Photoperiodic determination of gynoparae and males of damson-hop aphid Phorodon humuli

Abstract. Sexual morph production in Phorodon humuli is controlled by daylength. Wingless aphids reared from birth in short-day conditions (LD 12 : 12 h) and transferred when adult to long-day conditions (LD 18 : 6 h) produce only gynoparae and males until death some 7 weeks later, whereas those reared in long days and transferred to short days produce 20% wingless parthenogenetic females, 50% gynoparae and 30% of males in an overlapping sequence. No winged morph capable of re-infesting hop is produced. Less mature embryos are more sensitive to short days than older embryos because 100% of the former became gynoparae after 4 days of exposure of their mothers, and 59% when their mothers were exposed for the 4 days immediately preceding the birth of their offspring. Two days of exposure to short days switches 94% of young embryos from wingless to gynopara production when mature. The response to short days is irreversible. Wingless aphids reared from birth to adult in short days produce 30% fewer offspring than those reared to the same stage in long days and are male-biased, with 76–78% of their offspring being male.     

Photoperiodic induction of winged females in the black bean aphid. Aphis fabae

Abstract. The Photoperiodic of winged females (alatae) in the black bean aphid, Aphis fabae Scop. (Homopetera: Aphididae), is investigated in detail with emphasis on the interaction of the maternal and embryonic/young larval photoperiodic clocks. Previous work had shown that in uncrowded conditions the induction of gynoparae (winged females that produce sexual females) requires both prenatal and postnatal exposure to long-night (12 h) Photoperidic cycles: present results show that sole postantal exposure to long nights of any lenght does not induce wing formation in early-born aphids.
When aphids were exposed to experimental light-dark cycles postanatally only, their daughters developed as alate in long nights and as apterae in short nights: the critical night lenght (CNL) was 11:1 h. Additional prenatal exposure to experimental regimes resulted in a significantly shorter CNL (10.6 h). This difference could be accounted for by the fact that more experimental light-dark cycles were experienced in the latter case.
Apterous aphids transferred from LD 16:8 h to LD 12:12 h as either third-or fourth-stadium larvae, or young adults, switched for aptera-production to alata-production. The transition form aptera- to alata-production was rather abrupt in third-stadium transfers but more gradual when transfers occurred as fourth-stadium larvae and adults. Moreover, s the number of days required for 50% of the aphids to become alata-producers increased from 7–8 in third-stadium transfers, to 9–10 and 11–12 in the later transfers.     

Lipid composition in long-day and short-day forms of the black bean aphid,Aphis fabae

Long-day reared winged (alate) virginoparae from laboratory stock cultures which had been reared throughout larval development on bean stipules were significantly smaller (0.46+/-0.02 mg; mean+/-SEM) than short-day-reared gynoparae (0.69+/-0.04 mg; the winged autumn migrant) which completed development on intact beans. When winged virginoparae were raised from the third stadium on bean seedlings they grew larger (0.86+/-0.02 mg) but the gynoparae contained proportionally more total lipid (12.1+/-0.4%, gynoparae; 7.4+/-0.6%, stipule-reared virginoparae; 9.2+/-0.8%, seedling-reared virginoparae). Wingless aphids (apterae) were heavier, whether reared in short (0.99+/-0.03 mg) or long days (0.95+/-0.04 mg) but the lipid content was low (4.5+/-0.7% and 4.9+/-0.6%, respectively). The triacyl-, diacylglycerol and the phospholipid contents followed this trend but analysis of the fatty acid moieties of the triacylglycerides showed phenotypic differences. The ratios of myristic acid:palmitic acid were significantly higher in the winged forms than the wingless forms and were much higher in the gynoparae than the winged virginoparae. Short-day-reared wingless females also had a higher myristic acid:palmitic acid ratio than long-day-reared apterae, possibly reflecting the embryonic gynoparae maturing in their ovaries.     

