REGULATION OF FLIGHT CAPACITY AND CONTENTS OF ENERGY SUBSTANCES BY METHOPRENE IN THE MOTHS OF ORIENTAL ARMYWORM,MYTHIMNA SEPARATA*

Abstract The flight capacity and contents of energy substances in the moths of migratory Oriental armyworm, Mythimna separata (Walker)dosed with juvenile hormone analogue methoprene(JHM) were compared with the normal moth through day‐1 to day‐6 after the treatment. The flight capacity of JHM‐treated moths, as determined by a 12‐hours tethered‐flight test, was significantly lower than that of the controls, although it might differ with their age and sex. The moth's flight capacity between the treatments was insignificantly different from the first to 3rd day after the treatment but was significantly different from the 4th to 6th day either in the mean flight duration or in distance. Evaluation of the long fliers at the same stages between the treatments also supported this finding. Flight capacity of the JHM‐treated moths also differed with the sex. The JHM‐treated female did not fly less than the control from day‐1 to day‐3, but did significantly less from day‐4 to day‐6 after the treatment. The flight capacity of the JHM‐treated males, however, was not weaker than that of the control throughout the test period. Comparison of the levels of energy substances in the JHM‐treated adults and the controls showed that JHM may directly acted on metabolism of glyceride, which in turn affected flight capacity. Furthermore, higher levels of glyceride in the treated male adults raised their flight capacity to a stronger levels than the control in most of the days of tethered‐flight test period. Contents of glycogen, however, were insignificantly affected by JHM in both male and female adults. In might therefore be concluded that levels of JH is a critical factor regulationg migratory behavior in the army worm moth. In addition to affect metabolism of glyceride, other possible physiological roles that JH played in determining migrattory behavior are also discussed.     

Reproduction-flight relationship in the beet armyworm, Spodoptera exigua (Lepidoptera: Noctuidae)

To evaluate the relationship between reproduction and migratory flight, we examined ovaries, tethered flight, and egg laying in the beet armyworm, Spodoptera exigua (Hübner). Ovarian development in female S. exigua was classified into five stages in relation to the postemergence age of the moths. Oviposition peaked at ovarian stages III and IV. Mating accelerated ovarian development in this species, and thus the mated females developed faster than the virgin moths in ovaries. In virgin adults, flight capacity increased from the first day to the third day after emergence and remained at a high level until the moths were 7 d old. In contrast, the mated moths reached the maximum flight capacity in 2 d after emergence and maintained a high level until the age of 7 d. However, mating status could not influence the flight capacity of the beet armyworm, although there were general differences in flight capacity between virgin and mated adults at the same age. Moreover, significant correlations between the number of eggs laid and the flight capacity were not performed by the mated moths. These results showed that mating status and ovarian development did not seem to influence flight capacity in the beet armyworm, which was not in agreement with the definitions of the oogenesis-flight syndrome observed in many other migratory insects.     

飞行对粘虫体内甘油酯积累与咽侧体活性的影响

研究了粘虫Mythimna separata (Walker)成虫飞行对甘油酯的恢复、保幼激素的滴度及飞行肌降解的影响, 结果表明,不同日龄粘虫的飞行活动对其能源物质的积累及保幼激素分泌产生不同的影响。1日龄蛾的飞行对粘虫这两方面产生的影响最大,其飞行个体能源物质的积累明显高于未飞行的对照个体; 3日龄飞行个体的能源物质积累与对照相当; 但5日龄飞行个体则很难达到对照水平。1日龄飞行个体咽侧体活性在36 h后明显高于对照,60 h后已是对照的10倍,108 h达到其峰值; 3日龄飞行个体咽侧体的活性稍高于对照,但差别不显著; 5日龄飞行个体的咽侧体活性则稍低于对照。不同日龄飞行对飞行肌的降解也产生不同的影响。1日龄飞行个体的飞行肌在飞行后6 天已经明显低于对照。3、5日龄的飞行活动对其飞行肌降解的影响不明显。因此推测,粘虫咽侧体活化的关键时期可能在羽化后1～3 天之间。     

保幼激素类似物对粘虫飞行能力及能源物质含量的调控(英文)

