Trail-following behavior in the German cockroach (Dictyoptera: Blattellidae)

The trail-following behavior of the German cockroach, Blattella germanica (L.), was evaluated by comparing the distance between cockroach movement paths and preapplied "trails" of fecal extract. For each cockroach group tested (adult males, females, gravid females, and late instars), the mean perpendicular distance of the cockroach from the trail was significantly less than the distance from a control trail. The results indicated that the German cockroaches did exhibit trail-following behavior. Trail- following accuracy varied among the cockroach groups. The mean distance from the fecal trail ranged from 18.45 to 110.05 cm with adult males < or = adult females < or = late instars < gravid females. Very dilute fecal extract could still induce trail-following behavior in adult male cockroaches. A 5.6% concentration of fecal extract in methanol was able to induce trail-following behavior in 50% of the cockroaches. Although German cockroaches have demonstrated trail-following behavior, fecal trails are still not thought to be actively deposited. Rather, the passive distribution of fecal material within the home range results in the accumulation of trails along frequently traveled routes (i.e., between resources and the cockroach harborage).     

Toxicity of fatty acid salts to German and American cockroaches

The toxicity of fatty acid salts to German, Blattella germanica (L.), and American cockroaches, Periplaneta americana (L.), was evaluated. Potassium and sodium laurate caused up to 95% mortality of German cockroaches and 100% mortality of American cockroaches. Even-numbered potassium fatty acid salts, C8-C18 were assessed for toxicity at 0.125, 0.25, 0.5, 1, and 2% concentrations by a 30-s immersion of cockroaches. The more soluble of the fatty acid salts at 2% concentration caused 65-95% mortality of German cockroaches and 100% mortality of American cockroaches. Potassium oleate, C18, was most toxic to both German (LC50 = 0.36%) and American (LC50 = 0.17%) cockroaches. Fatty acid salt solutions on a substrate were tested by placing cockroaches in contact with treated floor tiles immediately after application (wet) or after the solutions had dried. Sodium laurate and potassium caprate caused mortality of German (62 +/- 17.4 and 58 +/- 12.6%, respectively) and American cockroaches (52 +/- 18.5 and 28 +/- 4.9%, respectively) on wet tiles, whereas potassium oleate caused mortality of German cockroaches (67 +/- 14.1%) only. Dry fatty acids caused no mortality among exposed cockroaches. Fatty acid salt solutions can be effective in killing German and American cockroaches but only when insects are thoroughly wetted with 1-2% fatty acid salt solutions.     

Evaluation of attractants for monitoring populations of the German cockroach (Dictyoptera: Blattellidae)

Lures that are used to attract German cockroaches, Blattella germanica (L.), to traps were compared in olfactometer assays in the laboratory and in trapping experiments in cockroach-infested homes and a swine farm. In olfactometer assays, AgriSense GP-2 was the most attractive lure, followed by peanut butter, and distiller's grain. Other lures, including Trapper tablet; Victor pheromone, a crude fecal extract that ostensibly contains B. germanica aggregation pheromone; and Victor food lure elicited upwind orientation from <50% of the test insects. Peanut butter and distiller's grain were equally attractive in trapping experiments in swine production barns and they captured significantly more cockroaches than the GP-2 tablet or the Victor pheromone lure; the commercial lures failed to attract significantly more cockroaches than the unbaited control traps. When tested against blank controls, cockroaches preferred to rest in shelters that contained the aggregation pheromone-based lure (Victor), but this lure was the least attractive to cockroaches in olfactometer assays. These results do not support claims that commercial crude fecal extracts attract cockroaches to traps, and they highlight a need for developing more attractive lures for detection of cockroaches and for monitoring populations.     

Influence of starvation and lighting on the movement behavior of the German cockroach (Blattodea: Blattellidae).

Computerized moving-image analysis measured the influence of starvation and lighting conditions on the locomotor activity of the German cockroach, Blattella germanica (L.). Starvation increased the distance traveled, velocity, and the proportion of time in motion of adult males and last instars but did not increase adult female movement. For females, males, and nymphs, starvation increased cockroach residence time around the source of water and harborage and decreased the distance traveled by females and nymphs around the observation arena's edge. Fluorescent light reduced the proportion of time in motion for males, but the velocity of males was greater in fluorescent light relative to infrared light. Adult males and females stopped more frequently in infrared light. Overall, males and nymphs were more mobile than females and they explored larger areas. The moving-image analysis technique is an efficient and accurate tool for observing movement of individual cockroaches and has applications in repellency, attractancy, and basic movement behavior studies.     

