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PubMed2002

IgE-binding epitopes of the American cockroach Per a 1 allergen.

Cockroach is one of the major indoor allergens for IgE-mediated allergic respiratory illnesses throughout the world. The American cockroach (Periplaneta americana) Per a 1 allergen is antigenically cross-reactive with the German cockroach (Blattella germanica) Bla g 1 allergen. The aim of this study was to identify linear B cell epitopes of Per a 1 that are recognized by human IgE. Per a 1 deletion mutants were generated from the recombinant Per a 1.0104 allergen (274 amino acid residues), and antigenicities were assessed by immunoblotting, enzyme-linked immunosorbent assay (ELISA), and binding inhibition. Human atopic sera were not able to recognize deletion mutants consisting of amino acids 1-77, 86-205, and 200-266. However, human sera did recognize the N-terminal mutant containing amino acids 1-87 and the C-terminal mutant containing amino acids 200-274, demonstrating positive IgE binding that was heterogeneously distributed among the different sera tested. Amino acid residues 78-85 and 267-274, containing internal repeats, were shown to be required for IgE binding to the Per a 1.0104 protein. Two peptides corresponding to these IgE-binding amino acid sequences were synthesized. Peptide 78-85 showed a positive IgE interaction with 80% of the sera, while peptide 267-274 was capable of IgE binding to all of the sera tested. Moreover, preincubation of atopic sera with IgE-positive recombinants and peptides resulted in marked inhibition of the IgE binding to purified Per a 1.0104 allergen. Amino acid sequences 78LIRALFGL85 and 267IRSWFGLP1274 of the American cockroach Per a 1.0104 allergen were involved in IgE binding. These findings will advance the understanding of the specific reactivity of the epitopes of cockroach allergens, thereby contributing to the development of specific immunotherapies for clinical use.

Allergens↗
PubMed2001

Myoinhibitory neuropeptides in the American cockroach.

A large number of myostimulatory neuropeptides from neurohaemal organs of the American cockroach have been described since 1989. These peptides, isolated from the retrocerebral complex and abdominal perisympathetic organs, are thought to be released as hormones. To study the coordinated action of these neuropeptides in the regulation of visceral muscle activity, it might be necessary to include myoinhibitors as well, however, not a single myoinhibitory neuropeptide of the American cockroach has been described so far. To fill this gap, we describe the isolation of LMS (leucomyosuppressin) and Pea-MIP (myoinhibitory peptide) from neurohaemal organs of the American cockroach. LMS was very effective in inhibiting phasic activity of all visceral muscles tested. It was found in the corpora cardiaca of different species of cockroaches, as well as in related insect groups, including mantids and termites. Pea-MIP which is strongly accumulated in the corpora cardiaca was not detected with a muscle bioassay system but when searching for tryptophane-containing peptides using a diode-array detector. This peptide caused only a moderate inhibition in visceral muscle assays. The distribution of Pea-MIP in neurohaemal organs and cells supplying these organs with Pea-MIP immunoreactive material, is described. Additionally to LMS and Pea-MIP, a member of the allatostatin peptide family, known to exhibit inhibitory properties in other insects, was tested in visceral muscle assays. This allatostatin was highly effective in inhibiting spontaneous activity of the foregut, but not of other tested visceral muscles of the American cockroach.

Animals↗
PubMed2003

Isolation of non-tuberculous mycobacteria from hospital cockroaches (Periplaneta americana).

This study attempted to isolate mycobacteria from hospital and household cockroaches from 90 hospitals and 40 households in Kaohsiung City and Kaohsiung County, South Taiwan. Among 203 cockroaches (139 Periplaneta americana and 64 Blattella germanica) collected from the hospitals, six Mycobacterium spp. were isolated and identified by polymerase chain reaction-restriction fragment length polymorphism analysis. In 12 cockroaches (P. americana): four Mycobacterium kansaii, three Mycobacterium xenopi, two Mycobacterium gordonae, one Mycobacterium hemophilium, one Mycobacterium fortuitum, and one Mycobacterium avium. However, no mycobacteria were obtained form the hospital B. germanica or 226 household cockroaches (123 P. americana and 103 B. germanica). As cockroach infestation occurs commonly in the hospital environment, they may potentially be implicated as a cause of hospital-acquired infections due to non-tuberculous mycobacteria.

