Characterisation of the arrestment responses of Trichogramma evanescens

Summary Contact kairomones and oviposition in a host egg stimulated arrestment behaviour in Trichogramma evanescens, characterised by a reduction in walking speed and increased turning. Previous oviposition experience did not influence a parasitoid's response to contact kairomones, but successive encounters with kairomone patches resulted in parasitoids habituating to the contact chemical. Oviposition on a kairomone patch did not reverse this habituation effect. It was concluded that contact kairomones and host eggs will both contribute independently to the duration of a patch visit. The selection of patches by T. evanescens will depend on its response to kairomones. Results from this study indicate that the application of contact kairomones to field crops will not necessarily increase the probability of parasitoids finding hosts.     

The use of visual information by house flies,Musca domestica (Diptera: Muscidae), foraging in resource patches

Summary House flies, Musca domestica, respond to visual contrasts on the substrate if a resource is associated with the contrasting patterns. Visible resource patch boundaries serve as a signal to flies that they are about to leave a rewarding patch. Searching flies respond to such visual information by walking along the resource patch boundary and turning back into the patch at its edge. This edge detection and response serve as a mechanism for flies with visual cues to stay in a rewarding patch and locate more resources within it. The intensity of their response correlates with the quality of the resource. In the absence of visual cues, patch shape affects foraging success; flies find more resources in circular than in linear resource distributions. The effects of visual cues, however, render patch shape unimportant. Various substrate contrasts are effective as resource information for flies: dark (e.g., green) figures on bright (e.g., white) backgrounds or bright figures on dark backgrounds. Responses to substrate contrasts measured in this study indicate that, over the short term, house flies can learn a visual cue associated with a food source.     

Impact of Host Plant Connectivity,Crop Border and Patch Size on Adult Colorado Potato Beetle Retention

Tagged Colorado potato beetles (CPB), Leptinotarsa decemlineata (Say), were released on potato plants, Solanum tuberosum L., and tracked using a portable harmonic radar system to determine the impact of host plant spatial distribution on the tendency of the pest to remain on the colonized host plant or patch. Results confirmed the long residency time on the host plant and showed that close connection of the plant to neighboring plants hastened dispersal between plants. Tracking walking CPB for over 6 h in small potato plots revealed that all types of mixed borders tested (potato/bare ground, potato/timothy and potato/woodland) acted as a strong barrier and retained beetles within the patch. In another experiment in potato patches surrounded by bare ground borders, tracked walking CPB displayed similar behaviour for up to four days. The distribution of turning angles in the CPB walking paths was not uniform and corresponded to beetles following the edge rows of potato patches in response to the crop border barrier or reversing their direction as they reached the end of a row and therefore a border. Patch size had no or little effect on beetle retention in the patch. The relative distribution of counts of tagged beetles detected among small (16 m²), medium (64 m²) and large size (256 m²) patches of potato four days after initial release remained similar to that of numbers released. Even though mixed crop borders were a strong barrier to walking CPB emigrating from potato patches, the departure rate of beetles over time was high. Results suggest that the effect of mixed borders is largely limited to dispersal by walking and does not apply to beetles leaving host patches by flight. The manipulation of crop borders and patch size seem to have limited potential for the management of CPB emigrating from potato fields.     

Volatile spider-mite pheromone and host-plant kairomone, involved in spaced-out gregariousness in the spider mite Tetranychus urticae