Use of fecal extract trails to enhance trap catch in German cockroach (Dictyoptera: Blattellidae) monitoring stations

An aqueous extract of German cockroach, Blattella germanica (L.), fecal material was evaluated for inducing trail-following behavior in German cockroaches. In arena tests the fecal extract was found to stimulate trail following in 74% of adult male cockroaches. Significantly fewer cockroaches (22%) followed water-treated (control) trails. Residual activity of the fecal extract trails was evaluated by bioassay after the trails had been stored in the refrigerator or in the open air. Although trails stored in the refrigerator showed no decline in activity after 14 d, those stored in the open air declined significantly after 3 d, inducing only 40% of adult male cockroaches to follow the trail. After 7 d the activity of trails stored in the open air was further reduced to 23%. The ability of fecal-extract trails to influence trap catch in monitoring stations was determined by bioassay. Paper trails treated with fecal extract or water were positioned between cockroach harborages and monitoring stations inside 122-cm2 arenas. The presence of the fecal extract-treated trails significantly enhanced trap catch. Mean catch in the traps with fecal extract trails was 28 cockroaches compared with a mean of 11 cockroaches in the control traps. The trap catch ratios of adults to nymphs in the treated and control treatments were not significantly different.     

Susceptibility of cockroaches (Dictyoptera: Blattellidae, Blattidae) to infection by Steinernema carpocapsae.

The susceptibility of American cockroaches, Periplaneta americana (L.); smoky brown cockroaches, P. fuliginosa (Serville); oriental cockroaches, Blatta orientalis L.; German cockroaches, Blattella germanica (L.); and brownbanded cockroaches, Supella longipalpa (F.), to Steinernema carpocapsae Weiser (All strain) was evaluated under laboratory conditions. A 1-ml water suspension containing 500,000 nematodes was placed on filter paper in a petri dish or the pad of a bait station. German, brown-banded, oriental, and smoky-brown cockroaches died within 1 d after placement in the petri dishes. The relative order for the LT50s were American greater than oriental greater than smoky-brown greater than brown-banded = German. All cockroaches actively groomed nematodes from legs and antennae of forced (petri dish) exposure. The LT50s for S. carpocapsae for nonforced (bait station) exposure were significantly greater than those for forced exposure. The LT50s were 3.25, 4.13, 9.86, and 11.38 d for brown-banded, German, oriental, and smoky-brown cockroaches, respectively. The relative order of the LT50s after forced (American greater than oriental greater than smoky-brown greater than German = brown-banded) and nonforced (American greater than smoky-brown greater than oriental greater than German greater than brown-banded) exposure to S. carpocapsae was inversely related to the moisture of their preferred habitats.     

Boric acid dust as a component of an integrated cockroach management program in confined swine production

Boric acid dust treatments were evaluated as a tool for the integrated management of the German cockroach, Blattella germanica (L.), in commercial confined swine production. The efficacy of boric acid dust was comparable to that of an organic residual insecticide, cyfluthrin, which is commonly used to control cockroaches in this environment. Fall treatments suppressed the cockroach population for longer durations than treatments in the Spring. Boric acid dust is an effective, inexpensive, and low risk (to animal and human health, and the environment) alternative for the management of cockroaches in livestock production systems.     

Cytochrome P450 CYP6L1 is specifically expressed in the reproductive tissues of adult male German cockroaches, Blattella germanica (L.)

A full-length cDNA encoding a new cytochrome P450, CYP6L1, was cloned from German cockroaches, Blattella germanica. CYP6L1 has an open reading frame of 1509 nucleotides with a deduced protein of 503 amino acids and molecular mass of 57 Kd. CYP6L1 is most similar to CYP6H1, a putative ecdysone 20-hydroxylase from Locusta migratoria. CYP6L1 mRNA was not detected in embryos nor nymphs, nor in adult females. CYP6L1 mRNA was detected only in the testes and accessory glands of male adult German cockroaches. Given that the testes and accessory glands are the most important components of the reproductive system in male insects, the expression of CYP6L1 mRNA exclusively in these tissues strongly suggests that CYP6L1 has a role in reproduction. Possible substrates for CYP6L1 are discussed.     

