Relative contribution of genetic and environmental factors to determination of wing morphs of the brown planthopper Nilaparvata lugens

Wing dimorphism is a fascinating feature of the ability of insects to adapt to environments. The brown planthopper (BPH) Nilaparvata lugens, a serious pest of rice, can switch between the long- and short-winged morphs. It has been known that environmental factors can affect the wing morph of BPH. However, it is still unclear whether the effect of environment is dependent on BPH genetic backgrounds or not. In the present study, we established the pure-bred lineages of short- and long-winged BPHs via multigenerational selection, and we found that survival and fecundity were similar between these 2 lineages. Wing morphs of the pure-bred lineages were almost fully dependent on genetics, but independent of the environmental factors, nymphal density and rice plant stage, 2 key factors affecting BPH wing morphs. In the unselected BPH population, short- and long-winged morphs were produced depending on those 2 environmental factors, indicating the contribution of environment to wing morph. In the wing-selected lineages, 4 developmental regulated genes of wing, NlInR1, NlInR2, NlAkt, and NlFoxo were expressed stably in the short-winged adults, but almost silenced in the long-winged adults. However, all these genes were expressed normally with a similar level in both the short- and long-winged adults in an unselected population except NlFoxo. The pure-bred lineages of long- and short-winged morphs exhibited different expression patterns of wing development-regulated genes, suggesting the genetic determination of wing morphs. Effects of environmental factors on wing morphs occurred only in the genetic mix population.     

丽斗蟋翅二型雌虫飞行肌和卵巢发育间的资源分配差异

丽斗蟋Velarifictorus ornatus具有明显的翅二型现象, 长翅型与短翅型雌虫的卵巢和飞行肌存在着生理权衡。本研究分别应用蒽酮比色法、 硫代磷酸香草醛法、 考马斯亮蓝染液对羽化后10 d内两型雌虫飞行肌与卵巢内糖原、 总脂及蛋白质含量进行了定量分析。结果表明： 成虫羽化后10 d内, 两型雌虫体重无明显差异(P>0.05), 但短翅型雌虫怀卵量明显多于长翅型雌虫, 而人工脱翅能够促进长翅型雌虫怀卵量增加(P<0.05)。短翅型雌虫飞行肌内蛋白质、 糖原及总脂含量在成虫羽化后10 d内无明显变化, 但长翅型雌虫飞行肌内蛋白质在成虫羽化后3 d时达到最大值564.4±87.5 μg/♀, 糖原与总脂含量分别于羽化后第5天达到最大值85.2±21.7 μg/♀和5 284.7±1 267.4 μg/♀。然后开始下降, 各实验处理天数内, 长翅型雌虫飞行肌内蛋白质、 糖原及总脂含量都显著多于短翅型雌虫(P<0.05)。相反, 各处理天数内, 短翅型雌虫卵巢内蛋白质、 糖原及总脂含量则明显多于长翅型雌虫(P<0.05), 同时虫龄对蛋白质、 糖原及总脂在两型雌虫飞行肌与卵巢内分配也产生明显影响(P<0.05)。人工脱翅能够促进长翅型雌虫卵巢内蛋白质、 糖原及总脂含量增加, 同时诱导飞行肌内蛋白质、 糖原及总脂含量降低, 其中总脂含量在脱翅后10 d时降为2 394.9±1 461.8 μg/♀, 只有最大值的一半, 而与短翅型雌虫相似(P>0.05), 表明总脂为丽斗蟋飞行的主要能源物质。外用保幼激素Ⅲ能够促进长翅型雌虫卵巢内蛋白质、 糖原及总脂含量增加(P<0.05), 但对飞行肌内三者含量无明显影响(P>0.05), 外用早熟素Ⅰ对短翅型雌虫卵巢内蛋白质、 糖原及总脂含量亦无明显影响(P>0.05)。上述结果表明, 丽斗蟋长翅型雌虫首先将获得的资源用于发育飞行所需的飞行肌, 短翅型雌虫则首先将所获得的资源用于发育繁殖所需的卵巢, 但长翅型雌虫飞行肌与卵巢间的资源分配方式受保幼激素的影响。     

