The photoperiodic responses and phenology of an English strain of the pea aphid Acyrthosiphon pisum

Abstract. 1. The response curves for the photoperiodic induction of the sexual forms (oviparae and males) differ significantly in an English clone of the pea aphid Acyrthosiphon pisum Harris. Male production is sharply peaked. The late summer scotophases (dark periods) that initiate male production are 1.0–1.5 h shorter than those that initiate ovipara production; the induction of males, but not oviparae, virtually ceases at scotophases longer than 12 h. This disparity suggests that there are two photoperiodic clocks.
2. All ovipara-producing aphids switch spontaneously to virginopara production part-way through the progeny sequence, irrespective of photoperiod. This may confer the ability to overwinter partheno-genetically under favourable climatic conditions. Since males are always born last in the progeny sequence this possibility would be pre-empted unless the terminal male sequences were suppressed by long winter scotophases.
3. The role of the photoperiodic response in determining the seasonal phenology was tested in the field by exposing a succession of laboratory-reared aphids to natural photoperiods, in late summer and autumn. Adult males and oviparae developed synchronously in early October, the difference in critical scotophase compensating for the late appearance of males in the progeny sequence. A large part of the 'civil twilight' is photoperiodically active.
4. Clones propagated outdoors by isolating aphids in each generation from the terminal sequence of virginoparae did not survive the whole winter of 1985/86. The chances of survival were reduced by severe weather and by the inability of the photoperiodic system to 'turn off' the production of early born oviparae which are, therefore, 'wasted'.     

Juvenile hormone effects on polymorphism in the pea aphid, Acyrthosiphon pisum

When reared in short days (LD 12:12) at 15°C, apterous Acyrthosiphon pisum gave birth to sexual females (oviparae) exclusively for the first eight days of larviposition. After this time they switched to the production of parthenogenetic females (viviparae). Topical application of juvenile hormones I, II and III to fourth instar or adult ovipara-producers induced the precocious appearance of parthenogenetic females in the progeny sequence. Various forms intermediate between oviparae and viviparae were also produced and repetitive JH-I treatments resulted in a few alatiform progeny. However, many of the JH induced apterous, parthenogenetic females appeared to be normal viviparae and were capble of reproduction. Thus, prenatal treatment of oviparous embryos with JH diverts development towards the viviparous form. JH-I treatment of long-day reared A. pisum had no effect on the type of progeny produced.

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Effets de l'hormone juvénile sur le polymorphisme d'Acythosiphon pisum

Résumé Quand il est élevé sous jours courts (LN 12/12) à 15°C, le type anglais vert d'Acyrthosiphon pisum ne donne naissance qu'à des femelles sexuées (ovipares) pendant la première partie de sa période de reproduction. Ensuite quelques types intermédiaires ovipares/vivipares peuvent apparaitre avant que les pucerons ne bifurquent spontanément vers la production de femelles parthénogénétiques (vivipares). L'application cutanée d'hormones juvéniles (JH I, II, et III) aux larves de quatrième stade ou à des adultes producteurs d'ovipares provoque l'apparition prématurée d'intermédiaires et de vivipares dans la descendance. Les différentes formes intermédiaires produites par des applications répétées de J.H. comprenaient des types ailés ou partiellement ailés. Cependant, les vivipares aptères induits par J.H. étaient morphologiquement normaux et beaucoup étaient capables de se reproduire. Des traitements semblables aux J.H. de vivipares élevées en jours longs (LN 16/8) n'ont pas eu d'effets sur le type de la descendance.On ne sait pas si l'action de JH exogène sur l'induction des vivipares est direct ou indirect. La reprogrammation des embryons, autrement destinés à se développer comme ovipares, est examinée en relation avec notre connaissance du contrôle endocrine du polymorphisme des pucerons.