Animals↗
PubMed2003

Abatement of cockroach allergen (Bla g 1) in low-income, urban housing: A randomized controlled trial.

BACKGROUND: Clinically relevant reductions in exposure to cockroach allergen, an important risk factor for asthma in inner-city households, have proven difficult to achieve in intervention trials. OBJECTIVE: This study investigated a method for the abatement of cockroach allergen in low-income, urban homes. The goal was to reduce mean Bla g 1 concentrations below the previously proposed thresholds for allergic sensitization and asthma morbidity. METHODS: A prerandomized, nonmasked trial with 16 intervention and 15 control homes was conducted. Study inclusion was based on 50 to 500 cockroaches trapped in a 3-day period. The interventions consisted of occupant education, placement of insecticide bait, and professional cleaning. Vacuumed dust and multiple swab samples were collected at 0, 1, 2, 4, and 6 months in intervention homes and at 0 and 6 months in control homes. Room maps containing cockroach and allergen data were used to guide and monitor the interventions. RESULTS: From 0 to 6 months among intervention homes, geometric mean Bla g 1 concentrations (U/g dust) decreased from 633 to 24 on kitchen floors (96% reduction), from 25 to 4.3 on living room floors/sofas (83% reduction), from 46 to 7.3 on bedroom floors (84% reduction), and from 6.1 to 1.0 in bedroom beds (84% reduction). These reductions, with the exception of that on the bedroom floor (P =.06), were statistically significant relative to changes in control homes. CONCLUSIONS: Substantial reductions in cockroach allergen levels can be achieved in inner-city homes. In this study, allergen levels were reduced below the sensitization threshold (2 U/g) in beds, arguably the most relevant site for exposure, and below the asthma morbidity threshold (8 U/g) on bedroom floors and living room floors/sofas. The level on kitchen floors, although reduced 96%, remained above the asthma morbidity threshold. Future studies will test the intervention's effectiveness in asthma prevention trials.

Allergens↗
PubMed2024

Solving the 250-year-old mystery of the origin and global spread of the German cockroach, Blattella germanica.

The origin of the German cockroach, Blattella germanica, is enigmatic, in part because it is ubiquitous worldwide in human-built structures but absent from any natural habitats. The first historical records of this species are from ca. 250 years ago (ya) from central Europe (hence its name). However, recent research suggests that the center of diversity of the genus is Asian, where its closest relatives are found. To solve this paradox, we sampled genome-wide markers of 281 cockroaches from 17 countries across six continents. We confirm that B. germanica evolved from the Asian cockroach Blattella asahinai approximately 2,100 ya, probably by adapting to human settlements in India or Myanmar. Our genomic analyses reconstructed two primary global spread routes, one older, westward route to the Middle East coinciding with various Islamic dynasties (~1,200 ya), and another younger eastward route coinciding with the European colonial period (~390 ya). While Europe was not central to the early domestication and spread of the German cockroach, European advances in long-distance transportation and temperature-controlled housing were likely important for the more recent global spread, increasing chances of successful dispersal to and establishment in new regions. The global genetic structure of German cockroaches further supports our model, as it generally aligns with geopolitical boundaries, suggesting regional bridgehead populations established following the advent of international commerce.

Animals↗
PubMed1997

Comparison of traps and development of a two-stage sampling plan for smokybrown cockroaches (Dictyoptera: Blattidae) outdoors.

Four commercial sticky traps were compared with jar traps for sampling smokybrown cockroaches, Periplaneta fuliginosa (Serville), outdoors. Lo-Line and Raid Roach Traps caught more cockroaches than jar traps, whereas the Mr. Sticky and Roach Motel caught < 40% of the cockroaches caught in jar traps. Sampling properties of traps (data collected over 4 yr and 3 cities) were examined retrospectively. Taylor's a and b coefficients were both significantly > 1 for jar traps, Raid Roach Trap, and Lo-Line trap. Estimating cockroach catch typical of Alabama (< 3 per trap) at single houses would require > 20 traps with a precision of 0.25 (i.e., standard error as a proportion of the mean). Taylor's power law was also used as a predictor for within-house (i.e., pooled within-house) and between-house catch variances. Coefficients for these predictors are presented that can be used in a fixed-precision 2-stage sampling plan to estimate cockroach catch from large areas.

