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

Multitrophic effects of nutrient addition in upland grassland

Although the effects of nutrient enhancement on aquatic systems are well documented, the consequences of nutritional supplements on soil food webs are poorly understood, and results of past research examining bottom-up effects are often conflicting. In addition, many studies have failed to separate the effects of nutrient enrichment and the physical effects of adding organic matter. In this field study, we hypothesised that the addition of nitrogen to soil would result in a trophic cascade, through detritivores (Collembola) to predators (spiders), increasing invertebrate numbers and diversity. Nitrogen and lime were added to plots in an upland grassland in a randomised block design. Populations of Collembola and spiders were sampled by means of pitfall traps and identified to species. Seventeen species of Collembola were identified from the nitrogen plus lime (N + L) and control plots. Species assemblage, diversity, richness, evenness and total number were not affected by nutrient additions. However, there was an increase in the number of Isotomidae juveniles and Parisotoma anglicana trapped in the N + L plots. Of the 44 spider species identified, over 80% were Linyphiidae. An effect on species assemblage from the addition of N + L to the plots was observed on two of the four sampling dates (July 2002 and June 2003). The linyphiid, Oedothorax retusus, was the only species significantly affected by the treatments and was more likely to be trapped in the control plots. The increased number of juvenile Collembola, and change in community composition of spiders, were consequences of the bottom-up effect caused by nutrient inputs. However, despite efforts to eliminate the indirect effects of nutrient inputs, a reduction in soil moisture in the N + L plots cannot be eliminated as a cause of the invertebrate population changes observed. Even so, this experiment was not confounded by the physical effects of habitat structure reported in most previous studies. It provides evidence of moderate bottom-up influences of epigeic soil invertebrate food webs and distinguishes between nutrient addition and plant physical structure effects. It also emphasises the importance Of understanding the effects of soil management practices on soil biodiversity, which is under increasing pressure from land development and food production.

Trophic interactions in a changing world: modelling aboveground-belowground interactions

The rate and scale of human-driven changes can exert profound impacts on ecosystems, the species that make them up and the services they provide that sustain humanity. Given the speed at which these changes are occurring, one of society's major challenges is to coexist within ecosystems and to manage ecosystem services in a sustainable way. The effect of possible scenarios of global change on ecosystem services can be explored using ecosystem models. Such models should adequately represent ecosystem processes above and below the soil surface (aboveground and belowground) and the interactions between them. We explore possibilities to include such interactions into ecosystem models at scales that range from global to local. At the regional to global scale we suggest to expand the plant functional type concept (aggregating plants into groups according to their physiological attributes) to include functional types of aboveground-belowground interactions. At the scale of discrete plant communities, process-based and organism-oriented models could be combined into "hybrid approaches" that include organism-oriented mechanistic representation of a limited number of trophic interactions in an otherwise process - oriented approach. Under global change the density and activity of organisms determining the processes may change non-linearly and therefore explicit knowledge of the organisms and their responses should ideally be included. At the individual plant scale a common organism-based conceptual model of aboveground-belowground interactions has emerged. This conceptual model facilitates the formulation of research questions to guide experiments aiming to identify patterns that are common within, but differ between, ecosystem types and biomes. Such experiments inform modelling approaches at larger scales. Future ecosystem models should better include this evolving knowledge of common patterns of aboveground-belowground interactions. Improved ecosystem models are necessary toots to reduce the uncertainty in the information that assists us in the sustainable management of our environment in a changing world. (C) 2004 Elsevier GmbH. All rights reserved.