Reproductive, morphological and behavioural affinities between the alate gynopara and virginopara of the aphid, Aphis fabae

ABSTRACT. 'Normal' alate virginoparae reared under long day conditions (LD 16:8) with postnatal crowding, and 'normal' gynoparae reared in un-crowded, short day conditions (LD 12:12) over two generations were compared in respect of the morph of the progeny, the frequency distribution of secondary rhinaria (placoid sensilla) on the third and fourth antennal segments, and preferred larviposition sites. Although in Aphis fabae the two morphs closely resemble one another morphologically, they differed in each of the above characters. Progeny sequences were analysed after transfer to the opposite photoperiodic regime at various times during postnatal development of the alate mothers. Transfer of presumptive gynoparae from short to long days induced the appearance of virginoparous progeny even when the transfer was made just after the final moult. Transfer of presumptive alate virginoparae from long to short days was effective in inducing oviparous offspring, only if the transfer was made during the first instar; oviparae were always first born. Intermediates between oviparae and viviparae were found at the time when the morph of the progeny changed. Transfer from long to short days also induced gynopara production but there appeared to be an intrinsic mechanism which prevented exclusive gynopara production by alates. When first instar, presumptive gynoparae were transferred to long days, the resulting adults would be described both reproductively and behaviourally as alate virginoparae; the opposite switch, of presumptive alate virginoparae to short days, resulted in adult gynoparae on the same criteria. However, the numbers of secondary rhinaria were influenced more by the prenatal than by the postnatal photoperiodic regime. There was an asymmetry of response in that a switch from short to long days had a greater effect on all three characters monitored than did the opposite switch from long to short days.     

Production of sexual morphs by apterous virginoparae of Myzus persicae (Hemiptera: Aphididae) in relation to pre- and postnatal exposure to short day conditions

The production of males and females by apterae of a holocyclic clone of Myzus persicae (Sulzer) was studied in the laboratory by transferring apterous virginoparae from long day to short day conditions at different stages of their pre- and post-natal development. Prenatal exposure was also examined in nine additional clones coming from different regions of Greece. By increasing the number of short day cycles at 17 degrees C, the aphids switched to male production earlier and produced more males. The highest mean number of male progeny was observed in apterae that were transferred to short day conditions 13-14 days prenatally. Apterae that were transferred to short day conditions after the fourth nymphal instar produced only females. Apterous virginoparae were produced by aphids that were transferred to short day conditions 0-1 days prenatally or postnatally. Aphids, given short days prenatally or in the first instar, produced alate female progeny that were all gynoparae. The later the instar aphids were transferred to short day conditions the fewer the gynoparae produced among their female progeny. A reduction in the total number of progeny and embryos per ovariole was observed as aphids were transferred to short day conditions in earlier stages of their development. The increase in male production, which follows the increase in the duration of long night exposure, was also observed in the additional eight out of nine examined clones.     

Juvenile hormone and photoperiodically controlled polymorphism in Aphis fabae: Postnatal effects on presumptive gynoparae

Long days (short nights) (LD 16:8) and high temperatures (> 15°C) have an apterizing effect on the short day (LD 12:12) induced, presumptive gynopara of Aphis fabae. Transfer of presumptive gynoparae to long days (15°C) or to 25°C (short days) at varying times during postnatal development demonstrate that the adult form is determined by the second day of the second instar, i.e. 5 days after birth at 15°C. Transfer on day 1 induces maximum apterization with the proportion of aphids affected decreasing with age at transfer.Apterization induced by long days immediately after birth can, to some extent, be cancelled by return to short days but only up to day 4. Thus long days are morphogenetically more potent than short days at the beginning of larval development. At temperatures above 15°C the proportion of aphids apterized increases almost linearly.Apterized insects can be distinguished from juvenilized insects in the fifth-instar. Topical application of juvenile hormone (JH) induces both apterization and juvenilization of presumptive gynoparae but at different times during larval development, JH treatment during the early-instars promotes apterization but induces little juvenilization, whereas maximum juvenilization, without apterization, is produced by middle-instar treatment. The apterizing effects of JH are, thus, not due to its neotenic action.The response profile of JH-induced apterization is similar to that observed with long days and 25°C. It is suggested that such conditions increase endogenous JH levels in A. fabae. The three naturally occurring JH's differ in activity in the order JH I > JH II > JH III. Both long-day and JH-apterized insects switch from the normal ovipara production of the adult gynopara to vivipara production.     

