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.     

Simultaneous monitoring of flight and oviposition of individual velvetbean caterpillar moths (by Wales, Barfield & Leppla, 1985): a critique

ABSTRACT. Flight durations of tethered female Anticarsia gemmatalis moths recorded in the laboratory were interpreted by Wales et al. as indicating a capacity for significant inter-reproductive displacements in the field. Indications that their data may not support this conclusion are discussed and evidence is presented that, in tethered Spodoptera exempta , prolonged flights starting early in the night represent migratory behaviour while those of similar durations starting after midnight do not.     

Flight capacity: genetic determination and physiological constraints in a migratory moth Mythimna separata

Abstract. Flight capacity of the oriental armyworn Mythimna separata (Walker) (Lepidoptera: Noctuidae) was estimated by a new tethered flight technique. Samples of a migrant population showed a substantial proportion of 'long fliers'. Heritability of flight capacity was found to be significant ( h ²= 0.27). Female moths from a line selected for long pre-reproductive period (PRP) showed greater flight capacity than those from a short PRP line. This difference was not apparent among males. Onset of calling behaviour inhibited the expression of flight capacity in females showing a clear 'oogenesis-flight syndrome'. These results are discussed in relation to seasonal migration in eastern China and the mechanisms involved in the migratory strategy of this species.     

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     

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.     

A technique for studying flight behaviour of tethered Spodoptera exempta moths

ABSTRACT. A new apparatus was developed for studies on flight behaviour of the African armyworm, Spodoptera exempta (Wlk.), to record the time and duration of all flights continuously for up to several days. A technique for attaching the mounting bracket to the pharate adult before eclosion allowed flight recording to commence within 1–2h of emergence without the need for anaesthesia. Results obtained using the apparatus indicate that migratory flight does not occur on the night of emergence. The time of take-off of tethered moths in the laboratory agrees closely with observations of flight behaviour in the field.     

The effect of adult feeding on lipid and protein reserves in African armyworm, Spodoptera exempta, moths before and during reproduction

ABSTRACT. Newly-emerged Spodoptera exempta (Walker) (Lepidoptera: Noctuidae) moths contain high levels of lipid held largely in the abdominal fat body, the quantity depending on the larval feeding conditions. There is a positive relationship between weight-related lipid content and moth weight, which is consistent for female but not for male moths, suggesting that larval feeding conditions producing large individuals allow the accumulation of quantitatively disproportionate lipid reserves. Male and female moths have comparable levels of abdominal protein.
Changes in the water content in starved moths or ones provided with distilled water or sucrose solution show that while starved individuals die rapidly from desiccation, water-fed moths regulate their water contents between narrow limits which are higher for females than for males. Sucrose-fed moths maintain higher, more variable water contents probably due to the phagostimulatory effect of the sugar.
Reproducing and unmated moths are able to supplement their lipid and, to a lesser extent, their protein reserves following carbohydrate uptake.
During both larval and early adult stages, the capacity to accumulate lipid reserves in excess of those apparently required for reproduction, suggests that these reserves also provide the main fuel for the prolonged flights of which migratory individuals are known to be capable.     

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。     

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。     

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.     

Quantifying interspecific variation in dispersal ability of noctuid moths using an advanced tethered flight technique

Dispersal plays a crucial role in many aspects of species' life histories, yet is often difficult to measure directly. This is particularly true for many insects, especially nocturnal species (e.g. moths) that cannot be easily observed under natural field conditions. Consequently, over the past five decades, laboratory tethered flight techniques have been developed as a means of measuring insect flight duration and speed. However, these previous designs have tended to focus on single species (typically migrant pests), and here we describe an improved apparatus that allows the study of flight ability in a wide range of insect body sizes and types. Obtaining dispersal information from a range of species is crucial for understanding insect population dynamics and range shifts. Our new laboratory tethered flight apparatus automatically records flight duration, speed, and distance of individual insects. The rotational tethered flight mill has very low friction and the arm to which flying insects are attached is extremely lightweight while remaining rigid and strong, permitting both small and large insects to be studied. The apparatus is compact and thus allows many individuals to be studied simultaneously under controlled laboratory conditions. We demonstrate the performance of the apparatus by using the mills to assess the flight capability of 24 species of British noctuid moths, ranging in size from 12–27 mm forewing length (~40–660 mg body mass). We validate the new technique by comparing our tethered flight data with existing information on dispersal ability of noctuids from the published literature and expert opinion. Values for tethered flight variables were in agreement with existing knowledge of dispersal ability in these species, supporting the use of this method to quantify dispersal in insects. Importantly, this new technology opens up the potential to investigate genetic and environmental factors affecting insect dispersal among a wide range of species.     

