The effect of past and current hosts on reproductive investment by the adult cowpea aphid Aphis craccivora

Abstract 1. A general index of accumulated investment that alate aphids make into reproduction with respect to environmental changes (specifically changes in host at adult moult) was investigated to determine whether and how reproduction in Aphis craccivora Koch changes once aphids migrate to a different host from the one on which they were born and reared. The aphids were monitored for 4 days to determine the immediate effects of a change in host.
2. A general index of accumulated investment defined as reproductive index was used; reproductive index is equal to the reproductive output (weight of nymphs born; actual fecundity) plus potential reproduction (weight of embryos within the ovarioles with red eye spots; future fecundity).
3. The results revealed that reproductive index is affected by adult weight. The host on which the aphids have been born and reared (source host) has no effect on reproduction other than its influence on adult weight.     

Effects of population density on alienicolae of Aphis fabae Scop.: V. Variations in size, relative wing length and numbers of antennal sensorla of field alatae

Size, relative wing length and numbers of antennal sensoria of alate Aphis fabae caught in a suction trap 12.2 m above ground and of migrants, flyers and non-flyers among alate A. fabae from field infestations on beans and beet are compared. The size of aphids in successive samples decreased throughout the period of colonization but, on average, beet aphids were bigger and broad bean aphids were smaller than field bean aphids. There were very few migrant alatae among the very large or the very small aphids from any of the crops, neither were aphids of these extreme sizes caught in the suction trap. Non-flyers and beet aphids generally had relatively shorter wings than migrants and bean aphids respectively. The numbers of antennal sensoria differed more between aphids from the three crops than between classes of alatae from the same crop, although trap aphids, presumed to be migrating from field beans had more sensoria than alatae taken from that crop. The mean dry mass of flyers was usually less than that of migrants of a similar size, although if the mass of nymphs deposited before flight is added, it is apparent that a flyer would be heavier than a migrant of the same size at maturity. This additional mass is thought to be due to increased development of the embryos in the ovarioles of flyers at this time and indicative of their greater adaptation for reproduction.     

Effects of pattern and timing of high temperature exposure on reproduction of the rose grain aphid, Metopolophium dirhodum

Metopolophium dirhodum (Walker) (Hemiptera: Aphididae) is one of the three most important cereal aphid species in Germany and central European countries. High temperature is known to play a detrimental role in the reproduction and survival of aphids. Detailed experiments were conducted to assess the effects of high temperatures (27, 29, 31, and 33 °C), exposure pattern (8 h day^?1 for 1, 2, 4, and 6 days), and exposure development stage (2nd, 3rd, 4th instar larvae, and adult) on the reproduction and longevity of M. dirhodum. All three factors have a significant influence on the aphids. Increasing the temperature to over 29 °C significantly decreases their lifetime fecundity and longevity. Mature aphids are more sensitive to high temperatures than younger ones. A high temperature pulse to treat aphids older than 3rd instar larvae greatly shortened their longevity and lessened their lifetime fecundity. Lifetime fecundity and longevity was found to be inversely related to exposure pattern. A high temperature pulse lasting longer than 1 week greatly reduced the lifetime fecundity and longevity. The limitation of the data collected from experiments at constant temperatures for aphid forecasting is discussed.     

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.     

Parasitism of aphid embryos by Aphidius smithi: some effects of extremely small host size

Embryos of the pea aphid, Acyrthosiphon pisum (Harris), were successfully parasitized by Aphidius smithi Sharma & Subba Rao while still inside their mother. Parasites from embryos were much smaller, laid fewer eggs, but survived as long as their counterparts that had developed in adult viviparous aphids. Lifetime fecundity was positively correlated with wasp size, as measured by dry weight at the time of death. A. smithi females produced the same offspring sex ratio at eclosion on both host classes, although emergence was significantly lower from mummified adult aphids than from embryos. The data are discussed with regard to progeny and sex allocation decisions by solitary hymenopterous parasites. It is suggested that, in A. smithi, the decision to lay a fertilized or unfertilized egg is influenced by the host's overall size, rather than by the size and quality of a smaller target area inside the host, such as embryos.     