本文报道了保幼激素类似物methoprene(JHM)对粘虫Mythimnaseparata(Walker)飞行能力及飞行能源物质含量的影响。总体而言 ,用JHM (30 μg/蛾 )处理过的成虫飞行能力比对照 (丙酮处理 )的弱 ,但因性别及处理后的时间变化而异 :处理雌蛾的飞行能力在处理后的 1d到 3d与对照没有显著差异 ,但在4d到 6d则显著低于对照 ;处理雄蛾的飞行能力一直比对照的强 ,尽管差异不显著。处理过的雌蛾飞行个体比对照的少 ,而处理过的雄蛾比对照的多。究其原因 ,主要是JHM处理过的雌蛾甘油酯含量在处理后的 3d后明显下降 ,而处理过的雄蛾甘油酯含量一直比对照的高。但无论是雌蛾还是雄蛾 ,处理的与对照的糖元含量均没有明显的差异。这些结果表明 ,作用于成虫体内甘油酯含量是保幼激素影响粘虫飞行能力的主要原因之一。最后 ,对保幼激素影响粘虫飞行能力的其它可能原因也进行了讨论     

稻纵卷叶螟的飞行和再迁飞能力

稻纵卷叶螟Cnaphalocrocis medinalis (Guenée)可以通过连续几夜的飞行实现远距离迁移, 这种变幻莫测的再迁飞能力给稻纵卷叶螟的预测造成了极大的困难, 明确其迁飞行为参数是灾变预警的前提。为此, 利用飞行磨装置测定了稻纵卷叶螟成虫的飞行能力和再迁飞能力。结果表明: 在生殖隔离条件下, 稻纵卷叶螟不同日龄和不同性别的成虫的飞行能力间没有显著差异, 雌雄虫的平均飞行时间分别为156.94±103.19 min和147.71±111.38 min。根据吊飞的累计飞行时间(accumulative flight duration, AFD)可将稻纵卷叶螟种群划分为居留型(AFD<40 min)、迁飞型(40 min≤AFD≤130 min)和强迁飞型(AFD>130 min) 3种类型; 且3种类型的个体比率分别为46.1%, 27.3%和26.6%, 其中居留型平均累计飞行时间为11 min, 迁飞型为82 min, 强迁飞型为232 min。稻纵卷叶螟具有很强的再迁飞能力, 其种群作一次迁飞的个体比率都大于90%, 2次(夜)再迁飞的比率达70%以上, 一般可进行4~5次(夜)再迁飞, 最多可达9次(夜)。成虫的补充营养对再迁飞能力没有显著影响, 但蜜水可增强成虫的飞行能力。这些研究结果为稻纵卷叶螟的虫源分析和迁入区预测提供了基本参数。     

Effect of physiological factors on the flight ability of Calliptamus italicus L

Calliptamus italicus L. is a major migratory pest that often causes serious agricultural losses in the desert/semi‐desert steppe of Central Asia. The present study aims to understand the physiological factors that affect migration of C. italicus by examining the relationships between flight capacity, energy accumulation, ovarian development and Juvenile hormone (JH) titre. The results show that flight capacity decreases with age, being greatest among 2‐day‐old males and lowest among 13‐day‐old locusts. There is no significant correlation between energy accumulation and flight capacity, although JH titre is negatively correlated with flight capacity. Energy accumulation and flight capacity first increases then decreases in 1–13‐day‐old females. Glycogen accumulation is significantly correlated with flight speed, and triglyceride accumulation is significantly correlated with flight distance, duration and speed. Changes in JH titre in 1–13‐day‐old females show double titre peaks, which are negatively correlated with flight capacity. Seven‐day‐old C. italicus have the highest glycogen and triglyceride accumulation, greatest flight capacity, grade II ovarian development and lowest JH titre. By contrast, 13‐day‐old C. italicus have the lowest triglyceride accumulation, lowest flight capacity, grade IV ovarian development and lowest JH titre. Taken together, these findings demonstrate that there is a trade‐off between ovarian development and flight capacity of C. italicus.     

