Relative strengths of top-down and bottom-up forces in a tropical forest community

We tested integrative bottom-up and top-down trophic cascade hypotheses with manipulative experiments in a tropical wet forest, using the ant-plant Piper cenocladum and its associated arthropod community. We examined enhanced nutrients and light along with predator and herbivore exclusions as sources of variation in the relative biomass of plants, their herbivores (via rates of herbivory), and resident predaceous ants. The combined manipulations of secondary consumers, primary consumers, and plant resources allowed us to examine some of the direct and indirect effects on each trophic level and to determine the relative contributions of bottom-up and top-down cascades to the structure of the community. We found that enhanced plant resources (nutrients and light) had direct positive effects on plant biomass. However, we found no evidence of indirect (cascading through the herbivores) effects of plant biomass on predators or top predators. In contrast, ants had indirect effects on plant biomass by decreasing herbivory on the plants. This top-down cascade occurred whether or not plant resources were enriched, conditions which are expected to modify top-down forces. Received: 9 August 1998 / Accepted: 1 December 1998     

Neural ‘bubble’ dynamics in two dimensions: foundations

An extension to two dimensions of recent results in continuum neural field theory (CNFT) in one dimension is presented here. Focus is placed on the treatment of receptive fields and of learning on afferent synapses to obtain topographic maps. Received: 26 March 1997 / Accepted in revised form: 16 December 1998     

Experimental tests of trophic dynamics: taking a closer look

The relative importance of bottom-up versus top-down forces, and the effect of productivity on community dynamics continue to be of much interest to ecologists. Trophic dynamic theories are difficult to test, as they require explicit knowledge of the many organisms involved, as well as the nature of the interactions between them. The Oksanen-Fretwell (OF) theory, which suggests that the relative roles of top-down and bottom-up factors vary with primary productivity, is well known in the literature, but is difficult to test rigorously. Recently, two experimental studies have tried to test OF theory. In this paper we discuss methodological problems associated with these studies that may weaken the conclusions drawn by the authors. Received: 21 April 1998 / Accepted: 30 July 1998     

Omnivory and the stability of simple food webs

Traditional ecological theory predicts that the stability of simple food webs will decline with an increasing number of trophic levels and increasing amounts of omnivory. These ideas have been tested using protozoans in laboratory microcosms. However, the results are equivocal, and contrary to expectation, omnivory is common in natural food webs. Two recent developments lead us to re-evaluate these predictions using food webs assembled from protists and bacteria. First, recent modelling work suggests that omnivory is actually stabilizing, providing that interactions are not too strong. Second, it is difficult to evaluate the degree of omnivory of some protozoan species without explicit experimental tests. This study used seven species of ciliated protozoa and a mixed bacterial flora to assemble four food webs with two trophic levels, and four webs with three trophic levels. Protist species were assigned a rank for their degree of omnivory using information in the literature and the results of experiments that tested whether the starvation rate of predators was influenced by the amount of bacteria on which they may have fed and whether cannibalism (a form of omnivory) occurred. Consistent with recent modelling work, both bacterivorous and predatory species with higher degrees of omnivory showed more stable dynamics, measured using time until extinction and the temporal variability of population density. Systems with two protist species were less persistent than systems with one protist species, supporting the prediction that longer food chains will be less stable dynamically. Received: 28 December 1997 / Accepted: 22 June 1998     

Changes in trophic level of Squatina guggenheim with increasing body length: relationships with type, size and trophic level of its prey

The occurrence of changes in the trophic level (TL) of sharks with growth has not been quantified until now. Here length-related changes on Squatina guggenheim Marini trophic level were determined, and shifts in type, size and trophic level of its prey were analysed. Sampling took place during five bottom trawl surveys conducted in the Argentine–Uruguayan Common Fishing Zone during spring (December/1995, October/1997) and fall (March/1997, March–April/1998, May–June/1998), using an Engel bottom-trawl net to capture the sharks. Three length groups were defined based on diet composition and using a cluster analysis (group I, 23–60 cm; group II, 61–80 cm; group III, 81–91 cm L _T). An ANOSIM procedure detected significant differences (P < 0.05) in the diet spectrum between the three length groups. The smallest sharks (group I) ingested fish prey ranging from 5 to 21 cm L _T, medium sharks (group II) fed on fish prey between 11 and 35 cm L _T, and largest sharks (group III) preyed on fish between 13 and 40 cm L _T. Diet structure of length groups were discriminated by almost the same prey taxa that characterized them. The increase of S. guggenheim body length promoted a decrease in the relative importance of small pelagic fishes. Contrarily, prey as medium benthopelagic fishes, medium pelagic squid and medium benthopelagic fishes showed an inverse tendency, indicating a broad diet spectrum of adults. Predator-length and prey-length relationship indicated a trend where 44.8% of S. guggenheim diet was integrated by prey <20% of their own body length and 32.8% of their diet was composed by prey >30% of their own length. The increase of mean prey weight was associated with the increase of predator weight and length. Smallest sharks (group I) were identified as secondary consumers (TL < 4) whereas medium sharks (group II) and largest sharks (group III) were placed as tertiary consumers (TL > 4). The study revealed an increase in S. guggenheim TL with shark growth as a consequence of changes on type, size and TL of prey ingested.     

