Sex pheromone titre in the glands of Spodoptera litura females: circadian rhythm and the effects of age and mating

The present study investigates the effects of age and mating status on the circadian variations of gland sex pheromone titre in female Spodoptera litura Fabricius. Similar to other nocturnal moths, S. litura females exhibit circadian variations of gland sex pheromone contents, with higher levels during scotophase and lower levels during photophase. The sex pheromone titre in the glands peaks during the first scotophase after eclosion and sharply declines afterwards. Higher pheromone contents during scotophase may facilitate female reproductive activities, and the negative relationship between pheromone titre and female calling is likely the result of pheromone release during female calling. Interestingly, the present study demonstrates that mated S. litura females have significantly higher sex pheromone titre in their pheromone glands (PGs) than virgin females. This finding contrasts with all previous studies of other insect species, in which mating generally reduces the sex pheromone titre in female PGs. In S. litura, mating and male accessory gland fluids can suppress female calling behaviours and re‐matings. These results suggest that the suppression of female calling behaviours by mating and male accessory gland fluids may significantly reduce the release of sex pheromones and thus result in higher sex pheromone titre in the PGs of mated females.     

Sublethal effects of permethrin on the chemical communication system of the pink bollworm moth,Pectinophora gossypiella

Topically applied sublethal doses of permethrin can interrupt chemical communication between the sexes of Pectinophora gossypiella by affecting both the signaler and the responder. The probability of calling by females is reduced when they are treated with doses of permethrin which are much less than the LD₅₀. Similarly, key stages in the behavioral response of males to sex pheromone are effectively blocked at these low doses. Males recover from these effects 4 days after treatment, but calling by females is still significantly reduced at this time. Chemical control of P. gossypiella populations with permethrin may not be limited to mortality, and potentially includes effective control of behavioral aspects of chemical communication.     

光强度对棉铃虫交配行为的影响

【目的】为阐明光强度对棉铃虫Helicoverpa armigera (Hübner)交配行为的影响。【方法】本实验设置4个光照强度(0,0.5,5.0和50.0 lx),观察记录不同光照强度下棉铃虫雌蛾的求偶行为;分别通过单个腺体性信息素提取法和解剖雌虫受精囊的方式,检测不同光照强度下棉铃虫的雌蛾性信息素滴度和交配率。【结果】在强光(50.0 lx)下,棉铃虫雌蛾求偶起始时间最长,求偶持续时间最短,求偶次数最少,雌蛾性信息素滴度始终处于较低水平,交配率也属于最低。在微光(0.5 lx)下,雌蛾求偶起始时间最短,求偶持续时间最长,求偶次数最多;雌蛾性信息素始终处于较低水平,但暗期后段求偶率高达40%。【结论】强光(50.0 lx)可以抑制棉铃虫雌蛾的求偶行为、性信息素的合成及交配;微光(0.5 lx)可以促进棉铃虫雌蛾的求偶行为;相对黑暗环境(0 lx),微光(0.5 lx)还可以促进棉铃虫快速(1 h)完成交配,微光(0.5 lx)对棉铃虫的交配行为具有重要意义;求偶和性信息素的合成没有必然联系。本研究可为探讨光对夜蛾交配行为的影响提供一定的理论基础,也可为利用物理、化学通讯信息调控夜蛾行为提供一定的参考。     

Long‐term rearing affects pheromone‐mediated flight behaviour of the Indian meal moth,Plodia interpunctella

The pest Plodia interpunctella (Hübner) is reared in many research laboratories. In a culture established in 1996, attraction of males to the female‐produced sex pheromone in flight tunnel assays gradually decreased after ≈15 years of rearing. A new culture was established to enable comparison with the old culture regarding traits associated with mate finding. Female calling activity, pheromone titre and male antennal response to pheromone components did not differ between cultures. In contrast, very few males from the old culture reached the pheromone source in flight tunnel assays compared with 61%–81% of males from the other culture. Our results highlight the importance of maintaining viable insect cultures for research purposes and suggest frequent evaluation of traits involved in chemical communication in such cultures to ensure reliable results in experiments.     

