Insect repellents and associated personal protection for a reduction in human disease

Personal protection measures against biting arthropods include topical insect repellents, area repellents, insecticide‐treated bednets and treated clothing. The literature on the effectiveness of personal protection products against arthropods is mainly limited to studies of prevention of bites, rather than prevention of disease. Tungiasis was successfully controlled by application of topical repellents and scrub typhus was reduced through the use of treated clothing. Successful reduction of leishmaniasis was achieved through the use of topical repellents, treated bednets and treated clothing in individual studies. Malaria has been reduced by the use of insecticide‐treated bednets (ITN), certain campaigns involving topical repellents, and the combination of treated bednets and topical repellents. Although area repellents such as mosquito coils are used extensively, their ability to protect humans from vector‐transmitted pathogens has not been proven. Taken together, the literature indicates that personal protection measures must be used correctly to be effective. A study that showed successful control of malaria by combining treated bednets and topical repellents suggests that combinations of personal protection measures are likely to be more effective than single methods. Implementation of successful programmes based on personal protection will require a level of cooperation commonly associated with other basic societal functions, such as education and food safety.     

Repellencies and toxicities of five ant-derived defensive compounds against the lone star tick, <Emphasis Type="Italic">Amblyomma americanum</Emphasis> (Acari: Ixodidae)

The lone star tick, Amblyomma americanum, is a vector of several important human and animal diseases. This tick species has rapidly expanded in its geographic distribution, and its aggressive behavior has increased the risk of tick-borne diseases in these new areas. Repellents are recommended by the Centers for Disease Control and Prevention (CDC) for protection against tick bites. DEET is the most common repellent, but public concerns over its safety have increased the need for alternative safe and efficacious tick repellents. Several naturally derived animal compounds have been tested against other species of ticks or other arthropod pests, but not against A. americanum. Based on EC₅₀ values obtained using a vertical paper bioassay, decylamine and MT-710 (a 2-tridecanone formulation) were found to be as repellent as DEET. Those two substances along with 2-tridecanone were also found to be as repellent as DEET when their EC₉₅ values were compared. Lone star ticks were more susceptible to the toxic effects of DEET in glass vial assays than all of the ant-derived defensive compounds/formulations. These results suggest that the ant-derived defensive compounds are likely more effective lone star tick repellents than DEET, but they are not as toxic as DEET towards the ticks. The suitability of these compounds for use as personal repellents, as well as at the landscape scale, should be explored.     

Pest Responses to Odors From Predators Fed a Diet of Target Species Conspecifics and Heterospecifics

ABSTRACT Research into the use of predator-odor-based repellents as a management tool has gained momentum during the past 30 years. Some studies have suggested that odors from a predator whose diet includes the target species are more effective than odors from a predator that does not consume the species. To evaluate this management tool in the Australian context and to determine the effect, if any, of predator diet on odor repellence, we tested eutherian and metatherian, predator fecal odors on phylogenetically separated pest species that currently occur in Australia. We evaluated fecal odors from tigers (Panthera tigris) and Tasmanian devils (Sarcophilus harrisii) as repellents for goats (Capra hircus) and eastern grey kangaroos (Macropus giganteus). We fed tigers and Tasmanian devils 2 diets: one of goat and one of eastern grey kangaroo. The test fecal odors were more effective than a control odor of carrier material and solvent at deterring goats (P < 0.001) and kangaroos (P = 0.02) from food. Tiger fecal odor was more effective than Tasmanian devil odor in deterring both goats (P = 0.001)and kangaroos (P = 0.03). We observed a decrease in the number of feeding events for goats when they were exposed to the odor from a tiger fed goat compared with all other predator-diet combinations (P < 0.001). We also observed a decrease in feeding events for kangaroos when exposed to the odor from a tiger fed kangaroo compared with all other predator-diet combinations. We observed signs of desensitization to the test odors in goats and habituation to the test odors in kangaroos over the experimental period. A better understanding of the factors involved in desensitization and habituation may increase the effectiveness of fecal odor-based repellents as a humane and nonlethal management tool for managers.     

Does concentrating chemical defenses within specific regions of marine sponges result in enhanced protection from predators?

