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.     

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.     

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.     

Embryonic growth rate and birth weight of the offspring of apterous and alate aphids: a cost of dispersal

Apterous and alate Sitobion avenae (Fab.) were dissected from the third instar onwards and the size and degree of development of their largest embryos recorded. From at least the third instar apterae had larger and more well developed embryos than alatae. Embryos in both morphs showed an exponential increase in volume with time during their mothers' nymphal development, but those in apterous mothers grew faster. Although alatae had a longer pre-reproductive development than apterae, it was not long enough to compensate for the lower embryo growth rates in alatae and as a consequence they initially produced smaller and fewer offspring than apterae. After 4 days of reproduction apterous and alate mothers produced similar sized offspring and at the same rate. The total fecundity of apterae was greater than that of alatae.

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Résumé La taille et l'état de développement des plus gros embryons de Sitobion avenae Fab ont été examinés par dissection d'ailés et aptères à partir du troisième stade. La taille des embryons des 2 morphes augmente pendant le développement larvaire des mères, mais ceux des mères aptères grossissent plus vite. Bien que la période précédant la reproduction des alates soit plus longue chez les ailés que chez les aptères, les embryons des ailés sont néanmoins incapables de l'emporter en taille sur les aptères, et les ailés produisent initialement moins de pucerons et plus petits que les aptères. Après 4 jours de reproduction, les mères ailées et aptères produisent des pucerons de même taille au même rythme. La fécondité totale des aptères est supérieure à celle des ailés; la différence est liée à la biologie des 2 morphes.

     

Alata-production by an aphid: The “interval timer” concept and maternal age effects

Throughout their lives alate Acyrthosiphon pisum aphids remained unable to produce more alatae in response to crowding. Similarly, the first-born progeny of most alatae were likewise unable to respond to crowding. In contrast, the first-born progeny of apterae responded exceptionally strongly to the crowding stimulus. Later-born progeny of both morphs responded similarly. These data are inconsistent with Lees' theory of “interval timers” since ability to produce alatae did not return simultaneously in all lines founded by alatae. The theory of interval timers must be either replaced or modified to take into account differences related to maternal age.     

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.     

Influence of diet composition and crowding on wing production by the aphid Myzus persicae

Two factors have been shown to affect alata-production in a Californian strain of Myzus persicae. Contact between adult apterous females increases the proportion of alatae in their progenies as, to a lesser extent, does contact between the larvae themselves. In the absence of behavioural interactions between the aphids, the relative proportions of apterae and alatae can be influenced by the diet the mothers and their larvae receive. By suitable dietary adjustments, it is therefore possible to increase or decrease the proportions of alatae among the progeny of isolated as well as of grouped aphids. In neither case is it necessary to isolate the larvae directly after their birth in order to observe the influence of diet composition on their wing dimorphism.     

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.     

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.     

Influence of temperature on the production of parthenogenetic and sexual females by Aphis fabae under short-day conditions

Leafdiscs of broadbean, Vicia faba, were used to raise successive generations of the black bean aphid, Aphis fabae, under short-day conditions (10 hr light per day) at different constant temperatures. The progeny of apterous virginoparae that were temporarily crowded consisted mainly of alatae at 11.5° and 15.5° C, alatae and apterae at 17.5°–25.5°, and apterae only at 28.5°. All of the apterae were virginoparae. At 11.5°–19.5° all of the alatae were gynoparae (produced at least one ovipara). From 22.5°–25.5° the proportion of gynoparae among the alatae declined from 96% to 8%. At 11.5° the gynoparae exclusively produced oviparae. At 15.5°–23.5° some gynoparae gave birth both to virginoparae and other gynoparae after depositing a number of oviparae. At 22.5° a few males were also produced by some gynoparae. Gynoparae therefore produced alate and apterous virginoparae, males, gynoparae, as well as oviparae under constant conditions favoring ovipara production. Alate virginoparae also produced some gynoparae at 23.5°.All the larvae engendered by crowded apterae at 17.5° that developed into alatae became gynoparae even if their entire post-natal development occurred at 28.5° (at which temperature no gynoparae were engendered). However, only few of the presumptively alatiform progeny of crowded apterae developed into alatae (and hence few gynoparae were produced) if the larvae were transferred to 28.5° within the first two instars.The ecological significance of certain aspects of the production of various morphs in relation to temperature is discussed.