The sublethal effects of tebufenozide on the precopulatory and copulatory activities of Choristoneura fumiferana and C. rosaceana

The sublethal effects of tebufenozide, an ecdysone agonist, on the reproductive biology of Choristoneura fumiferana (Clem) and of Choristoneura rosaceana (Harris) (Lepidoptera: Tortricidae), treated during the larval stage, were evaluated using two treatment methods: the force‐feeding method and the diet method. The percentage of mortality and the developmental time of survivors increased linearly with the concentration of tebufenozide used. This ecdysone analogue proved to be more toxic to C. fumiferana than to C. rosaceana. In C. rosaceana, the weight of males and females decreased proportionally with the dose ingested, but females were affected to a greater extent. This difference might be due to a greater consumption of the treated diet, or to a differential vulnerability to tebufenozide. Tebufenozide did not modify the pre‐copulatory activities associated with chemical communication in the females. However, the consumption of tebufenozide delayed ovarian maturation, causing a reduction in the fecundity of females. Treated males had smaller spermatophores and fewer eupyrene sperms in their bursa copulatrix and spermatheca, along with lower mating success. In C. fumiferana, tebufenozide delayed the females’ onset time of calling the first night after emergence, but did not affect the mean time spent calling or the production of the main component of the sex pheromone. The males showed significantly greater difficulty in executing oriented flight in a wind tunnel, although their mating success was not affected. We concluded that tebufenozide interferes with various aspects of the reproductive biology of males and females of C. fumiferana and C. rosaceana, including some pre‐copulatory behaviors associated with sex pheromone communication.     

Intraspecific acoustic communication and mechanical sensitivity of the tympanal ear of the gypsy moth Lymantria dispar

Tympanal ears of female gypsy moths Lymantria dispar dispar (L.) (Lepidoptera: Erebidae: Lymantriinae) are reportedly more sensitive than ears of conspecific males to sounds below 20 kHz. The hypothesis is tested that this differential sensitivity is a result of sex‐specific functional roles of sound during sexual communication, with males sending and females receiving acoustic signals. Analyses of sounds produced by flying males reveal a 33‐Hz wing beat frequency and 14‐kHz associated clicks, which remain unchanged in the presence of female sex pheromone. Females exposed to playback sounds of flying conspecific males respond with wing raising, fluttering and walking, generating distinctive visual signals that may be utilized by mate‐seeking males at close range. By contrast, females exposed to playback sounds of flying heterospecific males (Lymantria fumida Butler) do not exhibit the above behavioural responses. Laser Doppler vibrometry reveals that female tympana are particularly sensitive to frequencies in the range produced by flying conspecific males, including the 33‐Hz wing beat frequency, as well as the 7‐kHz fundamental frequency and 14‐kHz dominant frequency of associated clicks. These results support the hypothesis that the female L. dispar ear is tuned to sounds of flying conspecific males. Based on previous findings and the data of the present study, sexual communication in L. dispar appears to proceed as: (i) females emitting sex pheromone that attracts males; (ii) males flying toward calling females; and (iii) sound signals from flying males at close range inducing movement in females, which, in turn, provides visual signals that could orient males toward females.     

DIEL RHYTHMS OF CALLING BEHAVIOR AND SEX PHEROMONE PRODUCTION OF BEET ARMYWORM,SPODOPTERA EXIGUA (LEPIDOPTERA:NOCTUIDAE)*

Abstract Beet armyworm (BAW), Spodoptera exigua, is becoming one of more and more serious pests in China in recent years. As a part of research program of sex pheromone and its application of BAW in China, the hourly and daily variation of calling behavior and pheromone production of BAW females were investigated. Both calling behavior and titers of 4 sex pheromone components showed distinct diel rhythms, and the two peak periods were synchronous. In comparison, the calling activity lasted shorter period of time with a longer peak time, whereas the production of the sex pheromone lasted throughout the whole scotophase and part of the photophase with a very short peak time. The calling behavior began at the middle scotophase, reached the maximum at the middle‐later scotophase, and continued the maximal calling activity until the end of the scotophase. When the light was on, the calling percentage reduced sharply, and all females stopped calling 1 hour later. The variation patterns of the 4 pheromone components in the glands of the 3 day old moths were similar from one to another. From 0.5 h before to 4.5 h into scotophase, the titers increased slightly, but at 6.5 h they showed a significant increase up to the peak values of the scotophase. Soon at 8.5 h into the scotophase, they decreased significantly and thereafter gradually to undetectable level at 4.5 h into the next photophase. The daily change experiment showed that BAW females began calling at 0 scotophase, became fully active in calling at 3rd scotophase, and maintained the calling activity to 7th scotophase. There was no significant difference in pheromone titers among different day‐old moths.     

