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Biology and management of the smokybrown cockroach.

The smokybrown cockroach, Periplaneta fuliginosa (Serville), has become an increasingly important peridomestic pest throughout much of the southeastern United States, Japan, and southeast Asia. We provide a brief description of the species and clarify its worldwide distribution. Aspects of smokybrown cockroach behavior, biology, ecology, and physiology are discussed in the context of this species' successful exploitation of the urban environment. We describe smokybrown cockroach management based upon a model that relates house and landscape characteristics to cockroach population size. Finally, we illustrate the effectiveness of a model-based IPM system that increases control while reducing use of insecticides.

Animals↗

Pharmacology of skeletal muscle GABA-gated chloride channels in the cockroach Periplaneta americana.

The pharmacology of -aminobutyric acid (GABA)-gated chloride channels of the coxal levator (182c,d) muscle of the cockroach Periplaneta americana has been investigated and the data compared with similar findings for the cell body of the cockroach fast coxal depressor motor neurone (Df). Muscle GABA receptors resembled those of the motor neurone cell body in their sensitivity to picrotoxinin and insensitivity to bicuculline. However, muscle GABA receptors were insensitive to the neuronal GABA receptor agonists isoguvacine (10(-4) mol l-1) and 3-aminopropane sulphonic acid (10(-3 )mol l-1). The benzodiazepine flunitrazepam, which at 10(-6 )mol l-1 greatly enhances the amplitude of the motor neurone GABA-induced responses, failed to affect muscle responses to GABA when tested at the same and at a higher (10(-4 )mol l-1) concentration. The convulsant t-butylbicyclophosphorothionate was a weak antagonist of cockroach muscle GABA receptors, whereas several cyclodienes were much more effective antagonists. Thus, studies using a benzodiazepine and several convulsant antagonists reveal differences in the pharmacology of muscle and neuronal GABA receptors of the cockroach Periplaneta americana.

Animals↗

Attractiveness of insecticide baits for cockroach control (Dictyoptera: Blattellidae): laboratory and field studies.

Several insecticide bait formulations were evaluated for their attractiveness to cockroaches in olfactometer assays in the laboratory and in trapping experiments in the field. Included in the assays were bait stations, gels, pastes, and a powder that contained one of the following active ingredients: abamectin, boric acid, chlorpyrifos, or hydramethylnon. There were significant differences among the baits in their attractiveness to the German cockroach, Blattella germanica (L.). In trapping experiments, Avert powder (abamectin), Maxforce station and gel, and Siege gel (all hydramethylnon) were consistently attractive to B. germanica adults and nymphs. Laboratory olfactometer assays with adult males confirmed these results and showed that nymphs were as responsive as males whereas females were less responsive. Our bioassays also demonstrate that attractiveness of bait can be dramatically affected by the age of the bait. One week of aging significantly reduced the attractiveness of Avert powder in both laboratory and field assays. Aging, however did not diminish the attractiveness of Maxforce gel, indicating that the formulation may be critical for retention of attractiveness of baits. Baits that were most attractive to the German cockroach were also the most attractive to nymphs and adults of the brownbanded cockroach, Supella longipalpa (F.).

Animals↗

The effect of parasitism on host fecundity is dependent on temperature in a cockroach-acanthocephalan system.

The fitness of infected organisms can vary greatly depending on the temperature at which they find themselves. Understanding the role of temperature in the fitness of infected organisms can be crucial to population studies, epidemiological studies, and when screening for biological control agents. We measured the effect of parasitism on host survival and reproduction at 4 constant temperatures using the acanthocephalan parasite Moniliformis moniliformis and its intermediate host, the cockroach Supella longipalpa. Infection did not affect cockroach survival at any temperature. Infection had a negative impact on cockroach fecundity but only at higher temperatures (28 and 31 C) and only later in infection (>20 days postinfection). At lower temperatures, infected and uninfected cockroaches had similar fecundities throughout the duration of the experiment (120 days). This study, along with previous studies, suggests that researchers would do well to consider environmental variables when exploring the effects of parasitism.

Analysis of Variance↗

Allatostatin-like-immunoreactive neurons of the tobacco hornworm, Manduca sexta, and isolation and identification of a new neuropeptide related to cockroach allatostatins.