灰飞虱长、短翅型品系对高、低温的适应能力差异

【目的】灰飞虱Laodelphax striatellus种群存在长翅和短翅个体,其翅型分化受遗传和环境条件的影响。本研究旨在明确经连续定向选育的灰飞虱长、短翅型品系对高、低温的适应能力差异。【方法】在室内分别长期定向筛选63和65代建立灰飞虱长、短翅型品系,并在南京田间采集灰飞虱自然种群;分别在25℃, 30℃和35℃恒温条件下测定上述灰飞虱长、短翅型品系和自然种群的繁殖力、存活率和发育历期,测定若虫在5℃和-20℃下的存活率以及若虫和成虫的过冷却点,通过品系间存活率、发育历期、生殖力和过冷却点的比较分析,确定灰飞虱长、短翅型品系对高温和低温的耐受能力。【结果】在25℃, 30℃和35℃下,灰飞虱自然种群雌成虫的繁殖力均显著高于室内筛选后的长、短翅型品系,而长、短翅型品系雌成虫之间的繁殖力均无显著差异;长、短翅型品系的繁殖力在25℃和30℃之间均无显著差异,但自然种群在25℃下的繁殖力显著高于30℃条件下的繁殖力;35℃下长、短翅型品系基本无若虫产生。25℃下长翅型品系、短翅型品系和自然种群若虫的存活率无显著差异,但在30℃下,短翅型品系若虫的存活率显著低于自然种群,而与长翅型品系差异不显著;35℃下长、短翅型品系的若虫最长只能存活到3龄,且两品系间无显著差异,而自然种群可存活到5龄。25℃下,短翅型品系若虫的发育历期显著短于自然种群,而与长翅型品系无差异;但是在30℃下,短翅型品系若虫历期显著长于长翅型品系和自然种群。长翅型品系、短翅型品系和自然种群灰飞虱3-4龄若虫在5℃低温下的存活天数无显著差异,并且长翅和短翅型品系在-20℃极端低温下暴露10 min后其死亡率也无显著差异且均显著高于自然种群。3个品系(种群)成虫的过冷却点无显著差异,但5龄若虫的过冷却点表现为短翅型品系显著高于自然种群,而与长翅型品系无显著差异。【结论】经连续定向选育的灰飞虱长、短翅型品系对低温和高温的适应力基本相当,且低于自然种群。在高温条件下灰飞虱短翅型品系的发育速率低于长翅型品系。     

Mate choice by male water striders (Gerris lacustris): expression of a wing morph preference depends on a size difference between females

Natural selection maintains wing polymorphism in populationsof water striders. Long-winged females benefit from being ableto migrate because they avoid the risk of pool evaporation,but early in the season their fecundity is lower than that ofshort-winged females. We tested preferences of 18 long-wingedmales and 29 short-winged males in spring. Males preferred long-wingedfemales, and there was no difference between the two male morphs.The preference was only observed when the long-winged femalewas heavier; when the short-winged female was larger, the malesshowed no preference. There was a positive relationship betweenlong-winged female weight and number of eggs laid per day, andthe males behaved as if they were taking into account that large,long-winged females are more fecund. We suggest that males preferredlong-winged females because this preference increases the proportionof long-winged individuals in the autumn F2 generation.     

Fecundity in relation to wing-morph of three closely related species of the melanocephalus group of the genus Calathus (Coleoptera: Carabidae)

Summary In two successive years the fecundity of the carabid beetles Calathus (Neocalathus) cinctus, C. (N.) melanocephalus and C. (N.) mollis was studied in relation to wing-morph and temperature. Differences were found between the three species in both egg production and timing and length of the oviposition period. In all species the fecundity of laboratory bred beetles was significantly higher than that of females collected in the field. Long-winged females of both cinctus and melanocephalus had significantly higher egg production than short-winged females, and they also tended to produce eggs over a longer period. In mollis only the fecundity of the long-winged morph was established. The observed lower relative fitness of the short-winged morph in both cinctus and melanocephalus contradicts the supposed increase of the frequency of this morph in ageing, more or less isolated, populations of these species. The loss of long-winged genotypes, resulting from flight activities, is considered the most plausible cause of the increase of short-winged beetles in ageing populations. The higher fecundity of macropterous females makes them especially suited for (re)establishing populations.Communication No. 429 of the Biological Station WijsterPresent address and address for offprint requests: Kortenburg 31, NL-6871 ND Rentum     

Differential allocation of protein resources to flight muscles and reproductive organs in the flightless wing-polymorphic bug, <Emphasis Type="Italic">Pyrrhocoris apterus</Emphasis> (L.) (Heteroptera)