Chemical defenses are an effective mode of predator deterrence across benthic marine organisms, but their production may come with associated costs to the organism as limited resources are diverted away from primary processes like growth and reproduction. Organisms concentrating ecologically relevant levels of these defenses in tissues most at risk to predator attack may alleviate this cost while deterring predators. We addressed this hypothesis by investigating the deterrence of chemical extracts from the inner and outer regions of the sponges Aplysina fulva, Ircinia felix, and I. campana from a temperate hard-bottom reef in the South Atlantic Bight. Assays were conducted using natural fish assemblages and sea urchins. Although, A. fulva and I. felix have higher concentrations of defensive metabolites in the outer and inner regions, respectively, extracts from these regions did not display enhanced deterrency against fish or mobile invertebrate predators. Likewise, extracts from both regions of the sponge Ircinia campana, which has a uniform distribution of defensive chemicals throughout, did not differ in their ability to deter either group of predators. Since chemical defenses were effective deterrents at lower concentrations, secondary metabolite allocation patterns observed among these sponges are likely not driven by predation pressure from generalist fish and mobile invertebrate predators on these reefs. Alternatively, these patterns may be driven by other ecological stressors, another suite of predators, or may be more effective at deterring predators when combined with structural defenses.     

Strategies of chemical anti-predator defences in leaf beetles: is sequestration of plant toxins less costly than de novo synthesis?

The evolution of defensive traits is driven both by benefits gained from protection against enemies and by costs of defence production. We tested the hypothesis that specialisation of herbivores on toxic host plants, accompanied by the ability to acquire plant defensive compounds for herbivore defence, is favoured by the lower costs of sequestration compared to de novo synthesis of defensive compounds. We measured physiological costs of chemical defence as a reduction in larval performance in response to repeated removal of secretions (simulating predator attack) and compared these costs between five species synthesising defences de novo and three species sequestering salicylic glucosides (SGs) from their host plants. Experiments simulating low predator pressure revealed no physiological costs in terms of survival, weight and duration of development in any of study species. However, simulation of high predation caused reduction in relative growth rate in Chrysomela lapponica larvae producing autogenous defences more frequently, than in larvae sequestering SGs. Still meta-analysis of combined data showed no overall difference in costs of autogenous and sequestered defences. However, larvae synthesising their defences de novo demonstrated secretion-conserving behaviour, produced smaller amounts of secretions, replenished them at considerably lower rates and employed other types of defences (regurgitation, evasion) more frequently when compared to sequestering larvae. These latter results provide indirect evidence for biosynthetic constraints for amounts of defensive secretions produced de novo, resulting in low defence effectiveness. Lifting these constraints by sequestration may have driven some leaf beetle lineages toward sequestration of plant allelochemicals as the main defensive strategy.     

Experimental use of two standard tick collection methods to evaluate the relative effectiveness of several plant-derived and synthetic repellents against Ixodes scapularis and Amblyomma americanum (Acari: Ixodidae)

We used two standard tick collection methods to test the relative effectiveness of two natural product compounds (nootkatone and carvacrol, classified as an eremophilene sesquiterpene and a monoterpene, respectively, that are derived from botanical sources) with commercially-available plant-derived (EcoSMART Organic Insect Repellent, comprised of plant essential oils) and permethrin-based (Repel Permanone) repellents against Ixodes scapularis Say and Amblyomma americanum (L.). Cloth drags were equally effective in sampling both species of host-seeking nymphs, whereas CO, traps attracted primarily A. americanum. All four repellents performed well on drags, with nootkatone and Permanone Repel (100% repelled through 14 d) slightly more effective than carvacrol and EcoSMART (90.7% and 97.7% repelled at 14 d, respectively) at repelling I. scapularis nymphs. Although the same trend in percent repellency was noted in the CO2 trap trial against both A. americanum nymphs and adults, EcoSMART outperformed Permanone in repelling A. Americanum nymphs after 14 d in the drag trial. Generally, the effectiveness of all repellents tested declined over time. The use of tick drags and CO2 traps was rapid, inexpensive, and easy to use in determining the relative effectiveness of repellents in the field.     