The YXFGLamide C-terminus serves to define most members of a family of structurally related neuropeptides, the YXFGLamides. These peptides have been identified from the nervous system of various insects and include the allatostatins of cockroaches and crickets, the schistostatins of locusts, and the callatostatins of blowflies. The YXFGLamides have been shown to have various functions, including inhibition of juvenile hormone biosynthesis in cockroaches and crickets and inhibition of contraction of certain insect visceral muscles. We wanted to know if these peptides occur in Manduca sexta and what functions they might have. A new peptide, AKSYNFGLamide, was isolated and identified from M. sexta and has been named "lepidostatin-1"; this is the first YXFGLamide to be found in a lepidopteran, and there are indications that additional YXFGLamides occur in M. sexta. An antiserum to cockroach allatostatins (YXFGLamides) was shown to recognize lepidostatin-1 of M. sexta and was used to map YXFGLamide-immunoreactive neurons in larvae. Because immunoreactive interneurons were found to form an extensive neuropil, YXFGLamides probably function as neuromodulators in M. sexta. Neuroendocrine cells in the brain, abdominal ganglia, and their respective neurohemal organs were YXFGLamide immunoreactive and appear to release YXFGLamides as neurohormones. Immunoreactivity to YXFGLamides and M. sexta diuretic hormone were found to be colocalized and appear to be coreleased in these neuroendocrine cells, indicating that YXFGLamides may be involved in regulation of fluid transport. Innervation of the corpora allata by YXFGLamide-immunoreactive processes was very sparse, suggesting that this innervation does not play an important role in allatostasis. Many thoracic motor neurons were YXFGLamide immunoreactive, suggesting that YXFGLamides may have a myomodulatory or myotrophic function in larvae. However, this immunoreactivity disappeared early in metamorphosis and did not reappear in the adult. The YXFGLamide-immunoreactive neurons in the terminal abdominal ganglion were found to innervate the hindgut, indicating that YXFGLamides may be involved in the control of the rate of myogenic contractions of the larval hindgut.

Animals↗

Neural organization of ocellar pathways in the cockroach brain.

A large number of photoreceptors of insect ocelli converge onto a smaller number of second-order neurons. Second-order neurons exit the ocellus and project into the ocellar tract neuropil of the brain. Here, the anatomy and physiology of ocellar interneurons of the ocellar tract neuropil of the cockroach are described. The total number and gross morphologies of ocellar tract neurons were examined by extracellular cobalt impregnations into the neuropil. Morphology and physiology of individual neurons were examined using intracellular recording and stainings. Each ocellar tract neuropil contains at least 25 interneurons comprising: 1) four second-order neurons, 2) 15 third-order neurons that receive synapses from second-order neurons at the ocellar tract and project into a number of target neuropil areas of the brain, 3) two possible efferent neurons, 4) three third-order or efferent neurons, and 5) one neuron still to be characterized. The projection areas of ocellar third-order neurons include 1) visual, olfactory, and mechanosensory centers; 2) the mushroom body (a higher associative center); 3) the posterior slope, a premotor center from which descending brain neurons originate; and 4) the thoracic motor systems. By comparing the present results to those reported from other insects, I conclude that the cockroach ocellar system has two distinctive features. 1) The ratio of convergence at synapses between photoreceptors and second-order neurons is higher than those reported for other insects so far studied. 2) Ocellar signals are transmitted to various target neuropils by third-order neurons, whereas ocellar systems of all other insects possess pathways in which ocellar signals are transmitted to target neuropils by second-order neurons. The functional significance of these features of the cockroach ocellar system is discussed.

Animals↗

Descending influences on escape behavior and motor pattern in the cockroach.

The escape behavior of the cockroach is a ballistic behavior with well characterized kinematics. The circuitry known to control the behavior lies in the thoracic ganglia, abdominal ganglia, and abdominal nerve cord. Some evidence suggests inputs may occur from the brain or suboesophageal ganglion. We tested this notion by decapitating cockroaches, removing all descending inputs, and evoking escape responses. The decapitated cockroaches exhibited directionally appropriate escape turns. However, there was a front-to-back gradient of change: the front legs moved little if at all, the middle legs moved in the proper direction but with reduced excursion, and the rear legs moved normally. The same pattern was seen when only inputs from the brain were removed, the suboesophageal ganglion remaining intact and connected to the thoracic ganglia. Electromyogram (EMG) analysis showed that the loss of or reduction in excursion was accompanied by a loss of or reduction in fast motor neuron activity. The loss of fast motor neuron activity was also observed in a reduced preparation in which descending neural signals were reversibly blocked via an isotonic sucrose solution superfusing the neck connectives, indicating that the changes seen were not due to trauma. Our data demonstrate that while the thoracic circuitry is sufficient to produce directional escape, lesion or blockage of the connective affects the excitability of components of the escape circuitry. Because of the rapidity of the escape response, such effects are likely due to the elimination of tonic descending inputs.