Responses of invertebrate trophic level, feeding guild and body size to the management of improved grassland field margins

P>1. Management of lowland mesotrophic grasslands in north-west Europe often makes use of inorganic fertilizers, high stocking densities and silage-based forage systems to maximize productivity. The impact of these practices has resulted in a simplification of the plant community combined with wide-scale declines in the species richness of grassland invertebrates. We aim to identify how field margin management can be used to promote invertebrate diversity across a suite of functionally diverse taxa (beetles, planthoppers, true bugs, butterflies, bumblebees and spiders). 2. Using an information theoretic approach we identify the impacts of management (cattle grazing, cutting and inorganic fertilizer) and plant community composition (forb species richness, grass species richness and sward architecture) on invertebrate species richness and body size. As many of these management practices are common to grassland systems throughout the world, understanding invertebrate responses to them is important for the maintenance of biodiversity. 3. Sward architecture was identified as the primary factor promoting increased species richness of both predatory and phytophagous trophic levels, as well as being positively correlated with mean body size. In all cases phytophagous invertebrate species richness was positively correlated with measures of plant species richness. 4. The direct effects of management practices appear to be comparatively weak, suggesting that their impacts are indirect and mediated though the continuous measures of plant community structure, such as sward architecture or plant species richness. 5. Synthesis and applications. By partitioning field margins from the remainder of the field, economically viable intensive grassland management can be combined with extensive management aimed at promoting native biodiversity. The absence of inorganic fertilizer, combined with a reduction in the intensity of both cutting and grazing regimes, promotes floral species richness and sward architectural complexity. By increasing sward architecture the total biomass of invertebrates also increased (by c. 60% across the range of sward architectural measures seen in this study), increasing food available for higher trophic levels, such as birds and mammals.

Dicopper(II) complexes of a new di-para-xylyldioxatetraazamacrocycle and cascade species with dicarboxylate anions: thermodynamics and structural properties

The new dioxatetraazamacrocycle (L-1) was synthesized by a 2 + 2 condensation and characterized. Stability constants of its copper(II) complexes were determined by spectrophotometry in DMSO at 298.2 K in 0. 10 mol dm(-3) KClO4. Mainly dinuclear complexes are formed and the presence of mononuclear species is dependent on the counterion (Cl- or ClO4-). The association constants of the dinuclear copper(II) complexes with dicarboxylate anions [oxalate (oxa(2-)), malonate (mal(2-)), succinate (suc(2-)), and glutarate (glu(2-))] were also determined by spectrophotometry at 298.2 K in DMSO, and it was found that values decrease with an increase of the alkyl chain between the carboxylate groups. X-Band EPR spectra of the dicopper(II) complexes and of their cascade species in frozen DMSO exhibit dipole-dipole coupling, and their simulation, together with their UV-vis spectra, showed that the copper centres of the complexes in solution had square pyramidal geometries though with different distortions. From the experimental data, it was also possible to predict the Cu...Cu distances, the minimum being found at 6.4 angstrom for the (Cu2LCl4)-Cl-1 complex and then successively this distance slightly increases when the chloride anions are replaced by dicarboxylate anions, from 6.6 angstrom for oxa(2-) to 7.8 for glu(2-). The crystal structures of the dinuclear copper cascade species with oxa(2-) and suc(2-) were determined and showed one anion bridging both copper centres and Cu...Cu distances of 5.485(7) angstrom and 6.442(8) angstrom, respectively.

Cascade dicopper architectures of a dibenzodioxatetraazamacrocycle

The dibenzodioxatetraazamacrocycle [26]pbz(2)N(4)O(2) was characterised by single crystal X-ray diffraction and the protonation constants of this compound and the stability constants of its copper(II) and lead(II) complexes were determined by potentiometry in water at 298.2 K in 0.10 mol dm(-3) in KNO3. Mono- and dinuclear complexes were found for both metal ions, the dinuclear complexes being the main species in the 5-7.5 pH range for copper(II) and 7.5-8.5 for lead(II). As expected the values of the stability constants for the copper(II) complexes are lower than those for related macrocycles containing only nitrogen atoms. The presence of mono- and dinuclear copper complexes was also confirmed by electrospray ionization mass spectrometry. These results suggest that the symmetric macrocyclic cavity of [26]pbZ(2)N(4)O(2) has enough space for the coordination of two metal ions. Additionally, NMR spectroscopy showed that the dinuclear complex of lead(II) has high symmetry. The equilibrium constants of the dinuclear copper(II) complexes and dicarboxylate anions (oxalate, malonate and succinate) were also determined in 0.10 mol dm-3 aqueous KNO3 solution. Only species containing one anion, Cu(2)H(h)LA((2+h)), were found, strongly suggesting that the anion bridges the two copper(II) ions. The binding constants of the cascade species formed by [Cu-2[26]pbZ(2)N(4)O(2)(H2O)(4+) with dicarboxylate anions decrease with the increase in length of the alkyl chain of the anion, a fact which was attributed to a higher conformational energy necessary for the rearrangement of the macrocycle to accommodate the larger anions bridging the two copper(II) centres. The variation of the magnetic susceptibility with temperature Of [Cu-2(H-2[26]pbz(2)N(4)O(2))(oxa)(3)]-4H(2)O and [Cu-2([26]pbz(2)N(4)O(2))(suc)Cl-2] were measured and the two complexes showed different behaviour. (c) 2007 Elsevier Ltd. All rights reserved.

