Male production by apterous viviparae of the aphid Myzus persicae temporarily exposed to different scotoperiods

Male production by apterous viviparae of a holocyclic biotype of the green peach aphid, Myzus persicae, was induced by pre-natal and/or post-natal exposures to a long-night regime of 15 h dark per diem. When the apterae were exposed to three or more long nights immediately after birth and they subsequently developed under a short-night regime of 8 h dark per diem, they produced females (mostly alate viviparae) during the first 8–10 days of reproduction and a high proportion of males thereafter. When the apterae were exposed to two long nights immediately before their birth and to short nights thereafter, they produced relatively few females (mostly alate viviparae), and males were deposited already after 4–6 days of reproduction, i.e. 16–18 days after the first exposure to the long nights. The proportion of males among the progeny of these apterae was highest when the two prenatal exposures comprised scotophases of 11–15 h; under such long-night regimes many aphids switched to producing males exclusively. To achieve this effect, the two long scotophases had to be separated by a photophase of more than 1–2 h. Fewer males were produced and most of the apterae reverted to the production of females (apterous viviparae) when the duration of the two prenatal scotophases was 9 h 45 min-10 h 30 min, or 18 h and longer.One long night of 15–39 h could also induce temporary male production if the aphids were exposed to it late in the 4th larval instar or as teneral adults.     

Embryogenesis and oögenesis in alate virginoparae,gynoparae, and oviparae of the aphid Myzus persicae, in relation to photoperiod

The study reports on the effects of prenatal and/or postnatal exposures to short-night or long-night conditions, and of crowding, on embryogenesis and oögenesis in alate virginoparae, gynoparae, and oviparae of a holocyclic strain of the green peach aphid, Myzus persicae, from Yakima, Washington State.In alate virginoparae raised at a density of 10–20 per radish seedling in a short-night regime (8 hr darkness per diem), 3–4 embryos occurred in each of their 10 ovarioles, when the aphids attained adulthood. More than 30 larvae were deposited by most of these alatae. However, in young adult gynoparae, raised at these densities in a long-night regime (15 hr darkness per diem), only one viable embryo (a presumptive ovipara) occurred per ovariole. The follicle containing this embryo was followed by 1–2 abnormal follicles in each ovariole, and the number of larvae deposited by a gynopara was generally less than 10. In young adult oviparae similarly raised under a long-night regime, only one egg typically occurred in each of their 10 ovarioles, and the eggs deposited by an ovipara (only after it had mated) generally numbered less than 10. Alate virginoparae and gynoparae contained an additional embryo in some of their ovarioles when these morphs were raised at a lower density (1–5 per plant).Presumptive gynoparae partially developed the reproductive features of alate virginoparae when transferred to a short-night regime at birth; the converse was true when presumptive alate virginoparae were transferred to a long-night regime early in larval life. Oviparae maintained in short nights from before birth developed the appearance of apterous virginoparae but still produced eggs rather than embryos. However, their oögenesis was enhanced and eggs (10–20) were deposited by them without prior mating. Under all regimes tested, oviparae were always deposited early in the larviposition sequence of their alate mothers, and the number of oviparae deposited never exceeded 15.The possible involvement of juvenile hormone in the regulation of these events and the ecological significance of the results are discussed.     

Inhibition by kinoprene of photoperiod-induced male production by apterous and alate viviparae of the aphid Myzus persicae

Photoperiod-induced male production by apterous and alate viviparae of the green peach aphid. Myzus persicae, was reduced or prevented by treating the aphids with the JH analog kinoprene (Zoecon's insect growth regulator ZR 777). Concomitant with this reduction in the number of males was an increase in the production of females. This was most clearly demonstrated for apterous viviparae that had been raised from birth under weak long-night regimes of 10–10.5 hr dark day, and exposed to 0.1% kinoprene-treated radish seedlings for 2–3 days on reaching the fourth larval instar or adulthood.Male production by alate viviparae, whose apterous mothers were born and raised for 5 days under a long-night regime of 15 hr dark day and then maintained under a short-night regime of 8 hr dark day, was completely inhibited by exposing them to kinoprene-treated plants for 2–3 days when they had reached the fourth instar.It is concluded that the kinoprene treatment affects sex chromosome replication in the oögonia of an aphid so that these germ cells develop parthenogenetically into females (2n = 12 with two X chromosomes) rather than males (2n = 11 with one X chromosome). This may result either from a direct action of absorbed kinoprene on the oögonia or from an indirect action whereby the kinoprene stimulates the insect's hormonal system to produce a hormone that stimulates development of the oögonia into females.     