     

Body colour and genetic variation in winged morph production in the pea aphid

Aphids (Homoptera: Aphidoidea) produce a number of different phenotypes in their life-cycle, among which are winged (alate) and wingless (apterous) morphs. Lowe & Taylor (1964) and Sutherland (1969a, b) were the first to suggest that aphid clones differ in their propensity to produce the winged morph and that in the pea aphid (Acyrthosiphon pisum Harris), this propensity is linked to the colour of the phenotype. We tested for the occurrence of genetic variation in winged morph production by rearing individuals from red and green clones of pea aphid under wing-inducing (crowding) and control conditions, and scored the phenotypes of their offspring. Clones differed significantly in alate production and red clones produced on average a higher proportion of winged morphs than green clones. Importantly, however, there was considerable variation between clones of the same colour. Broad-sense heritabilities of winged morph production were 0.69 (crowding treatment) and 0.63 (control). Clones also differed in the number of offspring they produced. When exposed to the crowding stimulus, aphids deferred offspring production, resulting in a higher number of offspring produced in the crowding treatment than in the control.     

Effects of parasitism by Praon pequodorum on age-specific fecundity and population growth of the pea aphid, Acyrthosiphon pisum

The effects of parasitism by Praon pequodorum Viereck (Hymenoptera: Aphidiidae) on the survival, development and reproduction of apterous pea aphids, Acyrthosiphon pisum (Harris) (Homoptera: Aphididae), were studied in the laboratory. Ten cohorts of aphids differing in age from 0.5 h to 9 days were tested. At 20°C, pea aphids survived an average of 7 days after parasitization. Parasitized first and second instars died before reaching the adult stage. Third instars died as adults but without producing any offspring. Parasitized fourth instars and adult aphids produced a variable number of offspring; their mean total fecundity increased as an exponential function of the age at parasitization. Host age did not significantly affect the period from the beginning of parasitism to the end of reproduction. Equations are provided to estimate the length of the reproductive period, the total fecundity, the intrinsic rate of increase, and the doubling time as functions of aphid age at parasitization. The data are discussed with regard to host population growth. It is shown that a parasite's potential impact on an aphid population is critically dependent on its preference for and successful oviposition in early host instars.

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Résumé Les effets du parasitisme par P. pequodorum Viereck (Hym.: Aphidiidae), sur la survie, le développement et la reproduction des aptères de A. pisum, ont été examinés au laboratoire. L'étude a porté sur 10 cohortes de pucerons dont les âges différaient de 0,5 heure à 9 jours. A 20°C, les pucerons parasités ont survécu en moyenne pendant 7 jours après avoir été parasités. Leur évolution ultérieure dépend au stade auquel ils ont été parasités: au premier et second stades, ils sont morts avant de devenir adultes; au troisième stade, ils sont morts adultes avant de se reproduire; au quatrième stade et adultes, ils ont donné un nombre variable de descendants. La fécondité totale moyenne a crû comme une fonction exponentielle de l'âge au moment où ils avaient été parasités. L'âge de l'hôte n'a pas modifié significativement la durée de la période entre le moment où ils ont été parasités et la fin de la phase de reproduction. Des équations ont été établies en fonction de l'âge des pucerons au moment où ils ont été parasités, pour estimer la longueur de la période de reproduction, la fécondité totale, le taux intrinsèque d'accroissement et le temps de doublement des cohortes de pucerons. Les résultats sont discutés sous l'angle de la dynamique de la population. On en déduit que l'impact potentiel d'un parasite sur une population de pucerons dépend d'abord de ses préférences pour les premiers stades de l'hôte et du succès de sa ponte sur ces stades.

     