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Zusammenfassung Blattscheiben von Ackerbohnen, Vicia faba, wurden verwendet zur laufenden Zucht der Schwarzen Bohnenlaus, Aphis fabae, bei Kurztag (10/14) bei verschiedenen, konstanten Temperaturen. Die Nachkommen von apteren Virginoparen, die vorübergehend (6 h) eng gedrängt in Röhrchen verbracht waren (crowded), bestanden überwiegend aus Geflügelten bei 11,5° und 15,5°, aus Geflügelten und Ungeflügelten bei 17,5° bis 25,5° und nur aus Ungeflügelten bei 28,5°. Alle Ungeflügelten waren virginopar. Bei 11,5°–19,5° waren alle Geflügelten gynopar (d.h. erzeugten mindestens eine Ovipare). Von 22,5°–25,5° sank der Anteil der Gynoparen unter den Geflügelten von 96% auf 8%. Bei 11,5° erzeugten die Gynoparen ausschliesslich Ovipare. Bei 15,5°–23,5° gebaren einige Gynopare sowohl Virginopare wie Gynopare, nachdem sie einige Ovipare abgelegt hatten. Bei 22,5° wurden auch einige Männchen von einigen Gynoparen erzeugt. Gynopare erzeugten demnach geflügelte und ungeflügelte Virginopare, Männchen, Gynopare und Ovipare unter Bedingungen, welche die Oviparenproduktion begünstigten. Geflügelte Virginopare erzeugten bei 23,5° auch einige Gynopare.Alle Larven, die von Ungeflügelten nach Gedrängebehandlung bei 17,5° erzeugt wurden, entwickelten sich, soweit sie geflügelt wurden, zu Gynoparen, auch wenn ihre ganze postnatale Entwicklung bei 28,5° ablief (bei welcher Temperatur keine Gynopare erzeugt werden). Dagegen entwickelten sich nur wenige der präsumptiv geflügelten Nachkommen von Ungeflügelten mit Gedrängebehandlung zu Geflügelten (und dementsprechend wurden nur wenige Gynopare erzeugt), wenn die Larven innerhalb der ersten zwei Stadien in 28,5° überführt wurden.Die ökologische Bedeutung der Erzeugung der verschiedenen Morphen in Beziehung zur Temperatur wird diskutiert.

     

有翅型和无翅型大豆蚜卵巢和胚胎发育的差异

【目的】比较大豆蚜Aphis glycines有翅型和无翅型个体的卵巢发育,为从生理生化角度研究蚜虫翅型分化发生的繁殖适应性提供依据。【方法】在卵巢形态显微镜观察基础上,比较有翅型和无翅型大豆蚜卵巢小管数量、胚胎总数量、成熟胚胎数量以及卵巢小管中最大胚胎体积及其发育速率。【结果】有翅型和无翅型大豆蚜的3龄和4龄若蚜以及成蚜均有2个卵巢/雌,大多数蚜虫每个卵巢包含4条卵巢小管,可见一个卵巢包含5条卵巢小管的现象。有翅型和无翅型个体卵巢中胚胎总数量和成熟胚胎数量以及最大胚胎体积随发育时间增加均显著增大,但有翅型个体胚胎总数量和成熟胚胎数量以及最大胚胎体积均显著小于同一发育阶段无翅型个体。有翅型和无翅型的3龄和4龄若蚜以及成蚜卵巢中胚胎总数量与后足胫节长度均呈显著正相关关系。有翅型和无翅型大豆蚜卵巢小管中最大胚胎体积随发育时间均呈指数增长,但有翅型个体最大胚胎体积显著小于无翅型个体。【结论】有翅型大豆蚜相比于无翅型个体存在卵巢发育延迟现象,卵巢发育程度的差异可能与翅的发育相关。     

豆蚜有翅蚜产生的原因

本文研究拥挤、寄土质量、温度和蚜型等因子对大豆蚜(Ahis glyeines)有翅蚜产生的影响。结果表明：1.大豆蚜无翅胎生成好个体间的拥挤是有翅蚜产生的主要原因。在低密度下拥挤反应随密度增大而增强,但过度拥挤会导致反应的降低。无翅若好间的拥挤不能导致其本身发育为有翅胎生蚜。 2.寄主质量能改变无翅胎生成蚜对拥挤的反应。每笼2头经成熟叶片处理的无翅胎生成蚜后代中有姻蚜的比例高于幼嫩叶片和对照(无叶片)处理,且饥饿不能促进有翅蚜的产生。3.温度能影响有翅蚜的产生。较高的温度(30℃和25℃)较21℃对有翅胎生蚌的产生有较强的抑制作用。4.不同母蚜型产生有翊蚜的能力不同。有翅胎生蚜间的拥挤也能使其在后代中产生少量的有翊蚜,但对拥挤的敏感程度低于无翅胎生蚜。     