Sex pheromones and trail‐following pheromone in the basal termites Zootermopsis nevadensis (Hagen) and Z. angusticollis (Hagen) (Isoptera: Termopsidae: Termopsinae)

In the context of an evolutionary study of the chemical communication in termites, sex pheromones and trail‐following pheromones were investigated in two Termopsidae, Zootermopsis nevadensis and Z. angusticollis. In these species, in which the presence of sex‐specific pheromones has been demonstrated previously, the chemical structure of the female sex pheromone has now been identified as (5E)‐2,6,10‐trimethylundeca‐5,9‐dienal and the male sex pheromone as (+)‐ or (?)‐syn‐4,6‐dimethyldodecanal. The amount of sex pheromone was estimated at 5–10 ng per individual in females and 2–5 ng in males. Because these two sympatric species do not differ in their pheromonal chemical composition, reproductive isolation is probably mediated chiefly by differences in dispersal flight chronology. The trail‐following pheromone was shown to be composed of the same compound as the male sex pheromone, that is syn‐4,6‐dimethyldodecanal. The compound syn‐4,6‐dimethyldodecanal was 10 times more active than the racemic (+/?)‐syn + (+/?)‐anti‐4,6‐dimethyldodecanal in eliciting trail‐following. The amount of syn‐4,6‐dimethyldodecanal was estimated at 0.1–0.5 ng per pseudergate. Regarding the phylogenetic aspects, the nature of the female sex pheromone of Zootermopsis is structurally akin to the trail‐following pheromone of Mastotermes darwiniensis of Mastotermitidae and Porotermes adamsoni and Stolotermes victoriensis of Termopsidae. Interestingly, the nature of the trail‐following pheromone of the Termopsinae Zootermopsis is clearly different from that of the Porotermitinae P. adamsoni and the Stolotermitinae S. victoriensis, which mirrors recent molecular data on the paraphyly of Termopsidae. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100 , 519–530.     

Calling behavior of the female german cockroach,Blattella germanica (Dictyoptera: Blattellidae)

Virgin German cockroach females, Blattella germanica(L.), were observed, for the first time, to exhibit a characteristic calling behavior during which females emit a volatile sex pheromone. Under a photoperiod of 12L12D, the percentage of 7-day-old virgin females that exhibited this behavior peaked before the end of the scotophase in a similar pattern to the diel periodicity of mating. A clear relationship was evident between calling and stages of sexual receptivity during successive gonotrophic cycles. Females initiated calling 5–6 days after the imaginal molt, when their basal oocytes were 1.6 mm long. If not mated, females continued to exhibit bouts of calling during the next 3–4 days until 24 h before ovulation. Calling was completely suppressed by mating as well as the presence of an egg case in the genital atrium in both virgin and mated gravid females. We suggest that calling and the emission of a volatile sex pheromone serve to attract males from a distance as well as to potentiate responses to contact sex pheromone in aggregations.     

Sublethal effects of chlordimeform on chemical communication and other reproductive behaviors in the female cabbage looper moth (Lepidoptera: Noctuidae)

Insecticides can affect the complex coordination of activities associated with reproduction through their sublethal impact on the nervous system. Our objective was to document the effects of a sublethal dose (1% mortality) of chlordimeform on reproductive events in the female cabbage looper moth, Trichoplusia ni. A significantly higher percentage of females treated with chlordimeform initiated calling at some time during the scotophase. Chlordimeform stimulated pheromone emission early in the scotophase. However, late in the scotophase pheromone emission was significantly lower in chlordimeform-treated females when compared with control females. Overall, a lower amount of pheromone was detected in glands of chlordimeform-treated females. This effect was significant only during the second half of the scotophase. We suggest that chlordimeform stimulates pheromone transport to the gland surface and calling behavior resulting in depletion of pheromone reserves over the course of the scotophase. Chlordimeform also decreased the mating success of males. Oviposition and egg hatch were also affected by chlordimeform. Mated females treated with chlordimeform laid significantly fewer eggs than acetone-treated females. In addition, hatchability of eggs laid by mated female T. ni treated with chlordimeform was significantly lower than for eggs laid by control females.     