Morph determination in the bird cherry-oat aphid, Rhopalosiphum padi L

Both crowding and poor nutrition induce the appearance of emigrants in Rhopalosiphum padi L. No emigrants developed when the aphid was reared in isolation for six successive generations on actively growing leaves of bird cherry. However, emigrants developed as soon as the leaves ceased to grow or when the aphids became crowded. Crowding of mothers and postnatal crowding of the nymphs of apterous exules both influenced the induction of alate exules. The highest proportion of alate exules developed when both mother and offspring were crowded. Short day-length induced the appearance of gynoparae and males. For 50% production of gynoparae and males, over the range of temperature 10–18d?C, a 1.75 h reduction in day-length is required for every 4d? increase in temperature. At 18 d?C, even at day-lengths as short as 6 h, exposure for three generations is required before all offspring become gynoparae or males. Low temperature and short day-length was ineffective in inducing the development of gynoparae or males in the first three generations developing from the fundatrix.     

An interval timer controls the production of sexual morphs in Myzus persicae (Homoptera: Aphididae)

Abstract The interval timer mechanism was examined in two clones of Myzus persicae (Sulzer) originating from peach in Naousa, North Greece. Overwintering eggs were collected in April 1997 and after egg hatch the morphs of progeny were recorded at LD 10 : 14 h and 17 °C until the appearance of sexuals. In one clone, observations were continued until the 21st generation after egg hatch. In addition, the production of sexual morphs was examined in two other clones originating from Lehonia, Central Greece, when apterae (wingless females) were transferred from long to short days and 17 °C in the second and eighth generation of parthenogenetic rearing after their collection from peach trees. In clones from Naousa, the first gynoparae appear and comprise 10% of alate (winged) females in the second half of the third generation after egg hatch. The percentage of gynoparae increased in each consecutive generation, reaching 100% in fifth and seventh generations in clones Nb and Na, respectively. First males appeared later than gynoparae, in fifth and sixth generation in clones Nb and Na, respectively. In clone Nb, the number of males, after their appearance in fifth generation, was relatively constant from the sixth to the ninth generation (10.8–12.3). Then a significant increase was observed in the 10th generation, thereafter remaining relatively constant until the 21st generation (18.1–19.8). A similar trend in the number of males produced was observed in clones deriving from Lehonia.     

The time of determination of gynoparae and males in the bird cherry–oat aphid, Rhopalosiphum padi

Apterous exules of Rhopalosiphum padi which experienced short photoperiods only during the first half of their nymphal life all produced gynoparae which made up 57% of the offspring produced in the first 7 days of the mother's reproductive life. Short photoperiods during the second half of a mother's nymphal life did not induce the production of gynoparae. However, when short photoperiods were experienced throughout a mother's nymphal life significantly more gynoparae (82%) were produced. Ten per cent of the offspring of mothers that experienced short photoperiods only during their adult life developed into gynoparae. Of the offspring reared in short photoperiods, but born to mothers reared and kept in long photoperiods, 30% developed into gynoparae. When mothers were exposed to short photoperiods in their adult life and their offspring so treated, then 90% of the offspring developed into gynoparae. Males only occurred amongst the last offspring of mothers that experienced short photoperiods in either the first or the second half of their nymphal life.     

Photoperiodic regulation of the development of winged gynoparae in the aphid, Aphis fabae

ABSTRACT. In uncrowded rearing conditions, determination of the winged gynopara of the Aphis fabae Scopoli is dependent upon both pre- and postnatal short days (LD 12:12). Extension of the photophase to produce longday (LD 16:8) conditions localized on the head or abdominal region, showed that during the early larval instars of the parent generation (G1) there is a maternally-controlled induction of gynoparae. However, during later stages of embryonic development the presumptive gynoparae respond directly to photoperiod through the mother's abdominal wall. During early postnatal development, presumptive gynoparae (G2) remain sensitive to photoperiodic conditions; the photoperiodic receptors lie in the head. These results are discussed with respect to possible endocrine mediators of this photoperiodically controlled polymorphism.     

The cost of being able to fly in the milkweed-oleander aphid,Aphis nerii (Homoptera: Aphididae)

Summary In the wing dimorphic milkweed-oleander aphid,Aphis nerii, winged aphids begin reproducing about 1.5 days after wingless aphids. The longer maturation period is primarily due to slower development since even adult eclosion by winged aphids takes place after wingless aphids begin reproducing. The delay is not due to a post-eclosion, pre-reproductive flight since, beginning with the fourth instar, larval winged aphids were reared at a density of one per plant and the vast majority were not stimulated to fly under such low-density conditions. Thus, the ability to fly incurs a fitness cost in terms of delayed reproduction, irrespective of whether flight actually occurs. We did not observe a difference between morphs for lifetime fecundity, even though wingless aphids have larger abdomens than winged aphids and for both morphs there is a significant correlation between abdomen width and fecundity. Offspring produced by wingless aphids over the first four days of reproduction are larger than those produced by winged aphids, and the size difference at birth is maintained into adulthood. However, there are no differences in life history traits between these offspring, including maturation period and lifetime fecundity. Thus, reduced body size does not increase the cost of being able to fly, at least under the conditions of these experiments. The cost of being able to fly in this species should favor reduced production of winged individuals in populations that exploit more permanent host plants.     