Animals↗
PubMed1997

Evaluation of methods of insecticide application for control of smokybrown cockroaches (Dictyoptera: Blattidae).

The insecticide components of a previously developed integrated pest management system for smokybrown cockroaches, Periplaneta fuliginosa (Serville), were tested individually and at reduced application rates in 4 separate trials over 2 yr. A 2-bait combination (pellet and gel baits) did not significantly reduce cockroach abundance more than a single constituent bait alone. A targeted spray of trelamethrin was significantly less effective than a 2-bait combination. Residual activity of insecticides was determined with American cockroaches, P. americana (L.), in a laboratory bioassay. Chlorpyrifos wettable powder remained active < 12 d in the field, after which time its activity was not different from an experimental control (no insecticide residues), whereas chlorpyrifos pellet and hydramethylnon paste baits retained initial activity for > 27 d. Nematodes [Steinernema carpocapsae (Weiser)] applied as a spray solution did not effect the abundance of smokybrown cockroaches, and no cockroaches were recovered that exhibited nematode infection. Time needed to apply various insecticide treatments did not differ substantially (15-25 min per house), even for lower application rates.

Animals↗
PubMed2003

Improvement of German cockroach (Dictyoptera: Blattellidae) population control by fragmented distribution of gel baits.

The aim of this study was to test the efficiency of gels in relation to fragmentation of baits and population density against the German cockroach, Blattella germanica (L.). We hypothesized that the efficiency of cockroach control could be improved by fragmentation of gel baits, i.e., by distributing the same amount of bait in several small drops instead of one large one. Our results show that bait fragmentation increased the number of cockroaches gaining access to the gel. However, bait fragmentation increased cockroach mortality only at high population densities. Our results allow us to recommend modulation of bait application in relation to cockroach population density. At low population densities (42 individuals/m2), the recommended dose, one single 30-mg drop/m2, can be applied. At high population densities (> or = 208 individuals/m2), the same recommended dose would be more efficient if applied as several 3-mg drops.

Animals↗
PubMed2003

Origin and extent of resistance to fipronil in the German cockroach, Blattella germanica (L.) (Dictyoptera: Blattellidae).

Fipronil, a phenylpyrazole insecticide, was made available in 1999 in bait formulations for use against the German cockroach, Blattella germanica (L.). We have investigated resistance to fipronil in the descendants of cockroaches collected just before, or contemporaneously with, the introduction of fipronil baits. Cockroaches were obtained in two types of settings: homes that either had or had not been serviced by a pest management professional while occupied by their current residents. Thorough inspections by us turned up no evidence that fipronil had been used in any of the homes, and in addition, no residents claimed to have used baits containing fipronil. Resistance to fipronil was detected by topically dosing adult males with the LC99 of fipronil, the value of which was determined in a dose-response assay with males of an insecticide-susceptible strain. Fewer than 99 of 100 males of all field-collected strains died within 72 h of being treated. Moreover, substantial numbers of males survived doses three and 10-fold greater than the LC99. Regression analysis showed that 67% of the variation in the percentage of males that died after being treated with fipronil was explained by a linear relationship with the percentage that died after being treated with dieldrin. Therefore, it appears that resistance to fipronil in German cockroaches--whose ancestors had never been exposed to it--is attributable to enduring resistance to the cyclodienes, which were formerly used for cockroach control and have a similar mode of action as fipronil. Lastly, we found that insects resistant to topically administered fipronil were likewise resistant, and to a similar degree, to ingested fipronil.

Animals↗
PubMed2004

Allergenic characterization of tropomyosin from the dusky brown cockroach, Periplaneta fuliginosa.