Biotechnology and microbiology of coal degradation

For several years it has been known that fungi and bacteria can attack and even liquefy low rank coals. This review covers the progress in coal biotechnology and microbiology, mainly during the last decade, from describing the first effects to elucidating the mechanisms used by the microorganisms. More than one mechanism is responsible for microbial coal degradation/liquefaction: oxidative enzymes (peroxidases, laccases), hydrolytic enzymes (esterases), alkaline metabolites and natural chelators. Due to the heterogeneous structure of coal, which is described in one section, and for economic reasons the review focuses on the enzymatic depolymerization of brown coal. Approaches which seem not so promising are discussed (anaerobic, reductive pathways, chemical pretreatment). Finally the possible applications and products in this field are summarized, as lignite with a worldwide production of about 940 million tons a year will continue to play an important economic role in the future. Received: 19 October 1998 / Received revision: 16 December 1998 / Accepted: 21 December 1998     

Metal binding to the HIV nucleocapsid peptide

 Co(II) and Zn(II) binding constants have been measured for binding to the HIV-1 nucleocapsid N-terminal metal binding domain (residues 1–18), using competition titration methods and monitoring Co(II) binding by visible absorbance spectroscopy. Enthalpies for binding were directly measured by isothermal titration colorimetry. The results are compared with recent studies of related systems, including a study of Zn(II) binding by the full length protein. Received: 1 December 1998 / Accepted: 31 December 1998     

High-cell-density cultivation of microorganisms

High-cell-density cultivation (HCDC) is required to improve microbial biomass and product formation substantially. An overview of HCDC is given for microorganisms including bacteria, archae and eukarya (yeasts). Problems encountered by HCDC and their possible solutions are discussed. Improvements of strains, different types of bioreactors and cultivation strategies for successful HCDC are described. Stirred-tank reactors with and without cell retention, a dialysis-membrane reactor, a gas-lift reactor and a membrane cyclone reactor used for HCDC are outlined. Recently modified traditional feeding strategies and new ones are included, in particular those for unlimited growth to very dense cultures. Emphasis is placed on robust fermentation control because of the growing industrial interest in this field. Therefore, developments in the application of multivariate statistical control, artificial neural networks, fuzzy control and knowledge-based supervision (expert systems) are summarized. Recent advances using Escherichia coli– the pioneer organism for HCDC – are outlined. Received: 20 October 1998 / Received revision: 18 December 1998 / Accepted: 21 December 1998     

Influence of consumer mutual interference on the stabilization of a three-level trophic system

In this paper, we present a three-level (food–prey–predator) trophic food chain which includes consumer mutual interference (MIF). In contrast with other analyses, we consider the effect of both prey and predator MIF on the dynamics of a three-level trophic system. MIF is generally considered to exert a stabilizing effect on population dynamics based on the predator–prey model. However, results from analytical and numerical simulations utilizing a simple three-species food chain model suggest that while the addition of prey MIF to the model provides a stabilizing influence, as the chaotic dynamics collapse to a stable steady state, adding only predator MIF to the model can only stabilize the system at intermediate MIF values. The three-species trophic food chain is also stabilized when combination of both prey and predator MIF is added to the model. Our work serves to provide insight into the effects of MIF in the real world.     

Effect of parity on productivity and sustainability of Lotka–Volterra food chains

Hairston, Slobodkin, and Smith conjectured that top down forces act on food chains, which opposed the previously accepted theory that bottom up forces exclusively dictate the dynamics of populations. We model food chains using the Lotka–Volterra predation model and derive sustainability constants which determine which species will persist or go extinct. Further, we show that the productivity of a sustainable food chain with even trophic levels is predator regulated, or top down, while a sustainable food chain with odd trophic levels is resource limited, which is bottom up, which is consistent with current ecological theory.     