A model of animal movements in a bounded space

Most studies describing animal movements have been developed in the framework of population dispersion or population dynamics, and have mainly focused on movements in open spaces. During their trips, however, animals are likely to encounter physical heterogeneities that guide their movements and, as a result, influence their spatial distribution. In this paper, we develop a statistical model of individual movement in a bounded space. We introduced cockroaches in a circular arena and quantified accurately the behaviors underlying their movement in a finite space. Close to the edges, we considered that the animals exhibit a linear movement mode with a constant probability per unit time to leave the edge and enter the central zone of the arena. Far from the walls cockroaches were assumed to move according to a diffusive random walk which enabled us to overcome the inherent problem of the quantification of the turning angle distribution. A numerical model implementing the behavioral rules derived from our experiments, confirms that the pattern of the spatial distribution of animals observed can be reliably accounted for by wall-following behaviors combined with a diffusive random walk. The approach developed in this study can be applied to model the movements of animals in various environment under consideration of spatial structure.     

Estimating nutritional status of German cockroaches,Blattella germanica (L.) (Dictyoptera: Blattellidae), in the field

Nutritional status of German cockroaches from the field (HUD apartments) was estimated using uric acid content to measure amount of protein consumed, and respiratory quotient (RQ) to measure fat and carbohydrate metabolized. Initial trials demonstrated the stability of these two indicators as nymphal cockroaches grow and with timing of meals. Nutrient consumption (and presumed availability) was estimated by comparing uric acid content and RQ of nymphal cockroaches collected from kitchens of HUD apartments with those reared in the laboratory and provided a series of meridic diets. Uric acid content was linearly related to percentage of dietary protein (y=6.2x-32.07, r(2)=0.96) and RQ was linearly related to log(10)(% fat:% carbohydrate) (y=-0.148Log(x)+0.790, r(2)=0.68). Field-collected German cockroaches contained 10.9+/-7.7 to 22.9+/-5.1 &mgr;g/mg uric acid and RQ of 0.770+/-0.024 to 0.803+/-0.260. Comparatively, cockroaches provided rodent chow had greater uric acid content (125.1+/-9.6 &mgr;g/mg) and RQ (0.878+/-0.022). Employing linear calibration and these regressions, diet consumed by German cockroaches in the field was estimated at 7+/-3% to 9+/-3% protein and equivalent amounts of carbohydrates and fat as an energy source. German cockroaches in the field consume less protein and carbohydrates, and more fat compared to those provided a standard laboratory diet such as rodent chow. Diet available in the field is considered suboptimal, resulting in physiological stress; the biological implications of this stress are discussed.     

Relationship between the para-homologous sodium channel point mutation (g --> c at nucleotide 2979) and knockdown resistance in the German cockroach using multiplex polymerase chain reaction to discern genotype

Extensive use of pyrethroid insecticides for urban pest control has led to widespread pyrethroid resistance in the German cockroach. A mutation at nucleotide position 2979 (G to C, causing a leucine to phenylalanine change) in the S6 transmembrane segment of domain II of the para-homologous voltage-gated sodium channel has been previously identified in knockdown-resistant cockroaches and demonstrated by site-directed mutagenesis to reduce channel sensitivity to pyrethroids. In a recent survey, 83% of pyrethroid-resistant German cockroach populations were found to possess this mutation. A German cockroach strain with a low incidence of the L993F mutation was subjected to selection pressure with cypermethrin and subsequently evaluated over several generations for the knockdown resistance phenotype. Correspondingly, we determined the genotype of individual cockroaches of each population at the 2979 position of the para-homologous gene. Genotype was discerned by development of a polymerase chain reaction method that employed a mismatched primer-template set. A direct relationship was observed between mean knockdown time and the presence of the kdr mutation. Furthermore, individuals homozygous for the kdr mutation exhibited a significantly higher mean knockdown time than heterozygotes or wildtype cockroaches. This is the first report demonstrating the progressive expression of the kdr allele in response to insecticide selection pressure.     

Detection of insecticide resistance in the German cockroach (Dictyoptera: Blattellidae) with glue-toxin traps.

Glue that contained an insecticide was evaluated for its ability to yield useful toxicological data for German cockroaches, Blattella germanica (L.). Toxicities of three classes of insecticides were evaluated by topical applications (LD50) and exposure to insecticide-impregnated glue (LC50). Cockroaches that were resistant to topical insecticide applications were also resistant to the glue formulation. Reliability was greatest when mortality was scored 40 to 48 h after the cockroaches were placed on the glue. This method should be adaptable for insecticide resistance monitoring of German cockroaches.     