ABSTRACT. From the host plant-spider mite complex Phaseolus lunatus—Tetranychus urticae Koch a volatile chemical is emitted that acts as a kairomone for the predatory mite Phytoseiulus persimilis Athias-Henriot (Sabelis et al. , 1984a). This kairomone is apparently a byproduct of a vital physiological process and/or it has a function in the biology of the spider mite as well.
The spider mite—host plant complex also emits a volatile spider-mite dispersing pheromone. This is shown in the present study where spider mites were introduced into an odour patch on a horizontal screen in a vertical airflow olfactometer. When spider-mite infested leaves of Lima bean are offered, the spider mites walk mainly straight and soon reach the edge of the screen. On the other hand, when clean Lima bean leaves are offered, the mites walk tortuously most of the time and reach the edge of the screen much later. Artificially damaged plants elicit the same response as undamaged plants. Differences in spider-mite behaviour are observed in the vertical airflow olfactometer when odour of either clean or spider-mite infested leaves is offered. A comparison of the behaviour in these two situations with that when no odour was offered suggests that Lima bean leaves emit a volatile kairomone that activates T. urticae and makes them return after losing the stimulus. A Y-tube olfactometer experiment confirms the existence of this kairomone.
At a low ratio of dispersing pheromone to plant kairomone, the spider mites behave as if only kairomone is present, walking mainly tortuously. At a high ratio they disperse. No aggregation-pheromonal effect is observed.
The possibility that the spider-mite dispersing pheromone acts as a kairomone for P. persimilis is discussed.     

Volatile and Contact Chemicals Released by Nezara viridula (Heteroptera:Pentatomidae) Have a Kairomonal Effect on the Egg Parasitoid Trissolcus basalis (Hymenoptera: Scelionidae)

The responses of females of the egg parasitoid Trissolcus basalis (Wollaston) (Hymenoptera: Scelionidae) to volatile and contact chemicals from its host Nezara viridula (L.) (Heteroptera: Pentatomidae) were investigated in a Y-tube olfactometer and under open arena conditions. In the Y-tube tests, volatiles from virgin males and from females in a preovipositional state attracted T. basalis females, while volatiles from host virgin females did not. In an open arena, traces left by N. viridula adults in different physiological conditions function as contact cues inducing the wasps to remain longer in the arena and to change the pattern of their walking behavior. However, only contact kairomones from N. viridula mated females in a preovipositional condition induced an arrestment response characterized by an increase in patch searching time and turning rates and a reduction in linear speed. The chemical ecological implications of these results on this host–parasitoid association are discussed.     

Concentration effects and the response of Nemeritis canescens to a secretion of its host

An ovipositional response in the ichneumonid parasite Nemeritis canescens is elicited by a compound from the mandibular gland secretion of flour moth caterpillars. Exposure to that compound, even at concentrations too low to elicit an observable response, influences the parasites' subsequent response to a host larva or to the compound itself, increasing or decreasing that subsequent response to an extent that depends on the timing and on the amount of the compound at the initial exposure. When the same compound is deposited on the parasites' antennae, as by attacked host larvae, the parasites groom and oviposition is postponed.     

Host finding by Uscana lariophaga (Hymenoptera: Trichogrammatidae) in stored cowpea: the effect of distance, time interval, host patch size and spatial orientation

Host finding and parasitization by Uscana lariophaga Steffan, a potential biocontrol agent of the storage pest Callosobruchus maculatus (Fabricius), were investigated in stored cowpea. Host finding was shown to be a function of distance, time, host patch size and the spatial position of U. lariophaga relative to the host patch. Uscana lariophaga females were able to find hosts up to 75 cm horizontal distance from the host patch, which was the largest distance tested. The probability that a host patch was found when an individual U. lariophaga female was released at 2.5 cm horizontal distance from the host patch ranged from 0.6 after 2 h foraging time to 0.9 after 8 h foraging time. At 10 cm from the host patch, host finding probability ranged from 0.2 to 0.45 at these respective foraging times. Finding probabilities doubled compared to horizontal distances when U. lariophaga was released below the host patch, and halved when it was released above the host patch. The negative geotaxic response was shown not to be an artefact of the release method. The median net displacement rate in the direction of the host patch was two beans per hour (1.4 cm h(-1)). The results suggest that U. lariophaga females start searching for hosts regardless of the quality of the olfactory information they receive. Additional observations indicated that U. lariophaga is adapted to a host with a patchy distribution, which implies that host finding over larger distances is relevant for U. lariophaga.     

