Physiological responses of Uroleucon jaceae (L.) to seasonal changes in the quality of its host plant Centaurea jacea L.: multilevel control of adaptations to the life cycle of the host

Summary Exploiters of short-lived plants have evolved strategies in response to physiological changes that occur during the development of their hosts. The ability to adapt to host quality changes is necessary particularly if the mobility of an animal is poor or risk-constrained. In the plant-aphid system Centaurea jaceae-Uroleucon jaceae, the responses of the aphid to the seasonal changes in its host plant grown in poor and good quality soil were investigated. Coarse- and fine-tuned physiological reactions were observed in discrete aphid generations which were reared on plants grown either in a growth chamber or in a greenhouse. The number of ovarioles and developmental time depended on extrinsic factors (length of photoperiod) not directly related to plant quality. The reproductive investment of the aphids was also independent of their total dry weight. However, within the gonadal system high correlations were found at the embryonic level (e.g. the number of sclerotized embryos or the length of the oldest embryos per ovariole were highly correlated with gonadal dry weight). Aphids which were living on the 4-leaf stage of high-quality host plants showed a significantly higher investment in their gonads than aphids feeding on senescenting hosts. Factor analysis corroborated that aphids reared on poor-quality hosts revealed no grouping of variables measured, whereas those which were reared on high-quality plants showed clustering in respect of somatic (tibia length, dry weight of soma) and gonadal (number of sclerotized and unsclerotized embryos, length of the oldest embryos, ovariole number, gonad dry weight) factors. Three different levels of adaptive response to plant quality are distinguished: individual response to host quality, maternal influences on offspring and response to changes not specific to habitat (photoperiod). These different levels of regulation are thought to enable the aphids to adapt to a host of a given nutritional quality and anticipate deteriorating habitat quality simultaneously. It is concluded that physiological constraints in aphids are only revealed when aphids are exposed to severe nutritional stress for several generations.     

Generation specific life history traits of winged shape Sitobion avenae

When reared under constant conditions from egg hatch the proportion of offspring of Sitobion avenae (F.) (Aphididae: Macrosiphini) that developed into winged individuals declined from the second to fourth generation. The winged individuals produced in these three generations showed a progressive decrease in their wet and dry weights and an increase in the length of their largest embryo with generation. The trend in winged individual production and also the trends in their size and reproductive strategy appear to be programmed and are similar to the generation specific strategies that have been described in other species of aphid.     

Seasonality and reproduction in aphids

Aphids are short-lived and occupy habitats which vary in quality in both time and space. They exploit their ephemerally nutritious habitats by rapidly producing many small offspring when conditions are good, and fewer, larger offspring when conditions are poor, at which time they also divert more of their resources into fat storage. Aphids of each of the generations which make up a life cycle have specific reproductive strategies adapted to the conditions they are most likely to encounter, that is they anticipate the predictable seasonal trends in habitat quality.     

Seasonal changes in fat content,form, state of gonads and length of adult life in the sycamore aphid,Drepanosiphum platanoides (Schr.)

• 1 Even when reared under similar conditions of crowding and nutrition, first and second generation adult sycamore aphids differ in their morphology and physiology. Second generation adults have a smaller abdomen, a longer gut, greater fat reserves, less developed gonads, a longer reproductive diapause and live longer than first generation aphids.
• 2 Second generation nymphs reared in isolation develop into adults which have less well-developed gonads but a reproductive delay similar to that of first generation adults.
• 3 Crowding of second generation adults prolongs the reproductive diapause.

