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Activity rhythms involved in pheromone communication of spruce budworm Choristoneura fumiferana represented as normal curves with mean time (X) and SD. A Emergence from pupae (Kipp et al. 1995); BE11-14:Al pheromone titer of female gland (Delisle et al. 1999); C calling females (Delisle et al. 1999); D emission of E11-14:Al pheromone from female (Morse et al. 1982); and E male catch on virgin females in high-density population (Kipp et al. 1995). Calculation of R² and coefficients for corresponding normal curves explained in text

Activity rhythms involved in pheromone communication of spruce budworm Choristoneura fumiferana represented as normal curves with mean time (X) and SD. A Emergence from pupae (Kipp et al. 1995); BE11-14:Al pheromone titer of female gland (Delisle et al. 1999); C calling females (Delisle et al. 1999); D emission of E11-14:Al pheromone from female (Morse et al. 1982); and E male catch on virgin females in high-density population (Kipp et al. 1995). Calculation of R² and coefficients for corresponding normal curves explained in text

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Many insect species have circadian rhythms of pheromone production/titer, calling, emission, and response that are involved in intraspecific communication and impact pest management practices. Rhythms of pheromone biosynthesis, most studied in moths affecting forestry and agriculture, contribute to a periodicity of pheromone concentration or titer...

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... Virtually all life on the Earth experiences a 24 h circadian rhythm, which affects almost all behaviors, including sexual activity and mating. 1 Insects are no exception, with most species hitherto studied displaying circadian rhythm of mating behavior, sex pheromone titer, calling, and pheromone emission. 2 One notable exception to this chronobiology rule is the large black chafer Holotrichia parallela (Coleoptera: Scarabaeidae: Melolonthinae). As a result of a 4-year study, it was reported for the first time that the large black chafer (then named Lachnosterna morosa) remains in the soil during the daytime, coming to the surface every other evening soon after sunset. ...
... However, whether the ''every other day'' periodicity in H. parallela was formed by an endogenous, free-running, circabidian rhythm remained speculative until recently. In nature, cycles are synchronized by exogenous environmental cues (time givers, also known as zeitgebers), 2 and there are no known 2-day zeitgebers. Recently, Kawasaki and colleagues 7 elegantly addressed the 48 h periodicity in H. parallela through a comprehensive chronobiological study and confirmed that under 12 h light (photophase) and 12 h dark (scotophase), H. parallela appeared on the ground at the beginning of the scotophase every 2 days. ...
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Article Circabidian rhythm of sex pheromone reception in a scarab beetle Graphical abstract Highlights d HparOR14 is required for sex communication in the scarab beetle H. parallela d AlphaFold models and docking predict possible ligand-binding sites in HparOR14 d Transcripts of the sex pheromone gene exhibit an unusual 48 h (circabidian) rhythm d Male antennal responses to the sex pheromone are abolished in non-calling days In brief Wang et al. through studies on the scarab beetle Holotrichia parallela, discover the first sex pheromone receptor in Coleoptera. They also demonstrate that expression of the receptor gene exhibits an unusual 48 h periodicity, consistent with the male's electrophysiological responses to the sex pheromone. SUMMARY Animals have endogenous clocks that regulate their behavior and physiology. These clocks rely on environmental cues (time givers) that appear approximately every 24 h due to the Earth's rotation; thus, most insects exhibit a circadian rhythm. One notable exception is the scarab beetle, Holotrichia parallela, a severe agricultural pest in China, Japan, South Korea, and India. Females emerge from the soil every other night, reach the canopy of host plants, evert an abdominal gland, and release a pheromone bouquet comprising L-isoleucine methyl ester (LIME) and L-linalool. To determine whether this circa'bi'dian rhythm affects the olfactory system , we aimed to identify H. parallela sex pheromone receptor(s) and study their expression patterns. We cloned 14 odorant receptors (ORs) and attempted de-orphanizing them in the Xenopus oocyte recording system. HparOR14 gave robust responses to LIME and smaller responses to L-linalool. Structural modeling, tissue expression profile, and RNAi treatment followed by physiological and behavioral studies support that HparOR14 is a sex pheromone receptor-the first of its kind discovered in Coleoptera. Examination of the HparOR14 transcript levels throughout the adult's life showed that on sexually active days, gene expression was significantly higher in the scotophase than in the photophase. Additionally, the HparOR14 expression profile showed a circabidian rhythm synchronized with the previously identified pattern of sex pheromone emission. 48 h of electroantennogram recordings showed that responses to LIME were abolished on non-calling nights. In contrast, responses to the green leaf volatile (Z)-3-henexyl acetate remained almost constant throughout the recording period.
