Seasonal cycles and persistence in an acarine predator-prey system on cassava in Africa

We applied time series analysis and a mechanistic predator-prey model to long-term data of monthly population counts of the herbivorous pest mite Mononychellus tanajoa and its introduced phytoseiid predator Typhlodromalus aripo from a cassava field in Benin, West Africa. In this approach, we determined the extent to which the main features of the observed predator-prey fluctuations in cassava fields can be explained from biotic traits inherent to the biology of predator and prey, and the extent of the significance of abiotic factors in determining population levels. The time series analyses with cross-correlation showed that the period of predator-prey fluctuations coincided with the annual pattern of intense rainfall and onset of dry season. A pronounced M. tanajoa peak followed after a short lag (2 weeks) by a T. aripo peak coincided with a trough in rainfall intensity. Both the prey and predator had local and lower peaks that coincided with high rainfall intensity, but with a considerably longer lag (ca. 3 months) compared with the high peaks occurring at the onset of the dry season. Regression of log-transformed data series (over a 7-year period) showed that—except for the first year after predator release—M. tanajoa fluctuated around an almost time-invariant mean population density, while T. aripo densities showed a consistent decline over the full observation period. To explain observed trends and periodic components in the data-series of predator and prey densities, we review hypotheses that are based on (1) the annual patterns and trends in abiotic factors, (2) mechanisms endogenous to the predator-prey system and (3) a combination of exogenous and endogenous factors.     

Patterns of exclusion in an intraguild predator-prey system depend on initial conditions

1. When intraguild (IG)-prey are superior to IG-predators in competing for a shared resource, theory predicts coexistence of the IG-prey or the IG-predator with the resource depending on the productivity level: (a) resource and IG prey coexist when productivity is low; (b) IG-predator and resource coexist at high productivity; (c) if IG-prey and IG-predators can coexist, it is only at intermediate productivity levels. 2. We tested the existence of productivity-dependent regions of coexistence using an experimental system of two predatory mites and a shared food source (pollen). 3. At high levels of pollen supply (i.e. high productivity), the IG-predator excluded the IG-prey in most, but not all, cases. The same pattern of exclusion was observed at low productivity, at which the IG-prey was expected to exclude the IG-predator. Therefore, species composition could not be predicted by productivity levels. Instead, our results show that initial conditions affected strongly the outcome of the interaction. 4. We emphasize the need for theory on IG-predation that takes the effects of stage structure, initial conditions and transient dynamics into account.     

Can plants betray the presence of multiple herbivore species to predators and parasitoids? The role of learning in phytochemical information networks

In response to feeding by phytophagous arthropods, plants emit volatile chemicals. This is shown to be an active physiological response of the plant and the released chemicals are therefore called herbivore-induced plant volatiles (HIPV). One of the supposed functions of HIPV for the plant is to attract carnivorous natural enemies of herbivores. Depending on which plant and herbivore species interact, blends of HIPV show qualitative and quantitative variation. Hence, one may ask whether this allows the natural enemies to discriminate between volatiles from plants infested by herbivore species that are either suitable or unsuitable as a food source for the natural enemy. Another question is whether natural enemies can also recognise HIPV when two or more herbivore species that differ in suitability as a food source simultaneously attack the same plant species. By reviewing the literature we show that arthropod predators and parasitoids can tell different HIPV blends apart in several cases of single plant–single herbivore systems and even in single plant–multiple herbivore systems. Yet, there are also cases where predators and parasitoids do not discriminate or discriminate only after having learned the association between HIPV and herbivores that are either suitable or non-suitable as a source of food. In this case, suitable herbivores may profit from colonising plants that are already infested by another non-suitable herbivore. The resulting temporal or partial refuge may have important population dynamical consequences, as such refuges have been shown to stabilise otherwise unstable predator–prey models of the Lotka-Volterra or Nicholson-Bailey type.     

Spider mite web mediates anti-predator behaviour

Herbivores suffer significant mortality from predation and are therefore subject to natural selection on traits promoting predator avoidance and resistance. They can employ an array of strategies to reduce predation, for example through changes in behaviour, morphology and life history. So far, the anti-predator response studied most intensively in spider mites has been the avoidance of patches with high predation risk. Less attention has been given to the dense web produced by spider mites, which is a complex structure of silken threads that is thought to hinder predators. Here, we investigate the effects of the web produced by the red spider mite, Tetranychus evansi Baker & Pritchard, on its interactions with the predatory mite, Phytoseiulus longipes Evans. We tested whether female spider mites recognize predator cues and whether these can induce the spider mites to produce denser web. We found that the prey did not produce denser web in response to such cues, but laid more eggs suspended in the web, away from the leaf surface. These suspended eggs suffered less from predation by P. longipes than eggs that were laid on the leaf surface under the web. Thus, by altering their oviposition behaviour in response to predator cues, females of T. evansi protect their offspring.     