Pheromone Traps for the Dissemination of an Entomopathogen by the Damson-hop Aphid Phorodon humuli

Two types of sex pheromone-emitting trap, a simple louvred trap and a commercially-produced wasp trap (Waspy®) designed or modified as delivery systems for entomopathogens, were tested using an isolate of the pathogen Verticillium lecanii . Their efficiencies for capturing winged autumn females (gynoparae) and males of Phorodon humuli in a hop garden and a plum orchard were compared with each other and with catches by yellow-painted water traps. Gynoparae rarely entered either type of pathogen dissemination trap. Males only entered traps that released (1 R , S ,4a R , 7 S , 7a S )-nepetalactol, the sex pheromone of P. humuli . In the hop garden, approximately five times as many males entered the modified Waspy® as entered the louvred traps, whereas in the plum orchard similar numbers of males were caught in both traps. On average, males visited the modified Waspy® trap for 34.1 ±7.9 (SE) min. Males spent about 50% of their time in the arena where the entomopathogen would normally be present and a further 20% of the time on the inside of the arena lid alongside the pheromone source. Out of a sample of 16 live aphids removed from a trap dispensing V. lecanii , 9 died from infection by the pathogen and 5 of the 16 initiated colonies of the fungus after they walked on a sterile agar plate for five minutes. None of the 15 aphids collected from traps without V. lecanii became infected or initiated colonies of the fungus on agar plates.     

The photoperiodic control of wing development in the black bean aphid,Aphis fabae

Newly born presumptive gynoparae of Aphis fabae were transferred from their prenatal short-day rearing conditions (light-dark 12:12, 15°C) to a variety of postnatal photoperiodic regimes. Long days prevented wing formation and the majority of aphids developed into apterous or alate-apterous intermediate adults. Continued short days resulted, almost exclusively, in winged adults. The photoperiodic-response curve (T = 24) revealed a critical photoperiod of light-dark 13.5:10.5 and further investigations showed that morphogenesis depended more upon night- than day-length. Maximal apterization occurred with 8 or 9 h dark regardless of the length of the photophase but at photophases of 12 h or less the critical night length was reduced to 8.5 h. Night-interruption experiments revealed two peaks of photosensitivity when the scotophase was 12–14 h long but only a single peak was seen with longer dark periods. A series of experiments using early night interruptions followed by extended dark revealed apparent critical night lengths which decreased as the interruption was placed later in the scotophase. Resonance experiments involving 12 or 16 h photophases and extended scotophases at 15°C revealed long-day effects with scotophases shorter than critical and short-day effects with longer dark periods. However, similar regimes at 20°C produced three peaks of apterization 24 and 20 h apart indicating the possibility of a circadian element involved in the photoperiodic response. The results are compared with the photoperiodic responses of other insects.     

Host-selection behaviour by genetically identical aphids with different plant preferences

The behaviour of summer and autumn winged forms of the black bean aphid, Aphis fabae Scopoli (Homoptera: Aphididae), was compared on two plants utilized at different stages of the insect’s life cycle. Adult autumn migrants (gynoparae) are monophagous, colonizing spindle (Euonymus europaeus), whereas polyphagous summer winged aphids (alate virginoparae) are associated with a variety of herbaceous plants, including broad bean (Vicia faba). When aphids from a single clone were given access to a spindle leaf and a bean seedling in choice tests, many virginoparae settled and larviposited on both plant species over 24 h. By contrast, gynoparae showed a clear preference for spindle, with 93.5% of settled adults and 98.3% of larvae on this plant species. Close‐up video monitoring showed that gynoparae discriminated beans from spindle within a 5‐min period, whereas virginoparae behaved similarly on both plant species. For gynoparae, the major behavioural difference on the two plants appeared after a brief (epidermal) stylet penetration, with many insects taking flight within a few seconds of stylet withdrawal from bean. Factors detected during stylet insertion by gynoparae must therefore inhibit take‐off on spindle. Electrical recording experiments showed that aphids often punctured a cell membrane during brief probes on both plant species, and intracellular stylet activities always included a waveform associated with ingestion. When gynoparae puncture spindle cells their behaviour is probably modified by intracellular metabolites detected via gustation of ingested epidermal cell sap. These cues may inhibit the take‐off reflex which otherwise follows probing.     