Differences in the growth of dorsolongitudinal flight muscles and gonads in 1–28 days old long-winged (macropterous) and short-winged (brachypterous) adults of the firebug (Pyrrhocoris apterus L.) and the resource allocation to these organs were studied by means of total protein analysis. We found predominant allocation of food resources to flight muscles compared to reproductive organs in both macropterous males and females during the first 5 days of adult life. Subsequent histolysis of developed flight muscles coincided with increased total protein content in some reproductive organs. Initiation of intensive food intake after starvation or application of higher dose of methoprene on macropterous adults changed the resource allocation in favour of growth of reproductive organs and induced precocious histolysis of flight muscles. It indicates an involvement of juvenile hormone in wing morph-related differential allocation of resources in the bug. Increased total protein contents in the ovaries and accessory glands of starved macropterous females and males treated with methoprene, respectively, indicate that proteins derived from the methoprene-induced histolysis of the flight muscles are re-utilized for the growth of the reproductive organs. It is the first report of persistence of differential resource allocation to flight muscles and reproductive organs in the wing-polymorphic insects with non-functional macropterism.     

Cryptic female choice predicated on wing dimorphism in decorated crickets

Male decorated crickets, Gryttodes sigllatus, normally lackbind wings and are incapable of flight (short-winged males),but occasionally exhibit fully developed hind wings that makerudimentary flight possible (long-winged males). Long-wingedmales bear a cost of flight in the form of decreased inseminationsuccess, which arises as a consequence of two interrelated factors:(1) long-winged males exhibit a lower reproductive investmentrelative to short-winged males, as measured by the mass of amale's spermatophore and reproductive organs and (2) the postcopulatorybehavior of females favors males that maximize their reproductiveinvestment. Of particular importance to male mating successis the spermatophylax, a large gelatinous mass forming partof the spermatophore and consumed by the female after mating.Consumption of the spermatophylax keeps the female preoccupiedwhile sperm are discharged from the remaining portion of thespermatophore (sperm ampulla) into her repro ductive tract.The spermatophylax of long-winged males is significantly smallerthan that of short-winged males and consequently requires lesstime to consume. As a result, the sperm ampulla of long-wingedmales is frequently removed before its complete evacuation andsignificantly sooner than that of short-winged males. Becausethe spermatophore-removal behavior of females mediates the relativeinsemination success of short-winged and long-winged males,it can be considered a form of cryptic female choice     

Genetic variation in developmental times among four pure lines exhibiting specific wing form and body color in the brown planthopper, <Emphasis Type="Italic">Nilaparvata lugens</Emphasis> (Hemiptera: Auchenorrhyncha)

Developmental times were compared among four pure lines of wing-dimorphic Nilaparvata lugens generated by successive selections for the following designated characteristics: blackish short-winged (BS), blackish long-winged (BL), yellowish brown short-winged (YS) and yellowish brown long-winged (YL). When the nymphs were reared on rice seedlings at low density, the developmental time was in the order of BS = BL < YS = YL, while with an increase in rearing density, the time lengthened markedly in long-winged lines but only slightly in short-winged lines, rearranging the order to BS < BL = YS < YL. Among F₁ progeny of BS × YL, the developmental time in short-winged phenotypes was the same as those of BS lines at any of the densities tested, but the long-winged phenotypes emerged as adults at intermediate time between BS and YL lines at high density. This method of inheritance of developmental time and the differences in the duration observed among the four pure lines suggest that the genes regulating wing form and body color expression are also involved in the regulation of developmental time.     

Wing dimorphism in Gryllus rubens: genetic basis of morph determination and fertility differences between morphs

Summary The genetic basis of wing morph determination and fertility differences between wing morphs were studied in the wing-dimorphic cricket, Gryllus rubens. Using pair corsses, a significant effect of genotype on morph determination was documented in F₃ progeny of field-collected crickets. The effect of genotype was significantly stronger in females than in males. Results are consistent with an earlier study (Zera and Tiebel 1988) showing that wing development is more strongly buffered from environmental variation in females. Segregation patterns were consistent with a polygenic mode of inheritance and provided no evidence for the existence of genes of major effect, maternal effects, or sex linkage. Only a weak morph x sex association was observed. These results contrast those of Walker (1987) where sex-linked loci of major effect on morph determination were identified in crosses between long-wing and short-wing-selected strains of G. rubens. Short-winged female G. rubens began ovipositing earlier and oviposited significantly more eggs than long-winged females during the first 24 days after adult eclosion. The greater reproductive output of the short-winged morph was due entirely to greater oviposition during the first two weeks after adult eclosion. Preliminary results indicate that flight may further accentuate the reduced reproductive output of long-winged versus short-winged females. These data provide the foundation for investigating the endocrine basis of morph-associated fertility variation in G. rubens and its mechanistic relationship with morph determination.     