粘虫飞行对生殖及寿命的影响

该文报道了粘虫Mythimna separata（Walker）成虫飞行后产卵、交配及寿命的研究结果。1日龄成虫飞行6 h、12 h、18 h、24 h后的产卵前期均显著短于对照的,其中飞行6 h、12 h的比对照的短2天以上,产卵量均比对照的高。对1～5日龄成虫分别飞行23.5 h后的研究结果表明,1日龄飞行的产卵前期和上述结果相一致。2～4日龄飞行的与对照的没有显著差异,但产卵量则随飞行日龄的延迟而逐渐减少。5日龄飞行的产卵前期显著延长,产卵量已不到对照的一半。所有经过飞行的成虫产卵高峰日比对照的早1天。不同日龄成虫飞行时间、距离与成虫产卵量的关系为：1～3日龄飞行时间、距离长的个体产卵量也高;但4～5日龄的成虫飞行时间与距离越长,其产卵量越少,表现出明显的卵子发生飞行拮抗症(oogenesis-flight syndrome)。除了5日龄飞行的成虫交配率有所下降以外,所有经过飞行的成虫产卵历期、交配率及寿命与对照的没有显著差异。最后,根据这些结果,对粘虫迁飞的起飞时期,迁飞在粘虫生殖、种群动态及成灾规律中的作用进行了讨论。     

Tethered flight of the potato moth, Phthorimaea operculella

ABSTRACT. The effects of sex, age and mated state on average flight speed, duration and distance were determined for potato moths, Phthorimaea operculella (Zeller) (Lepidoptera, Gelechiidae), tethered to flight mills. Moths were classified as non-fliers (NFs), good-fliers (GFs) and remaining-fliers (RFs) on the basis of their performance over the first two flights. Some moths flew for over 5 h non-stop, while others tethered overnight flew between 20 and 30 km. Speed, duration and distance flown were greater during the first flight. First flight duration and distance flown by females decreased with age, whereas no trend was evident for males. Mated males and females flew slower first flights with increasing age, whereas virgin moths showed no marked trend. The analysis of fliers and NFs revealed that GFs were heavier than both RFs and NFs, GFs were faster than RFs, the percentage of NFs increased with age especially for mated females, and the percentage of GFs decreased with age. Age and mated state are important factors influencing flight performance especially for female moths. The relevance of these results to the field situation and the possible application of tethered flight to tests of potato moth quality are discussed.     

Increased juvenile hormone levels after long-duration flight in the grasshopper, Melanoplus sanguinipes

Although, in many insects, migration imposes a cost in terms of timing or amount of reproduction, in the migratory grasshopper Melanoplus sanguinipes performance of long-duration flight to voluntary cessation or exhaustion accelerates the onset of first reproduction and enhances reproductive success over the entire lifetime of the insect. Since juvenile hormone (JH) is involved in the control of reproduction in most species, we examined JH titer after long flight using a chiral selective radioimmunoassay. JH levels increased on days 5 and 8 in animals flown to exhaustion on day 4 but not in 1-h or non-flier controls. No difference was seen in the diel pattern of JH titer, but hemolymph samples were taken between 5 and 7 h after lights on. Treatment of grasshoppers with JH-III mimicked the effect of long-duration flight in the induction of early reproduction. The increased JH titer induced by performance of long-duration flight is thus at least one component of flight-enhanced reproduction. To test the possibility that post-flight JH titer increases are caused by adipokinetic hormone (AKH) released during long flights, a series of injections of physiological doses of Lom-AKH I were given to unflown animals to simulate AKH release during long flight. This treatment had no effect on JH titers. Thus, although AKH is released during flight and controls lipid mobilization, it is not the factor responsible for increased JH titers after long-duration flight.     

