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Thickening of the integument as a mechanism of resistance to insecticides is a well recognised phenomenon in the insect world and, in recent times, has been found in insects exhibiting pyrethroid-resistance. Resistance to pyrethroid insecticides in the common bed bug, Cimex lectularius L., is widespread and has been frequently inferred as a reason for the pest's resurgence. Overexpression of cuticle depositing proteins has been demonstrated in pyrethroid-resistant bed bugs although, to date, no morphological analysis of the cuticle has been undertaken in order to confirm a phenotypic link. This paper describes examination of the cuticle thickness of a highly pyrethroid-resistant field strain collected in Sydney, Australia, in response to time-to-knockdown upon forced exposure to a pyrethroid insecticide. Mean cuticle thickness was positively correlated to time-to-knockdown, with significant differences observed between bugs knocked-down at 2 hours, 4 hours, and those still unaffected at 24 hours. Further analysis also demonstrated that the 24 hours survivors possessed a statistically significantly thicker cuticle when compared to a pyrethroid-susceptible strain of C. lectularius. This study demonstrates that cuticle thickening is present within a pyrethroid-resistant strain of C. lectularius and that, even within a stable resistant strain, cuticle thickness will vary according to time-to-knockdown upon exposure to an insecticide. This response should thus be considered in future studies on the cuticle of insecticide-resistant bed bugs and, potentially, other insects.

Emerging infestations of bed bugs are affecting normal human lifestyle globally. This study has been designed to optimize the rearing conditions for Cimex lectularius L. (Hemiptera), to support the scientific research on them. Bed bugs have been projected onto three different temperature (20 °C, 25 °C, and 30 °C) and relative humidity (50%, 70%, and 90%) conditions to check their overall growth and survival rate. Adult mortality, weight loss, egg laying, percentage hatching, hatching initiation and completion, nymph mortality, and molting have been evaluated to optimize the best conditions. The temperature at 25 °C with 90% RH showed minimum mortality for adults (female 13.33 ± 3.33% and male 6.67 ± 3.33%) and nymphs (13.33 ± 3.33%), while maximum egg laying (40.33 ± 1.86), with highest percentage hatching (98.23 ± 0.58%). At 30 °C with 90% RH, hatching initiation and completion (5.19 ± 0.12 days and 7.23 ± 0.16 days) as well as molting initiation and completion (3.73 ± 0.12 days and 7.00 ± 0.24 days) were found to be fastest. Thus, it can be concluded that 25 °C with 90% RH is ideal for rearing of adults and 30 °C with 90% RH is appropriate for rapid growth of nymphs.

Understanding movement and dispersal of the common bed bug (Cimex lectularius L.) under field conditions is important in the control of infestations and for managing the spread of bed bugs to new locations. We investigated bed bug movement within and between apartments using mark-release-recapture (m-r-r) technique combined with apartment-wide monitoring using pitfall-style interceptors. Bed bugs were collected, marked, and released in six apartments. The distribution of marked and unmarked bed bugs in these apartments and their 24 neighboring units were monitored over 32 days. Extensive movement of marked bed bugs within and between apartments occurred regardless of the number of bed bugs released or presence/absence of a host. Comparison of marked and unmarked bed bug distributions confirms that the extensive bed bug activity observed was not an artifact of the m-r-r technique used. Marked bed bugs were recovered in apartments neighboring five of six m-r-r apartments. Their dispersal rates at 14 or 15 d were 0.0-5.0%. The estimated number of bed bugs per apartment in the six m-r-r apartments was 2,433-14,291 at 4-7 d after release. Longevity of bed bugs in the absence of a host was recorded in a vacant apartment. Marked large nymphs (3rd- 5th instar), adult females, and adult males continued to be recovered up to 57, 113, and 134 d after host absence, respectively. Among the naturally existing unmarked bed bugs, unfed small nymphs (1st- 2nd instar) were recovered up to 134 d; large nymphs and adults were still found at 155 d when the study ended. Our findings provide important insight into the behavioral ecology of bed bugs in infested apartments and have significant implications in regards to eradication programs and managing the spread of bed bugs within multi-occupancy dwellings.