Influence of elevated CO2 on interspecific interactions at higher trophic levels

We report the results of a study investigating the influence of elevated CO2 on species interactions across three trophic levels: a plant (Brassica oleracea), two aphid herbivores (the generalist Myzus persicae and the specialist Brevicoryne brassicae), and two natural enemies (the coccinellid Hippodamia convergens (ladybird) and the parasitoid wasp Diaeretiella rapae). Brassica oleracea plants reared under elevated CO2 conditions (650 ppmv vs. 350 ppmv) were larger and had decreased water and nitrogen content. Brevicoryne brassicae reared on plants grown in elevated CO2 were larger and accumulated more fat, while there was no change in M. persicae traits. Fecundity of individual aphids appeared to be increased when reared on plants grown in elevated CO2. However, these differences were generally lost when aphids were reared in colonies, suggesting that such changes in plant quality will have subtile effects on aphid intraspecific interactions. Nevertheless, CO2 treatment did influence aphid distribution on plants, with significantly fewer M. persicae found on the shoots, and B. brassicae was only found on senescing leaves, when colonies were reared on plants grown in elevated CO2. We reared B. brassicae and M. persicae in competition on plants grown at both the CO2 concentration treatments. We found a significantly lower ratio of M. persicae: B. brassicae on plants grown under elevated CO2 conditions, strongly suggesting that increasing CO2 concentrations can alter the outcome of competition among insect herbivores. This was also reflected in the distribution of the aphids on the plants. While the CO2 treatment did not influence where B. brassicae were found, fewer M. persicae were present on senescing leaves under elevated CO2 conditions. Changes in plant quality resulting from the CO2 treatments did not appear to alter aphid quality as prey species, as the number consumed by the ladybird H. convergens, and the number parasitised by the parasitoid wasp D. rapae, did not change. To our knowledge, this study provides the first empirical evidence that changes in host plant quality mediated by increasing levels of CO2 can alter the outcome of interspecific competition among insect herbivores.

Organocatalytic enantioselective construction of nitrocyclohexanes containing multiple chiral centres via a cascade reaction

The stereoselective construction of complex molecules with multiple stereogenicity in a single step represents an extremely useful, but challenging approach to complexity in chemical synthesis. The development of organocatalytic cascade processes has proven useful in these studies, but reports where four or more stereocentres are created in a single step from just two achiral reagents are rare. Herein we report the development of a novel asymmetric domino Michael-Michael reaction between nitrohex-4-enoates and nitro-olefins to generate cyclohexanes of high complexity, including one with a quaternary centre, and one with five contiguous stereocentres. This methodology provides access to a range of useful nitrocyclohexane derivatives, including a novel class of a-lycorane-like structures.