Effect of flight on the production of alatae by the vetch aphid, Megoura viciae

Apterous virginoparae and alate virginoparae that had flown and not flown of Megoura viciae were crowded and the morphs of the resulting offspring compared. Flying alatae that had flown produced few if any alate offspring whereas alatae which did not fly produced alatae as readily as do apterae.

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Résumé Certaines femelles virginipares ailées de Megoura viciae ne volent jamais. Des expériences ont été réalisées en vue de préciser le type de descendants qu'engendrent ces femelles.La descendance de virginipares ailées qui n'ont pas volé, de virginipares ailées qui ont volé, enfin de virginipares aptères, est étudiée et comparée dans des conditions expérimentales strictes.Le pourcentage moyen d'ailés produits par ces divers types de femelles virginipares est de 22,3% par les virginipares aptères, 1,1% par les virginipares ailées qui ont volé et 21,8% par les virginipares ailées qui n'ont pas volé. Ces derniers sont donc susceptibles de produire des ailés au même titre que les aptères. Le vol, d'une façon quelconque, est cause de l'inhibition de la production d'ailés par les virginipares ailées.

     

INFLUENCE OF DAYLENGTH ON THE PRODUCTION OF PARTHENOGENETIC AND SEXUAL FEMALES OF APHIS FABAE AT 17.5°

When apterous virginoparae of Aphis fabae and their progenies were raised at 17.5^o on leaf discs of Vicia faba under photoperiods of 12 h or less per day, more alatae occurred among the progenies than under longer photoperiods. The alatae developed exclusively into gynoparae at 4, 10 and 12 h light per day and almost all did so at 2 h. At 13 h 45 min, 14 h, 16 h, and 24 h all the alatae developed into virginoparae. At transitional photoperiods ranging from 13 h 6 min to 13 h 28 min, both alate morphs occurred. They could both be deposited by a single apterous virginopara and produce intermorphs containing eggs and embryos. Under most light regimes, a proportion of the gynoparae (generally < 10%) produced some apterous virginoparae, gynoparae, and males after producing a number of oviparae. Alate virginoparae produced almost three times as many larvae as did the gynoparae.By transferring alatiform larvae produced by apterae raised at short day (SD: 10 h light) to long day (LD: 16 h light) or vice versa, a strong prenatal influence of SD on gynopara production was demonstrated. SD also had a postnatal influence on presumptive virginoparous alatiforms, if they were transferred to SD already in the 1 st instar. Alatiform larvae could be influenced by SD or LD as late as in the 3d instar if they and their mothers had been maintained under a weak short (13 h 6 min) or a weak long day (13 h 45 min).The results are discussed in relation to possible hormonal mechanisms of morph determination.

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Zusammenfassung Wenn ungeflügelte, virginopare Aphis fabae und ihre Nachkommen gehalten wurden bei 17,5^o auf Blattscheiben von Vicia faba bei Fotoperioden von 12 oder weniger Stunden je Tag, so entstanden mehr Geflügelte unter den Nachkommen als unter längeren Fotoperioden. Die Geflügelten entwickelten sich ausschliesslich zu Gynoparen bei 4, 10 und 12 h Licht je Tag und fast alle bei 2 h. Bei 13 h 45 min, 14, 16 und 24 h entwickelten sich alle Geflügelten zu Virginoparen. Bei Übergangsfotoperioden zwischen 13 h 6 min und 13 h 28 min kamen beide geflügelte Morphen vor. Diese konnten beide von einer einzigen ungeflügelten Virginopara abgesetzt werden und Zwischenformen mit Eiern und Embryonen erzeugen. Bei den meisten Fotoperioden erzeugte ein Teil der Gynoparen (meist < 10%) einige ungeflügelte Virginopare, Gynopare und Männchen, nachdem sie eine Anzahl von Oviparen erzeugt hatten. Geflügelte Virginopare erzeugten fast dreimal so viele Larven als wie Gynopare es tun.Wenn man alatiforme Larven, deren ungeflügelte Mütter bei Kurztag (KT: 10 h Licht) gehalten waren, in Langtag (LT: 16 h Licht) übertrug oder umgekehrt, so zeigte sich ein starker pränataler Einfluss des KT auf die Gynoparenerzeugung. KT hatte auch einen postnatalen Einfluss auf präsumptive virginopare geflügelte Formen, wenn diese schon im 1. Larvenstadium übertragen wurden. Alatiforme Larven konnten selbst noch im 3. Stadium durch KT oder LT beeinflusst werden, wenn sie und ihre Mütter bei schwachem Kurz- (13 h 6 min) oder schwachem Lang- (13 h 45 min) Tag gehalten waren.Die Ergebnisse werden in Hinsicht auf mögliche hormonale Mechanismen der Morphenbestimmung diskutiert.