Bartonella quintana, the etiologic agent of trench fever and other human diseases, is transmitted by the feces of body lice. Recently, this bacterium has been detected in other arthropod families such as bed bugs, which begs the question of their involvement in B. quintana transmission. Although several infectious pathogens have been reported and are suggested to be transmitted by bed bugs, the evidence regarding their competence as vectors is unclear. Bed bugs at the adult and instar developmental stages were fed three successive human blood meals inoculated with B. quintana bacterium from day one (D1) to D5; subsequently they were fed with pathogen-free human blood until the end of the experiment. Bed bugs and feces were collected in time series, to evaluate their capacities to acquire, multiply and expel viable B. quintana using molecular biology, immunohistochemistry and cultures assays. B. quintana was detected molecularly in 100% of randomly selected experimentally infected bed bug specimens (D3). The monitoring of B. quintana in bed bug feces showed that the bacterium was detectable starting on the 3rd day post-infection (pi) and persisted until day 18±1 pi. Although immunohistochemistry assays localized the bacteria to the gastrointestinal bed bug gut, the detection of B. quintana in the first and second instar larva stages suggested a vertical non-transovarial transmission of the bacterium. The present work demonstrated for the first time that bed bugs can acquire, maintain for more than 2 weeks and release viable B. quintana organisms following a stercorarial shedding. We also observed the vertical transmission of the bacterium to their progeny. Although the biological role of bed bugs in the transmission of B. quintana under natural conditions has yet to be confirmed, the present work highlights the need to reconsider monitoring of these arthropods for the transmission of human pathogens.

Recent resurgence of the bedbug Cimex lectularius is a global problem on the public health. On account of the worldwide rise of insecticide-resistant bedbug populations, exploration of new approaches to the bedbug control and management is anticipated. In this context, gene silencing by RNA interference (RNAi) has been considered for its potential application to pest control and management, because RNAi enables specific suppression of target genes and thus flexible selection of target traits to be disrupted. In this study, in an attempt to develop a control strategy targeting reproduction of the bedbug, we investigated RNAi-mediated gene silencing of vitellogenin (Vg), a major yolk protein precursor essential for oogenesis. From the bedbug transcriptomes, we identified a typical Vg gene and a truncated Vg gene, which were designated as ClVg and ClVg-like, respectively. ClVg gene was highly expressed mainly in the fat body of adult females, which was more than 100 times higher than the expression level of ClVg-like gene, indicating that ClVg gene is the primary functional Vg gene in the bedbug. RNAi-mediated suppression of ClVg gene expression in adult females resulted in drastically reduced egg production, atrophied ovaries, and inflated abdomen due to hypertrophied fat bodies. These phenotypic consequences are expected not only to suppress the bedbug reproduction directly but also to deteriorate its feeding and survival indirectly via behavioral modifications. These results suggest the potential of ClVg gene as a promising target for RNAi-based population management of the bedbug.

Sublethal heat stress may weaken bed bug infestations to potentially ease control. In the present study, experimental populations exposed to 34, 36 or 38°C for 2 or 3 weeks suffered significant mortality during exposure. Among survivors, egg production, egg hatching, moulting success and offspring proliferation decreased significantly in the subsequent 7 week recovery period at 22°C. The overall population success was negatively impacted by increasing temperature and duration of the stress. Such heat stress is inadequate as a single tool for eradication, but may be included as a low cost part of an integrated pest management protocol. Depending on the time available and infestation conditions, the success of some treatments can improve if sublethal heat is implemented prior to the onset of more conventional pest control measures.

Many insect species display daily variation of sensitivity to insecticides when they are exposed to the same concentration at different times during the day. To date, this has not been investigated in bed bugs. To address this, we explored circadian rhythms in insecticide susceptibility, xenobiotic metabolizing (XM) gene expressions, and metabolic detoxification in the common bed bug, Cimex lectularius. An insecticide susceptible Monheim strain of C. lectularius was most tolerant of deltamethrin during the late photophase at ZT9 (i.e. nine hours after light is present in the light-dark cycle (LD) cycle) and similarly repeated at CT9 (i.e. nine hours into the subjective day in constant darkness (DD)) suggesting endogenous circadian involvement in susceptibility to deltamethrin. No diel rhythm was observed against imidacloprid insecticide despite significant daily susceptibility in both LD and DD conditions. Rhythmic expressions of metabolic detoxification genes, GSTs1 and CYP397A1 displayed similar expression patterns with total GST and P450 enzyme activities in LD and DD conditions, respectively. The oscillation of mRNA levels of GSTs1 and CYP397A1 was found consistent with peak phases of deltamethrin susceptibility in C. lectularius. This study demonstrates that circadian patterns of metabolic detoxification gene expression occur within C. lectularius. As a consequence, insecticide efficacy can vary dramatically throughout a 24 hour period.