Olfactory orientation responses of the eucalyptus woodborer, Phoracantha semipunctata, to host plant in a wind tunnel

The eucalyptus woodborer, Phoracantha semipunctata Fabricius (Coleoptera: Cerambycidae), attacks mainly species of Eucalyptus (Myrtaceae). This study investigated walking and flight behaviour of P. semipunctata males and females exposed to an odour plume originating from a log of E. globulus placed vertically in the upwind end of a wind tunnel. In control experiments, beetles were exposed to a PVC drainpipe in the same position as the log, providing a visual stimulus without host‐tree odour. No statistical differences were found between behavioural responses of either sex when exposed to the log or PVC pipe. No beetles landed on the PVC pipe, whereas 49% of the beetles exposed to host‐tree odour plume landed on the log. Beetles aged over 24 days after emergence from the host tree were more responsive than beetles aged 20–24 days, and accounted vor 86% of the beetles that landed on the log. While walking, host‐tree odour affected the behaviour of the beetles that landed on the log as follows: upwind movement and path linearity increased, whereas turning rate, stopping frequency, mean stopping time and time to take‐off flight decreased. During flight, host‐tree odour affected the behaviour of the beetles that landed on the log as follows: increased upwind flight, turning rate, flight time, flight distance, and decreased flight speed. For beetles that never lost contact with the odour plume, flight progressed upwind with narrow zigzags, and showed higher directedness upwind, path linearity, faster flight speed and lower turning rate than for beetles that lost contact with the odour plume. After loosing contact with the plume, beetles tended to decrease their upwind progression, exhibiting a sharp turn or quick counterturns followed by crosswind or downwind excursions. This led to regaining contact with the odour plume and resumed upwind progression at higher speed provided they flew within the boundaries of the plume. The results showed that host‐tree odour affects both walking and flight behaviour of P. semipunctata beetles, inducing a more directed upwind movement and landing on the visual stimulus of a tree trunk.     

Lipid-glass adhesion in giga-sealed patch-clamped membranes.

Adhesion between patch-clamped lipid membranes and glass micropipettes is measured by high contrast video imaging of the mechanical response to the application of suction pressure across the patch. The free patch of membrane reversibly alters both its contact angle and radius of curvature on pressure changes. The assumption that an adhesive force between the membrane and the pipette can sustain normal tension up to a maximum Ta at the edge of the free patch accounts for the observed mechanical responses. When the normal component of the pressure-induced membrane tension exceeds Ta membrane at the contact point between the free patch and the lipid-glass interface is pulled away from the pipette wall, resulting in a decreased radius of curvature for the patch and an increased contact angle. Measurements of the membrane radius of curvature as a function of the suction pressure and pipette radius determine line adhesion tensions Ta which range from 0.5 to 4.0 dyn/cm. Similar behavior of patch-clamped cell membranes implies similar adhesion mechanics.     

Differences in egg wastage by superparasitism,contrastingVenturia (=Nemeritis) canescens searching singly versus searching in groups

A statistical and graphic study is presented of the wastage of eggs by Venturia (=Nemeritis) canescens when searching singly and in groups of 10 among hosts at four different host densities in laboratory universes as described byHuffaker andMatsumoto (preceding paper of this journal). The host insect was the fluour moth Anagasta kühniella and the host densities used were 10, 30, 100 and 200 per universe. Intensity of egg wastage due to superparasitim varied significantly according to host density, and between the two parasite densities employed, 1 and 10, using both F-tests and chi-square tests. Plots of k-factor analysis on this egg wastage showed high negative correlations with host density, and the raw data for single parasites was well represented by a parabola while that for the grouped parasites departed from this relationship only at the lowest host density.     