Effectiveness of odour repellents on red deer (<Emphasis Type="Italic">Cervus elaphus</Emphasis>) and roe deer (<Emphasis Type="Italic">Capreolus capreolus</Emphasis>): a field test

Chemical repellents are promoted as a method to reduce ungulate–vehicle collisions and ungulate browsing damages to agricultural and forestry resources. We tested the effectiveness of two odour repellents (Mota FL and Wolf Urine) on the foraging behaviour and area avoidance of free-ranging roe deer (Capreolus capreolus) and red deer (Cervus elaphus). The effects of the repellents were assessed by comparing deer visitation rates to sand arenas before and after application of repellents and visitation rates to control arenas. Neither of the tested products reduced deer visitation rates. Rapid habituation to olfactory stimuli and lack of sensitivity to predator odours may explain the ineffectiveness of the repellents to alter the behaviour of the deer. The results indicate that the tested products have no effects on roe deer and red deer behaviour and suggest that the effectiveness of the chemical area repellents as a measure to reduce deer–vehicle collision risk and browsing damages is questionable.     

Biochemical and evolutionary aspects of arthropod predation on ferns

Summary The widely held assumption that very few arthropods feed on ferns was questioned following field observations of arthropod damage on ferns in the state of Veracruz, Mexico. The extent and type of damage was recorded and it was found that in a measured locality, ferns were no less attacked than the angiospermous flora. As chemistry and arthropod host relationships have been shown to be so closely intertwined, plants collected in the field were analysed for both condensed tannins and cyanogenic glycosides, compounds known to be effective deterrents in temperate climates. Although all ferns tested contained tannins these did not appear to inhibit predation. Cyanogenic glycosides were present in only 3% of the fern species analysed, and it is, therefore, unlikely that they play a significant role as defensive compounds in the ferns examined.A literature search revealed a large number of ferns cited as being arthropod hosts. Approximately 420 named species of arthropods have been recorded, the majority of which are from the orders Coleoptera, Hymenoptera, Lepidoptera, and Hemiptera. Both evolutionary primitive (sawflies) and advanced (moths) arthropods are reported to be present on ferns suggesting possible coevolution of arthropods and ferns both before and after the radiation of angiosperms.     

Vertical T-maze Choice Assay for Arthropod Response to Odorants

Given the economic importance of insects and arachnids as pests of agricultural crops, urban environments or as vectors of plant and human diseases, various technologies are being developed as control tools. A subset of these tools focuses on modifying the behavior of arthropods by attraction or repulsion. Therefore, arthropods are often the focus of behavioral investigations. Various tools have been developed to measure arthropod behavior, including wind tunnels, flight mills, servospheres, and various types of olfactometers. The purpose of these tools is to measure insect or arachnid response to visual or more often olfactory cues. The vertical T-maze oflactometer described here measures choices performed by insects in response to attractants or repellents. It is a high throughput assay device that takes advantage of the positive phototaxis (attraction to light) and negative geotaxis (tendency to walk or fly upward) exhibited by many arthropods. The olfactometer consists of a 30 cm glass tube that is divided in half with a Teflon strip forming a T-maze. Each half serves as an arm of the olfactometer enabling the test subjects to make a choice between two potential odor fields in assays involving attractants. In assays involving repellents, lack of normal response to known attractants can also be measured as a third variable.     

The chemical defenses of millipedes (diplopoda): Biochemistry,physiology and ecology

Millipedes are some of the earliest examples of terrestrial animals, and fossils from the early Carboniferous Period indicate they were also some of the earliest prey. These fossils record ozopores, the openings of chemical defense glands, occurring along the length of the body. The secretions of these glands may consist of topical irritants, repellents, antifeedants, or, in the case of the large and widespread Order Polydesmida, hydrogen cyanide (HCN) gas that can be fatal to other arthropods or even small vertebrates in a confined environment. Müllerian mimicry rings may develop in which unrelated species of millipedes that co-occur closely resemble one another, while participating in a completely differently patterned ring in another part of their geographic range. Chemistry is not the only defense of millipedes. Polyxenids carry tufts of entangling setae, many species rely on crypsis and the ability to roll into a smooth, resistant sphere or coil, and still others have spikey projections that may deter soft-mouthed predators. Nevertheless, chemical defense has been of significance in helminthomorph millipedes at least since the Lower Carboniferous. The distribution of classes of chemical defense compounds follows major phylogenetic groupings. Evidence suggests that the defensive glands of Glomerida arose independently of those of the Helminthomorpha.     