光周期对两种色型豌豆蚜种群参数的影响

为探讨光周期对红绿两种色型豌豆蚜种群增长的影响,于室内模拟光周期22L∶2D、16L∶8D、10L∶14D和4L∶20D四个水平,研究了红绿两种色型豌豆蚜的发育、繁殖和生命表。结果表明:相对于其它光周期,在光周期22L∶2D时,红绿两种色型豌豆蚜若蚜存活率最高,繁殖力最强;其红色型豌豆蚜种群增长内禀增长率(r_m=0.3263)高于绿色型豌豆(r_m=0.2863)。光周期16L∶8D时,红色型豌豆蚜的若蚜存活率(72.22%)高于绿色型豌豆蚜(69.45%),而成蚜寿命(6.58d)较短;但绿色型豌豆蚜内禀增长率(r_m=0.2648)高于红色型豌豆蚜(r_m=0.2249),其种群繁殖力强。光周期10L∶14D时,红色型豌豆蚜与绿色型豌豆蚜相比,若蚜存活率(80.55%)高,内禀增长率(r_m=0.2490)高,种群增长力强,而绿色型豌豆蚜成蚜寿命(12.71d)较长。光周期4L∶20D时,两种色型豌豆蚜若蚜存活率最低,繁殖力最弱,成蚜寿命显著延长;红色型豌豆蚜若蚜存活率(64.67%)高于绿色型(35.50%),成蚜没有产生后代;绿色型豌豆蚜内禀增长率(r_m=-0.0366),其种群为负增长。综上,短光照条件(4L∶20D)延长了两种色型豌豆蚜的若蚜发育历期及成蚜寿命,降低了若蚜存活率、繁殖力及生命表各项参数;长光照条件(22L∶2D)缩短了两种色型豌豆蚜的若蚜发育历期及成蚜寿命,若蚜存活率高、繁殖力强;红色型豌豆蚜更适宜光周期10L∶14D,而绿色型豌豆蚜更适宜光周期16L∶8D。     

醉马草水浸液对豌豆蚜触杀活性及种群增长的影响

为探讨醉马草水浸液对豌豆蚜触杀活性及种群增长的影响,采用带虫浸叶法比较不同生育期醉马草带菌（E+）和不带菌（E-）水浸液对豌豆蚜触杀活性及种群生命表,测定了豌豆蚜的死亡率及触杀后对其生殖期,平均繁殖力,繁殖率及生命表参数的影响。结果表明,不同生育期醉马草带菌（E+）水浸液触杀豌豆蚜后对其各项指标均有显著影响。在苗期时,E+水浸液触杀豌豆蚜后校正死亡率最高,繁殖力最低,内禀增长率（r_m=0.145 d^-1）和净生殖率（R₀=4.802头）均为最小值。在成熟期时,E+水浸液触杀豌豆蚜24 h、48 h和72 h后校正死亡率分别为26.15%,19.01%,9.07%;繁殖期（3.87 d）,平均繁殖力（8.80头）,繁殖率（1.40%）,内禀增长率（r_m=0.208 d^-1）和净生殖率（R₀=8.820头）。在枯黄期时,E+水浸液触杀豌豆蚜后校正死亡率最低,繁殖力最强,内禀增长率（r_m=0.247 d^-1）和净生殖率（R₀=13.647头）均为最大值。不同生育期醉马草E-水浸液触杀豌豆蚜后对其种群繁殖无显著影响,与对照差异不显著（P>0.05）。综上,苗期醉马草E+水浸液对豌豆蚜有较好的触杀效果,校正死亡率高,且触杀后当代繁殖力减弱,种群扩建时间延长,不利于其种群繁殖和增长;故苗期醉马草E+水浸液具有很好的杀虫潜力,所采用水浸液方法制备简单,成本低,可为新型植物源农药研发提供重要理论依据。     

High variation in host adaptation among clones of the pea aphid,Acyrthosiphon pisum on peas,Pisum sativum

The performance of one clone of the pea aphid,Acyrthosiphon pisum (Harris), was assessed on 37 different cultivars and species ofPisum L. In addition, random samples of 36 pea aphid clones collected on alfalfa and clover were tested on a selection of fivePisum sativum L. cultivars. Aphid performance was evaluated in terms of the mean relative growth rate (MRGR) during the first five days of life or other life history variables. The MRGR of the first-mentioned pea aphid clone differed little between cultivars. No significant differences in MRGR were found between wild and cultivatedPisum species or between modern and oldP. sativum cultivars. There was considerable variation in host adaptation among the 36 pea aphid clones within each sampled field. The pea aphid clones showed no consistent pattern in performance on four of the five pea cultivars i.e. there was a significant pea aphid genotype —pea genotype interaction. On one of the cultivars all clones performed well. Pea aphid clones collected from red clover generally performed relatively poorly on pea cultivars, in contrast to the pea aphid clones collected on alfalfa. There was no difference in performance between the two pea aphid colour forms tested. Possible reasons for the high variation and the observed adaptation patterns are discussed. The fact that all clones were collected in two adjacent fields indicates thatA. pisum shows high local intraspecific variability in terms of host adaptation.     