Animals↗

The shadow response of the cockroach periplaneta americana.

We show that the "escape response" (running) of the cockroach Periplaneta americana is interrupted when the insect runs through a shadow. Shadows were cast by holding an opaque plastic sheet so that the cockroach ran beneath it. In control experiments, a similarly positioned clear plastic sheet was ineffective. Removal of the shadow stimulus allowed running to continue. The likelihood that cockroaches would resume running after removal of the shadow stimulus decreased with the time they were allowed to remain in shadow.

Adaptation, Biological↗

A model of pattern generation of cockroach walking reconsidered.

Cockroaches that have been decapitated or that have cut thoracic connectives can show rhythmic bursting in motoneurons to intrinsic leg muscles. These preparations have been studied as models for walking and to evaluate the functions of leg proprioceptors. The present study demonstrates that headless cockroaches walk extremely poorly and slowly with considerable discoordination of motoneuronal activity, these preparations show rhythmic motoneuron bursting that is similar to righting responses (attempts to turn upright) of intact animals when placed on their backs, and bursting is inhibited when a headless animal is turned or turns itself upright. Thus, rhythmic motoneuron activity of these preparations is most probably attempted righting rather than walking. It is concluded that the headless cockroach is useful for understanding the motor mechanisms underlying righting and walking but is not of value in assessing the functions of proprioceptive feedback.

Animals↗

Correlation of a resistance-associated Rdl mutation in the German cockroach, Blattella germanica (L), with persistent dieldrin resistance in two Danish field populations.

Danish Blattella germanica (L) populations carry the resistance-associated mutation A302S within the Resistance to dieldrin (Rdl) gene. The mutation has remained in field populations long after the discontinuation of dieldrin for cockroach control. The mutation has also persisted in our laboratory strains with high and intermediate frequencies for more than 8 years without selection. The toxicity of dieldrin was tested by topical application to male cockroaches in the susceptible strain DPIL-SUS and two field strains, Zo960302 and Su960304, which were 1270- and 15-fold resistant to dieldrin at LD50. We report the sequencing of exon 7 of the B. germanica Rdl gene and the finding of the putative resistance-associated A302S mutation. We have developed and implemented a PCR-based diagnostic method with the detection of a restriction endonuclease polymorphism, which allows for easy discrimination of susceptible, resistant and heterozygote genotypes. The frequency of the resistance-associate allele A302S was 0.97 and 0.38 in the Zo960302 and Su960304 populations, respectively. The cockroach Rdl A302S allele confers high dieldrin resistance in homozygotes and intermediate resistance in heterozygotes, and its presence is responsible for the persisting dieldrin resistance in Danish populations of B. germanica.

Amino Acid Sequence↗

Interaction of dinotefuran and its analogues with nicotinic acetylcholine receptors of cockroach nerve cords.

To investigate the action of dinotefuran (MTI-446, 1-methyl-2-nitro-3-(tetrahydro-3-furylmethyl)guanidine), a recently developed insecticide, on insect nicotinic acetylcholine receptors (nAChRs), we determined the potencies of the compound and 15 analogues in inhibiting the specific binding of [3H]epibatidine (EPI), a nAChR agonist, and [3H]alpha-bungarotoxin (alpha-BGT), a competitive nAChR antagonist, to the nerve cord membranes of American cockroaches (Periplaneta americana). Racemic dinotefuran inhibited [3H]EPI binding with an IC50 of 890 nM and [3H]alpha-BGT binding with an IC50 of 36.1 microM. Scatchard analysis indicated that the dinotefuran inhibition of [3H]EPI binding was a competitive one. Slight structural modification caused a drastic reduction in potency; only four analogues were found to be equipotent to or more potent than dinotefuran. Chloropyridinyl and chlorothiazolyl neonicotinoid insecticides displayed two or three orders of magnitude higher potency than dinotefuran. There was a good correlation between the IC50 values of tested compounds obtained with [3H]EPI and those obtained with [3H]alpha-BGT. A better correlation was observed between 3-h knockdown activities (KD50) against German cockroaches (Blattella germanica) and IC50 values obtained from [3H]EPI assays than between 24-h lethal activities (LD50) and IC50 values. While the results indicate that dinotefuran and its analogues interact with the ACh-binding site in cockroach nAChRs, it remains to be elucidated why they displayed lower potencies than those expected based on their insecticidal activities.