Interactions Between Two Neotropical Phytoseiid Predators on Cassava Plants and Consequences for Biological Control of a Shared Spider Mite Prey: a Screenhouse Evaluation

The issue of introducing single or multiple natural enemy species for classical biological control has been an area of intense inquiry by ecologists and biological control practitioners. This is particularly relevant to classical biological control of cassava green mite Mononychellus tanajoa (Bondar) (Tetranychidae) in Africa, as this pest mite is shared by several natural enemies in the Neotropics (its area of origin), two of which have been introduced and established widely in Africa. We conducted two screenhouse experiments using the two neotropical phytoseiid predatory mites, Typhlodromalus aripo DeLeon and Typhlodromalus manihoti Moraes, to determine the effects of single and two-predator species on population dynamics of the two predators and on suppression of M. tanajoa populations. The two predators are thought to be complementary in their impact on their shared prey M. tanajoa, due to similarities in their preference for this prey and to differences in their spatial distribution and foraging activities on cassava. The two predator species were released alone or together at low and at high initial densities of M. tanajoa. In all cases, predator releases resulted in significant suppression of M. tanajoa, but the degree of suppression did not differ among single and two-species releases with one exception: at high initial density of M. tanajoa, releases of T. aripo alone had less impact than that of either T. manihoti alone or of the two species together. Typhlodromalus aripo also appeared to be inferior as a competitor of T. manihoti: at low initial density of M. tanajoa, the proportion of T. aripo in the two-predator release treatments gradually declined and was strikingly lower than in the single species release, probably due to intraguild predation on its larvae by T. manihoti. However, T. aripo persisted longer than T. manihoti after elimination of M. tanajoa. On the basis of this study under semi-natural conditions, it appears that either species is sufficient for controlling M. tanajoa populations, with T. manihoti being more efficient at high initial prey densities and T. aripo at low initial prey densities. At high prey density, T. manihoti increased to large numbers and outcompeted T. aripo. Relevance of these findings to larger spatial scale and under natural conditions is discussed.     

Within- and between-population variation for Wolbachia-induced reproductive incompatibility in a haplodiploid mite

Wolbachia pipientis is a bacterium that induces cytoplasmic incompatibility (CI), the phenomenon in which infected males are reproductively incompatible with uninfected females. CI spreads in a population of hosts because it reduces the fitness of uninfected females relative to infected females. CI encompasses two steps: modification (mod) of sperm of infected males and rescuing (resc) of these chromosomes by Wolbachia in the egg. Infections associated with CI have mod+ resa+ phenotypes. However, mod- resc+ phenotypes also exist; these do not result in CI. Assuming mod/resc phenotypes are properties of the symbiont, theory predicts that mod- resc+ infections can only spread in a host population where a mod+ resc+ infection already occurs. A mod- resc+ infection spreads if the cost it imposes on the infected females is lower than the cost inflicted by the resident (mod+ resc+) infection. Furthermore, introduction of a mod- Wolbachia eventually drives infection to extinction. The uninfected population that results can be recolonized by a CI-causing Wolbachia. Here, we investigated whether variability for induction of CI was present in two Tetranychus urticae populations. In one population all isofemale lines tested were mod-. In the other, mod+ resc+ and mod- resc+ isofemale lines coexisted. We found no evidence for a cost difference to females expressing either type (mod-/-). Infections in the two populations could not be distinguished based on sequences of two Wolbachia genes. We consider the possibility that mod- is a host effect through a population dynamics model. A mod- host allele leads to infection extinction in the absence of fecundity differences. Furthermore, the uninfected population that results is immune to reestablishment of the (same) CI-causing Wolbachia.     