粘虫迁出与迁入种群的行为和生理特性比较

系统比较了田间粘虫Mythimnaseparata (Walker)迁出与迁入种群在飞行能力、 飞行肌干重、卵巢发育、交配以及保幼激素滴度等指标的差异。结果显示,迁出种群的飞行潜 力显著强于迁入种群,雌蛾中胸背纵肌平均干重（6.53 mg）也显著高于迁入种群（5.92 mg ）,但卵巢发育程度显著低于迁入种群;迁出种群的交配率和平均交配次数均显著低于迁入种 群;早期迁入的雌蛾卵巢发育级别和交配率相对较低,并随迁入时间的延长而增加。迁出与迁入 种群雌蛾的保幼激素滴度也存在显著差异,两者的JHⅡ滴度平均分别为4.63 ng/mg和8.69 ng/mg。这些结果说明,粘虫迁飞前后的行为、生理和生殖特性具有明显的变化,迁出种群具有较强的飞行潜力,生殖系统的发育受到暂时性抑制;而迁入种群飞行能力下降,卵巢发育级别 较高,交配率增加。同一世代早期迁入的雌蛾卵巢发育程度较低,表明粘虫迁飞是发生在成虫 发育的初期,迁飞过程中卵巢发育受到暂时性抑制,迁入以后便迅速发育并开始繁殖。粘虫迁 飞的行为和生理特性与“卵子发生 飞行拮抗综合症候(oogenesisi flight syndrome)”相符 。内分泌激素（保幼激素）在其迁飞与生殖的转换过程中起着重要的调控作用。     

Longevity,flight ability and reproductive performance of the diamondback moth,Plutella xylostella (L.) (Lepidoptera: Yponomeutidae), related to adult body size

Plutella xylostella in the temperate zone shows a clear seasonal change in adult body size. In the laboratory, large and small moths were produced during immature stages at 15°C and 25°C, respectively. These moths were then used to evaluate longevity, age-specific flight ability, flight ability of mated and unmated females, and the influence of flight experience on the subsequent reproductive success. The large moths lived longer and displayed a greater flight ability over 3 weeks. Irrespective of body size, unmated females flew for a longer time than mated females, and flight experience affected their subsequent reproductive success. Females of both sizes mated and laid eggs soon after emergence, without any obvious pre-reproductive period. More flight experience did not delay oviposition, but did reduce egg production. It is likely that large moths with a longer adult life span and greater flight ability are better fitted for long-distance flight and more fecund than small ones. These experimental results may explain why long-distance migration ofP. xylostella is mostly seen during cool seasons, when relatively large moths with long forewing appear in the field.     

粘虫成虫在气流场中飞行行为的观察研究

通过改进悬吊测飞技术、室内风洞和野外雷达相结合的观测方法,研究了粘虫在气流场中的飞行行为特征及其与气流的关系。直筒风洞自由飞行观测的结果表明,粘虫蛾对气流有明显行为反应,表现为头部迎风起飞和迎风飞行的特性;在3.0～5.5 m/s风速下,有92%～94% 的个体可一次逆风飞行通过2 m长的风洞;当风速≥6.0 m/s时,有71.9%的蛾子沿螺旋状的飞行轨迹逆风通过风洞。环形风洞悬吊飞行测试的结果表明,粘虫可逆风飞行的最大风速为7.2 m/s;在风速≤4 m/s条件下,90%以上个体头部迎风飞行或头部朝向与风向成一定的夹角,侧逆风飞行。 雷达观测发现粘虫在空中迁飞过程中具有成层现象,并有较强的秋季回迁定向行为,其头部总是朝向西南;迁飞的最终位移与风向及风速大小有关,迁飞位移速度是飞行速度与风速的矢量和。     

Long flights in Cydia pomonella L. (Lepidoptera: Tortricidae) measured by a flight mill: influence of sex, mated status and age

Abstract. The flight capacity of Cydia pomonella L. (Lepidoptera: Tortricidae) was measured in the laboratory by using computer-linked flight mills. Codling moths showed a large variation in flight capacity between individuals. We defined arbitrarily a longest single flight (LSF) of more than 5 km as an index for long-flyers. About 16.7% of virgin and 10.0% of mated males and 20.0% of virgin and 7.4% of mated females undertook such flights. Based on the LSF and the total distance flown (TDF^we concluded that males and females have little or no difference in flight capacity and that both the within-and between-habitat flights are similar in number and magnitude for both sexes. In the field, females are therefore potentially able to undertake flights of up to 11 km, as reported for males by other authors. This ability was highest at ages of 2–7 days after emergence, i.e. the first third of their lifetime, for virgin and mated male moths and for virgin female moths. Mated females showed peak flight capacity between 1 and 3 days after eclosion, which corresponded with the major egg-laying period. Few long flights were undertaken before oviposition. These findings do not agree with the oogenesis flight syndrome described by other authors, and this theory is believed not to apply to C.pomonella. Our laboratory results are discussed in relation to field experiments in general and hypotheses are developed about the significance of long-flyers for this species.     