Household arthropods are one of the most common causes of allergic diseases. Four species of cockroaches are found to reside in Korean homes, but published work deals almost exclusively with the German and American cockroaches. This study was undertaken to investigate the cross-reactive allergenic components of the dusky brown cockroach, Periplaneta fuliginosa. Enzyme-linked immunosorbent assay (ELISA) inhibition and immunoblot analyses for the dusky brown cockroach were performed with Blattella germanica and Dermatophagoides farinae allergic sera. cDNA encoding tropomyosin, which is a well known cross-reactive pan-allergen, was cloned by reverse transcriptase PCR, and recombinant protein was produced by using a pET-28b expression system. Native tropomyosin was purified by ammonium sulfate fractionation and electroelution. The immunoglobulin E (IgE) reactivities of native and recombinant tropomyosins were compared by an ELISA inhibition study. All 30 sera tested showed P. fuliginosa-specific IgE, and the IgE-binding reactivity of the P. fuliginosa extract was inhibited as much as 79.4% by a B. germanica extract and as much as 63.3% by a D. farinae extract. The deduced amino acid sequence of cloned cDNA was identical with that of Periplaneta americana tropomyosin (98.5% nucleotide sequence identity). Seven of 26 (26.9%) allergic sera had IgE specific for recombinant protein, and the maximum inhibition of P. fuliginosa-specific IgE achieved with recombinant tropomyosin was 37.7% at an inhibitor concentration of 10 microg/ml. Native tropomyosin inhibited the binding of IgE to the P. fuliginosa, B. germanica, and D. farinae extracts by 65.0, 51.8, and 39% at an inhibitor concentration of 1 microg/ml. P. fuliginosa appears to possess allergens that are highly cross-reactive with allergens of B. germanica and D. farinae. Tropomyosin was found to be a major allergenic component accounting for the cross-reactivity between cockroaches and dust mites.

Adolescent↗
PubMed1995

Maximum single leg force production: cockroaches righting on photoelastic gelatin

Integrating studies of mechanics, neural control and isolated muscle function are possible using arthropod legs. To evaluate leg performance, we measured the ground reaction forces generated by individual legs of the six-legged cockroach Blaberus discoidalis (3.1 g), during an emergency behavior, righting or over-turning. We used a photoelastic method to measure the forces generated by individual legs simultaneously. A gelatin track placed between crossed polarizing filters was illuminated from below, and a high-speed video camera recorded the stress-induced optical signals from above. The size and skew of the optical patterns were found to be related to the magnitude and direction of the force. We discovered that the ground reaction forces generated during the righting behavior of the death-head cockroach were eight times greater than those observed during high-speed running, supporting the possibility that relative leg forces (leg force per unit body weight) during running and maximal leg activity differ more in small arthropods than in larger vertebrates. Non-geometric scaling of relative leg force (i.e. scaling to less than body mass-0.33), along with the reduced force-generating ability of a single leg in animals with many legs, may help to explain why the maximum relative leg force production by six-legged cockroaches, as well as by some other small insects, can be similar to the relative single leg forces produced by two- and four-legged vertebrates that are almost 1000 times more massive. Leg number and body mass alone, however, appear to be insufficient to explain the variation observed in relative leg force production at a given body mass, because enormous diversity in musculo-skeletal parameters exists. The maximal relative leg force of the cockroach B. discoidalis during righting was at the low end of a 100-fold variation observed for smaller insects wedging (pushing through a small crevice) and pulling loads. Thus, this cockroach can be characterized as a moderately strong insect with the capacity for relatively high speed. Results from the present study question the predictive strength of the simple geometric scaling arguments involving a strength:weight ratio as they are applied to small arthropods and encourage further consideration of the importance of leg number, muscle force production and mechanical advantage in the derivation of general principles of leg performance.

Journal Article↗
PubMed2002

[Enterobacteria isolated from cockroaches (Periplaneta americana) captured in a Brazilian hospital].

OBJECTIVE: To isolate and identify microorganisms from cockroaches that were captured in a public hospital and to test the antimicrobial susceptibility of these microorganisms. METHODS: Cockroaches were captured in the morning and at night. They were placed in flasks rinsed with 70% alcohol, transferred to sterilized flasks, and then taken to the laboratory. Only cockroaches captured whole and live were utilized for the study. After being immobilized at 0 degree C, each cockroach was placed in a test tube with sterile saline solution (0.8%) and then homogenized. The resulting solution was then placed in the following five culture media: MacConkey agar, nutrient broth, brain-heart infusion agar, Sabouraud agar, and mannitol. The cultures were examined using a stereomicroscope, and colony-forming units were counted. The disk diffusion test was used to determine antimicrobial susceptibility. RESULTS: We found a 56% prevalence of enterobacteria and an 18% prevalence of coagulase-negative staphylococci. Fifteen species of enterobacteria were identified. The most frequent were Klebsiella pneumoniae (17%), Enterobacter aerogenes (14%), Serratia marcescens (13%), Hafnia alvei (12%), Enterobacter gergoviae and Enterobacter cloacae (each 9%), and Serratia spp. (6%). Both the enterobacteria and the coagulase-negative staphylococci showed significant resistance to antimicrobials, including oxacillin. CONCLUSIONS: The prevalence of enteropathogenic bacteria and coagulase-negative staphylococci isolated from Periplaneta americana cockroaches in the studied hospital reflects the weakness of the measures adopted both for vector control and for antimicrobial use. The results show the need to implement effective health-institution programs focusing on hygiene and the rational use of antimicrobials.