Resistance of different lucerne cultivars to the pea aphid Acyrthosiphon pisum: influence of phloem composition on aphid fecundity

Sugars and amino acids were analysed in the phloem sap (sampling by the exudation method) of four clones of lucerne (Medicago sativa L.) characterized by their resistance to the pea aphid, Acyrthosiphon pisum (Harris) (Homoptera: Aphididae). Differences in the qualitative amino acid balance are shown and a negative correlation between the reproductive rate of the aphid and the sugars/amino acids ratio is underlined. To demonstrate any causal relationship between these two variables, the reproduction of A. pisum on artificial diets copying the sap of the two extreme clones (resistant and susceptible) was measured. The results show that the resistance is not a simple nutritional effect; however, the amino acid balance contributes to the resistance exhibited by some cultivars.

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Résumé L'analyse (sucres et acides aminés) de la sève de 4 cultivars de luzerne (Medicago sativa) caractérisés pour leur résistance au puceron du pois (Acyrthosiphon pisum (Harris)) (Homoptera: Aphididae), est réalisée sur des exsudats de phloème. La teneur en sucres (exprimée en équivalent saccharose) montre d'importantes différences: rapport de 1 à 3 respectivement entre les clones sensible et résistant. La teneur globale en acides aminés exsudés varie en sens inverse, entraînant une corrélation négative pour les 4 clones étudiés entre le rapport sucre/acides aminés (0,6 à 3,6) et la résistance mesurée par le taux net de reproduction à 14 jours (78,5 à 11,2 larves/femelle). L'analyse des acidses aminés révèle de plus des différences qualitatives entre cultivars, notamment pour certains acides aminés essentiels méthionine, histidine, lysine, aromatiques.Le rôle de l'équilibre en acides aminés et en sacharose dans la résistance de certains cultivars au puceron du pois est apprécié par des mesures de la reproduction de A. pisum sur des milieux artificiels reproduisant le rapport sucre/acides aminés et/ou le spectre des acides aminés des 2 clones extrêmes (sensible et résistant). La résistance n'apparaît pas être le résultat d'une simple différence de la qualité nutritionnelle de la sève; la balance en acides aminés doit cependant participer pour une part à la résistance au puceron.

     

The elimination of intracellular microorganisms from insects: an analysis of antibiotic-treatment in the pea aphid (Acyrthosiphon pisum)

Antibiotics are routinely used to eliminate intracellular prokaryotic microorganisms from a wide range of insect species, but concerns about deleterious effects of antibiotic therapy on the insect host are seldom addressed. Here, the impact of antibiotic therapy in the symbiosis between the pea aphid Acyrthosiphon pisum and bacteria of the genus Buchnera is reviewed. Antibiotic-treatment produces aposymbiotic (i.e. symbiont-free) aphids, but does not depress the mitochondrial complement, the assimilation of dietary amino acids or the incorporation of amino acids into protein in these insects and does not impair osmoregulation, feeding rate and the capacity to penetrate plant tissues. It is concluded that the general malaise associated with aposymbiotic aphids is not attributable to a direct effect of the antibiotic. However, an important implication of this study is that aposymbiotic insects exhibit substantial metabolic adjustments to loss of the symbiosis; they are not simply aphids from which the symbiotic bacteria have been removed.     

Genetic variation in asexual populations of two aphids in the genus Acyrthosiphon, from an Australian lucerne field

Twelve lines of Acyrthosiphon kondoi Shinji and Acyrthosiphon pisum (Harris) were sampled from a small lucerne field in Australia and tested to assess genetic variation by measuring their responses to three lucerne cultivars. Seven or eight life history traits were assessed for each species. Differences in the responses of these traits to the cultivars revealed that three to seven genetically distinct clones were represented in the samples of 12 lines for each species. We conclude that the populations of these two aphids are variable in Australia, although they were recently introduced and reproduce asexually. The responses of the aphids to the three cultivars are described. Because of genetic variation in host plant responses of the aphids, cultivars with potential aphid resistance must be exposed to a large number of clones to confirm the presence of useful aphid resistance.