The impact of patch encounter rate on patch residence time of female parasitoids increases with patch quality

Abstract.  1. For animal species that forage on patchily distributed resources, patch time allocation is of prime importance to their reproductive success. According to Charnov's marginal value theorem (MVT), the rate of patch encounter should influence negatively the patch residence time: as the rate of patch encounter decreases, the patch residence time increases. Moreover, the MVT predicts that animals should stay longer in high quality patches.
2. Using the aphid parasitoid Aphidius rhopalosiphi (Hymenoptera: Aphidiinae), the effects of these two factors (patch encounter rate and host density) were combined in order to test if the increment in patch residence time for a given decrease in patch encounter rate was larger for high quality patches than for low quality patches.
3. The results show a significant effect of the interaction between the two factors. In high host density patches, parasitoids spent more time if they experienced a low patch encounter rate, while in low host density patches, patch encounter rate had no significant effect on the patch residence time. This suggests that the response of A. rhopalosiphi females to patch encounter rate varied with host density in the patch. Moreover, the same interaction effect was observed for the number of ovipositor contacts on aphids.
4. Parasitoid females can use patch encounter rate to estimate patch density in the habitat but the effect of this estimate on their patch residence time is modulated by patch quality. Staying longer in a patch when patches are rare is more advantageous when the fitness gained by doing so is large. In low quality patches, the expected fitness gain is small and the female may gain more by leaving and taking her chance at finding another patch.     

The particulate basis of the resistance of a parasitoid to the defence reactions of its insect host.

Small, membrane-bound particles, about 130 nm in diameter, have been isolated from that region of the genital tract of the ichneumon Nemeritis known as the calyx. These particles have been shown to possess the ability to confer resistance to encapsulation by Ephestia, the natural host of Nemeritis, upon a surface which would otherwise be encapsulated.     

Arrestment responses of the predatory mite, Phytoseiulus persimilis, to steep odour gradients of a kairomone

ABSTRACT. The response of the predatory mite, Phytoseiulus persimilis Athias-Henriot, to steep gradients of a volatile kairomone emitted by its prey, Tetranychus urticue Koch, was studied in a vertical air flow chamber. The orientation to wind direction was eliminated by using an olfactometer that had an air stream approaching the predator from below a gauze screen upon which the predator walked. The steep gradient of odour was obtained by putting a cylinder filled with prey-infested leaves vertically below the screen. Starved predators were arrested in the odour patch by walking more slowly and tortuously than well-fed predators. The latter mites did not show a significant ortho- or klinokinetic response to the presence of odour. Both well-fed and starved predators showed a chemotactic response to steep gradients at the border of the circular odour patch. Predators that happened to walk out of the patch, frequently turned back to it. This response is presumably based on idiothetic information about the predator's immediately previous walking directions, because it occurred in the odour-free zone after passing the steep gradient of prey odour. Right-about turns can help the predator to stay in static odour plumes with steep gradients at the borders. This type of plume is present only close to the odour source. Further away from the source the odour plume tends to move to and fro due to variation in wind direction. For the predator to keep track of these snaking plumes the right-about turns are unlikely to be of any value because the response is of short duration and because the response to a moving plume appeared to be inadequate; by moving the cylinder below the screen (and consequently the odour patch) it was found that the predator turned back even if the odour gradient was made to pass the predator in the same direction as that of the predator's movement.     

Response of fauna in seagrass to habitat edges,patch attributes and hydrodynamics

This study has investigated the taxon‐specific responses of fauna to patch edges, and how these relate to patch attributes (patch size, seagrass biomass and water depth), and hydrodynamics in the seagrass habitat. Faunal abundances were sampled at the edge, 2 m in from the edge, and in the middle of 10 seagrass patches of variable size in Port Phillip Bay, Australia. Five of nine taxa showed edge effects. There were higher abundances at the edge compared with the middle for porcellid harpacticoids, and an increase in abundance from the edge to the middle of the patches for tanaids and isopods. For caprellid and gammarid amphipods, the edge effect varied across patches. Changes in current within the patch and patch size were related to the variability in the edge effect pattern of caprellids. None of the measured environmental variables (seagrass biomass, current and water depth) or patch size had a role in the variable edge effect pattern of gammarid amphipods. At the patch level, the distribution of six of nine taxa in this study, namely isopods, polychaetes, ‘other harpacticoids’, porcellid harpacticoids, cumaceans and gammarid amphipods, was related to differences in average water depth, average seagrass biomass and patch size. Our study indicates that the faunal response to edges cannot be generalized across seagrass habitat, and the implications of habitat area loss will vary depending on the taxon under consideration.     