     

Juvenile hormone and photoperiodically controlled polymorphism in Aphis fabae: Prenatal effects on presumptive oviparae

All three naturally occurring juvenile hormones (JH's) were shown to have effects on the parthenogenetic/gamic polymorphism of Aphis fabae; they mimicked long day conditions by inducing parthenogenetic forms. When topically applied to fourth instar gynoparae, JH caused the appearance of oviparous/viviparous intermediate morphs in the progeny. JH induced both wing development and embryogenesis in embryonic, presumptive oviparae. Embryogenesis was induced by lower doses of JH. Adult, embryo-containing alatae produced by treatment with high JH doses were capable of flight, and whilst reluctant to reproduce, their few viable progeny were oviparae. They did, however, differ from normal gynoparae in size, occasional presence of scent plaques on the metathoracic tibiae, numbers of secondary rhinaria on the antennae and morphogenetic response to postnatal rearing in long day conditions. The presumptive, oviparous embryos most sensitive to JH treatment were shown to be ca 323 μm in length, close to the stage where their germaria differentiate as parthenogenetic or gamic. Similar effects were observed in the progeny of JH-treated, teneral adult gynoparae but there was no effect on the morph of progeny of long day, alate virginoparae. The JH's differed in potency in the order JH I > JH II > JH III. The treatment of fourth instar gynoparae also induced a terminal batch of apparently normal viviparous progeny in a number of aphids. This result was obtained even at JH doses below threshold for the appearance of oviparous/viviparous intermorphs.     

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.     

The rôle of the corpus allatum in prenatal wing determination in Megoura viciae

Crowding stimulates alatae production by the apterous morph of Megoura viciae, but it has no effect on the volume and ultrastructure of the maternal corpus allatum. In adult apterae the corpus allatum appears to be inactive. It is therefore considered that the maternal corpus allatum is not involved in the prenatal control of alary polymorphism in this aphid.     

Reproductive strategy of winged and wingless morphs of the aphids Sitobion avenae and Metopolophium dirhodum

The reproduction of apterous and alate morphs of the aphids Sitobion avenae and Metopolophium dirhodum is compared on the basis of fecundity in 5- and 10-day periods of adult life. Apterae of both species are consistently more fecund than alatae of comparable weight, producing about three more nymphs on average in any 5-day period. The reproductive differences are related to the number and quality of embryos at eclosion and to ovulation rates, both of which in turn appear to be linked to wing-muscle maintenance. These relationships between weight, embryos and reproduction may be used to predict a newly moulted adult aphid's fecundity, a method which may facilitate the assessment of resistance to aphids in new cereal varieties, by obviating lengthy recording of reproduction. The strategies by which alatae of these and other aphid species minimize the difference between their fecundity and that of apterae are discussed.     

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.     

The effects of nutrition and density on the production of alate Elatobium abietinum on Sitka spruce

Summary Observations over a period of 10 years showed that, in Northeastern Scotland, alatae of E. abietinum regularly appeared in mid-May, the timing being unrelated to aphid density. The peak number of alatae produced was, however, correlated with aphid density. Following an initially high level the proportion of alatae dropped to virtually nil by mid-June, whilst over the same period the aphid population density increased. Amino acid levels in spruce needles were considerably higher during the period of alate formation than they were at the termination of alate production. It is suggested that a high amino acid level was the main factor controlling the formation of alatae and that population density affected the proportions of these alatae only when nutritional levels were favourable for alate formation.