     

The role of food quality and competition in shaping the seasonal cycle in the reproductive activity of the sycamore aphid

The hypothesis that seasonal changes in sycamore aphid,Drepanosiphum platanoidis (Schr.), recruitment are determined by changes in food quality and aphid population density was tested. There was no clear association between the reproductive activity of the sycamore aphid and the seasonal changes in specific amino acids or groups of amino acids in extracts of sycamore,Acer pseudoplatanus L., leaves. Seasonal changes in reproductive activity tracked the changes in total amino acids of the leaf tissue of the host, but with a short time delay. High numbers of adult aphids appeared to depress reproductive activity. A regression analysis of the results revealed that total amino acids the previous week and current numbers of adult aphids significantly affected sycamore aphid reproductive activity. The results of this analysis support the above hypothesis, that the marked seasonal changes in the total quantity of amino acids in sycamore leaves and intraspecific competition for this resource, through its effect on adult weight, shape the seasonal cycle in the reproductive activity of the sycamore aphid.     

一种室内饲养麦蚜的方法

介绍利用水培麦苗大量饲养麦蚜的技术。在不补充营养液的条件下麦苗可以维持10~15d,一般在饲养条件下可以1周换1次麦苗。小麦种子在经过24h浸种后,平铺在底部垫有滤纸的直径9cm的培养皿中,在25℃,4d后就可以接种麦蚜。适宜的试样条件为温度在18~25℃,相对湿度50%~70%,光周期为17hr∶7hr,饲养1代大约需要5~9d时间,每皿可养麦蚜500头以上。经过40代以上的连续饲养,蚜虫体重和每蚜产幼蚜数量没有明显降低趋势。     

Maintenance of soldier-producing aphids on an artificial diet

In our efforts to understand the biology of soldier-producing aphids, we attempted to maintain them in the laboratory using a chemically defined artificial diet. The ability of 16 species from the subfamilies Eriosomatinae and Hormaphidinae, most of which are soldier-producing species, to survive on the artificial diet was examined. Some species neither fed nor grew on the diet, whereas other species accepted the diet, grew to some extent, and managed to produce a small number of short-lived offspring. Although they performed poorly on the diet in general, aphid performance was correlated with the stage in the life cycle and the developmental stadium in that aphids of the gall generation tended to accept the diet and survive on it, whereas aphids of the non-gall generation did not. Also, old insects tended to perform better on the diet than young nymphs. Notably, only one species, Tuberaphis styraci, a gall-forming aphid that produces 2nd instar sterile soldier, showed good performance on the diet. Insects collected from galls (generation G0) survived on the diet, grew well, and produced many progeny. Three successive generations (G1, G2 and G3) were produced on the diet. Developmental period, adult body size, and age of first reproduction were almost constant through G0, G1 and G2 whereas fecundity, adult longevity and daily offspring production declined as the generations proceeded. These results are comparable to previous studies in which pest aphids have been maintained on similar artificial diets for several generations. Therefore, it is suggested that the artificial-diet rearing system will provide a useful tool to investigate various biological aspects of the soldier-producing eusocial aphid, T. styraci.     

A comparison of maternal effects and current environment on vital rates of Aphis nerii, the milkweed–oleander aphid

Abstract.  1. Non-Mendelian maternal effects, the effects of maternal phenotype or environment on offspring phenotype, have been documented in numerous taxa. By affecting offspring vital rates (birth, death, and movement), maternal effects have the potential to influence population dynamics. However, relatively few studies have directly linked maternal phenotype or environment to offspring vital rates. Additionally, even fewer studies have compared the magnitude of across-generation effects (i.e. maternal effects) to within-generation effects.
2. Because of their telescoping generations, aphids can be strongly influenced by maternal effects. The effects of maternal density and maternal host-plant species on offspring survival, fecundity, and alate formation were investigated experimentally in Aphis nerii , the milkweed–oleander aphid.
3. Additionally, the relative strength of maternal effects were compared with those operating within a generation. Therefore, in another set of experiments, the effects of current density and host-plant species (within-generation effects) on aphid vital rates were examined.
4. While maternal effects were present, within-generation effects were much stronger and more strongly influenced aphid vital rates. Within a generation, aphids exhibited density-dependent survival, fecundity, and alate formation and these effects varied among host-plant species.
5. These results indicate that while maternal effects have the potential to affect population dynamics, this potential is not always met. Additionally, the current environment, not the environment of previous generations, more strongly impacts population dynamics.     