Diel periodicity and influence of age and mating on female sex pheromone titre in Estigmene acrea (Lep., Arctiidae)

Abstract:  Calling behaviour, diel periodicity, and effect of age and mating on female sex pheromone titre in Estigmene acrea (Drury) were studied under laboratory conditions. Forty-five per cent of females started calling during the first scotophase, but the highest number of calling females was observed during the second, third and fourth scotophases. Calling behaviour occurred from the third hour after dark until just before the end of the scotophase. However, females exhibited a bimodal pattern of calling with the first peak occurring between 4 and 6 h and a second peak at 10 h after the onset of scotophase. The mean onset of calling time differed significantly with age. Older females showed a tendency to call longer, but there was no significant difference. The amount of (Z,Z)-3,6-cis-9, 10-epoxyheneicosadiene in females was quantified from the first scotophase following emergence, until the fifth scotophase. Glands of 0-day-old females presented a higher content of pheromone compared with that found in glands of 1-, 2-, 3- and 4-day-old females. Pheromone titre was determined at 2-h intervals throughout the third scotophase and photophase. (Z,Z)-3,6-cis-9,10-epoxyheneicosadiene was found in the gland during the scotophase as well as the photophase. However, there was no consistent pattern of pheromone production throughout the scotophase or photophase. Mated females of E. acrea produced significantly less pheromone than virgin females.     

Temporal distribution of PBAN‐like immunoreactivity in the hemolymph of Mamestra brassicae females in relation to sex pheromone production and calling behavior

A direct enzyme‐linked immunosorbent assay has been developed and applied to the analysis of PBAN immunoreactivity in female hemolymph of the cabbage armyworm, Mamestra brassicae. PBAN‐IR determinations have been carried out with third scotophase insects at different times of the photoperiod. The rhythm of calling and the pattern of pheromone production by third scotophase females at different times of the photoperiod have also been determined. PBAN‐IR and calling are well correlated. However, whereas pheromone titers decrease, both PBAN‐IR levels and percentage of calling females remain high in the last hours of the scotophase. These results are discussed in the context of the regulation of sex pheromone biosynthesis in M. brassicae. Arch. Insect Biochem. Physiol. 40:80–87, 1999. © 1999 Wiley‐Liss, Inc.     

The effects of mating,exogenous juvenile hormone and a juvenile hormone analogue on pheromone titre,calling and oviposition in the omnivorous leafroller moth (Platynota stultana)

Mating in Platynota stultana resulted in the termination of calling, the gradual reduction of pheromone in the pheromone glands to non-detectable levels (<0.1 ng/♀) within 14 h, and oviposition of the first batch of eggs 20–24 h after copulation. Decapitation of virgin females resulted in a similar decline in pheromone titre, and also eliminated oviposition and calling. Pheromone production appears to be controlled via the head. Mating probably terminates neural or hormonal input required for pheromone production and/or removes neural or hormonal inhibition of pheromone degradation. A juvenile hormone analogue (ZR-512) and juvenile hormones I, II and III applied exogenously to virgin females elicited oviposition comparable to mated females and terminated calling within 48 h. The juvenile hormone analogue also appeared to block pheromone production in virgin females. These results suggest that juvenile hormone may be involved in the switch from virgin to mated behaviour in this species.     

Determination of the amount of sex pheromone emitted by individual virgin females of the white grub beetle, <Emphasis Type="Italic">Dasylepida ishigakiensis</Emphasis> (Coleoptera:Scarabaeidae) at different stages of reproductive life

Virgin females of the white grub beetle, Dasylepida ishigakiensis Niijima et Kinoshita, emit the sex pheromone, 2-butanol. To investigate the relationships between pheromone emission and ovarian development in this beetle, virgin adult females were kept at 18°C for various lengths of time. The sex pheromone released by individual females at first calling was collected at 23°C and the amounts determined. Females started developing visible oocytes at day 44 and emitting sex pheromone at day 50. Almost all females at day 52 or later possessed eggs, indicating that diapause had ended. A high positive correlation was found between the amounts of pheromone emitted and levels of ovarian development. A large variation of pheromone emission was encountered even among individuals of the same age. Females retained the ability to emit sex pheromone as long as they were virgin. These results may provide important information regarding the timing and duration of application of synthetic sex pheromone in an attempt to control this pest through disrupting the male-female communication.     