Pea aphid dropping behavior diminishes foraging efficiency of a predatory ladybeetle

Antipredator defensive behaviors are a well‐studied and often crucial part of prey life histories, but little has been done to quantify how such behaviors affect natural enemies, their foraging, and their effectiveness as biological control agents. We explored how the generalist predatory coccinellid Harmonia axyridis Pallas (Coleoptera: Coccinellidae) affects the dropping behavior of the pea aphid, Acyrthosiphon pisum (Harris) (Homoptera: Aphididae), and in turn, how that defensive behavior affects the foraging efficiency of the predator. Experimental arenas that allowed or prevented pea aphid dropping were compared to determine how dropping influences the foraging of multiple life stages of H. axyridis: second instars, fourth instars, and adults. Dropping reduced predation on aphids by all ladybeetle life stages. Despite older predators inducing more dropping, aphid dropping reduced predation by approximately 40% across all ladybeetle life stages. Aphid dropping and predator consumption of aphids were both correlated with how much the predator moved, which also increased with predator life stage. We suggest that the high rates of dropping induced by H. axyridis and the subsequent decrease in H. axyridis foraging efficiency may partially explain why H. axyridis is less effective at controlling pea aphids than it is at controlling other aphid species that do not drop.     

Morphological and behavioral variation of <Emphasis Type="Italic">Rasbora paviei</Emphasis> (Cyprinidae,Cypriniformes) from lotic and limnic habitats (Central Vietnam)

Morphological and behavioral traits of Rasbora paviei from river and lake environments are studied. For investigations, the fish were caught from the middle reaches of the Cai River and from the inshore zone of the Kam Lam Reservoir (Khanh Hoa Province, Central Vietnam). Phenotypic differences are found in river and lake fish groups. Experimentally, differences in tactics and efficiency of defensive behavior are found in specimens from different habitats in the presence of a predator—Lepisosteus sp. (Lepisosteidae). In river specimens of Rasbora, the defensive reaction is more efficient in comparison with specimens from the reservoir. The river specimens of Rasbora used predominantly the tactics of group defense from the predator and stayed in a school longer than the lake specimens.     

Predator-mediated genotypic shifts in a prey population: experimental evidence

We demonstrate the effect of fish predation on genotype frequencies in a laboratory population composed of two Daphnia magna clones, with historically contrasting exposures to fish predation. The two clones differed in their responsiveness to predation via differential avoidance/escape behavior. The clone which coexists with fish in nature is more responsive to the presence of a fish predator, while the clone not exposed to fish predation does not exhibit the defensive reaction. Fish caused a rapid (within 18 h) and significant shift in Daphnia clonal composition, from 1:1 to 8:1, in favor of the responsive clone. Genotype-specific defensive abilities (modus defendi) can contribute greatly to the phenomenon of genotype replacement under selective predation.     

REDUCTION OF THE PECTORAL SPINE AND GIRDLE IN DOMESTICATED CHANNEL CATFISH IS LIKELY CAUSED BY CHANGES IN SELECTION PRESSURE

Locked pectoral spines of the Channel Catfish Ictalurus punctatus more than double the fish's width and complicate ingestion by gape‐limited predators. The spine mates with the pectoral girdle, a robust structure that anchors the spine. This study demonstrates that both spine and girdle exhibit negative allometric growth and that pectoral spines and girdles are lighter in domesticated than in wild Channel Catfish. This finding could be explained by changes in selection pressure for spine growth during domestication or by an epigenetic effect in which exposure to predators in wild fish stimulates pectoral growth. We tested the epigenetic hypothesis by exposing domesticated Channel Catfish fingerlings to Largemouth Bass Micropterus salmoides predators for 13 weeks. Spines and girdles grow isometrically in the fingerlings, and regression analysis indicates no difference in proportional pectoral growth between control and predator‐exposed fish. Therefore a change in selection pressure likely accounts for smaller pectoral growth in domesticated Channel Catfish. Decreasing spine growth in older fish suggests anti‐predator functions are most important in smaller fish. Additionally, growth of the appendicular and axial skeleton is controlled differentially, and mechanical properties of the spine and not just its length are an important component of this defensive adaptation.     