黄地老虎飞行能力研究

黄地老虎Agrotis segetum是一种重要的农业迁飞性害虫,研究飞行生物学对明确其迁飞机理有重要意义。本项研究利用昆虫飞行磨系统对黄地老虎的飞行能力进行了测定。对实验室种群飞行能力的测定结果表明:1日龄成虫的飞行能力最弱;3日龄飞行能力最强,其24 h平均飞行时间、平均飞行距离和平均飞行速率分别达到12.16±0.74 h、48.94±3.40 km和3.76±0.12 km/h。雌蛾与雄蛾间的各项飞行参数均无显著差异,但处女成虫平均飞行距离和平均飞行速率均显著高于已交配个体。对渤海湾野外迁飞种群飞行能力的测定显示,早季节北迁个体和晚季节回迁个体的平均飞行时间和平均飞行距离无显著差异,但皆显著低于实验室种群。黄地老虎具有较强的飞行能力,日龄和交配是影响其飞行能力的关键因素。     

The migration syndrome in the African armyworm moth, Spodoptera exempts: allocation of resources to flight and reproduction

Abstract. Selection for the capacity for prolonged tethered flight in Spodoptera exempta resulted in heavier moths, with significantly larger abdominal glyceride glycerol contents in females of two flight-selected strains (87% and 49% higher than in the comparable non-selected strain) and in males of one of them (80% higher). All flight-selected strains contained individuals of both sexes with very high abdominal glyceride levels. There was no significant relationship between abdominal total glyceride glycerol after flight and flight duration for non-selected moths, but an inverse linear relationship was evident in both sexes from a flight-selected strain. Oxygen consumption during tethered flights by flight-selected moths ranged from 28.2 to 56.6 ml O₂g^-1h^-1.
Using these and previous data, notional energy budgets were calculated to account for flight, reproduction and resting metabolism for non-selected and flight-selected S. exempta flown on the flight balances and provided thereafter with distilled water. The results confirm the trade-off between flight and reproduction reported by Gunn et al. (1989). A similar approach using data for two non-selected strains from the field in Kenya indicated genetic variation in migratory potential, reflected both in pre-reproductive period and resources available for flight. We conclude that elevated glyceride levels are a component of the migratory syndrome in S. exempta and that this is the major factor underlying the curvilinear relationship between flight duration and fecundity obtained by Gunn et al. (1989).     

一日龄饥饿对粘虫成虫卵巢发育和飞行能力的影响

为了明确羽化后0～24 h (1日龄)饥饿处理对迁飞型粘虫Mythimna separata (Walker)生殖系统和飞行系统发育的影响,对1日龄饥饿处理后粘虫的卵巢发育级别、产卵前期、飞行肌和飞行能力进行了系统研究。结果表明, 1日龄饥饿对粘虫成虫卵巢发育级别、产卵前期、中胸背纵肌干重、飞行能力等均有显著的影响。饥饿后成虫卵巢发育级别显著高于对照,产卵前期显著短于对照。成虫中胸背纵肌干重从饥饿后96 h开始下降,至120 h时饥饿粘虫(5.1 mg)显著低于对照(5.86 mg)。成虫飞行能力从饥饿后96 h开始显著低于对照,此时饥饿处理粘虫蛾的飞行距离平均为22.03 km,飞行时间平均为4.07 h,显著低于同时期的对照粘虫的飞行距离(42.3 km)和飞行时间(7.99 h),尽管在饥饿后72 h内差异不显著。实验结果显示,1日龄饥饿可以显著抑制粘虫飞行系统的发育,加速飞行向生殖的转变。     

Laboratory evaluation of flight activity of the common cutworm, Spodoptera litura (Lepidoptera: Noctuidae)

Abstract  The flight activity of Spodoptera litura in tethered conditions is evaluated using a computer-mediated flight-mill in the laboratory. The 3–4-day-old moths fly longer and farther than those of other ages. Male and female moths exhibit similar flight activity. Mating status does not influence the flight duration and distance of 2-day-old females. However, these two flight parameters with a 6-day-old mated female is significantly lower than that of unmated ones. The optimum temperature for flight ranged from 16–24°C, whereas the optimum RH ranged from 60%–100%. During 72-h period, the total flight duration and distance of 1-day-old male and female moths were 19.6 h (± 5.8) and 83.3 km (± 28.4), and 24.0 h (± 7.0) and 105.4 km (± 37.4), respectively. These results indicate that S. litura has a great potential to undertake long-distance migratory flights.     