Age-dependent rates of infection of cassava green mites by a fungal pathogen in Brazil

Age-specific effects of invertebrate pathogens on their hosts can greatly influence the population dynamics in such interactions. Explanations for such differences are usually sought within differing intrinsic susceptibilities of the host life stages but we present data which indicate that host size, behaviour and life history may be the overriding factors determining age-specific effects of a fungal pathogen, Neozygites floridana (Entomophthorales: Neozygitaceae) on spider mites (Mononychellus tanajoa Bondar, Acari: Tetranychidae). Epizootics of N. floridana in spider mites are characterised by much greater relative mortality of adult females compared with other life stages (ca. 99%), despite similar physiological susceptibilities. We present empirical data that demonstrate encounter rates of mites with N. floridana increasing with life stage during an epizootic on cassava in northeastern Brazil. Estimates of the size, walking speeds and patterns, and life history of different life stages (and adult sexes) were used to calculate expected relative encounter rates which were found not to be different from the observed values (although not testable for larvae). This helps explain the different apparent susceptibility of host life stages in the field. Given the low ecological susceptibility of younger life stages to this pathogen, we predict that the interaction time between host and pathogen, determined by climatic conditions, will be critical in determining the degree of host population control in an epizootic. We further hypothesise that such variation in ecological susceptibility to pathogens can generate selection pressures on basic host traits, contributing to the sessile nature of many microarthropods. This revised version was published online in July 2006 with corrections to the Cover Date.     

Adaptive learning in arthropods: spider mites learn to distinguish food quality

Many herbivorous arthropods have been shown to possess learning capabilities, yet fitness effects of learning are largely unknown. In this paper, we test whether two-spotted spider mites (Tetranychus urticae) learn to distinguish food quality in choice tests, and whether this results in fitness benefits. Food consisted of cucumber plants with one of three degrees of feeding damage: undamaged (no mites), mildly damaged (infested by a mite strain adapted to tomato) and heavily damaged (infested by a mite strain adapted to cucumber). Mites were subjected to one choice test in a greenhouse and three sequential choice tests on leaf disks. Thereafter, individual mite performance was measured as oviposition rate over four days. In the course of the three small-scale choice tests, preference shifted towards less damaged food. The performance tests showed that learning was adaptive: mites learned to prefer the food type that yielded the higher oviposition rate. Interestingly, innate preferences in the greenhouse tests were close to those shown after learning in the small-scale tests. Given that both strains of mites had not experienced cucumber for several years, we hypothesize that the preference in the greenhouse was due to avoidance of mite odours rather than odours of damaged plants. Through its effect on foraging behaviour, adaptive learning may promote the evolution of host plant specialization in herbivorous arthropods.     

State-dependent and odour-mediated anemotactic responses of the predatory mite Phytoseiulus persimilis in a wind tunnel

Anemotaxis in the predatory mite Phytoseiulus persimilis (both well-fed and starved), has previously been studied on a wire grid under slight turbulent airflow conditions yielding weak, yet distinct, gradients in wind speed and odour concentration (Sabelis and Van der Weel 1993). Such conditions might have critically influenced the outcome of the study. We repeated these experiments, under laminar airflow conditions on a flat surface in a wind tunnel, thereby avoiding variation in wind speed and odour concentration. Treatments for starved and well-fed mites were (1) still-air without herbivore-induced plant volatiles (HIPV) (well-fed mites only), (2) an HIPV-free air stream, and (3) an air stream with HIPV (originating from Lima bean plants infested by two-spotted spider mites, Tetranychus urticae). Well-fed mites oriented in random directions in still-air without HIPV. In an air stream, starved mites always oriented upwind, whether plant odours were present or not. Well-fed mites oriented downwind in an HIPV-free air stream, but in random directions in an air stream with HIPV. Only under the last treatment our results differed from those of Sabelis and Van der Weel (1993).     

ImageParser: a tool for finite element generation from three-dimensional medical images

Background

The finite element method (FEM) is a powerful mathematical tool to simulate and visualize the mechanical deformation of tissues and organs during medical examinations or interventions. It is yet a challenge to build up an FEM mesh directly from a volumetric image partially because the regions (or structures) of interest (ROIs) may be irregular and fuzzy.

Methods

A software package, ImageParser, is developed to generate an FEM mesh from 3-D tomographic medical images. This software uses a semi-automatic method to detect ROIs from the context of image including neighboring tissues and organs, completes segmentation of different tissues, and meshes the organ into elements.

Results

The ImageParser is shown to build up an FEM model for simulating the mechanical responses of the breast based on 3-D CT images. The breast is compressed by two plate paddles under an overall displacement as large as 20% of the initial distance between the paddles. The strain and tangential Young's modulus distributions are specified for the biomechanical analysis of breast tissues.

Conclusion

The ImageParser can successfully exact the geometry of ROIs from a complex medical image and generate the FEM mesh with customer-defined segmentation information.