Sex pheromone production and perception in the mating disruption-resistant strain of the smaller tea leafroller moth, Adoxophyes honmai

Mating disruption is an environmentally safe plant protection strategy that uses a synthetic copy of an insect pheromone to interfere with sexual communication and hence reproduction. To date, a number of pest moths have been controlled with applications of formulated pheromones as mating disruptants. Recently, however, the first example of resistance to mating disruption was documented in one of the major tea pests in Japan, the smaller tea leafroller moth, Adoxophyes honmai Yasuda (Lepidoptera: Tortricidae). To avoid other such cases, it is important to elucidate the mechanism(s) by which the disruptant lost its effectiveness. To this end, we imposed further selection by rearing field‐collected resistant insects with a synthetic pheromone in the laboratory. After more than 70 generations of selection, a strain with quite strong resistance was established, males of which could find and copulate with their mates even in the presence of 1 mg l^?1 of disruptant. Although the mating ability of this strain was greatly increased, the composition and blend ratio of the sex pheromone produced and emitted by females were not obviously changed in comparison with those of females sensitive to mating disruption. However, male response to the pheromone blend was markedly broadened after selection so that resistant males could locate a synthetic pheromone source even when it lacked a pheromone component that is normally necessary for attraction. Males capable of locking onto off‐ratio pheromone blends may be better able to find calling females in pheromone‐treated environments than narrowly tuned males because of greater capability of overcoming sensory imbalance.     

Conspecific females promote calling behavior in the noctuid moth,Pseudaletia adultera

Changes in female calling behavior in response to the presence of conspecific pheromones (pheromone autodetection) have been demonstrated in a number of moth species. However, the observed changes vary between species, and several ecological and adaptive explanations for autodetection have been proposed. We studied the effect of conspecific females on the calling behavior of the noctuid moth Pseudaletia adultera (Schaus) (Lepidoptera: Noctuidae, Hadenini), by comparing the age of first calling, as well as the onset and pattern of calling, when females were held individually or in the presence of conspecifics. Grouped females 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. We also demonstrated that female antennae respond to each of the three main components of the sex pheromone – (Z)‐11‐hexadecen‐1‐ol, (Z)‐11‐hexadecen‐1‐yl acetate, and (Z)‐11‐hexadecenal – and that the response patterns differed from those of male antennae. By calling more and extending her calling window in presence of conspecific females, a female may increase her chances of accessing males. However, the potential benefits need to be considered within an ecological context, considering factors such as migration, oviposition, and foraging.     

枣镰翅小卷蛾性信息素通讯系统

采用单个雌蛾性信息素腺体分析技术对枣镰翅小卷蛾Ancylis sativa Liu雌蛾性信息素的组分和组分间的精确比例进行了测定,结果表明,枣镰翅小卷蛾雌蛾的性信息素系统由二个顺反异构体组分组成,即反-9-十二碳烯醋酸酯(E9-12∶Ac)和顺-9-十二碳烯醋酸酯(Z9-12∶Ac)组成,E9-12∶Ac与Z9-12∶Ac的比例为6.5∶3.5;雌蛾产生和释放性信息素具有时辰节律性,在光周期14L∶10D、温度为21℃时,性信息素产生的高峰期为进入黑暗期6.5 h;不同日龄雌蛾产生的性信息素有差异,1日龄最低,2日龄最高,3～5日龄居中,不同日龄雌蛾产生的性信息素组分间的比例无显著差异;对3个世代雌蛾产生的性信息素的量及组分间的比例的研究表明,越冬代含量最高,为(10.1±7.0)ng/头,第2代为(9.5±4.6)ng/头,第1代仅为(1.4±1.0)ng/头,3个世代性信息素组分间的比例无显著差异,在63.1%～64.3%(E)范围内。     