Effects of maternal diet on offspring fitness in the bird cherry‐oat aphid

1. Maternal and offspring diet effects on life‐history traits of the bird cherry‐oat aphid Rhopalosiphum padi were tested on three wheat varieties. Using nine reciprocal combinations of wheat varieties, the effects of previous experience (maternal diet effect) on the aphid's response to resistant and susceptible varieties (offspring diet effect) were tested. Batis was susceptible, and Xiaoyan22 and Ww2730 were both resistant, but with different mechanisms. 2. Aphids produced the most alatae in the treatments with the most resistant maternal diet variety Xiaoyan22. The fecundity (F) and intrinsic rate of increase (r_m) of these alatae were at their greatest in the most resistant offspring diet variety, but these traits were not influenced in the apterae. 3. There were significant interactions in the alatae production and apterae life‐history traits, such as r_m, development time, weight gain, and mean relative growth rate, between the maternal and offspring diet varieties. The interactions in apterae responses between varieties, some of which were reciprocal, indicated phenotypic plasticity in these parthenogenetic aphids. 4. Rhopalosiphum padi produced more alatae on the most resistant variety; the alatae would disperse and were more fecund. The growth responses of the apterae showed phenotypic plasticity to the different combinations of maternal and offspring diet varieties. The phenotypic plasticity would allow R. padi to better utilise the variable environments represented by the small wheat plots of different varieties in China.     

The effect of plant phenology on the induction of alatae and the development of populations of Metopolophium dirhodum (Walker), the rose-grain aphid, on winter wheat

In field experiments apterous Metopolophium dirhodum produced significantly more alatae when reared on mature plants (milky ripe stage) than on young plants. Nymphal and adult survival and relative growth rate were little affected by the developmental stage of the host plant. Relative growth rate and fecundity were strongly related to temperature. Laboratory experiments confirmed that the developmental stage of wheat had no significant effect on survival or relative growth rate of the aphids. However, laboratory grown plants did not produce the levels of alate production seen in field crops and in the field experiments. It is suggested that in investigations into the effects of the host plant on cereal aphids field grown plants should be used whenever possible. Alate production is proposed as the major plant induced factor affecting the development of M. dirhodum populations on wheat in the field. Alatae developed on mature plants even when the number of aphids was very low.     

Transgenic Bt maize and Rhopalosiphum padi (Hom., Aphididae) performance

Abstract.  1. The population abundance and age structure of Rhopalosiphum padi , one of the most common maize aphid species, on transgenic Bt (expressing the Cry1Ab protein) and non-Bt isogenic maize was studied in commercial plots during three crop seasons.
2. A higher density of aphids, particularly alates and young nymphs, occurred in Bt plots at very young maize development stages, corresponding to the settlement period, in the 3 years studied. Possible causes are discussed. After this period, there were no differences between Bt and non-Bt maize.
3. Mortality, development, and reproduction of the offspring of alate forms of R. padi and the offspring of different generations of apterous forms fed with Bt maize were evaluated in the laboratory under controlled conditions.
4. The developmental and pre-reproductive times of the offspring of the first generation of alatae were shorter and the intrinsic rate of natural increase ( r _m ) higher when aphids fed on Bt maize. The opposite occurs with the offspring of the first generation of apterous mothers, which have lower nymphal and adult mortality, shorter developmental and pre-reproductive times, a higher effective fecundity rate, and greater r _m , when fed on non-Bt maize. The differences in aphid development on the two cultivars may be linked to changes in host-plant quality due to pleiotropic effects of the genetic modification.
5. No differences on aphid mortality, developmental and pre-reproductive times, fecundity, and r _m were found between the offspring of apterous aphids maintained on Bt or non-Bt maize for several generations.     

Levels and components of resistance to Amphorophora idaei in raspberry cultivars containing different resistance genes