Animals↗

Multiple feedback loops in the flying cockroach: excitation of the dorsal and inhibition of the ventral giant interneurons.

1. In a tethered cockroach (Periplaneta americana) whose wings have been cut back to stumps, it is possible to elicit brief sequences of flight-like activity by puffing wind on the animal's body. 2. During such brief sequences, rhythmic bursts of action potentials coming from the thorax at the wingbeat frequency, descend the abdominal nerve cord to the last abdominal ganglion (A6). This descending rhythm is often accompanied by an ascending rhythm (Fig. 2). 3. Intracellular recording during flight-like activity from identified ascending giant interneurons, and from some unidentified descending axons in the abdominal nerve cord, shows that: (a) ventral giant interneurons (vGIs) remain silent (Fig. 3); (b) dorsal giant interneurons (dGIs) are activated at the onset of the flight-like activity and remain active sporadically throughout the flight sequence (Fig.4); (c) some descending axons in the abdominal nerve cord show rhythmic activity phase-locked to the flight rhythm (Fig. 5). 4. Also during such brief sequences, the cercal nerves, running from the cerci (paired, posterior, wind sensitive appendages) to the last abdominal ganglion, show rhythmic activity at the wingbeat frequency (Fig. 6). This includes activity of some motor axons controlling vibratory cercal movements and of some sensory axons. 5. More prolonged flight sequences were elicited in cockroaches whose wings were not cut and which flew in front of a wind tunnel (Fig. 1B). 6. In these more prolonged flight sequences, the number of ascending spikes per burst was greater than that seen in the wingless preparation (Fig. 8; compare to Fig. 2). Recordings from both ventral and dorsal GIs show that: in spite of the ongoing wind from both the tunnel and the beating wings, which is far above threshold for the vGIs in a resting cockroach, the vGIs are entirely silent during flight. Moreover, the vGIs response to strong wind puffs that normally evoke maximal GI responses is reduced by a mean of 86% during flight (Fig. 9). The dGIs are active in a strong rhythm (Figs. 11 and 12). 7. Three sources appear to contribute to the ascending dGI rhythm (1) the axons carrying the rhythmic descending bursts; (2) the rhythmic sensory activity resulting from the cercal vibration; and (3) the sensory activity resulting from rhythmic wind gusts produced by the wingbeat and detected by the cerci. The contribution of each source has been tested alone while removing the other two (Figs. 13 and 14). Such experiments suggest that all 3 feedback loops are involved in rhythmically exciting the dGIs (Fig. 15).

Action Potentials↗

Insecticidal and neuroexcitant actions of DDT analogs on the cockroach, Periplaneta americana.

The American cockroach (Periplaneta americana) was used to test 14 DDT analogs for their ability to cause a) death and b) repetitive firing in the abdominal nerve cord. P,p' -DDT, p,p' -DDD, p,p' -methoxychlor, and o,p -DDT were all found to induce repetitive firing in the nerve cord after a twice threshold stimulus, but only p,p' -DDT, p,p' -DDD, and p,p' -methoxychlor produced mortality within 4 days from injection of the pesticides. Also, cockroaches exposed to sublethal doses of p,p' -DDT and p,p' -methoxychlor were found to exhibit repetitive firing in the abdominal nerve cord three weeks after injection of the pesticide. The 'cause and effect' relationship between neuroexcitation and lethality is therefore questioned, at least in the case of cockroaches.

Action Potentials↗

When males whistle at females: complex FM acoustic signals in cockroaches.