Observations on the morphs of Macrosiphum avenae and Metopolophium dirhodum on cereals during the summer and autumn

From June to early August 1970, populations of Macrosiphum avenae and Metopolophium dirhodum on marked tillers of field barley were compared with the numbers of alatae trapped at crop height and at 12.2 m. There were always more M. dirhodum than M. avenae on the tillers. Only apterae were produced until mid-June when both alatae and apterae occurred; after mid-July only alate M. avenae were found. Until mid-June most of the flying alatae were caught at 12.2 m as they migrated from spring hosts to cereals. Thereafter, more alate M. avenae were trapped at 12.2 m than at crop level, whereas numbers of alate M. dirhodum were usually comparable at both heights. Although crop and flying populations occasionally showed temporal similarities, insufficient is known about their field distribution and the factors affecting their alate production and flight activity to interpret this relationship. In the autumn, two consecutive reproductive phases of M. dirhodum occurred on winter wheat grown in pots outdoors. Initially, apterous virginoparae and alatae, probably sexuparae, were produced, whereas only alate males appeared during the second phase. In contrast, M. avenae deposited mainly apterous virginoparae although some oviparae developed in October to lay scattered, probably infertile eggs on the tillers.     

桃蚜不同蚜型DNA多态性的RAPD研究

采用RAPD方法,对全周期桃蚜的有翅产雌性母蚜、无翅性母蚜、雄蚜、卵、干母、干雌、有翅迁移蚜等蚜型的DNA遗传多态性进行了分析。结果表明：卵的DNA多态性最大,性蚜次之,孤雌生殖蚜最小;卵与其它蚜型之间在遗传上具最大差异,其中与孤雌生蚜的差异大于与性蚜的;干母、干雌和迁移蚜之间的遗传关系最近。     

Precocious metamorphosis of the aphid Myzus persicae induced by the precocene analogue 6-methoxy-7-ethoxy-2,2-dimethylchromene

For up to three days after being treated with the precocene analogue 6-methoxy-7-ethoxy-2,2-dimethylchromene, adult green peach aphids (Myzus persicae) gave birth to offspring (alate and apterous virginoparae, males) which underwent precocious metamorphosis. Only a few precocious gynoparae and no precocious oviparae were recorded. The precocious aphids became adultoid in the third or fourth-instar as a result of corpus allatum destruction. They were able to develop mature embryos, but could not deposit them because of incomplete differentiation of the reproductive tract. The treatment did not induce the production of alate virginoparae in the experimental clone of Myzus persicae. However, a few males were born late in the reproductive sequence of treated apterae. The corpora allata of the treated adults appeared on histological examination to be unaffected by precocene.     

Biological studies onTrichogrammatoidea armigera Nagaraja a new dimorphic egg parasite ofHeliothis armigera (Hübner) in India