Differential regulations of wing and ovarian development and heterochronic changes of embryogenesis between morphs in wing polyphenism of the vetch aphid

SUMMARY In wing polyphenisms that produced alternative wing morphs depending on environmental conditions, the developmental regulations to balance between flight and reproductive abilities should be important. Many species of aphids exhibit wing polyphenisms, and the development of wing and flight muscles is thought to incur costs of reproductive ability. To evaluate the relationship between flight and reproduction, the fecundity and the wing- and ovarian development in the parthenogenetic generations were compared between winged and wingless aphids in the vetch aphid Megoura crassicauda . Although no differences in offspring number and size were detected, the onset of larviposition after imaginal molt was delayed in winged adults. The comparison of growth in flight apparatus revealed that, after the second-instar nymphs, the flight-apparatus primordia of presumptive wingless aphids were degenerated while those of winged nymphs rapidly developed. In the ovaries of winged line, the embryo size was smaller and the embryonic stages were delayed from third to fifth instars, although these differences had disappeared by the time of larviposition. It is therefore likely that the delay in larviposition in winged aphids is due to the slower embryonic development. The correlation between embryo size and developmental stage suggests that the embryos of winged aphids are better developed than similarly sized embryos in wingless aphids. These heterochronic shifts would facilitate the rapid onset of larviposition after the dispersal flight. This developmental regulation of embryogenesis in the aphid wing polyphenism is suggested to be an adaptation that compensates the delay of reproduction caused by the wing development.     

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.     

Simple signals indicate which period of the annual cycle drives declines in seasonal populations

For declining wild populations, a critical aspect of effective conservation is understanding when and where the causes of decline occur. The primary drivers of decline in migratory and seasonal populations can often be attributed to a specific period of the year. However, generic, broadly applicable indicators of these season‐specific drivers of population decline remain elusive. We used a multi‐generation experiment to investigate whether habitat loss in either the breeding or non‐breeding period generated distinct signatures of population decline. When breeding habitat was reduced, population size remained relatively stable for several generations, before declining precipitously. When non‐breeding habitat was reduced, between‐season variation in population counts increased relative to control populations, and non‐breeding population size declined steadily. Changes in seasonal vital rates and other indicators were predicted by the season in which habitat loss treatment occurred. Per capita reproductive output increased when non‐breeding habitat was reduced and decreased with breeding habitat reduction, whereas per capita non‐breeding survival showed the opposite trends. Our results reveal how simple signals inherent in counts and demographics of declining populations can indicate which period of the annual cycle is driving declines.     

Adaptive allocation of resources and life-history trade-offs in aphids relative to plant quality

The need to allocate a limited amount of energy between different life-history traits is a fundamental assumption in life-history theory. However, it has often turned out to be extremely difficult to measure the competing processes that contribute to costs or benefits for individual organisms. The present investigation begins by analysing how an aphid clonal lineage adapts its reproductive investment to moderate changes in host plant quality (e.g. during the life cycle of its host). Using Centaurea jacea and Uroleucon jaceae as a model plantaphid system, I show that reproductive investment can be far more complex than indicated by dry or wet mass of the gonads alone. The number of embryos of a particular size class or developmental state present in the reproductive system of an aphid is highly flexible and is influenced by the quality of the host plant. Next, the effects of a particular reproductive investment on survival during periods of food deprivation are analysed for aphids originating from host-plants of different qualities. When food stress is severe the ability to rapidly resorb and reallocate resources committed to offspring is important for survival. However, this ability is limited. I argue that, in periods of food stress, young, unsclerotized embryos might serve as a kind of energy buffer similar to a fat body and are therefore not relevant to cost-benefit calculations. However, embryos that are beginning to sclerotize within the ovarioles are not available for resorption and resource reallocation. They compete for nutrients with their mother and contribute to costs. Therefore, it is suggested that the reproductive investment of an aphid should not be equated with reproductive costs in a general al way. The dynamics of adaptive resource allocation and resorption are a key feature of an aphid's life history, and the implications for life-history theory are discussed.     