Deriving global flood hazard maps of fluvial floods through a physical model cascade

Global flood hazard maps can be used in the assessment of flood risk in a number of different applications, including (re)insurance and large scale flood preparedness. Such global hazard maps can be generated using large scale physically based models of rainfall-runoff and river routing, when used in conjunction with a number of post-processing methods. In this study, the European Centre for Medium Range Weather Forecasts (ECMWF) land surface model is coupled to ERA-Interim reanalysis meteorological forcing data, and resultant runoff is passed to a river routing algorithm which simulates floodplains and flood flow across the global land area. The global hazard map is based on a 30 yr (1979–2010) simulation period. A Gumbel distribution is fitted to the annual maxima flows to derive a number of flood return periods. The return periods are calculated initially for a 25×25 km grid, which is then reprojected onto a 1×1 km grid to derive maps of higher resolution and estimate flooded fractional area for the individual 25×25 km cells. Several global and regional maps of flood return periods ranging from 2 to 500 yr are presented. The results compare reasonably to a benchmark data set of global flood hazard. The developed methodology can be applied to other datasets on a global or regional scale.

Remarks concerning the double cascade as a necessary mechanism for the instability of steady equivalent-barotropic flows

n a recent paper, Petroniet al. claim that a necessary condition for the instability of two-dimensional steady flows is a «double cascade» of energy and enstrophy respectively to larger and to smaller scales of motion. It is shown here that the analytical reasoning employed by Petroniet al. is flawed and that their conclusions are incorrect. What is true is that in any scale interaction (whether an instability or not), neither energy nor enstrophy can be transferred in one spectral direction only, but this result is extremely well known.

CASCADE: An Agent Based Framework For Modeling The Dynamics Of Smart Electricity Systems

The Complex Adaptive Systems, Cognitive Agents and Distributed Energy (CASCADE) project is developing a framework based on Agent Based Modelling (ABM). The CASCADE Framework can be used both to gain policy and industry relevant insights into the smart grid concept itself and as a platform to design and test distributed ICT solutions for smart grid based business entities. ABM is used to capture the behaviors of diff erent social, economic and technical actors, which may be defi ned at various levels of abstraction. It is applied to understanding their interactions and can be adapted to include learning processes and emergent patterns. CASCADE models ‘prosumer’ agents (i.e., producers and/or consumers of energy) and ‘aggregator’ agents (e.g., traders of energy in both wholesale and retail markets) at various scales, from large generators and Energy Service Companies down to individual people and devices. The CASCADE Framework is formed of three main subdivisions that link models of electricity supply and demand, the electricity market and power fl ow. It can also model the variability of renewable energy generation caused by the weather, which is an important issue for grid balancing and the profi tability of energy suppliers. The development of CASCADE has already yielded some interesting early fi ndings, demonstrating that it is possible for a mediating agent (aggregator) to achieve stable demandfl attening across groups of domestic households fi tted with smart energy control and communication devices, where direct wholesale price signals had previously been found to produce characteristic complex system instability. In another example, it has demonstrated how large changes in supply mix can be caused even by small changes in demand profi le. Ongoing and planned refi nements to the Framework will support investigation of demand response at various scales, the integration of the power sector with transport and heat sectors, novel technology adoption and diffusion work, evolution of new smart grid business models, and complex power grid engineering and market interactions.

Endothelin-1 and fibroblast growth factors stimulate the mitogen-activated protein kinase signaling cascade in cardiac myocytes. The potential role of the cascade in the integration of two signaling pathways leading to myocyte hypertrophy.

Maximally effective concentrations of endothelin-1 (ET-1), acidic FGF (aFGF), or 12-O-tetradecanoylphorbol-13-acetate (TPA) activated mitogen-activated protein kinase (MAPK) by 3-4-fold in crude extracts of myocytes cultured from neonatal rat heart ventricles. Maximal activation was achieved after 5 min. Thereafter, MAPK activity stimulated by ET-1 or aFGF declined to control values within 1-2 h, whereas activation by TPA was more sustained. Two peaks of MAPK activity (a 42- and a 44-kDa MAPK) were resolved in cells exposed to ET-1 or aFGF by fast protein liquid chromatography on a Mono Q column. One major and one minor peak of MAPK kinase (MAPKK) was stimulated by ET-1 or aFGF. Cardiac myocytes expressed protein kinase C (PKC)-alpha, -delta, -epsilon and -zeta as shown immunoblotting. Exposure to 1 microM TPA for 24 h down-regulated PKC-alpha, -delta, and -epsilon, but not PKC-zeta. This maneuver wholly abolished the activation of MAPK on re-exposure to TPA but did not affect the response to aFGF. The effect of ET-1 was partially down-regulated. ET-1 stimulated phospho[3H]inositide hydrolysis 18-fold, whereas aFGF stimulated by only 30%. Agonists which initially utilize dissimilar signaling pathways may therefore converge at the level of MAPKK/MAPK and this may be relevant to the hypertrophic response of the heart.