Levels and components of resistance to Amphorophora idaei in raspberry cultivars containing different A. idaei resistance genes were studied under infestation tunnel, glasshouse and laboratory conditions. Each test consistently ranked raspberry cultivars, with increasing levels of resistance in the order, non-resistant (cv. Mailing Jewel), resistant cultivars containing minor genes, major gene A₁, gene A₁₀ (red raspberry) and gene A₁₀ (black raspberry) respectively. Resistance was expressed in three different ways; decreased alatae settling and feeding, decreased apterae settling and decreased aphid fecundity and rate of nymphal development. Following exposure to a large population of alatae, significantly fewer aphids settled on A₁, A₁₀ and to a lesser extent, minor gene-containing cultivars, compared to cv. Mailing Jewel. More alatae settled on the top than the bottom zone of non-resistant, minor gene resistant and two of four A₁-containing cultivars; alatae settling was low on all zones of A₁₀-containing raspberries. Aphid fecundity and nymph development patterns on different cultivars and resistance classes were similar to those found for alatae settling. After 7 days reproduction more than 30% of the nymphs developed to third or fourth instar on cv. Mailing Jewel, whilst on minor gene and major gene-containing cultivars the total number of nymphs and the proportion of later instars decreased. On resistant cultivars the nymphs were found mainly on the middle and bottom leaf zones, compared to cv. Mailing Jewel. A rapid (48 h) screening test using floating leaflets was developed and, on the basis of apterae settling, ranked cultivars reliably; it was particularly effective in distinguishing moderate and strong resistance to strain 1 A. idaei. Gene A₁-containing cultivars bred in England were much more resistant to strain 1 A. idaei than those bred in Scotland. Possible reasons for this difference are discussed, in relation to modified screening procedures and to the control of the viruses transmitted by this aphid vector.     

Positive effects of an arbuscular mycorrhizal fungus on aphid life history traits

Two generations of two aphid species (Myzus ascalonicus and M. persicae) were reared on Plantago lanceolata plants, with and without root colonization by the arbuscular mycorrhizal fungus, Glomus intraradices. Life history traits of the aphids measured were nymphal development time, teneral adult weight, growth rate, total fecundity, adult longevity and duration of post-reproductive life. For both aphids in both generations, mycorrhizal colonization increased aphid weight and fecundity, while other traits were unaffected. The increases were consistent between generations. In a second experiment, M. persicae was reared on plants with and without the fungus, under varying N and P regimes. The results of N addition were inconclusive because there was high aphid mortality. However, under P supplementation, positive effects of the mycorrhiza on aphid growth were seen at low and medium P levels, while at high P levels these effects disappeared. The positive effects of mycorrhizal colonization reported here are contrary to the majority of previous studies with chewing insects, which have reported negative effects. A number of possible mechanisms for this apparent discrepancy are discussed. Received: 1 February 1999 / Accepted: 22 March 1999     

Essential versus alternative foods of insect predators: benefits of a mixed diet

Although many predatory insects appear to be opportunistic generalists in their selection of prey, only a subset of prey species may in fact serve as “essential foods” capable of supporting immature growth and adult reproduction. It has been suggested that other, “alternative foods” serve only to maintain the predator when essential foods are not available, but little research has evaluated the significance of a mixed diet of essential and alternative foods for predator growth or reproduction. Here we test the general hypothesis that although alternative prey may be inadequate to support reproduction when consumed alone by adult predators, consumption of such prey may enhance the predator's reproductive output when the predator also has access to essential prey. We compared egg production by two aphidophagous lady beetles, Coccinella septempunctata and C. transversoguttata, provided with diets of aphids (essential prey) and weevils (alternative prey). As predicted, female predators produced greater numbers of eggs when a diet of pea aphids in limited number was supplemented by alfalfa weevil larvae. The predators laid no eggs when provided only with weevils or only with sugar. But once aphids were added to the diet, females of C. transversoguttata (but not C. septempunctata) laid eggs in greater numbers when they had fed previously on weevils than on sugar. Females of both species also produced eggs in modest numbers when provided with both weevils in excess and sugar, but this diet supported a lower rate of egg production than did a diet of weevils in excess plus a limited number of aphids. Although C. septempunctata has a longer history of association with the alfalfa weevil than does C. transversoguttata, the former species was not more effective in exploiting this alternative prey in support of reproduction. The tendency of generalist predators such as adult lady beetles to consume alternative as well as essential prey probably enhances considerably their ability to capitalize on short-lived and scattered opportunities as they seek out suitable sites in which to reproduce. Received: 11 May 1998 / Accepted: 1 May 1999     

Effects of the maternal and pre‐adult host plant on adult performance and preference in the pea aphid,Acyrthosiphon pisum

Abstract 1. The taxon known as the pea aphid, Acyrthosiphon pisum, is composed of a series of host plant associated populations and is widely used as a model system to explore ecological speciation and the evolution of specialisation. It is thus important to know how maternal and pre‐adult experience influences host plant utilisation in this species. 2. The relative importance of the maternal and pre‐adult host plant for adult fecundity and host preference was investigated using three aphid clones collected from Lathyrus pratensis and maintained on Lathyrus or Vicia faba. 3. No significant effects of the maternal host plant on offspring fecundity were detected. 4. The host plant on which the aphid grew up influenced adult fecundity, although in a complex way that depended on both the adult host plant species and when after transfer to the test plant fecundity was assessed. 5. All three clones preferred to colonise Lathyrus over Vicia, and this preference was stronger for aphids raised on Lathyrus. 6. The significance of the results for studies of the evolution of specialisation and speciation that employ A. pisum is discussed.     