Animals↗
PubMed2006

Profile of antimicrobial resistance of bacteria isolated from cockroaches (Periplaneta americana) in a Brazilian health care institution.

Cockroaches carry microorganisms to sterilized materials, equipment and non-contaminated food in hospitals and in homes. Hence, they are regarded as major microbial vectors. We isolated and identified microorganisms from living cockroaches captured from a Brazilian health care institution and we determined the antimicrobial susceptibility profile of the isolates. Following collection, cockroaches were immobilized at 4 degrees C for 10 to 20 minutes, immersed in 0.8% saline solution and then homogenized. Next, the suspension was seeded in culture media: brain and heart agar infusion, MacConkey agar, Naito agar, and they were incubated for up to 48 hours at 37 degrees C. Sabouraud agar cultures were incubated at ambient temperature. The microbial colonies were analyzed with a stereomicroscope to count and phenotypically identify the colony-forming units (cfu). The disc-diffusion method was used to determine the profile of susceptibility to antimicrobials. Among the 103 cockroaches analyzed, fungi were found in 97%, enterobacteria in 74.6% and coagulase-negative staphylococci (CNS) in 25.40%. Among the enterobacteria, 96% were resistant to gentamicin, 84% to ampicillin, 75.3% to caphalothin, 66.7% to ampicillin-sulbactam, 50% to aztreonam and 30% to chloramphenicol. Among the CNS, 61% were resistant to oxacillin. We concluded that cockroaches carry organisms associated with nosocomial infections and that are resistant to antimicrobials. This fact contributes to the epidemiological chain, complicating therapeutics, and consequently, medical costs are increased.

Animals↗
PubMed2000

Laboratory and field performance of an imidacloprid gel bait against German cockroaches (Dictyoptera: Blattellidae).

An experimental 2.15% imidacloprid gel bait containing approximately 44% water was evaluated in laboratory and field studies against the German cockroach, Blattella germanica (L). In continuous exposure tests, toxicity and presumably bait consumption varied with cockroach stage, deprivation of competitive food, and temperature. The LT50 values for cockroaches provided with competitive food ranged from approximately 0.9 h for adult females to 190 h for small nymphs. The LT50s for cockroaches not provided competitive food ranged from approximately 1.7 h for adult females to approximately 31 h for adult males. The LT50s decreased exponentially with temperature between 10 and 30 degrees C. Even though the bait was significantly more repellent (approximately 38%) than an untreated control (approximately 14%) when tested in Ebeling choice boxes, performance index values were positive and increased to nearly 100 (indicating high mortality and low repellency) after 14 d. When applied at 15-45 g per kitchen, the bait significantly reduced German cockroach trap catch in infested homes during a 4-wk period. There was a approximately 50% reduction after 1 wk and approximately 80% reduction 4 wk after treatment.

Animals↗
PubMed2001

Decapitation impacting effect of topically applied chlorpyrifos on acetylcholinesterase and general esterases in susceptible and resistant German cockroaches (Dictyoptera: Blattellidae).