Trade-off between flight and reproduction in the African armyworm moth, Spodoptera exempta

ABSTRACT. The existence of a trade-off between flight and reproduction in the migratory noctuid, Spodoptera exempta (Wlk.), was examined in experiments in which female moths were flown on flight balances and then provided with distilled water or sucrose solution. For individuals flight-tested from the night of eclosion (night 0) through night 1 and fed only distilled water, there was a linear, negative relationship between weight-related fecundity and flight duration. When flight was recorded through night 2, a curvilinear relationship was obtained suggesting that highly active moths suffer a smaller decrease in fecundity than predicted from its rate of decline with increasing flight duration in less persistent fliers. The hypothesis that this reflects an energetically expensive, initial phase of each flight, followed by a less costly, cruising phase is examined. Fecundity was independent of flight duration in females provided with sucrose after flight but was related to moth weight, as it was in all unflown moths. Effects of flight and feeding on longevity and mating frequency are also reported.
There is a clear trade-off between flight and fecundity in S.exempta which is only evident in moths denied access to a source of carbohydrate after flight. The results indicate the importance of nectar sources in the field in allowing restoration of lipid reserves depleted during migratory flight. An important implication of this conclusion is that lipid is the resource limiting to fecundity in this species and not protein as is generally supposed for leaf-feeding insects. This might be expected in highly active species as lipid is the resource for which the energetically expensive functions of flight and reproduction must compete.     

A flight cylinder bioassay as a simple,effective quality control test for Cydia pomonella

Assessment of quality of the sterile male insects that are being mass‐reared for release in area‐wide integrated pest management programmes that include a sterile insect technique component is crucial for the success of these programmes. Routine monitoring of sterile male quality needs to be carried out both in the mass‐rearing facility and in the field. Simple bioassays that can be conducted in the laboratory and that would be surrogates for laborious field tests would be a very cost‐effective way of monitoring sterile male field performance. Simple flight cylinders were used to assess whether these could detect differences in quality of male codling moth Cydia pomonella. The number of male and female codling moths that flew out of the cylinders was influenced by cylinder diameter, cylinder height and number of hours following the initiation of the test. The flight cylinder bioassay was capable of detecting differences in quality of codling moths induced by irradiation when moths were shipped, but no differences were found in flight ability when the moths were not transported. The tests also confirmed that handling and shipment reduced quality more for irradiated than for non‐treated codling moth, and that insect quality was significantly influenced by larval rearing protocols. The flight cylinder bioassay was therefore successful in detecting differences in codling moth quality induced by various treatments that had been identified previously by more complex laboratory bioassays and field trials. Treatment differences were most likely detected when flight cylinders were 16 cm high.     

甘蓝夜蛾Mamestra brassicae Linnaeus飞行能力研究

【目的】测定日龄、性别、交配等因素对农业迁飞性害虫甘蓝夜蛾Mamestra brassicae Linnaeus飞行能力的影响,为其迁飞行为研究和异地测报提供理论依据。【方法】在气温为(24±1)℃、相对湿度为75%±5%的实验室环境下,利用飞行磨系统对甘蓝夜蛾实验室种群1~5日龄未交配个体、已交配的3日龄个体以及野外迁飞种群分别进行连续24 h吊飞测试,记录其累计飞行时间、累计飞行距离、平均飞行速率等参数。【结果】甘蓝夜蛾实验室种群的飞行能力随日龄增长而变化,初羽化时即具有较强的飞行能力,2~3日龄达到峰值,4~5日龄的飞行能力显著下降;雌性甘蓝夜蛾的飞行能力强于雄性甘蓝夜蛾;交配的甘蓝夜蛾平均飞行速率显著小于处女甘蓝夜蛾,但二者的平均飞行时间和飞行距离均无显著差异。此外,渤海湾迁飞种群中晚季节南迁种群的飞行能力显著强于早季节北迁种群和过渡种群。【结论】日龄和交配状态是影响甘蓝夜蛾飞行能力的关键因素。