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.

     

Comparing wing loading,flight muscle and lipid content in ant‐attended and non‐attended Tuberculatus aphid species

Although all Tuberculatus aphids possess wings, some species associated with ants exhibit extremely low levels of dispersal compared with those not associated with ants. Furthermore, phylogenetic interspecific comparisons find significantly higher wing loading (i.e. higher ratio of body volume to wing area) in ant‐attended species. This observation indicates that ant‐attended species may allocate more of their body resources to reproductive traits (i.e. embryos) rather than flight apparatus (i.e. wings, flight muscle and lipid). The present study focuses on two sympatric aphid species and aims to investigate the hypothesized trade‐off in resource investment between fecundity and the flight apparatus; specifically, the ant‐attended Tuberculatus quercicola (Matsumura) and non‐attended Tuberculatus paiki Hille Ris Lambers. Species differences are compared in: (i) morphology, (ii) embryo production, (iii) triacylglycerol levels and (iv) wing loading and flight muscle. The results show that T. quercicola has a larger body volume, higher fecundity and higher wing loading compared with T. paiki, which has a smaller, slender‐shaped body, lower fecundity and lower wing loading. No significant difference is found between the species with respect to the percentage of triacylglycerol content in dry body weight. The flight muscle development is significantly lower in T. quercicola than in T. paiki. These results indicate that the additive effect of higher wing loading and the lower amount of flight muscle development in T. quercicola may increase the physical difficulty of flight, and hence be responsible for its lower dispersal ability. The trade‐off between fecundity and dispersal documented in wing‐dimorphic insects may therefore be applicable to T. quercicola, which has fully developed wings.     

Drought stress and cereal aphid performance

The performance of clones of Rhopalosiphum padi and Sitobion avenae from England and Spain was examined on drought-stressed tillering winter wheat in an environment chamber at 14 ± 1°C. Two different levels of drought stress and an unstressed control were established by different watering regimes which resulted in drought-stressed plants being smaller at the end of the experiment. The effect of drought stress to plants on aphid performance was not significantly different between the clones tested. Drought stress had no effect on aphid development time, nymphal mortality, the weight of teneral adults and the number of embryos in teneral adults up to the onset of reproduction in the first F₁ generation. The subsequent reproductive capacity, as measured by the effective and potential fecundity, and the reproductive rate, were much reduced on drought-stressed plants. However, there was only a small decrease in the intrinsic rate of increase (r_m). Overall the clone of R. padi from Spain performed better than that from England, the development and prereproductive times being shorter and the fecundity higher in the Spanish clone, giving a higher r_m. There were no differences in the fecundity and the r_m between the Spanish and the English clones of S. avenae. The proportion of the F₂ generation that was alate differed greatly between clones, and only the English S. avenae produced significantly more alatae on drought-stressed than on unstressed plants.     

Effect of dietary amino acids on the size and alary polymorphism of Aphis fabae

Aphis fabae was reared on synthetic diets omitting individual amino acids and amino acid groups. Methionine and histidine deletions induced apterae formation and prevented the aphids from reaching the adult stage, whereas arginine, leucine, lysine, and proline deletions induced alatae formation. The omission of alanine, cysteine, phenylalanine, proline, serine, and tyrosine reduced the size attained by adult alatae and together with methionine and histidine, these amino acids are considered to be essential for normal growth by A. fabae. A delay in parturition occurred in aphids reared on certain deficient diets and there is an association between the size attained by alatae and the delay in parturition.     

Assessment of methods for testing varietal resistance to aphid s in cereals

The adult weight of three species of cereal aphids on barley was correlated with the number of large embryos and the total number of embryos within newly moulted adults. The relationships between embryo number and weight were constant for each species of aphid on all seven varieties of barley under identical conditions. The weights of aphids of each species on seven varieties of barley were significantly different from each other in each experiment. Pot size and plant age had a significant effect on the weights of Rhopalosiphum padi. The validity of these tests compared with previous methods is discussed.     