Responses of the tropical bont tick, Amblyomma variegatum (Fabricius), to its aggregation-attachment pheromone presented in an air stream on a servosphere

Male Amblyomma variegatum ticks feeding on a host release a mixture of o -nitrophenol and methyl salicylate which serves to attract conspecifics. The behavioural responses of A. variegatum on a servosphere to these volatiles presented in an air stream are detailed here. In still air, ticks walked on all eight legs, but with long halts. In contrast, the air stream caused continuous walking and induced a reaching response where the forelegs actively sampled the air. Such reaching increased the angular velocity and reduced walking speed, effects that were amplified in the presence of vapours from o -nitrophenol and methyl salicylate in the air flowing over the ticks. Vapour from a 1:1 mixture of o -nitrophenol and methyl salicylate was attractive over a 10⁴-fold concentration range providing an increase in upwind displacement of 20–40%, significantly higher than the natural ratio where o -nitrophenol vapour predominates. Although the responses to o -nitrophenol vapour were variable when presented alone, this chemical was consistently attractive when delivered with steer hair odour – unattractive on its own. Moreover, the upwind walk to this combination did not cause a change in speed or angular velocity. This supports the hypothesis that the response to the pheromone is enhanced by host odour. Accepted: 16 October 1999     

Host frass as arrestant chemicals in locating host Mythimna separata by the tachinid fly Exorista japonica

We investigated the response of the tachinid fly, Exorista japonica (Townsend), to host frass or its extracts in order to clarify the host location mechanisms of female flies in a potential host habitat. Host searching time in a patch and the number of patch visits were analyzed by using a frass-containing patch which was excreted by host larvae, Mythimna separata (Walker) (Lepidoptera: Noctuidae), and patches to which host frass extracts were applied. E. japonica females were arrested in response to the host-frass-containing patch after contacting the frass with their front tarsi, thereby spending most of the time to search the patch and to revisit the host-frass patch. While host-searching time in the patch by the females was longest at their first visit of a patch with host frass, searching time decreased with successive visits. The female flies also exhibited area-restricted searching with methanol extracts of the host frass. Area-restricted searching activity increased with the concentration of host-frass extract, i.e., total searching time in the patch and the number of patch visits varied in a dose-dependent manner. E. japonica females likely employ chemicals in host frass as arrestants in host location.     

Formation and maintenance of aggregations in walleye pollock,Theragra chalcogramma,larvae under laboratory conditions: role of visual and chemical stimuli

Synopsis Although planktonic marine fish larvae are often distributed in aggregations, the role of behavioral responses to environmental factors in these aggregations is not well understood. This work examines, under laboratory conditions, the influence of visual and chemical stimuli in the formation and maintenance of aggregations in walleye pollock,Theragra chalcogramma, larvae. Larvae were exposed to a horizontal gradient of light (visual stimulus), prey scent (chemical stimuli: squid/copepod and rotifer) or prey density (visual & chemical stimuli: rotifers). While larvae did not respond to prey scent, they did respond to a gradient of light or prey, which resulted in the formation and maintenance of aggregations. Larvae moved into and remained in a zone of higher light intensity (0.56 versus 0.01 mol photons m^-2 s^-1). Once encountering a patch of prey, larvae remained aggregated within the patch to feed. In nature, movement of walleye pollock larvae in response to selected environmental factors (e.g., gravity, light, temperature, turbulence) may serendipitously bring them into contact with prey patches, where they then could remain to feed as long as light intensity remained at or above levels necessary for feeding.     