A new species ofTrichogrammatoidea which is being described byH. Nagaraja asT. armigera, has been reared from eggs ofHeliothis armigera onPolianthes tuberosa and from those of an unidentified Lepidopteron onCajanus cajan. In the laboratory the parasite was successfully bred onCorcyra cephalonica, Achaea janata, Gnorimoschema operculella andPlutella xylostella; it did not show any perceptible preference for any of these hosts. It also parasitised eggs ofSpodoptera litura, but although development proceeded to the adult stage, adults failed to emerge, suggesting unsuitability of this host. The males ofT. armigera are found to exhibit dimorphism — one form being alate and the other typically apterous. The apterous males were almost exclusively produced only by fertilised females and in the progeny of virgin females these forms were extremely rare (1 apterous male: 1,500 winged males). Among the progeny of a single mated female, an apterous male developed invariably in association with a female, but this rule did not apply when a singleCorcyra egg was parasitised by more than one parental female. However, in no case did an apterous male alone emerge from a single host egg. The biology ofT. armigera has been studied at 25°C±1°C. and R.H. 75%, usingC. cephalonica eggs. The parasite completed its life-cycle in 7–9 days — the egg, larval and pupal periods occupying, 1, 2–3 and 4–5 days, respectively. When fed honey, the average longevity of females was 7 (max. 11) days, of alate males 6 (max. 10) days and of apterous males 1 (max. 2) day. The maximum fecundity was 118 while the average was about 63. From 2–26 (average 9) eggs were parasitised per day. The sex-ratio was 62% females: 38% males (35% alate and 3% apterous). An alate male during its lifetime inseminated upto 10 (Av. 9) females while an apterous one inseminated up to 4 (Av. 3) females.     

Precocene II-induced alate production in isolated and crowded alate and apterous virginoparae of the aphid, Macrosiphum euphorbiae

Precocene II was applied at doses ranging from 0.05 to 0.70 μg per individual to newly moulted adult alate and apterous virginoparae of Macrosiphum euphorbiae kept isolated or in groups of 10 per plant, under an 18L:6D photoperiodic regime. While isolated controls of both morphs produced exclusively apterous progeny, alatae virginoparae were produced in generally dose-dependent proportions by precocene-treated individuals. Grouped controls of both morphs produced alatiform progeny as expected, but in precocene—treated groups, the proportions of alate progeny generally increased as a function of dose. The overall proportions of alate offspring produced, and numbers of days after treatment when morph production was affected, were generally greater for alatae than for apterae, indicating a greater sensitivity to precocene in alatae. However during the first few days after treatment, the alatizing effect of precocene was stronger for apterae, suggesting that the first embryos produced by alatae were irreversibly determined as apterae.In an experiment where isolated alatae and apterae received 0.5 μg of precocene II at different ages ranging from 1 to 13 days after the adult moult, the alatizing effect of the compound, measured by the persistence of alate production, varied with age and morph. While in alatae, the persistence decreased more or less regularly with age, in apterae it initially increased to a maximum in the middle of reproductive life, and subsequently decreased. The results provide support for the hypothesis that juvenile hormone is involved in regulating alary dimorphism in M. euphorbiae.     

The induction of normal and teratoid viviparae by a juvenile hormone and kinoprene in two species of aphids

ABSTRACT. Topical application of a juvenile hormone (JH1) and the JH mimic, kinoprene, to short-day, ovipara-producers of Megoura viciae usually leads to the production of oviparous/viviparous intermediate forms in the progeny sequence, in place of the expected sexual females (oviparae). The ovaries of these abnormal forms may contain embryos rather than the haploid yolky eggs of oviparae; 'mixed' ovaries containing both haploid eggs and embryos are also observed. The intermediates range in form from winged (alate) to wingless (apterous). The fully alate individuals usually contain only parthenogenetic ovaries but differ from the naturally occurring alate viviparae in that they are invariably infertile, have fewer antennal sense organs and often bear pheromone releasing glands on the metathoracic tibiae. The hormonally induced production of normal viviparae is difficult in this species but has been achieved by rearing short-day aphids on kinoprene-treated bean plants.
In Aphis fabae similar oviparous/viviparous teratomorphs have been reported and, in addition, single JH1 treatments were shown to induce normal viviparae at the end of the progeny sequence. Multiple applications, beginning prenatally and continuing through the postnatal development of the gynopara (winged ovipara-producer), showed that the numbers of viviparae born were related to the earliness of the treatment and to the dosage. The results are compared with the effect of a switch in photoperiod and discussed in relation to the endocrine control of aphid polymorphism.     

HOST ALTERNATION IN APHIS FABAE SCOP.