Patterns of diversity for aphidiine (Hymenoptera: Braconidae) parasitoid assemblages on aphids (Homoptera)

We used aphids (Aphidae) as a representative hemimetabolous host family to investigate patterns of parasitoid (Aphidiine) assemblage size. The aphidiine assemblages from 477 aphid species were used to estimate average assemblage size and the influence of eight ecological and taxonomic variables. Aphids species support an average of 1.7 aphidiine species. Aphid subfamily and invasion status (native or exotic) were the most important determinants of parasitoid richness, explaining 28% of the deviance in aphidiine assemblage size. Aphids within the largest aphid subfamily, the Aphidinae, support larger parasitoid assemblages than those in other subfamilies. Parasitoid diversity was also highest on exotic aphid hosts (within the Aphidinae) and on hosts in developed habitats (agricultural or urban), though the latter effect is weak. Patterns related to aphid food plant architecture were influenced by an interaction with aphid invasion status; parasitoid diversity drops with increasing architectural complexity on exotic aphids, whereas the diversities on native aphid hosts are similar on different plant types. Weak effects were also found for aphid food plant alternation (whether or not aphids switch hosts seasonally) and climate (annual range in temperature); alternating aphids support more parasitoids than non-alternating hosts, and parasitoid assemblage size is lowest in warm climates. Taxonomic isolation of aphids at the generic level showed no significant relationship with parasitoid diversity. Finally, in contrast to parasitoid assemblages on holometabolous hosts, sample size effects were weak for aphids, possibly due to the narrow host ranges of aphidiines. Received: 22 November 1997 / Accepted: 7 March 1998     

Mating or ovipositing? A crucial decision in the life history of the cabbage aphid parasitoid Diaeretiella rapae (M’Intosh)

1. The reproductive fitness of a parasitoid depends on its mating and ovipositing success. Virgin haplodiploid females can reproduce, but produce only males, and may diminish fitness by producing more male offspring than required. Therefore, females must decide on whether to mate or oviposit first. 2. This study was conducted to assess the mating versus ovipositing decision and its impact on the reproductive fitness of Diaeretiella rapae (Hymenoptera: Aphididae), an endoparasitoid of the cabbage aphid Brevicoryne brassicae (Hemiptera: Aphididae). 3. When newly emerged females were given a choice between mating and ovipositing, about 62% of D. rapae females preferred to mate before ovipositing. Those females who oviposited before mating parasitised only 10% of the available aphids. After mating, females superparasitised their hosts with fertilised eggs, which resulted in a highly female‐biased sex ratio in the offspring. 4. Mating success was very high (91%) in the presence of hosts (cabbage aphid nymphs) compared with that in the absence of aphids. However, mating success was not influenced by the quality (size) of the hosts present in the mating arena, despite a parasitoid preference for larger hosts during oviposition. The time between pairing and mating was also shorter in the presence of host aphids. The mean number of aphids parasitised and the parasitism rate were significantly greater after mating.     

Facultative symbionts are associated with host plant specialization in pea aphid populations

The pea aphid, Acyrthosiphon pisum, shows significant reproductive isolation and host plant specialization between populations on alfalfa and clover in New York. We examine whether specialization is seen in pea aphids in California, and whether fitness on alternative host plants is associated with the presence of bacterial symbionts. We measured the fitness of alfalfa- and clover-derived aphids on both types of plants and found no evidence for specialization when all aphid lineages were considered simultaneously. We then screened all aphids for the presence of four facultative bacterial symbionts: PAR, PASS, PABS and PAUS. Aphids with PAUS were host-plant specialized, having twice as many offspring as other aphids on clover, and dying on alfalfa. Other aphids showed no evidence of specialization. Additionally, aphids with PABS had 50% more offspring than aphids with PASS when on alfalfa. Thus, specialist and generalist aphid lineages coexist, and specialization is symbiont associated. Further work will resolve whether PAUS is directly responsible for this variation in fitness or whether PAUS is incidentally associated with host-plant specialized aphid lineages.     