Male cockroaches of the species Elliptorhina chopardi expel air through a pair of modified abdominal spiracles during courtship. This air expulsion simultaneously produces air and substrate-borne vibrations. We described and compared in details these two types of vibrations. Our analysis of the air-borne signals shows that males can produce three categories of signals with distinct temporal and frequency parameters. "Pure whistles" consist of two independent harmonic series fast frequency modulated with independent harmonics that can cross each other. "Noisy whistles" also possess two independent voices but include a noisy broad-band frequency part in the middle. Hiss sounds are more noise-like, being made of a broad-band frequency spectrum. All three call types are unusually high in dominant frequency (>5 kHz) for cockroaches. The substrate-borne signals are categorised similarly. Some harmonics of the substrate-borne signals were filtered out, however, making the acoustic energy centered on fewer frequency bands. Our analysis shows that cockroach signals are complex, with fast frequency modulations and two distinct voices. These results also readdress the question of what system could potentially receive and decode the information contained within such complex sounds.

Abdomen↗

Effects of neck and circumoesophageal connective lesions on posture and locomotion in the cockroach.

Few studies in arthropods have documented to what extent local control centers in the thorax can support locomotion in absence of inputs from head ganglia. Posture, walking, and leg motor activity was examined in cockroaches with lesions of neck or circumoesophageal connectives. Early in recovery, cockroaches with neck lesions had hyper-extended postures and did not walk. After recovery, posture was less hyper-extended and animals initiated slow leg movements for multiple cycles. Neck lesioned individuals showed an increase in walking after injection of either octopamine or pilocarpine. The phase of leg movement between segments was reduced in neck lesioned cockroaches from that seen in intact animals, while phases in the same segment remained constant. Neither octopamine nor pilocarpine initiated changes in coordination between segments in neck lesioned individuals. Animals with lesions of the circumoesophageal connectives had postures similar to intact individuals but walked in a tripod gait for extended periods of time. Changes in activity of slow tibial extensor and coxal depressor motor neurons and concomitant changes in leg joint angles were present after the lesions. This suggests that thoracic circuits are sufficient to produce leg movements but coordinated walking with normal motor patterns requires descending input from head ganglia.

Action Potentials↗

Computational model of the cockroach escape behavior: winner and losers in a population code.

I present a comprehensive biologically oriented computational model to account for the escape response of the cockroach on the ground. This model is an expansion of previous work that accounted only for discriminating left from right wind directions [Ezrachi et al. (1999) Biol Cybern 81: 89-99]. The model is composed of computational elements describing the biological processes taking place in the various neurons and includes input which emulates empirical data. With this model it is possible to obtain escape behavior that resembles natural behavior. The model is used to address an ongoing debate as to whether the cockroach's turn direction is determined by computations carried out by the entire neuronal population (PC) or rather by a "winner-take-all" (WTA) mechanism. I suggest that the computation mechanism that underlies the cockroach escape response is composed of both PC and WTA principles. Based on the properties of the suggested new mechanism I denote it a "Darwinian population code."

Animals↗

Isolation and structure of two gastrin/CCK-like neuropeptides from the American cockroach homologous to the leucosulfakinins.

Perisulfakinin, a peptide with sequence similarity to gastrin and cholecystokinin, was isolated from the corpora cardiaca of the American cockroach. Its sequence was determined to be Glu-Gln-Phe H-Asp-Asp-Tyr(SO3H)-Gly-His-Met-Arg-Phe-amide. The peptide induced hindgut contractions in the same species at concentrations as low as 250 pM. A related non-sulfated peptide was also isolated and sequenced; it was found to be identical with non-sulfated leucosulfakinin II (pGlu-Ser-Asp-Asp-Tyr-Gly-His-Met-Arg-Phe-amide). This peptide did not stimulate hindgut contractions. The structures of the cockroach peptides of the leucosulfakinin family are thus much more conserved than the cockroach hypertrehalosemic hormones.

Amino Acid Sequence↗

The effect of biotic and abiotic implants on the recognition of Blatta orientalis cuticular transplants by the cockroach Periplaneta americana.

The immune system of the American cockroach, Periplaneta americana, was activated to recognise as "non self" xenogeneic tissue normally treated as "self". Activation was accomplished by injecting into the insect haemocoele material known to elicit an encapsulation response such as Blaberus craniifer cuticular tissue and 6B-Sepharose beads. In these insects, cuticular skingrafts from Blatta orientalis, a closely related species of cockroach, were rejected by more than half of the recipients. There was no rejection of skingrafts by naive insects showing that the immune system of the cockroach had been triggered non-specifically to recognise tissue previously treated as "self".

Animals↗