Observations on the effect of photoperiod on the control of polymorphism in Psylla pyricola

ABSTRACT. Psylla pyricola Forster (Homoptera) occurs as two adult morphs, a summer form (f. typica) and a winter form (f. simulans). The latter is larger, darker and overwinters in ovarian diapause. Eggs laid by either morph under LD 12:12 (short days) or under LD 18:6 or 16:8 (long days) gave f. typica when reared under long days. When reared under short days, almost all were f. simulans. No interforms were recorded. When the photoperiod was switched during the nymphal stage (whether from long to short days, diapause induction; or from short to long days, diapause termination), nymphs were progressively less sensitive with increasing age, with no morph changes when the switch occurred in the fifth instar. Nymphs derived from f. simulans reared under long days tended to be more sensitive to a change to short days in the early instars than when the nymphs were first reared under short days and then switched to long days. Interforms were frequent when the switch occurred during the nymphal stage, particularly in the third instar. Wing lengths varied within morphs depending on the timing of the switch. The largest f. simulans were derived from f. typica when maintained under short days, and the smallest f. typica from either morph when maintained under long days, as would be expected under field conditions. No f. simulans were found under field conditions in the first f. typica generation, but a small percentage were present in the second generation. It is concluded that, even within a given generation, not all P. pyricola are equally sensitive to photoperiod, but that photoperiod is one of the major factors controlling the change to f. simulans in the autumn. Whilst under laboratory conditions photoperiod can also affect the switch to f. typica in the spring, in the field it may be less important than low temperatures.     

Some effects of photoperiod on larviposition and fresh weight-gain in Myzus persicae

ABSTRACT. The larviposition of adult apterous Myzus persicae previously entrained to LD 18:6 showed a marked rhythm both under LD 12:12 (on plants reared in LD 18:6) and under LL (on plants reared in LL). No rhythm was detectable in larviposition by adults reared in LL. Larviposition peaked towards the end of the photophase in LD 12:12. Fresh-weight gain also showed a circadian rhythm with the greatest weight increases during the photophase. Re-entrainment from LD 12:12 to DL 12:12 was not complete after 10 days. It is concluded that the changes in the light cycle did not affect the aphids through the plant.     

Differential photoperiodic responses in genetically identical winged and wingless pea aphids, Acyrthosiphon pisum, and the effect of day length on wing development

Abstract. Winged and wingless individuals of a pink clone of the pea aphid, Acyrthosiphon pisum (Harris), showed differences in the response curves for photoperiodic induction of both males and sexual females (oviparae). The critical night length (CNL) for ovipara induction in winged aphids was 0.75 h shorter than in wingless aphids, whereas the CNL for male induction in winged aphids was 1.0h longer than in wingless aphids. This means that in winged aphids the CNL for male induction in winged aphids was 0.5 h longer than that for ovipara induction, while in wingless aphids the CNL for male induction was 1.0–1.5 h shorter than that for ovipara induction, and also the shapes of the curves differed.
Winged aphids were produced by wingless mothers which were crowded as young adults. However, when young adults were crowded in long nights, winged aphids were not produced, and the CNL for wing inhibition was between 9.5 and 10h. This effect of photoperiod on wing induction was maternal.     