Reproductive diapause and the bacterial symbiosis in the sycamore aphid Drepanosiphum platanoidis

1. Parthenogenetic adults of the sycamore aphid Drepanosiphum platanoidis exhibited reproductive diapause for 4–6 weeks in the summer. 2. The diapausing aphids had small gonads (accounting for just 13% of the total aphid protein content) bearing small (< 0.5 mm long) and developmentally immature embryos. There was no evidence of embryo resorption. 3. The diapausing aphids had significantly depressed essential amino acid content and elevated glutamine content, relative to reproductively active D. platanoidis. 4. The reproductive characteristics and amino acid titres of the diapausing aphids resemble those of aphids lacking functional symbiotic bacteria. Uncoupling of maternal and embryo growth and suppression of bacterial function are proposed as key elements in the diapause of D. platanoidis.     

Suppression of autumnal sexual morph production in spring by a seasonal timer in an aphid

Photoperiod is the major seasonal cue for phenotypic plasticity in the regulation of development and reproduction. Aphids have a peculiar mechanism in which sensitivity to the photoperiod is lost for a few months through successive generations after hatching from diapause eggs, and therefore, the mechanism responsible for the photorefractory period is termed “a seasonal timer”. However, whether the seasonal timer has adaptive significance has never been investigated. Here, we show that the photorefractory generation avoids an unseasonal sexual morph production under short days in spring in the pea aphid, Acyrthosiphon pisum (Harris). When the seasonal timer operated, the overwintered generation, stem mothers, produced only parthenogenetic progenies under natural photoperiods and temperatures in both warm and cold springs. After repeated parthenogenesis following diapause, however, aphids with an expired seasonal timer produced sexual females and males typical of autumn under natural photoperiods and temperatures in a warm spring. These sexual morphs are thought to have a reproductive disadvantage compared to parthenogenetic morphs in the following summer. We thus conclude that the seasonal timer is adaptive for avoiding sexual morph production in spring. The present study shows, for the first time, that a photorefractory period lasting over generations is a trait of ecological importance.     

Competition and body size in plants: the between-species trade-off for maximum potential versus minimum reproductive threshold size

Aims According to traditional theory, superior competitive ability in plants generally requires relatively large plant body size. Yet even within the most crowded vegetation, most resident species are relatively small; species size distributions are right-skewed at virtually every scale. We examine a potential explanation for this paradox: small species coexist with and outnumber large species because they have greater 'reproductive economy', i.e. they are better equipped—and hence more likely—to produce offspring despite severe size suppression from intense competition.Methods Randomly placed plots within old-field vegetation were surveyed across the growing season. Within each plot, the largest (MAX) and smallest (MIN) reproductive individuals of each resident species were collected for above-ground dry mass measurement. We tested three hypotheses: (i) smaller resident species (with smaller MAX size) have generally smaller reproductive threshold sizes; (ii) smaller resident species have greater 'reproductive economy', i.e. a smaller MIN relative to MAX reproductive plant size; and (iii) MIN size predicts plot occupancy (species abundance within the community) better than MAX size.Important findings The results supported the first and third, but not the second hypothesis. However, we could not reject the hypothesis that smaller species have greater reproductive economy, as it was not possible to record data for the largest potential plant size for each species—since even the largest (MAX) plants collected from our sampled plots were subjected to competition from neighbours under these natural field conditions. Importantly, contrary to conventional competition theory, more successful species (in terms of greater plot occupancy) had smaller minimum not larger (or smaller) maximum reproductive sizes. These results suggest that a small reproductive threshold size, commonly associated with relatively small potential body size, is generally more effective in transmitting genes into future generations when selection from neighbourhood crowding/competition is intense—at least within natural old-field vegetation. Accordingly, we propose a simple conceptual model that represents the basis for a fundamental paradigm shift in the predicted selection effects of crowding/competition on plant body size evolution.     