Guns and butter: a no cost defense against predation for Chrysomela confluens

Summary Chrysomela confluens produces a salicylaldehyde-based defensive secretion which is very effective against generalist predators and apparently produced at no cost. If no cost defenses are common, then one of the basic assumptions in the plant-herbivore literature, i.e. tradeoffs among defense, reproduction, and growth, must be reconsidered. We examined the effectiveness of this defense by exposing defended larvae and larvae whose secretion had been removed to a generalist predator. Larvae which had their secretions intact were attacked by only 7% of the ants which encountered them, and none of these larvae suffered serious damage. In contrast, those which had been milked of their secretions immediately prior to exposure were attacked in 48% of such encounters, and two-thirds of the larvae were killed. Larvae which had been milked 24 or 72 h before exposure, then allowed to regenerate their defenses, were attacked at rates indistinguishable from larvae that had not been milked. Thus regenerated defenses are just as effective as original defenses. We also tested the hypothesis that the cost of defense production and maintainence would be reflected in reductions in developmental rates and final adult mass and increases in leaf consumption rate. We found that larvae which were milked daily of their secretions manifested no measurable cost of recharging reservoirs. Milked larvae grew and fed at the same rates as their control sibs, and became adults of equal or slightly larger size. The liberation of glucose from salicin, a precursor present in leaves of salicaceous hosts, during the production of salicylaldehyde apparently provides enough of an energetic benefit to offset the cost of maintaining an effective defense. Consistent with this hypothesis, we did not find that milked larvae compensated for increased nutritional or salicin demands by increasing their feeding rates. Although this patterns is familiar to chemical ecologists it is generally unappreciated in the plant-herbivore literature. It is likely that many arthropod herbivore defensive systems come at little or no cost, given the intimacy of association between herbivores and their food plants. Sequestration of host plant defensive chemicals which eliminates the cost of synthesis is common in arthropods. The de novo synthesis of chemical defenses may be less costly than expected if it is integrated into other parts of an insects metabolism. Calculations based on the bond energies or molecular constitution of the compounds will not yield a complete perception of cost. Tests over the life of the herbivore, coupled with an understanding of the herbivore's metabolism, are necessary.     

Soldier bug predation on swallowtail caterpillars (Lepidoptera: Papilionidae): Circumvention of defensive chemistry

All larvae in the family Papilionidae possess osmeteria, eversible glands that release defensive chemicals upon disturbance. These chemicals have been shown to be repellent to ants, mantids, and other predaceous arthropods with chewing mouthparts. In this study, we demonstrate that the pentatomid Podisus maculiventris,an important predator of the black swallowtail Papilio polyxenes,is capable of piercing the body wall of the caterpillar and consuming body fluids without eliciting osmeterial eversion. Tenebrio molitorlarvae coated with osmeterial secretions of Papilio polyxeneswere universally rejected by Podisus maculiventris,suggesting that the predator is in fact repelled by the swallowtail defensive chemistry. By circumventing the defensive behavior of the caterpillar, the pentatomid is able to utilize an otherwise chemically unsuitable prey species.     

Evolution towards oscillation or stability in a predator–prey system

We studied a prey–predator system in which both species evolve. We discuss here the conditions that result in coevolution towards a stable equilibrium or towards oscillations. First, we show that a stable equilibrium or population oscillations with small amplitude is likely to occur if the prey''s (host''s) defence is effective when compared with the predator''s (parasite''s) attacking ability at equilibrium, whereas large-amplitude oscillations are likely if the predator''s (parasite''s) attacking ability exceeds the prey''s (host''s) defensive ability. Second, a stable equilibrium is more likely if the prey''s defensive trait evolves faster than the predator''s attack trait, whereas population oscillations are likely if the predator''s trait evolves faster than that of the prey. Third, when the adaptation rates of both species are similar, the amplitude of the fluctuations in their abundances is small when the adaptation rate is either very slow or very fast, but at an intermediate rate of adaptation the fluctuations have a large amplitude. We also show the case in which the prey''s abundance and trait fluctuate greatly, while those of the predator remain almost unchanged. Our results predict that populations and traits in host–parasite systems are more likely than those in prey–predator systems to show large-amplitude oscillations.     