五种蜚蠊的生物学特性和综合治理

介绍了分布广泛和危害严重的室内 5种常见蜚蠊 ,德国小蠊、美洲大蠊、黑胸大蠊、澳洲大蠊和褐斑大蠊的生物学特性和行为习性以及德国小蠊为何能成为室内优势种的原因。蜚蠊的综合治理系统包括 :检查和调查、环境治理、非化学防治、杀虫剂防治、昆虫信息化合物和昆虫生长调节剂的防治和其它防治方法。对残留和非残留杀虫剂和剂型的基本概念和应用做了解释 ,并分别介绍了对每种蜚蠊的具体治理措施。     

Overexpression of the cytochrome c oxidase subunit I gene associated with a pyrethroid resistant strain of German cockroaches, Blattella germanica (L.)

A cytochrome c oxidase subunit I gene (COXI) was identified and isolated as a differentially expressed gene between insecticide susceptible ACY and resistant Apyr-R German cockroach strains using PCR-selected subtractive hybridization and cDNA array techniques. The cDNA sequence of COXI has an open reading frame of 1533 nucleotides encoding a putative protein of 511 amino acid residues. Northern blot analysis indicated that levels of COXI expression were similar in three life stages (eggs, nymphs, and adults) of the susceptible ACY strain. The expression of COXI in the resistant Apyr-R strain was developmentally regulated, with low expression in eggs, an increase (approximately 1.4-fold) in nymphs, and rose to a maximum (approximately 3-fold) in both adult females and males. Comparison of COXI expression between ACY and Apyr-R strains indicated that there was no difference in the eggs of the two strains, but expression was higher (approximately 1.5-fold) in nymphs and much higher (approximately 3- to 4-fold) in adult males and females of the Apyr-R strain. The levels of COXI mRNA showed about 1.4- and 1.7-fold increase in the abdomen tissues compared with the head+thorax tissues of ACY and Apyr-R strains, respectively. Although expression patterns of COXI in head+thorax and abdomen tissues were similar (i.e. lower in the head+thorax tissues and higher in the abdomen tissues) in both the ACY and Apyr-R strains, the expression of COXI was about 2.5-fold higher in the head+thorax and approximately 3-fold higher in the abdomen tissues of the Apyr-R strain compared with the corresponding ACY samples. The overexpression of COXI in resistant German cockroaches merits the investigation of the importance of the gene in insecticide resistant German cockroaches.     

Nutritional status and insecticide tolerance in German cockroaches (Orthoptera: Blattellidae).

The interrelationship of two extrinsic factors-nutrition and pesticide exposure-was studied in German cockroaches, Blattella germanica (L.). Amount of dietary protein had no effect on susceptibility of male German cockroaches to propoxur and chlorpyrifos; however, the source of protein produced significant changes in insecticide susceptibility. Increased total body lipids, carbohydrates, and uric acid may have resulted in increased tolerance to these insecticides. Three days of food or water deprivation (or both) caused significant differences in susceptibility to chlorpyrifos and propoxur.     

Adaptive contraction of diet breadth affects sexual maturation and specific nutrient consumption in an extreme generalist omnivore

Animals balance their intake of specific nutrients, but little is known about how they do so when foraging in an environment with toxic resources and whether toxic foods promote adaptations that affect life history traits. In German cockroach (Blattella germanica) populations, glucose aversion has evolved in response to glucose‐containing insecticidal baits. We restricted newly eclosed glucose‐averse (GA) and wild‐type (WT) female cockroaches to nutritionally defined diets varying in protein‐to‐carbohydrate (P : C) ratio (3 : 1, 1 : 1, or 1 : 3) or gave them free choice of the 3 : 1 and 1 : 3 diets, with either glucose or fructose as the sole carbohydrate source. We measured consumption of each diet over 6 days and then dissected the females to measure the length of basal oocytes in their ovaries. Our results showed significantly lower consumption by GA compared to WT cockroaches when restricted to glucose‐containing diets, but also lower fructose intake by GA compared to WT cockroaches when restricted to high fructose diets or given choice of fructose‐containing diets. Protein intake was regulated tightly regardless of carbohydrate intake, except by GA cockroaches restricted to glucose‐containing diets. Oocyte growth was completely suppressed in GA females restricted to glucose‐containing diets, but also significantly slower in GA than in WT females restricted to fructose‐containing diets. Our findings suggest that GA cockroaches have adapted to reduced diet breadth through endocrine adjustments which reduce requirements for energetic fuels. Our study illustrates how an evolutionary change in the chemosensory system may affect the evolution of other traits that govern animal life histories.     