Antennal regulation of migratory flight in the neotropical moth Urania fulgens

Migrating insects use their sensory systems to acquire local and global cues about their surroundings. Previous research on tethered insects suggests that, in addition to vision and cephalic bristles, insects use antennal mechanosensory feedback to maintain their airspeeds. Owing to the large displacements of migratory insects and difficulties inherent in tracking single individuals, the roles of these sensory inputs have never been tested in freely migrating insects. We tracked individual uraniid moths (Urania fulgens) as they migrated diurnally over the Panama Canal, and measured airspeeds and orientation for individuals with either intact or amputated flagella. Consistent with prior observations that antennal input is necessary for flight control, 59 per cent of the experimental moths could not fly after flagella amputation. The remaining fraction (41%) was flight-capable and maintained its prior airspeeds despite severe reduction in antennal input. Thus, maintenance of airspeeds may not involve antennal input alone, and is probably mediated by other modalities. Moths with amputated flagella could not recover their proper migratory orientations, suggesting that antennal integrity is necessary for long-distance navigation.     

Downwind migration of the African army worm moth, Spodoptera exempta, studied by mark-and-capture and by radar

Abstract. 1. About 166,000 African armyworm moths, Spodoptera exempta (Walk.), were marked at an emergence site near Nairobi when they fed at night on trees baited with dyed molasses.
2. Six marked moths were captured in pheromone traps, including one at 90 km after flying for only one night, and another at 147 km.
3. Moth flight trajectories deduced from radar and from marking showed that migration was downwind.
4. During migration, moths become dispersed; hence the high densities that lead to outbreaks must be produced by concentration.
5. Some moths were ready to mate on the same night they completed their long-distance flight.     

The use of video equipment to record in three dimensions the flight trajectories of Heliothis armigera and other moths at night

ABSTRACT A simple method of recording stereoscopically the behaviour of moths in nocturnal, low-altitude flight in the field is described. Sample results, showing the flight of Heliothis armigera (Hiibner) and other noctuids at 2–11 m above a crop of pigeon-pea in India are presented. Flight trajectories, reconstructed in three dimensions, demonstrated that most of the moths flew close to the horizontal, in straight lines, and that their average air (flight) speed was 5.0 ± 1.9 m s^-1. There was no evidence of mass climbing flight away from the crop.     

棉铃虫飞翔的能源物质及消耗

研究结果表明棉铃虫Helicoverpa armigera(Hubner)初羽化个体的能源储备主要为甘油酯,1日龄成虫甘油酯和糖原的含量分别为每头3.6365mg和0.3658mg,通过成虫期的取食,二者的含量分别于羽化后4d和5d达到高峰(6.5707mg和2.3500mg)。对吊飞个体的测定表明甘油酯和糖原是棉铃虫飞翔的能源物质,蛋白质含量和飞行活动无明显的相关关系。4日龄成虫在飞行磨上吊飞66h,飞翔时间37.9943h,累积飞行216.8650km。累积消耗能量218.72J,其中甘油酯占85.87%,糖原占14.13%。     

二点委夜蛾飞行行为特征

【目的】本研究旨在阐明二点委夜蛾的飞行行为特征,丰富二点委夜蛾飞行生物学理论,提高其预测预报水平。【方法】利用昆虫飞行磨被动吊飞系统和主动飞行监测系统,系统研究了二点委夜蛾Athdtis lepigone(M(o|¨)schler)被动飞行能力和主动飞行意愿。【结果】成虫具有较强的被动飞行潜力。室内连续吊飞80 h,雌雄蛾最远飞行距离分别达106.71 km和148.32 km,最长飞行时间分别达43.05 h和40.01h,最快飞行速度分别达7.60 km/h和8.14 km/h。雄蛾飞行潜力显著强于雌蛾,体现在飞行距离和飞行时间显著高于雌蛾,但飞行速度差异不显著。成虫蛾龄显著影响成虫飞行能力。对不同蛾龄成虫吊飞12 h的结果表明,1日龄即具备一定的飞行能力,之后逐渐增强,3日龄时飞行能力最强,雌雄蛾平均飞行距离分别为29.61 km和27.55 km,飞行时间分别为10.04 h和9.46 h,平均飞行速度分别达2.76 kn/h和2.46km/h,4日龄成虫飞行能力开始下降,但不同性别间成虫飞行能力差异不显著。蛾龄间飞行能力差异主要是由于不同蛾龄成虫的强、弱飞行个体比例不同。二点委夜蛾主动飞行呈现明显的节律行为,飞行活动主要集中在暗期(19:00—次日5:00),在光期(5:00—19:00)基本不飞行。成虫初羽化(1日龄)主动飞行意愿增强,之后飞行活动减少,但产卵开始时主动飞行活动又开始增强,到7日龄达到峰值。【结论】二点委夜蛾成虫具有较强的飞行能力,其飞行能力受蛾龄,雌雄等因素影响;飞行具有明显的节律性。     

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.     

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.     

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。