Repulsion of bacteria from marine surfaces.

Organic compounds are capable of repelling motile bacteria from marine surfaces. The most effective compounds were acrylamide and benzoic and tannic acids. These were active at concentrations that were not toxic to the bacteria. Repellents were incorporated in nontoxic paints and applied to metal panels. Treated panels immersed in seawater developed a bacterial film of only 10(6) bacteria per cm6 after 12 days compared with untreated panels, which had 5 times 10(12) bacteria per cm2 after the same period. Field studies confirmed the effectiveness of these repellents. The use of biological repellents provides a new approach to the control of marine fouling.     

Host plant and the predator Podisus nigrispinus: when the defense compounds of the plant affect the third trophic level

Many herbivorous insects can overcome chemical plant defenses, using the plant's defensive products for their own good, as a defense against predators. Eucalyptus spp. (Myrtaceae), recently introduced in Brazil, are rich in secondary compounds; however, there are reports that these plants have been suffering from population outbreaks of defoliating Lepidoptera in Brazil. The predator Podisus nigrispinus (Dallas) (Hemiptera: Pentatomidae) has been used against herbivorous insects in eucalyptus plantations, but little is known about its establishment in the field. This study aims to investigate whether the effectiveness of this predator may be affected indirectly by compounds of eucalyptus plants, when compared to guava, Psidium guajava L., a Brazilian native species of Myrtaceae. Thus, we evaluated the performance of P. nigrispinus on larvae of Thyrinteina arnobia (Stoll) (Lepidoptera: Geometridae) reared on eucalyptus (exotic species) or guava plants (native species). Podisus nigrispinus performance (reproduction and survival) was better on larvae fed on guava than on larvae fed on eucalyptus. It is possible that the negative effect on the predator's development occurred because of the plants’ secondary compounds appropriated by caterpillars, due to the short coevolutionary history between eucalyptus and the predator. The data suggest that the chemical compounds that could help the plant's defenses against herbivores may also affect their natural enemies, especially when the interaction between plant and natural enemy involves an exotic plant recently introduced into the insect's habitat.     

Strike Fast,Strike Hard: The Red-Throated Caracara Exploits Absconding Behavior of Social Wasps during Nest Predation

Red-throated Caracaras Ibycter americanus (Falconidae) are specialist predators of social wasps in the Neotropics. It had been proposed that these caracaras possess chemical repellents that allow them to take the brood of wasp nests without being attacked by worker wasps. To determine how caracaras exploit nests of social wasps and whether chemical repellents facilitate predation, we: (1) video recorded the birds attacking wasp nests; (2) analyzed surface extracts of the birds'' faces, feet, and feathers for potential chemical repellents; and (3) inflicted mechanical damage on wasp nests to determine the defensive behavior of wasps in response to varying levels of disturbance. During caracara predation events, two species of large-bodied wasps mounted stinging attacks on caracaras, whereas three smaller-bodied wasp species did not. The “hit-and-run” predation tactic of caracaras when they attacked nests of large and aggressive wasps reduced the risk of getting stung. Our data reveal that the predation strategy of caracaras is based on mechanical disturbance of, and damage to, target wasp nests. Caracara attacks and severe experimental disturbance of nests invariably caused wasps to abscond (abandon their nests). Two compounds in caracara foot extracts [sulcatone and iridodial] elicited electrophysiological responses from wasp antennae, and were also present in defensive secretions of sympatric arboreal-nesting Azteca ants. These compounds appear not to be wasp repellents but to be acquired coincidentally by caracaras when they perch on trees inhabited with Azteca ants. We conclude that caracara predation success does not depend on wasp repellents but relies on the absconding response that is typical of swarm-founding polistine wasps. Our study highlights the potential importance of vertebrate predators in the ecology and evolution of social wasps.