Aphid colonization of spring cereals

In 1970-1, Metopolophium dirhodum, Rhopalosiphum padi and Sitobion avenae were the commonest alatae trapped from April/May to August, with most in July and early August. The first alatae appeared in the Rothamsted survey suction trap 0–34 days before aphids were found on the cereals, but during May and June no relationship was found between the numbers trapped and the number on the crop. Most species occurred first near the sheltered edge of the crop, but M. dirhodum was widespread over the field. Most infestations were quickly dispersed by the movements of older morphs; adults only stayed in one place for about 2 days. Alate M. dirhodum moved more often than apterae, but both morphs of S. avenae moved equally often and more frequently between larvipositions than did those of M. dirhodum. Apterae deposited more nymphs in a ‘group’ than alatae, and M. dirhodum deposited more than S. avenae. Few ‘groups’ persisted for more than a week. Although M. dirhodum occupied the crop area faster than S. avenae, all 0–3 m lengths of row sampled being infested within 2–5 wk of their first appearance, most or all of the tillers were colonized only in late July 1970.     

Effects of photoperiod and diet on diapause tendency,maturation and fecundity in Harmonia axyridis (Coleoptera: Coccinellidae)

The effects of photoperiodic conditions of larval development and adult maturation (L : D = 12 : 12 vs. 18 : 6) and different diets (sugar solution, frozen eggs of Sitotroga cerealella, different numbers of aphids Myzus persicae, and their combinations) on survival, reproductive maturation and fecundity of Harmonia axyridis were studied in laboratory conditions. The fundamental aim of the work was to distinguish between cue effect of diet (neurohormonal triggering of reproduction) and direct effect of diet (nutritional maintenance of reproduction). When adults were kept under short‐day conditions, the proportion of ovipositing females decreased and the duration of the pre‐oviposition period increased. Moreover, a strong reaction to the direction of changes in the day length was demonstrated: when larvae and pupae developed at long day and adults were transferred to short day, the proportion of ovipositing females was much lower than in individuals that were permanently kept under short‐day conditions. The percentage of ovipositing females, the rate of their reproductive maturation and the average daily fecundity gradually increased in the following succession of diets: ‘sugar + 5 aphids per day < sugar + eggs < sugar + eggs + 5 aphids per day < sugar + 100 aphids per day’. However, dissection showed that most of the non‐laying females fed on these diets (particularly those kept under long‐day conditions) have started reproductive maturation, while even first stages of oogenesis were not found in females fed on sugar solution alone. We conclude that cue effect of diet (reproductive activation) can be achieved almost independently of the number of prey consumed, while nutritional effects (the rate of reproductive maturation and fecundity) are sensitive both to the quality and quantity of food.     

Effect of different diets on the growth and development of Harmonia axyridis (Pallas)

Harmonia axyridis (Pallas) is an important natural enemy of crop pests. Feeding H. axyridis through artificial diets (ADs) is an important means to achieve large-scale production. In this study, the effects of different diets on the growth, development and reproduction of H. axyridis were comprehensively evaluated, and the reasons for the reduced reproductive capacity of H. axyridis fed ADs were preliminarily explored. Artificial diets were prepared using pig liver and pork as the main ingredients. Larval duration, larval survival rate, egg production and other traits were comprehensively evaluated. Harmonia axyridis fed ADs during both the larval and adult periods completed growth and development from larva to adult but had a prolonged development duration (6.13-day extension) and reduced survival rate (15.3% reduction) in the larval stage. In addition, adults could not lay eggs. Next, individuals of H. axyridis were fed Megoura japonica (Matsumura) in the larval stage and an AD, adequate aphids (AA) or an artificial diet plus 5 (AD5), 10 (AD10) or 25 (AD25) aphids in the adult stage. The adults in the AD group produced a small number of eggs (103.3 eggs). When aphids were added to the diet, egg production gradually increased with the number of aphids added. The AD25 group exhibited no significant difference in egg production compared with that in the AA group (983.8 eggs and 1,158.3 eggs, respectively). Anatomical observations of the H. axyridis adults in these five groups revealed that ovary development was slowest in the AD group. In addition, we found that the levels of vitellogenin and juvenile hormone were significantly lower in the AD group than in the AA group. These results provide a foundation for formulating ADs for H. axyridis and improving the large-scale reproduction of this species.