Body Size and Termite Evolution

Termites are a monophyletic lineage within the paraphyletic Blattaria, with xylophagous cockroaches in the genus Cryptocercus as sister group. Given this ancestry, termite divergence involved a substantial leap in body plans, as they are pale, fragile, and miniaturized relative to most cockroaches. Here I suggest that the evolutionary transition to an altricial morphotype in termites is grounded in the economics of utilizing a wood diet, and occurred via a series of sequential steps associated with modifications in social behavior. The chief benefit of an altricial morphotype is suggested to be nitrogen made available by decreasing individual body size and by dispensing with a heavy, melanized cuticle. The primary costs lie in increased vulnerability to environment hazards, including predators, pathogens, and desiccation. Termites tipped the evolutionary scale via cooperative behaviors that mitigate the cost or amplify the benefit of a small, fragile morphotype, and were present in rudimentary form in their cockroach relatives. These include building behavior, cooperative feeding, allogrooming, and most notably, trophallaxis. It was the directed circulation of nitrogenous reserves via trophallactic feeding among units of the superorganism rather than their progressive storage in a large, heavily armored body that was the foundation of termite evolutionary success.     

The role of antennal hair plates in object-guided tactile orientation of the cockroach (Periplaneta americana)

The searching behavior of blinded cockroaches was examined under unrestrained conditions, in an arena, and on a treadmill. When cockroaches searching in a circular arena touched a stationary object (metal pole) with their antennae, they frequently approached the object more closely, and then climbed up it. Similar orientation behavior was observed in tethered animals in open loop conditions, walking on a Styrofoam ball. In these restrained cockroaches, a single antenna sufficed to distinguish the angular positions of an object, in the horizontal plane (0 degrees, 45 degrees, and 90 degrees). A group of mechanosensitive hairs on the basal segment of the antenna (scapal hair plate) appears to play a major role in antennal object detection in the horizontal plane, as gauged by shaving off these scapal hair plates. In unrestrained cockroaches, shaving the scapal hair plate increased the time needed to approach an object. Under tethered conditions, the ability to turn towards and to establish antennal contact with a test object was significantly impaired.     

Method of insecticide delivery affects horizontal transfer of fipronil in the German cockroach (Dictyoptera: Blattellidae).

Horizontal transmission of insecticide occurs when foragers contact or ingest an insecticide, return to the aggregation or nest, and translocate the insecticide to the shelter and its vicinity. Relatively more sedentary members of the population then contact or eat the translocated insecticide and die. We evaluated three different methods of delivering fipronil to adult male German cockroaches, Blattella germanica (L.), for their potential to cause such secondary mortality in various developmental stages of the cockroach. Adult males topically treated with 5 ng of fipronil (approximately LD99) caused low mortality in untreated nymphs and no mortality in untreated adults within the same aggregation. Males exposed to residual fipronil on a glass surface translocated more insecticide, resulting in higher mortality of cockroaches they contacted, but only early instars were affected and no adult mortality was observed. Ingested fipronil bait, however, was most effectively translocated, and caused high mortality of untreated adults and nymphs. Ingestion of fipronil also caused greater secondary kill compared with a topical application of 25 ng, approximately the same amount recovered from the exterior of males that ingested 1 mg of 0.05% fipronil bait. Secondary mortality in the untreated population was significantly affected by the duration of contact between the treated and untreated cockroaches, the quantity and freshness of excretions from the treated insects, and the accessibility of the secretions to untreated cockroaches. The mechanisms that cause secondary kill may include ingestion of excreted fipronil residues, cannibalism of bait-fed cockroaches, as well as contact with fipronil-contaminated substrates.     

Inconsistency in the expression of locomotor and ERG circadian rhythms in the German cockroach, Blattella germanica (L.).

ERG recordings from German cockroaches showed that the amplitude of light-evoked responses have a circadian rhythmicity in adult males that coincided with the locomotor circadian rhythm. The peak of the response occurred during the subjective night, and the circadian period was less than 24 h under DD condition. In contrast, although the locomotor circadian rhythm was masked by the development of ovaries and pregnancy in females, their visual responses displayed circadian rhythmicity. This inconsistency in expression of locomotor and visual sensitivity circadian rhythms in females implied separate pacemakers for these two overt rhythms. After severing the optic nerves, changes in ERG amplitude of the operated cockroaches still displayed a circadian rhythm under DD condition, demonstrating that the visual sensitive pacemaker was located in the eye and independent from the locomotor pacemaker.