Experiments on the feeding preferences and relative fecundity of Aphis fabae among leaves of different ages and kinds were extended to the comparison of different forms of the aphid: apterous and alate virginoparae and gynoparae.
All three forms showed a preference for the primary host, Euonymus , over a secondary host, sugar beet, but this preference was strongest in the gynoparae, weaker in the alate virginoparae and weakest in the apterae. The relative fecundity of the aphids on the two kinds of leaf paralleled their feeding preferences most closely in the gynoparae, less so in the alate virginoparae and least in the apterae.
All three forms also showed some preference for growing over mature leaves of the same kind, but this preference was strongest in the apterous virginoparae, weaker in the alate virginoparae and weakest in the gynoparae. The relative fecundity of the aphids on the two ages of leaf paralleled their feeding preferences most closely in the apterae, less so in the alate virginoparae and least in the gynoparae.
The physiological, ecological and evolutionary significance of the results is discussed in the light of the dual discrimination theory of aphid host selection.     

Wing development in parasitized male and female Sitobion fragariae

Abstract.Different stages of presumptive winged morphs (males, gynoparae and alate virginoparae) of the blackberry‐cereal aphid, Sitobion fragariae , were exposed to attack by Aphidius ervi . Even though the mechanism influencing wing development in the three aphid morphs differs, the effects of parasitism were similar. Alatiform structures were completely inhibited in all three morphs when the initial attack took place in their first or early second stadium. The disruption of wing development also resulted in apterous/alate‐intermediate forms when aphids were attacked from first (males and gynoparae) or early second (alate virginoparae) up to the fourth larval stadium. The fact that wing development was still disrupted when aphids with well developed wingbuds were parasitized indicates that the early stages of parasitization were influential. Thus, the morphogenetic effects may be exerted by the parasitoid egg or calyx fluid.     

Photoperiodic determination of the male and female sexual morphs of Myzus persicae

The chronology of the photoperiodic determination of sexual morphs in holocyclic Myzus persicae was studied in the laboratory by transfer of synchronized batches of aphids between long-day (16 hr) and short-day (10 hr) régimes at a constant temperature of 20°C. The length of exposure to the short-day régime was measured in terms of the number of long dark-periods received by the aphids. The photoperiodic response extended over four generations (P, G₁, G₂, and G₃ respectively). When P generation aphids were given short days from the fourth instar, alate viviparae and males appeared successively in generation G₂, oviparae in G₃. Increasing the number of long dark-periods received during the development of G₂ embryos had a cumulative effect on the number which developed into alate viviparae. Determination of all the first-born G₂ aphids as alatae occurred only if their mothers had been exposed to the short-day régime throughout larval development. Alate viviparae gave birth to oviparae if they received a minimum of 4 long dark-periods, starting from a late stage in their embryonic development. The critical stage for ovipara determination in the G₃ embryo was on the sixth or seventh day after ovulation, more than half-way through embryonic development. G₂ aphids were determined as males before ovulation if their parents received 4 (in some circumstances only 3) long dark-periods. In the clones studied, male determination, once initiated in the G₁ parent, could not be reversed by later back-transfer to the long-day régime.     

榆梨绵蚜一些型与卵的新发现和记述（同翅目：蚜总科：瘿绵蚜科）

对榆梨绵蚜Ｅｒｉｏｓｏｍａ ｌａｎｕｇｉｎｏｓｕｍ ｄｉｌａｎｕｇｉｎｏｓｕｍ Ｚｈａｎｇ,１９８０的干母,无翅干雌,有翅干雌,无翅侨蚜,有翅性母,雌,雄性蚜放卵进行了描述,除有翅干雌外,其余各型与卵均为首镒记述。所有研究用标本均保存在中国科学院动物研究所昆虫标本馆内。     