Foraging for mates in the hyperparasitic wasp, Dendrocerus carpenteri: impact of unfavourable weather conditions and parasitoid age

Males of the aphid hyperparasitoid Dendrocerus carpenteri (Curtis) were attracted by a sex pheromone released by conspecific females. The intensity of this cue, and thus female attractiveness, depended both on the female's mating status and her age. Only virgin females younger than 2 h were consistently recognized as mates by foraging males. Male age did not influence foraging and mating success. Empty mummies, from which females had emerged within the previous 10 min were attractive to males and examined intensively. Rain reduced the searching success of males, although the host plant Vicia faba provided sheltered places. Wind did not reduce mating success but prevented both sexes from leaving the host plant. Since the time of female attractiveness seems to be very limited, wind may have an enormous effect on the mating success of D. carpenteri in the field and thus on the population dynamics of this species. Received: 5 October 1998 / Accepted: 16 December 1998     

The polymerization ratchet model explains the force-velocity relation for growing microtubules

The polymerization of filamentous proteins generates mechanical forces which drive many cellular processes. Dogterom and Yurke measured the force-velocity relation generated by a single microtubule. They found that the force is generally in the range predicted by the “polymerization ratchet” mechanism, but the force-velocity relationship decreased faster than that theory predicted. Here we generalize the polymerization ratchet model to take into account the “subsidy effect” that arises because a microtubule consists of 13 protofilaments. With this generalization the model fits the experimental data well. The biological implications of the polymerization ratchet model are discussed. Received: 18 May 1998 / Revised version: 4 November 1998 / Accepted: 30 November 1998     

Stoichiometry in producer-grazer systems: Linking energy flow with element cycling

All organisms are composed of multiple chemical elements such as carbon, nitrogen and phosphorus. While energy flow and element cycling are two fundamental and unifying principles in ecosystem theory, population models usually ignore the latter. Such models implicitly assume chemical homogeneity of all trophic levels by concentrating on a single constituent, generally an equivalent of energy. In this paper, we examine ramifications of an explicit assumption that both producer and grazer are composed of two essential elements: carbon and phosphorous. Using stoichiometric principles, we construct a two-dimensional Lotka-Volterra type model that incorporates chemical heterogeneity of the first two trophic levels of a food chain. The analysis shows that indirect competition between two populations for phosphorus can shift predator—prey interactions from a (+, −) type to an unusual (−, −) class. This leads to complex dynamics with multiple positive equilibria, where bistability and deterministic extinction of the grazer are possible. We derive simple graphical tests for the local stability of all equilibria and show that system dynamics are confined to a bounded region. Numerical simulations supported by qualitative analysis reveal that Rosenzweig’s paradox of enrichment holds only in the part of the phase plane where the grazer is energy limited; a new phenomenon, the paradox of energy enrichment, arises in the other part, where the grazer is phosphorus limited. A bifurcation diagram shows that energy enrichment of producer—grazer systems differs radically from nutrient enrichment. Hence, expressing producer—grazer interactions in stoichiometrically realistic terms reveals qualitatively new dynamical behavior.     

Constraints on food chain length arising from regional metacommunity dynamics

Classical ecological theory has proposed several determinants of food chain length, but the role of metacommunity dynamics has not yet been fully considered. By modelling patchy predator-prey metacommunities with extinction-colonization dynamics, we identify two distinct constraints on food chain length. First, finite colonization rates limit predator occupancy to a subset of prey-occupied sites. Second, intrinsic extinction rates accumulate along trophic chains. We show how both processes concur to decrease maximal and average food chain length in metacommunities. This decrease is mitigated if predators track their prey during colonization (habitat selection) and can be reinforced by top-down control of prey vital rates (especially extinction). Moreover, top-down control of colonization and habitat selection can interact to produce a counterintuitive positive relationship between perturbation rate and food chain length. Our results show how novel limits to food chain length emerge in spatially structured communities. We discuss the connections between these constraints and the ones commonly discussed, and suggest ways to test for metacommunity effects in food webs.     

The eggs and larvae of the giant mudskipper, <Emphasis Type="Italic">Periophthalmodon schlosseri</Emphasis>, collected from a mudflat in Penang,Malaysia

 Eggs of the giant mudskipper, Periophthalmodon schlosseri were collected from a burrow in Penang, Malaysia, in November 1998, and hatched larvae were reared in the laboratory. The eggs were demersal with adhesive filaments and elliptical in shape (0.83–1.43 mm in long-axis diameter). Newly hatched larvae (2.1–2.6 mm in notochord length) possessed a yolk sac. The number of myomeres was 10 + 17 = 27. The mouth and anus were already opened. The larvae started feeding one day after hatching and completely absorbed the yolk by the third day at a water temperature of 24.5–28.0°C. Received: April 9, 2002 / Revised: October 25, 2002 / Accepted: December 10, 2002     

Effects of invasive macrophyte on trophic diversity and position of secondary consumers

Invasive species are one of the widespread stressors of aquatic ecosystems. Several studies document food web effects of invasive fish, but little information is available on the effects of invasive macrophytes. We studied differences in food chain length as well as trophic position and trophic diversity of fish and odonates in lakes dominated by native plants or invasive Eurasian watermilfoil. Trophic position and food chain length were determined using baseline-adjusted δ¹⁵N isotope signatures. Trophic diversity, or isotope niche width, was estimated from convex hull area analysis. Results show that trophic position of secondary consumers was not affected by the invasive macrophyte, whereas trophic diversity was greater in watermilfoil-dominated lakes. The direction of isotopic niche expansion was different in fish and odonates, suggesting potential decoupling in predator–prey interactions. This study shows that dominant non-native macrophytes may cause significant changes in food web structure of invaded ecosystems. Trophic diversity may be a more sensitive indicator of environmental stress than trophic position and has the potential to be used for assessment of invasive species impacts and restoration success.     