褐飞虱成虫体内磁性物质检测

地磁定向是昆虫远距离迁飞定向的重要机制之一.本研究以褐飞虱Nilaparvata lugens长翅型和短翅型成虫为研究对象,利用MPMS-7型号超导量子干涉磁强计(磁场范围为±4.8 mA/m,温度范围为1.9 ～ 400 K)检测虫体内的磁性物质,明确其体内的分布状况.结果表明:褐飞虱长翅型雄成虫整个虫体的温度退磁曲...     

植物病媒昆虫的翅型分化

翅多型现象是昆虫非遗传多型性的一种表现,包括不具飞行能力的短翅型或无翅型,以及可以进行长距离迁飞的长翅型或有翅型。翅多型现象常发生在可以携带病原并将其传播给植物宿主的媒介昆虫中,对植物病害的时空分布与暴发有重要影响。本文从翅型分化的遗传规律、诱导因素、分子机制和伴随翅型分化的其他生理表现4个方面,对植物病原主要传播媒介蚜虫和飞虱的翅型分化研究进行综述和梳理。昆虫翅型分化的诱导因素主要包括温度、湿度和光周期等非生物因素以及虫口密度、宿主营养、病毒等生物因素;而其内在的分子机制大多是通过胰岛素/胰岛素样生长因子信号(IIS)通路、c-Jun氨基末端激酶(c-Jun NH₂-terminal kinase, JNK)信号通路、Wingless和嗅觉受体SaveOrco等调控。翅型分化的同时伴随着生理状态的变化,表现为短翅型具有更强的繁殖能力和长翅型含有更丰富的飞行肌结构成分。目前,昆虫翅型分化的研究尚不够完善,有许多需要解答的问题,如找到胰岛素/胰岛素样生长因子信号通路中真正发挥功能的靶基因,JNK如何调控翅型分化以及虫媒病毒影响媒介昆虫翅型的分子机理。本综述可为控制虫...     

Diversities in morphological characteristics and seed germination behavior in fruits ofSalsola komarovii Iljin

Two different types of dispersal units (called fruits in this study) were observed inSalsola komarovii Iljin. One is a fruit which has dark brown lignified tepals with long wings and a green seed and falls easily from the mother plant (long-winged type). Another has light brown lignified tepals with short wings and a yellow seed and attaches tightly to the mother plant (short-winged type). This difference of fruit type appeared independently from maturity of fruit. Seeds in the short-winged fruits were in dormancy for a longer period of time than those in the long-winged fruits. The germination rate was significantly higher in the seeds of long-winged fruits. The dormancy in seeds of the long-winged fruits was effectively terminated by reducing the temperature but the effect of chilling was very weak in seeds of the short-winged fruits. It was concluded from these observations that there exists a dimorphism in the fruits of, or a heterocarpy in,S. komarovii. The plants grown under water stress produced mostly short-winged fruits and those grown under well-watered conditions bore fruits of both types. Exogenously applied abscisic acid (ABA) tended to produce the short-winged fruits, suggesting that the heterocarpy was, at least partly, regulated by ABA.     

Effects of wing-pad removal and corpus allatum implantation on development of wings, flight muscles, and related structures in the striped ground cricket, Allonemobius fasciatus

ABSTRACT. Removal of a hindwing pad from late instar nymphs of the striped ground cricket, Allonemobius fasciatus DeGeer (Gryllidae) inhibits the formation of long-winged adults. All nymphs become short-winged when their hindwing pad is removed during the sixth (penultimate) instar or during the first 2 days of the seventh instar. When a hindwing pad is removed on day 4 of the seventh instar or later, there is no effect on wing development and at least 50% of the crickets emerge as long-winged adults as in controls.
The period sensitive for removal of a hindwing pad varies among different structures, e.g. elytra, hindwings, flight muscles, and the acrotergite of the first abdominal segment. As a result, some intermediate forms are produced when a hindwing pad is removed in the middle of the seventh instar.
Implantation of corpora allata (CA) into late instar nymphs also inhibits formation of long-winged adults. When CA are implanted on day 2 or 3 of the seventh instar, some of the resulting adults have abnormally short elytra. Implantation of CA at an earlier stage produces supernumerary (eighth instar) nymphs.     