The effect of topically applied chlorpyrifos on acetylcholinesterase and other esterases in heads and decapitated bodies of CSMA and Crawford German cockroaches was examined with spectrophotometric enzyme assay and native polyacrylamide gel electrophoresis. The toxicity of chlorpyrifos was greatly reduced in decapitated CSMA male cockroaches with LD50 value 17.1-fold higher than that of normal CSMA cockroaches. Acetylcholinesterase activity from heads was significantly higher in the Crawford compared with the CSMA strain and did not change until 24 h after chlorpyrifos in vivo treatment in both strains. The p-nitrophenyl butyrate (NPB) esterase activities from both heads and decapitated bodies of the resistant Crawford strain were significantly greater than the susceptible CSMA strain. The p-NPB esterase activity was significantly inhibited by chlorpyrifos in vivo treatment, and total p-NPB esterase activity was significantly reduced in decapitated bodies compared with heads of both strains. Native polyacrylamide gel electrophoresis (PAGE) analysis of extracts solubilized with Triton X-100 from heads and decapitated bodies revealed five major esterase bands and an acetylcholinesterase (AChE) band with a high capability of hydrolyzing alpha-naphthyl butyrate and acetylthiocholine, respectively. In the heads of susceptible CSMA male cockroaches, the activity of mobile isozymes d1 and d2 was completely inhibited at 24 h after chlorpyrifos application, and isozyme e was partially inhibited. In contrast, isozymes c1 and c2 from the decapitated bodies of CSMA cockroaches were mostly affected at 24 h after the topical application of chlorpyrifos. The activities of acetylcholinesterase and esterase isozymes a and b from the decapitated body remained uninhibited in both strains. Inhibition of isozymes d1 and d2 seems to be more important in chlorpyrifos intoxication than acetylcholinesterase.

Acetylcholinesterase↗
PubMed1995

Cold-induced amnesia blocks escape deficits in the cockroach (Periplaneta americana).

For three consecutive days, four groups of adult cockroaches (Periplaneta americana) were yoked to cockroaches receiving escapable shock. The four groups of subjects received either inescapable shock followed by exposure to 12 minutes of cold (2 degrees C), inescapable shock followed by no exposure to cold, no-shock followed by exposure to cold, or no-shock followed by no exposure to cold. 24 hr. later all five groups were run on a different escape task. The cockroaches exposed to inescapable shock followed by no exposure to cold showed the usual escape deficit. The cockroaches exposed to inescapable shock followed by exposure to cold did not alter from the escapable shock cockroaches or the controls.

Animals↗
PubMed2006

The aminergic control of cockroach salivary glands.

The acinar salivary glands of cockroaches receive a dual innervation from the subesophageal ganglion and the stomatogastric nervous system. Acinar cells are surrounded by a plexus of dopaminergic and serotonergic varicose fibers. In addition, serotonergic terminals lie deep in the extracellular spaces between acinar cells. Excitation-secretion coupling in cockroach salivary glands is stimulated by both dopamine and serotonin. These monoamines cause increases in the intracellular concentrations of cAMP and Ca(2+). Stimulation of the glands by serotonin results in the production of a protein-rich saliva, whereas stimulation by dopamine results in saliva that is protein-free. Thus, two elementary secretory processes, namely electrolyte/water secretion and protein secretion, are triggered by different aminergic transmitters. Because of its simplicity and experimental accessibility, cockroach salivary glands have been used extensively as a model system to study the cellular actions of biogenic amines and to examine the pharmacological properties of biogenic amine receptors. In this review, we summarize current knowledge concerning the aminergic control of cockroach salivary glands and discuss our efforts to characterize Periplaneta biogenic amine receptors molecularly.

Animals↗
PubMed2003

A unique mushroom body substructure common to basal cockroaches and to termites.

The mushroom bodies of the cockroach Periplaneta americana are made up of intrinsic neurons (class I and class II Kenyon cells) with dendrites in a dorsal calyx and axons that bifurcate into medial and vertical lobes. Here, we describe a substructure of the cockroach mushroom bodies composed of a previously unrecognized class of Kenyon cells with distinct morphologies. The embryonically produced class III Kenyon cells form a separate accessory calyx below the calyx proper. The medial branches of class III Kenyon cell axons form the previously described "gamma bulb," whereas the vertical branches leave the vertical lobe to form a toroidal "lobelet" around the posterior surface. Taking advantage of the morphologically and immunochemically distinct nature of the lobelet, we have attempted to determine the distribution of this unique structure in other insects of the taxon Dictyoptera (cockroaches, mantises, and termites). Our data indicate that the lobelet is present only in basal cockroaches and in termites, supporting existing theories of a close phylogenetic relationship between these groups. Higher termites possess a duplicated lobe structure due to immense elaboration of the processes of class III Kenyon cells. The degree of complexity in the mushroom body lobes of termites agrees with current taxonomic arrangements of the Isoptera based on non-neural morphological and DNA sequence analyses. It thus appears that the evolution of the Dictyoptera has been accompanied by increasing complexity of the mushroom bodies, achieved in part through the further specialization and elaboration of a subset of Kenyon cells.

Animals↗