Number of ovarioles and fecundity in the black bean aphid, Aphis fabae

The number of ovarioles in the gonads of the host alternating aphid, Aphis fabae Scop., is not constant. The apterous generations on spindle, emigrants and initial apterous colonists of the secondary host each have a characteristic range of number of ovarioles; subsequent generations in the yearly life cycle have twelve, except for alate exules which have a range and the lowest average number of ovarioles. Ovariole number is programmed and is not a function of adult weight or food quality. The generations with the largest number of ovarioles have the greatest fecundity, highest reproductive rate and smallest offspring, and develop early in the season when the environment is favourable. Later generations have a low fecundity and proportionately more fat, a possible adaptation to the harsher environment these aphids experience.

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Zusammenfassung Die Anzahl der Ovariolen in den Eierstöcken ist bei der wirtswechselnden Bohnenlaus, Aphis fabae, nicht konstant. Bei Läusen der ungeflügelten Generationen auf Pfaffenhütchen, sowie bei Migrantes und den ersten apteren Aphiden auf dem Sekundärwirt variiert die Anzahl der Ovariolen in einem charakteristischen Bereich. Die folgenden Generationen im jährlichen Zyklus haben stets zwölf Ovariolen, ausgenommen alate Exsules, bei denen ebenfalls ein charakteristischer Bereich nachzuweisen war, allerdings mit einem sehr niedrigen Durchschnitt. Die Anzahl der Ovariolen ist programmiert. Ihre Anzahl ist konstant und nicht vom Gewicht der Imagines oder der Nahrungsqualität abhängig. Die Generationen mit der höchsten Anzahl von Ovariolen sind am fruchtbarsten, sie haben die höchste Reproduktionsrate und gebären die kleinsten Läuse. Sie entwickeln sich frühzeitig, wenn die Bedingungen günstig sind. Spätere Generationen zeigen eine niedrigere Fruchtbarkeit, lagern jedoch verhältnismäßig mehr Fett ein, wahrscheinlich eine Anpassung an die schlechteren Bedingungen, denen diese Läuse ausgesetzt sind.

     

An investigation of the differential performance of clones of the aphid Sitobion avenae on two host species

Individuals of the grain aphid Sitobion avenae were collected from wheat and cocksfoot stands around Hampshire, UK, during March and April 1994. Eight wheat and eight cocksfoot aphid clones were chosen on the basis of readily distinguishable RAPD-PCR fingerprint profiles. The performances (weight, fecundity and survival) of successive generations of each of these clones were then carefully monitored in the laboratory as new generations of aphids were transferred either to winter wheat or to cocksfoot in planned sequences. Even those clones that were originally caught on the same host showed significant variability in performance. Clones generally performed better on their host of origin than they did on the alternate host, and they performed less well on the alternate host compared to the clones that had originated there. A comparison of the performance of third generation aphids with first generation aphids showed that the experience of the mother in the second generation often influences the subsequent performance of their offspring. As the sequence of host transfers had more effect on the performance of wheat clones than cocksfoot clones, it is likely that wheat clones are more specialised, such that wheat is a satisfactory host for cocksfoot clones but not vice versa. The study provides evidence of genetic variation in performance on host and evidence for clonal adaptation to particular host species. This adaptation may well be a major cause of the observed consistent genetic differentiability of populations of S. avenae found on wheat and roadside grasses in early spring in southern England.