禾谷缢管蚜复眼外部形态的扫描电镜观察

【目的】通过比较禾谷缢管蚜Rhopalosiphum padi两种生活史(全周期型和不全周期型)及5种蚜型(有翅孤雌蚜、无翅孤雌蚜、雄蚜、雌性蚜和产雌性母)成蚜复眼外部形态,分析了其视觉能力差异,为进一步探索禾谷缢管蚜生殖转换及寄主搜寻机制提供参考。【方法】利用扫描电镜技术,观察全周期型和不全周期型禾谷缢管蚜无翅孤雌蚜及其低温短日照(12℃, 光周期8L∶16D)诱导条件下全周期型所产5种蚜型成蚜的复眼外部形态。【结果】禾谷缢管蚜成蚜复眼着生于头部触角基部,呈卵圆形,小眼间无感觉毛。全周期型和不全周期型禾谷缢管蚜无翅孤雌蚜的复眼大小(长轴直径及短轴直径)、复眼弧高、小眼数量和小眼直径均没有显著差异。而这5个复眼指标在低温短日照诱导全周期型禾谷缢管蚜产生的5种蚜型的成蚜中均存在显著性差异,其中,雄蚜、有翅孤雌蚜和产雌性母的复眼区域较大,小眼数量较多;雌性蚜的复眼区域最小,小眼数量也最少。【结论】综合以上指标结果,推测5种蚜型的视觉能力由大到小是：雄蚜＞有翅孤雌蚜＞产雌性母＞无翅孤雌蚜＞雌性蚜。复眼和小眼的这些差异可能与禾谷缢管蚜生殖转换及寄主搜寻存在一定的联系。     

Electroantennogram responses of the three migratory forms of the damson-hop aphid, Phorodon humuli, to aphid pheromones and plant volatiles

Electroantennogram (EAG) responses were recorded from alate fundatrigeniae (spring migrants), gynoparae (the winged female form that produces sexual females) and males, the three migratory forms of the damson-hop aphid, Phorodon humuli (Schrank). EAG responses of gynoparae and males showed typical dose response characteristics to (E)-2-hexenal, (-)-R-carvone, hexanenitrile and (1RS,4aR,7S,7aS)-nepetalactol, the sex pheromone of this species. The 34 plant volatiles elicited broadly similar EAG response profiles in the three migratory forms. Green leaf volatiles produced large responses in all forms; however, the relative order of responsiveness varied. EAG responses to isomers of the monoterpene carvone differed between forms, with males being most, and spring migrants least, responsive. The hop-plant volatile and aphid alarm pheromone, (E)-beta-farnesene, evoked similar EAG responses in all forms. By contrast, males were most responsive to the three sex pheromone components, (-)-(4aS,7S,7aR)-nepetalactol, (+)-(4aS,7S,7aR)-nepetalactone and (1RS,4aR,7S,7aS)-nepetalactol. Males were no more responsive to their own sex pheromone, (1RS,4aR,7S,7aS)-nepetalactol, than to the other aphid sex pheromone components tested. Spring migrants and gynoparae also responded to the three sex pheromone components. This study indicates that migratory forms of P. humuli detect a wide range of volatile compounds, and that they are equally well-adapted for the detection of volatiles associated with host and non-host plants and with other species of aphid.     

Responses of holocyclic and anholocyclic Rhopalosiphum padi populations to low‐temperature and short‐photoperiod induction

The different life cycles of aphid species make these organisms good models for studying the short‐term consequences of sex. The bird cherry‐oat aphid Rhopalosiphum padi has a wide geographic distribution and correspondingly different life cycles. In this study, the life cycles of R. padi collected from six different regions in China were characterized experimentally by comparing the responses of holocyclic and anholocyclic populations to low‐temperature and short‐photoperiod induction. Clones collected from Chuzhou, Taian, and Taigu consistently reproduced via obligate parthenogenesis, whereas clones from Hami and Baicheng were holocyclic in their response, and those from Lanzhou were both holocyclic and anholocyclic. Prolonged exposure to low temperature and a short photoperiod (LS) had negative effects on the offspring of anholocyclic aphids with regard to adult lifespan, total longevity, and fecundity compared with aphids maintained at a normal temperature and a long photoperiod (NL). Holocyclic LS R. padi had longer developmental times at all nymph stages, a shorter adult lifespan, shorter total longevity, and a lower fecundity than NL counterparts. The adult prereproduction period of gynoparae was significantly longer than that of virginoparae, and the total longevity of gynoparae was significantly shorter than that of virginoparae. Moreover, the net reproductive and gross reproduction rates, as well as the total fecundity, were roughly fivefold higher in virginoparae than in gynoparae, indicating that there is the short‐term cost of sex. When maintained on their secondary host (Triticum aestivum), gynoparae, males, and oviparae produced by holocyclic populations could survive, and gynoparae produced oviparae. However, under NL conditions, oviparae could not produce overwintering eggs on the secondary host, whereas a few overwintering eggs were generated by oviparae under LS conditions. Taken together, these results illuminate the complexity of insect responses and contribute to a complete understanding of the aphid life cycle and its evolution.