Assessment of the applicability of the Hertzian contact theory to edge-loaded prosthetic hip bearings

The components of prosthetic hip bearings may experience in-vivo subluxation and edge loading on the acetabular socket as a result of joint laxity, causing abnormally high, damaging contact stresses. In this research, edge-loaded contact of prosthetic hips is examined analytically and experimentally in the most commonly used categories of material pairs. In edge-loaded ceramic-on-ceramic hips, the Hertzian contact theory yields accurate (conservatively, <10% error) predictions of the contact dimensions. Moreover, the Hertzian theory successfully captures slope and curvature trends in the dependence of contact patch geometry on the applied load. In an edge-loaded ceramic-on-metal pair, a similar degree of accuracy is observed in the contact patch length; however, the contact width is less accurately predicted due to the onset of subsurface plasticity, which is predicted for loads >400N. The Hertzian contact theory is shown to be ill-suited to edge-loaded ceramic-on-polyethylene pairs due to polyethylene's nonlinear material behavior. This work elucidates the methods and the accuracy of applying classical contact theory to edge-loaded hip bearings. The results help to define the applicability of the Hertzian theory to the design of new components and materials to better resist severe edge loading contact stresses.     

Interspecific Relationships Between the Solitary Ectoparasitoid, Eupelmus vuilleti (Crw.) (Eupelmidae), and Its Sympatric Species, Dinarmus basalis (Rond.) (Pteromalidae), in the Presence of Their Host, Callosobruchus maculatus Pic (Coleoptera Bruchidae)

The eupelmid Eupelmus vuilleti CRW. and the pteromalid Dinarmus basalis Rond. are sympatric in West Africa. The reproduction of E. vuilleti and D. basalis females was analyzed in 10, 20, and 40 host patches when present alone or together. In 10- and 20-hosts patches, the presence of E. vuilleti affected the reproduction and offspring survival chances of D. basalis. In 40-hosts patch, only the offspring survival of D. basalis was affected by the presence of E. vuilleti. In contrast, the presence of D. basalis did not influence the reproduction of E. vuilleti and the survival chances of its offspring at all host densities tested. Multiparasitism did not occur at random. In a choice test, E. vuilleti showed a preference for hosts previously parasitized by D. basalis on healthy hosts. This attraction is mediated by chemical markers deposited by D. basalis females on the surface of the seed. At high host density, the likelihood of encountering a seed containing a host previously parasitized by D. basalis was lower for E. vuilleti, affecting moderately D. basalis reproductive success.     

Host plant finding in the specialised leaf beetle Cassida canaliculata: an analysis of small-scale movement behaviour

Abstract.  1. Host plant finding in walking herbivorous beetles is still poorly understood. Analysis of small-scale movement patterns under semi-natural conditions can be a useful tool to detect behavioural responses towards host plant cues.
2. In this study, the small-scale movement behaviour of the monophagous leaf beetle Cassida canaliculata Laich. (Coleoptera: Chrysomelidae) was studied in a semi-natural arena ( r = 1 m). In three different settings, a host ( Salvia pratensis L., Lamiales: Lamiaceae), a non-host ( Rumex conglomeratus Murr., Caryophyllales: Polygonaceae), or no plant was presented in the centre of the arena.
3. The beetles showed no differences in the absolute movement variables, straightness and mean walking speed, between the three settings. However, the relative movement variables, mean distance to the centre and mean angular deviation from walking straight to the centre, were significantly smaller when a host plant was offered. Likewise, the angular deviation from walking straight to the centre tended to decline with decreasing distance from the centre. Finally, significantly more beetles were found on the host than on the non-host at the end of all the trials.
4. It is concluded that C. canaliculata is able to recognise its host plant from a distance. Whether olfactory or visual cues (or a combination of both) are used to find the host plant remains to be elucidated by further studies.