Physiological trade-offs between reproduction,flight capability and longevity in a wing-dimorphic cricket,Modicogryllus confirmatus

Physiological and morphological comparisons were made between the short-winged (SW) and long-winged (LW) morphs of a cricket, Modicogryllus confirmatus, to determine the cost of the flight capacity and the physiological mechanisms underlying trade-offs between different life history traits related to migration and reproduction. Both wingmorphs grew at a similar rate and no consistent correlation was found between nymphal development and adult body size. The metathoracic muscles at adult emergence represented 4.2% of the wet body weight in the SW morph and 10.5% in the LW morph. Fat content of the body at adult emergence was positively correlated to dry body weight, but no significant difference was found in mean fat content between the two wingmorphs or between sexes after body weight was adjusted. SW females fed ad libitum produced significantly more eggs than LW females during the first 20days of adult life. Egg production was not correlated to either body size or the duration of nymphal development. LW adults lived longer than SW ones when kept with water alone or when given various amounts of food on day 1 and otherwise kept with water alone. In females, a highly significant correlation was found between longevity and egg production, indicating the presence of a trade-off. LW females mainly allocated the energy from food to flight muscle development and general maintenance of the body rather than to egg production, whereas SW females used it for egg production and longevity. LW females that had been de-alated at adult emergence histolyzed the flight muscle and used the energy from food for egg production almost exclusively. These results suggest that energy allocation and trade-offs after adult emergence may play crucial roles in the functional differentiation of the two wingmorphs.     

Diversity in seed germination behavior in relation to heterocarpy inSalsola komarovii Iljin

Seed germination behavior in two types of dimorphic fruits inSalsola komarovii Iljin (Chenopodiaceae) was studied. The germination percentage was much higher in the long-winged type than in the short-winged type when the fruits were tested for germinability. If lignified perianths were removed from the fruits, seeds of both types germinated at higher percentages and the difference in the germination percentage between two types decreased. The extract of the perianths inhibited the seed germination. Abscisic acid was detected in the extract of the perianths by enzyme-linked immunosorbent assay and its level was higher in the short-winged type than in the long-winged type. Germinability in both types was lost one year after harvest when the fruits were stored at room temperature and the germination was tested with fruits. When the seeds were tested, however, the short-winged type germinated even two years after the harvest; the germinability of the short-winged type lasted one year longer than that of the long-winged type. Light irradiation was not required for the seed germination in either type regardless of presence or absence of the perianths. No clear difference between the two types was observed either in optimal temperature for the germination or in strength of the resistance to salinity.     

Seasonal influence on reproduction in chimpanzees of gombe national park

Although wild chimpanzees are not seasonal breeders, there are seasonal effects on several aspects of chimpanzee reproduction. I examined the seasonal incidence of anogenital swelling in cyclic, pregnant, and acyclic female chimpanzees in Gombe National Park, May 1975–April 1992, and surveyed important reproductive events to determine whether there is a seasonal effect. I analyzed data by season (wet vs. dry) and seasonal quarter;early dry season = May–July;late dry = August–October;early wet = November–January;late wet = February–April. When data for the 17 years are combined, the percentage of females in each reproductive state remains consistent throughout the year. In a given month, 30–35% of subjects were in the cyclic category, 11–15% were pregnant, and 54–61% were acyclic. Cyclic females showed full swelling more often during the late dry season. Pregnant females exhibited anogenital swelling more often during the late dry and early wet seasons. Acyclic females also exhibited a seasonal effect with more anogenital swelling during the late dry season. There is no seasonal difference in frequency of live births (dry, 20;wet, 23). However, the timing of conception showed a seasonal effect (dry, 32;wet, 16). Consistent with earlier reports, the onset of postpartum cycles is highly seasonal;30 occurred during dry season, 9 during wet season. The occurrence of first full swellings for young females is also concentrated in the late dry season. It appears that the dry season is a time of great change for Gombe chimpanzee reproductive physiology. Previous studies indicated that seasonal changes in food availability play a role in increasing group size during the dry season and social contact between females can enhance cyclicity. Accordingly, I suggest that seasonal changes in diet may play a role, either directly (food content) or indirectly (social contact), to alter reproductive physiology.     