... In laboratory settings, sex dimorphism aids in distinguishing living adult insects for volatile collection and behavioral bioassays, while diel activity assists in conducting these bioassays effectively (Zarbin et al. 1999(Zarbin et al. , 2009Ambrogi and Zarbin 2008) . It is well established that pheromone production and bioactivity are influenced by specific photoperiods (Fonseca et al. 2010;Levi-Zada and Byers 2021). Ultimately, the information obtained from our research may serve as a valuable tool in the development of an integrated management program for cassava insect pests in Brazil. ...
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... To attract a mate, moth females of most species produce and release sex pheromones from specific glands at the tip of the abdomen during calling behavior (Valles et al. 1992;Pires et al. 1994;Foster and Anderson 2022). Calling behavior and pheromone production are regulated by several factors, mainly, photoperiod, environmental temperature, conspecifics females calling, and female age Soofbaf et al. 2007;Kawazu et al. 2011;Rehermann et al. 2016;Levi-Zada and Byers 2021). For example, Pseudaletia adultera (Schaus) (Lepidoptera: Noctuidae) females in the presence of conspecifics started calling at a lower age, a higher percentage of females called during the scotophase, and they called longer compared to females held in isolation (Rehermann et al. 2016). ...
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... Regardless of the outcome of these experiments, our data (Fig. 3) indicate that female S. grossa do manipulate their webs' attractiveness in response to perceived mate competition. These results are intriguing because, to date, timed pheromone production and dissemination are known only in insects 62 . Although we demonstrate that female spiders adjust the pheromone titer on their webs, and thus their webs' attractiveness, according to perceived social context, we did not experimentally demonstrate but only infer the preferential attraction of males to webs constructed in high-web-density settings. ...
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... The central clock that drives rhythmic behavior is usually located in the brain 21 . The biological clock plays important roles in the regulation of insect activities, including sex pheromone production 22 , mating 23,24 and egg laying 25,26 . Within the intestine, circadian rhythms can alter the gut microbe composition 27,28 . ...
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... Therefore, the sex pheromone communication behavior of both sexes has obvious circadian rhythm characteristics. 2,8 For example, before mating, female moths biosynthesize and release sex pheromones rhythmically through pheromone glands (PGs) at the end of the abdomen. 9 The calling rhythm of female Spodoptera littoralis is circadian, persistent for at least 4 days once it had been entrained, and regulated by light. ...
... Calling activity can reach its highest value in the scotophase at a specific time. 2,8 Some studies have also confirmed that light, temperature, and chemical signals can regulate the expression of circadian clock genes and sexual communication behavior in moths. 6,36 However, it is unclear whether Per genes can regulate the sex pheromone titer and calling behavior of female moths. ...
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... Chemicals with ester functional groups such as species-specific pheromones, host plant odorants, and xenobiotic insecticides are important for insect physiology, biochemistry, and ecology (Xu and Turlings, 2018;Levi-Zada and Byers, 2021;Mdeni et al., 2022). Carboxylesterases (CXEs) belong to a superfamily that includes diverse functional enzymes (Oakeshott et al., 1999;Oakeshott et al., 2005). ...
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... However, during other hourly periods, one or fewer male adults (with a proportion of less than 1.5%) observed responding to the sex pheromone lures could be random rather than intentional. Tuta absoluta male adults typically responded to female cues at dawn and in the early morning (peaking at 1 h after sunrise); the courtship behavior of female adults likely occurs synchronously, while mating behaviors should begin after or at the same time [32,41,42]. ...
... A higher proportion of male than female adults (p = 0.001) was detected in the second half of the night (from 01:30 to 06:30), which could be related to the following (i.e., 06:30-07:30) high proportion response to the sex pheromone traps. The lower proportion of female adults (p = 0.001) observed in the same period of time might be related to sex pheromone production or biosynthesis in the so-called female "calling" behavior [42]. ...
... Courtship, mating, and oviposition are the most important life activities of adult sexually reproducing insects. Related behaviors, especially the response to sex pheromone lures (or female cues) [42] and flying toward light traps [22,38], vary between species, even between those in the same family, e.g., the potato tuberworm moth, Phthorimaea operculella (Lepidoptera: Gelechiidae) [45]. Males responding to sex pheromone (false female) lures and male and female adults flying toward light traps displayed obvious diel rhythms. ...
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... Responses to these chemical signals vary according to environmental factors, previous experience, physiological status, or combinations of all of these [1]. Attraction to chemical stimuli has been the subject of study for its application in pest monitoring and control [2][3][4][5][6][7][8][9]. Pheromone attractants, speciesspecific by definition, may function with sexual or physiologic specificity to aid detection, population monitoring, and eradication programs [10]. ...
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