Bed bugs (Cimex lectularius) are blood-feeding insects poised to become one of the major pests in households throughout the United States. Resistance of C. lectularius to insecticides/pesticides is one factor thought to be involved in its sudden resurgence. Despite its high-impact status, scant knowledge exists at the genomic level for C. lectularius. Hence, we subjected the C. lectularius transcriptome to 454 pyrosequencing in order to identify potential genes involved in pesticide resistance. Using 454 pyrosequencing, we obtained a total of 216,419 reads with 79,596,412 bp, which were assembled into 35,646 expressed sequence tags (3902 contigs and 31744 singletons). Nearly 85.9% of the C. lectularius sequences showed similarity to insect sequences, but 44.8% of the deduced proteins of C. lectularius did not show similarity with sequences in the GenBank non-redundant database. KEGG analysis revealed putative members of several detoxification pathways involved in pesticide resistance. Lamprin domains, Protein Kinase domains, Protein Tyrosine Kinase domains and cytochrome P450 domains were among the top Pfam domains predicted for the C. lectularius sequences. An initial assessment of putative defense genes, including a cytochrome P450 and a glutathione-S-transferase (GST), revealed high transcript levels for the cytochrome P450 (CYP9) in pesticide-exposed versus pesticide-susceptible C. lectularius populations. A significant number of single nucleotide polymorphisms (296) and microsatellite loci (370) were predicted in the C. lectularius sequences. Furthermore, 59 putative sequences of Wolbachia were retrieved from the database. To our knowledge this is the first study to elucidate the genetic makeup of C. lectularius. This pyrosequencing effort provides clues to the identification of potential detoxification genes involved in pesticide resistance of C. lectularius and lays the foundation for future functional genomics studies.

The global population growth of the bed bug, Cimex lectularius (L.), is attributed to their cryptic behavior, diverse insecticide resistance mechanisms, and lack of public awareness. Bed bug control can be challenging and typically requires chemical and non-chemical treatments. One common non-chemical method for bed bug management is thermal remediation. However, in certain instances, bed bugs are known to survive heat treatments. Bed bugs may be present after a heat treatment due to (i) abiotic factors associated with the inability to achieve lethal temperatures in harborage areas for a sufficient time period, (ii) re-infestation from insects that escaped to cooler areas during a heat treatment or (iii) development of physiological resistance that allows them to survive heat exposure. Previous research has investigated the optimal temperature and exposure time required for either achieving complete mortality or sublethally affecting their growth and development. However, no research has examined bed bug populations for their ability to develop resistance to heat exposure and variation in thermo-tolerance between different bed bug strains. The goals of this study were: i) to determine if bed bugs could be selected for heat resistance under a laboratory selection regime, and ii) to determine if bed bug populations with various heat exposure histories, insecticide resistance profiles, and geographic origins have differential temperature tolerances using two heat exposure techniques (step-function and ramp-function). Selection experiments found an initial increase in bed bug survivorship; however, survivorship did not increase past the fourth generation. Sublethal exposure to heat significantly reduced bed bug feeding and, in some cases, inhibited development. The step-function exposure technique revealed non-significant variation in heat tolerance between populations and the ramp-function exposure technique provided similar results. Based on these study outcomes, the ability of bed bugs to develop heat resistance appears to be limited.

Bed bugs (Cimex lectularius L.) (Hemiptera: Cimicidae) are obligate hematophagous ectoparasites, and, therefore, must locate suitable hosts to ensure survival and reproduction. Their largely nocturnal activity suggests that chemosensory and thermosensory cues would play critical roles in host location. Yet, the importance of olfaction in host attraction of bed bugs remains unclear. We developed and validated a Y-tube, two-choice olfactometer and tested its suitability for investigating attraction to human odors (from skin swabs). Olfactometer orientation significantly affected the percentage of bed bugs that were activated by human odors, with significantly more bed bugs responding when the olfactometer was oriented vertically (bug introduced at bottom of the olfactometer) compared with all other orientations. Starved (7–10 d) adult males, mated females, and nymphs responded (47–77% moved up the olfactometer and made a choice) when human odors were present in the olfactometer, while starved, unmated females did not respond. Skin swabs from all five human participants elicited high response rates (65–82%), and bed bugs from four different populations responded to skin swabs (40–82% response rate). However, in all assays including those resulting in relatively low response rates, bed bugs exhibited >90% preference for human odors over blank controls. These results provide strong evidence that bed bugs can respond and orient towards human odors, independently of all other host cues. Furthermore, the validated olfactometer should enable rapid and efficient evaluations of bed bug behavioral responses to semiochemicals.