Using an Ecosystem Modeling Approach to Explore Possible Ecosystem Impacts of Fishing in the Beibu Gulf,Northern South China Sea

Using the Ecopath with Ecosim software, a trophic structure model of the Beibu Gulf was constructed to explore the energy flows and provide a snapshot of the ecosystem operations. Input data were mainly from the trawl survey data collected from October 1998 to September 1999 and related literatures. The impacts of various fishing pressure on the biomass were examined by simulation at different fishing mortality rates. The model consists of 20 functional groups (boxes), each representing organisms with a similar role in the food web, and only covers the major trophic flows in the Beibu Gulf ecosystem. It was found that the food web of the Beibu Gulf was dominated by the primary producers path, and phytoplankton was the primary producer mostly used as a food source. The fractional trophic levels ranged from 1.0 to 4.02, and the marine mammals occupied the highest trophic level. Using network analysis, the ecosystem network was mapped into a linear food chain, and six discrete trophic levels were found with a mean transfer efficiency of 11.2%. The Finn cycling index was 9.73%. The path length was 1.821. The omnivory index was 0.197. The ecosystem had some degree of instability due to exploitation and other human activities, according to Odum’s theory of ecosystem development. A 10-year simulation was performed for each fishery scenario. The fishing mortality rate was found to have a strong impact on the biomass. By keeping the fishing mortality rate at the current level for all fishing sectors, scenario 1 had a drastic decrease in the large fish groups. The biomass of the small and medium pelagic fish would increase to some extent. The biomass of the small and low trophic level species, jellyfish, prawns and benthic crustaceans would be stable. The total biomass of the fishery resources would have a 10% decrease from the current biomass after 10 years. In contrast, the reduced fishing mortality rate induced the recovery of biomass (scenarios 2–4). In scenario 2, the biomass of the large demersal fish and the large pelagic fish would increase to over 16 times and 10 times, respectively, of their current level. In scenario 4, the biomass of the large pelagic fish would increase to over 3 times of its current level. The total biomass of the fish groups, especially the high trophic level groups, would become significantly higher after 10 years, which illustrates the contribution on biomass recovery by relaxing the fishing pressure. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users. Author contributions: Xiaoping Jia designed research; Zuozhi Chen and Yongsong Qiu performed research; Zuozhi Chen, Yongsong Qiu, and Shannan Xu analyzed data; and Zuozhi Chen and Shannan Xu wrote the article.     

A mathematical model of the adaptive control of human arm motions

This paper discusses similarities between models of adaptive motor control suggested by recent experiments with human and animal subjects, and the structure of a new control law derived mathematically from nonlinear stability theory. In both models, the control actions required to track a specified trajectory are adaptively assembled from a large collection of simple computational elements. By adaptively recombining these elements, the controllers develop complex internal models which are used to compensate for the effects of externally imposed forces or changes in the physical properties of the system. On a motor learning task involving planar, multi-joint arm motions, the simulated performance of the mathematical model is shown to be qualitatively similar to observed human performance, suggesting that the model captures some of the interesting features of the dynamics of low-level motor adaptation. Received: 20 September 1994 / Accepted in revised form: 18 November 1998     

Soil biota responses to long-term atmospheric CO2 enrichment in two California annual grasslands

Root, arbuscular-mycorrhizal (AM), soil faunal (protozoa and microarthropods), and microbial responses to field exposure to CO₂ for six growing seasons were measured in spring 1997 in two adjacent grassland communities. The grasslands showed contrasting root responses to CO₂ enrichment: whereas root length was not affected in the sandstone grassland, it was greater in the serpentine grassland, as was specific root length. AM fungal hyphal lengths were greater in the sandstone, but were unaffected in the serpentine community. This lent support to the hypothesis that there may be a tradeoff in resource allocation to more fine roots or greater mycorrhizal extraradical hyphal length. AM root infection was greater in both communities at elevated CO₂, as was the proportion of roots containing arbuscules. Our data on total hyphal lengths, culturable and active fungi, bacteria, and protozoa supported the hypothesis that the fungal food chain was more strongly stimulated than the bacterial chain. This study is one of the first to test these hypotheses in natural multi-species communities in the field. Received: 23 October 1998 / Accepted: 19 February 1999