Functional specialisation and polyphenism in aphid olfactory sensilla

Electroantennogram (EAG) responses to the aphid sex pheromone components, (1R,4aS,7S,7aR)-nepetalactol and (4aS,7S,7aR)-nepetalactone, and a plant volatile, (E)-2-hexenal, were investigated at three different positions (5/6th, 4/5th and 3/4th inter-segmental regions) along the antennae of four different morphs in two host-alternating aphid species, Aphis fabae Scopoli and Rhopalosiphum padi (L.). Position-dependent and morph-specific EAG responses were elicited in both species. The nepetalactol and nepetalactone isomers elicited large EAG responses at all three recording positions in males of both species, such that primary rhinaria as well as secondary rhinaria appeared to respond. Asexual female morphs showed relatively smaller EAG responses to these compounds. The secondary rhinaria, which have been reported as sex pheromone receptors in males, were not very different in their number and distribution between gynoparae and alate virginoparae, but the gynoparae showed significantly larger EAG responses to nepetalactol and nepetalactone. The alate virginoparae showed EAG responses that were similar to those of apterous virginoparae, which lack secondary rhinaria. Taking the EAG response profiles together with the distribution of the secondary rhinaria, it is suggested that the function of secondary rhinaria differs between the morphs. Secondary rhinaria appear to detect sex pheromone components in males and gynoparae but not in the alate virginoparae. If they are functional in the latter morph, they are likely to play a role in detecting specific, but as yet unknown, volatile compounds. Some 30 plant volatiles were tested but none evoked an EAG response that could be allocated to the secondary rhinaria. In contrast to the very different EAG response profiles to the pheromone compounds between morphs, EAG responses to (E)-2-hexenal were similar in all forms and both species. These findings suggest that this plant volatile was detected only by the two primary rhinaria, which are common to all morphs. The present study showed that EAG responses were not a simple summation of receptor potentials between recording and reference electrodes in aphids. The localised distribution pattern of olfactory receptor neurones around the recording electrode was also likely to contribute to the EAG responses.     

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.     

用微卫星引物PCR分析棉蚜不同蚜型的DNA多态性

用微卫星引物PCR方法, 对棉蚜Aphis gossypii干母、干雌、孤雌蚜(有翅迁飞孤雌蚜和无翅孤雌蚜)、性母、性雌蚜和雄蚜的DNA多态性进行分析。 结果表明：① 棉蚜的有性蚜型和无性蚜型之间在遗传上有较大的差异; ② 性母与性雌蚜和雄蚜的遗传关系都十分接近,说明同一性母既产性雌蚜,也产雄蚜;③ 干母和干雌的遗传关系很近, 但孤雌蚜已与二者有分化。     

Embryogenesis and the production of males by apterous viviparae of the green peach aphid Myzus persicae in relation to photoperiod

Apterous viviparae of a holocyclic strain of Myzus persicae were given various numbers of prenatal and postnatal exposures to male-inducing, long-night scotophases (15 hr darkness per diem). The sex of the embryos in various positions within their ovarioles was determined from chromosome preparations and correlated with the sequence with which female and male larvae were deposited. An abnormal follicle was found frequently between the female and male embryos in the ovarioles of aphids raised under long nights.Apterous viviparae exposed to long nights from birth or earlier usually produced several females and then switched over to exclusive male production. The more prenatal exposures to long nights given to the aphids, the fewer females were deposited by them and the earlier they started to produce males. When apterae were continuously exposed to long nights from 2 days or more before their birth, some, and occasionally all, their ovarioles were solely androparous. In general, male-producing apterae deposited considerably fewer larvae (female as well as male) than apterae that produced females exclusively. The 1–2 day pause in larviposition that typically occurs between the deposition of the female and male larvae was attributed to a slower rate of development of the male embryos. The results are discussed in relation to the hormonal changes that may be induced by the photoperiodic treatments.     

Key to Indian species of Periphyllus including four new species and hitherto unknown morphs from North-west Himalaya (Homoptera: Aphididae)

ABSTRACT. Four new species of Periphyllus; P.aceriphaga Chakrabarti & Mandal, P.caesium Chakrabarti & Saha, P.garhwalensis Chakrabarti & Mandal and P.pallidus Chakrabarti & Saha are described. Hitherto unknown apterous viviparous female of P.villosii Chakrabarti and four dimorphs are also recorded. Keys to distinguish the species of apterous viviparous females, alate viviparous females, alate males and dimorphs are provided.