Attractiveness and competitiveness of irradiated light brown apple moths

The sterile insect technique (SIT) potentially provides a socially acceptable approach for insect eradication of new pest incursions. The light brown apple moth, Epiphyas postvittana (Walker) (Lepidoptera: Tortricidae), was discovered in Berkeley (CA, USA) in 2006, leading to an incursion response that included this technology. In this study, we assessed factors affecting mating success from a bisex release of irradiated moths: effects of radiation dose on male multiple mating, male flight competition, female sex pheromone titre and attractiveness of irradiated females to males, and identification of successful mating in vineyards of either irradiated or wild males (identified by isotope analysis of spermatophores from sentinel females). There was a significant negative relationship between male radiation dose and mating frequency. In head‐to‐head flights of irradiated males against non‐irradiated males to a pheromone lure in a wind tunnel, irradiated males reached the lure first only 31% of the time. With increasing radiation dose, the production of the major sex pheromone component in females, (E)‐11‐tetradecenyl acetate, dropped, from 0.7 ± 0.1 ng per female in non‐irradiated females to 0.2 ± 0.07 ng per female when irradiated at 300 Gy. Male catch was reduced to 11% of control females in traps containing females irradiated at 300 Gy. Isotope analysis of spermatophores found in the bursa copulatrix of females indicated that mating success of irradiated males inside the live (entry‐only) traps containing virgin females was lower (13.1 ± 3.3%) than suggested by male catch (21.2 ± 3.8%) in pheromone traps, the current standard for assessing field competitiveness. Impacts of irradiation on male and female moth fitness should be taken into account to improve estimates of irradiated to wild male E. postvittana overflooding ratios needed for population suppression.     

Male position and calling effort together influence male attractiveness in leks of the medfly,Ceratitis capitata (Diptera: Tephritidae)

Despite the close association between lek mating systems and the study of female mate choice, male mating success in leks is often associated with other aspects of sexual selection as well as female choice of male display traits. Males of the medfly Ceratitis capitata form leks on the undersides of leaves of their host plants. By experimentally creating artificial leks, we show that male success at attracting females depends not only on male calling effort (pheromone production dispersed by wing movement), but also on the position of the male within a lek. Males in the highest position in the artificial lek (closest to the light) attracted more females, and received more visits from those females. In our experiment, we deliberately minimized the visual cues that females approaching a male could use and, under these conditions, found no associations between male attractiveness and male size, weight or fluctuating asymmetry, either of the wings or sex setae (a pair of bilateral supra‐fronto‐orbital bristles). The latter result contrasts with earlier studies showing a significant negative association between sex setae fluctuating asymmetry and mating success. Accordingly, we conclude that symmetry of the male sex setae has no role in nonvisual communication (e.g. through directing pheromone plumes). Mating patterns associated with this insect are therefore every bit as complex as those in vertebrate leks. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 95 , 479–487.     

蜀柏毒蛾生殖行为及性信息素产生与释放节律

为了探索蜀柏毒蛾Parocneria orienta Chao性信息素产生和释放规律, 为利用性信息素监测和防治蜀柏毒蛾奠定基础, 本研究在野外及室内温度22±1℃、 相对湿度75%~80%、 光周期14L∶10D条件下观察研究了蜀柏毒蛾成虫的羽化、 求偶、 交尾、 产卵行为, 触角电位反应测定处女雌蛾性信息素产生与释放的时辰节律。结果表明： 蜀柏毒蛾羽化行为全天可见, 主要集中在1:00-5:00, 占总羽化量的44.94%, 7:30-11:00进行婚飞和交尾, 交尾高峰期出现在8:30左右, 交配时间少则2 h, 多则8 h, 求偶、 交配均发生在光期。随着日龄的增加, 召唤时间前移并且延长, 1日龄的处女雌蛾交尾时间较短; 雌蛾羽化当天就可交尾, 2日龄雌蛾交尾率最高, 达36.67%。雌蛾分多处产卵, 雌蛾一生最高产卵量达402粒, 最低产卵量为78粒。羽化当天的雌蛾体内性信息素含量较低, 第2天最高, 以后逐日下降; 2日龄蜀柏毒蛾处女雌蛾性信息素的产生量从7:00起逐渐增加, 8:30-9:30时最高, 9:30后逐渐减小。雄蛾对处女雌蛾腺体提取物的触角电位反应在8:30-9:00最强, 说明8:30-9:00是雌蛾产生和释放性信息素的高峰期。蜀柏毒蛾的羽化、 求偶、 交尾及性信息素的产生与释放存在一定的时辰节律, 野外处女雌蛾诱蛾试验证实了性信息素释放与交配行为在时辰节律上的一致性。