Genetic variations in the pre-feeding and pre-oviposition periods among four pure lines in the brown planthopper, <Emphasis Type="Italic">Nilaparvata lugens</Emphasis> (Hemiptera: Auchenorrhyncha:Delphacidae)

In the brown planthopper, Nilaparvata lugens Stål, traits related to ovarian development were compared among four pure lines predominantly expressing the designated characteristics: blackish short-winged (BS), yellowish brown short-winged (YS), blackish long-winged (BL), and yellowish brown long-winged (YL). The pre-oviposition period was longer in the following order: BS < YS < BL = YL line. The pre-feeding period, namely, the time of non-feeding after adult emergence, which was estimated from dry weight loss during starvation, was also longer in the same order: BS < YS < BL = YL. Vitellogenin, the precursor of yolk protein, first appeared in the hemolymph about 1½ days later after the initiation of feeding in respective lines. One-day’s starvation did not have any effect on the timing of vitellogenin’s appearance in the long-winged lines (BL, YL), but delayed it by 1 day in the short-winged lines (BS, YS). Further, we showed that the synthesis of vitellogenin mRNA was induced in the adults even before feeding by being topically applied with juvenile hormone III (JH III). These results suggest that feeding first triggers the increase of JH III titer, which activates vitellogenin synthesis, and enhances ovarian development. Thus, genetic differences in the pre-feeding periods cause the differences in the timings of physiological events relating to ovarian development among the four lines.     

昆虫翅型分化的调控及翅多型性的进化

翅多型现象普遍存在于各昆虫类群,一些学者就昆虫翅多型进行了大量的研究工作。根据昆虫翅型的分化,可划分为长翅型和短翅型,长翅型具飞行能力,而短翅型不能飞行。一些昆虫种类,如蚜虫,出现无翅个体,被称为无翅型。除飞行能力外,长翅型和短翅型在行为、生理等方面也存在差异。文章主要就环境因素对翅型分化的影响、翅多型的内分泌控制机理、翅多型的遗传机制及其进化等作一概述。     

Dealing with time constraints on development: the effect of food availability

Abstract.  1. Life-history theory predicts that organisms should speed up development in response to time constraints. However, acceleration of development carries energetic costs that have to be compensated, e.g. by an increase in foraging rate. For the wing dimorphic water strider Gerris lacustris (L.), the hypothesis was tested that the adjustment of development to time constraints is limited by the availability of food resources.
2. Six cohorts of larvae hatched increasingly late in the season were reared under two feeding regimes. For each cohort and experimental group the physiological time (in degree-days) of larval development was estimated.
3. In both high- and low-food groups there was a significant reduction of physiological time for development towards the end of the season. Furthermore, within cohorts, physiological development time was always lower in the high-food group than in the low-food group. However, there was no significant interaction effect between food treatment and cohort.
4. The results demonstrate that G. lacustris has the flexibility to adjust development to time constraints. In addition, 20% of the 'low-food individuals' developed into the short-winged morph while all of the 'high-food individuals' became long-winged. The limitation of food may thus lead to a reduced allocation of energy into the development of the flight apparatus. This may explain the strong increase in short-wingedness at the end of the season in natural populations, which are highly food limited.     

De-alation, flight muscle histolysis, and oocyte development in the striped ground cricket, Allonemobius fasciatus

ABSTRACT. Removal of hindwings from long-winged females of the striped ground cricket, Allonemobius fasciatus , DeGeer (Gryllidae), induces flight muscle histolysis and oocyte development. Such females develop oocytes as rapidly as do short-winged forms, while intact long-winged females retain their flight muscles and develop few oocytes.
Flight muscle histolysis occurs in starved long-winged females when they are de-alated. However, such females fail to mature oocytes. Implantation of corpora allata (CA) into long-winged females results in flight muscle histolysis as well as oocyte maturation even if their hindwings remain intact, indicating that flight muscle histolysis can take place without de-alation. It is likely that the CA are responsible for both flight muscle histolysis and oocyte development, and that CA activity is enhanced by de-alation.