

The potential of combining Pasteuria penetrans and neem (Azadirachta indica) formulations as a management system for root-knot nematodes on tomato

Initial applications of 10(4) spores g(-1) of Pasteuria penetrans, and dried neem cake and leaves at 3 and 2% w:w, respectively, were applied to soil in pots. Juveniles of Meloidogyne javanica were added immediately to the pots (500, 5,000 or 10,000) before planting 6-week-old tomato seedlings. The tomatoes were sampled after 64 days; subsequently a second crop was grown for 59 days and a third crop for 67 days without further applications of P. penetrans and neem. There was significantly less root-galling in the P. penetrans combined with neem cake treatment at the end of the third crop and this treatment also had the greatest effect on the growth of the tomato plants. At the end of the third crop, 30% of the females were infected with P. penetrans in those treatments where spores had been applied at the start of the experiment. The effects of neem leaves and neem cake on the nematode population did not persist through the crop sequences but the potential for combining the amendments with a biological control agent such as P. penetrans is worthy of further evaluation.