Regulation of mitogen-activated protein kinase cascade in adult rat heart preparations in vitro.

The regulation of mitogen-activated protein kinase (MAPK) and MAPK kinase (MEK) was studied in freshly isolated adult rat heart preparations. In contrast to the situation in ventricular myocytes cultured from neonatal rat hearts, stimulation of MAPK activity by 1 mumol/L phorbol 12-myristate 13-acetate (PMA) was not consistently detectable in crude extracts. After fast protein liquid chromatography, MAPK isoforms p42MAPK and p44MAPK and two peaks of MEK were shown to be activated > 10-fold in perfused hearts or ventricular myocytes exposed to 1 mumol/L PMA for 5 minutes. The identities of MAPK or MEK were confirmed by immunoblotting and, for MAPK, by the "in-gel" myelin basic protein phosphorylation assay. In retrogradely perfused hearts, high coronary perfusion pressure (120 mm Hg for 5 minutes), norepinephrine (50 mumol/L for 5 minutes), or isoproterenol (50 mumol/L for 5 minutes) stimulated MAPK and MEK approximately 2- to 5-fold. In isolated myocytes, endothelin 1 (100 nmol/L for 5 minutes) also stimulated MAPK, but stimulation by norepinephrine or isoproterenol was difficult to detect. Immunoblotting showed that the relative abundances of MAPK and MEK protein in ventricles declined to < 20% of their postpartal abundances after 50 days. This may explain the difficulties encountered in assaying the activity of MAPK in crude extracts from adult hearts. We conclude that potentially hypertrophic agonists and interventions stimulate the MAPK cascade in adult rats and suggest that the MAPK cascade may be an important intracellular signaling pathway in this response.

Activation of the mitogen-activated protein kinase cascade by pertussis toxin-sensitive and -insensitive pathways in cultured ventricular cardiomyocytes.

The involvement of pertussis toxin (PTX)-sensitive and -insensitive pathways in the activation of the mitogen-activated protein kinase (MAPK) cascade was examined in ventricular cardiomyocytes cultured from neonatal rats. A number of agonists that activate heterotrimeric G-protein-coupled receptors stimulated MAPK activity after exposure for 5 min. These included foetal calf serum (FCS), endothelin-1 (these two being the most effective of the agonists examined), phenylephrine, endothelin-3, lysophosphatidic acid, carbachol, isoprenaline and angiotensin II. Activation of MAPK and MAPK kinase (MEK) by carbachol returned to control levels within 30-60 min, whereas activation by FCS was more sustained. FPLC on Mono Q showed that carbachol and FCS activated two peaks of MEK and two peaks of MAPK (p42MAPK and p44MAPK). Pretreatment of cells with PTX for 24 h inhibited the activation of MAPK by carbachol, FCS and lysophosphatidic acid, but not that by endothelin-1, phenylephrine or isoprenaline. Involvement of G-proteins in the activation of the cardiac MAPK cascade was demonstrated by the sustained (PTX-insensitive) activation of MAPK (and MEK) after exposure of cells to AlF4-. AlF4- activated PtdIns hydrolysis, as did endothelin-1, endothelin-3, phenylephrine and FCS. In contrast, the effect of lysophosphatidic acid on PtdIns hydrolysis was small and carbachol was without significant effect even after prolonged exposure. We conclude that PTX-sensitive (i.e. Gi/G(o)-linked) and PTX-insensitive (i.e. Gq/Gs-linked) pathways of MAPK activation exist in neonatal ventricular myocytes. FCS may stimulate the MAPK cascade through both pathways.