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Résumé Douze lignées d'Acyrthosiphon kondoi Shinji et d'A. pisum Harris ont été échantillonnées dans un petit champ de luzerne australien et leur variabilité génétique examinée en comparant les réponses à 3 cultivars de luzerne. De 7 à 8 caractères biologiques avaient été retenus pour chaque espèce. Les différences observées entre les réponses suivant la distribution de ces caractères chez les pucerons, ont montré que 3 à 7 clones génétiquement distincts étaient représentés dans les 12 lignées de chaque espèce. Nous en concluons que les populations de ces deux espèces de pucerons sont polymorphes en Australie, quoiqu'elles se multiplient d'une façon asexuée et aient été récemment introduites. Les réactions des pucerons aux 3 cultivars sont indiquées. Par suite de la variabilité génétique des réactions des plantes hôtes aux pucerons, des cultivars à résistance potentielle élevée devraient être exposés à un grand nombre de clones de pucerons pour confirmer la présence d'une résistance utilisable.

     

Learning by the parasitoid wasp, Aphidius ervi (Hymenoptera: Braconidae), alters individual fixed preferences for pea aphid color morphs

Learning, defined as changes in behavior that occur due to past experience, has been well documented for nearly 20 species of hymenopterous parasitoids. Few studies, however, have explored the influence of learning on population-level patterns of host use by parasitoids in field populations. Our study explores learning in the parasitoid Aphidius ervi Haliday that attacks pea aphids, Acyrthosiphon pisum (Harris). We used a sequence of laboratory experiments to investigate whether there is a learned component in the selection of red or green aphid color morphs. We then used the results of these experiments to parameterize a model that examines whether learned behaviors can explain the changes in the rates of parasitism observed in field populations in South-central Wisconsin, USA. In the first of two experiments, we measured the sequence of host choice by A. ervi on pea aphid color morphs and analyzed this sequence for patterns in biased host selection. Parasitoids exhibited an inherent preference for green aphid morphs, but this preference was malleable; initial encounters with red aphids led to a greater chance of subsequent orientation towards red aphids than predicted by chance. In a second experiment, we found no evidence that parasitoids specialize on red or green morphs; for the same parasitoids tested in trials separated by 2 h, color preference in the first trial did not predict color preference in the second, as would be expected if they differed in fixed preferences or exhibited long-term (> 2 h) learning. Using data from the two experiments, we parameterized a population dynamics model and found that learning of the magnitude observed in our experiments leads to biased parasitism towards the most common color morph. This bias is sufficient to explain changes in the ratio of aphid color morphs observed in field sites over multiple years. Our study suggests that for even relatively simple organisms, learned behaviors may be important for explaining the population dynamics of their hosts.     

Juvenile hormone titres and winged offspring production do not correlate in the pea aphid, Acyrthosiphon pisum

Pea aphids, Acyrthosiphon pisum, reproduce parthenogenetically and are wing-dimorphic such that offspring can develop into winged (alate) or unwinged (apterous) adults. Alate induction is maternal and offspring phenotype is entirely determined by changes in the physiology and environment of the mother. Juvenile hormones (JHs) have been implicated in playing a role in wing differentiation in aphids, however until recently, methods were not available to accurately quantify these insect hormones in small insects such as aphids. Using a novel LC-MS approach we were able to quantify JH III in pea aphids that were either producing a high proportion of winged morphs among their offspring or mainly unwinged offspring. We measured JH III titres by pooling the hemolymph of 12 or fewer individuals (1 μL hemolymph) treated identically. Levels of JH ranged from 30 to 163 pg/μL. While aphids in the two treatments strongly differed in the proportion of winged morphs among their offspring, their JH III titres did not differ significantly. There was also no correlation between JH III titre and the proportion of winged offspring in induced aphids. This supports earlier findings that wing dimorphism in aphids may be regulated by other physiological mechanisms.     

Instar-specific defense of the pea aphid,Acyrthosiphon pisum: Influence on oviposition success of the parasiteAphelinus asychis (Hymenoptera: Aphelmidae)

The parasite Aphelinus asychisWalker (Hymenoptera: Aphelinidae) oviposits in all four instars of the pea aphid, Acyrthosiphon pisum(Harris) (Homoptera: Aphididae). Searching females display a highly stereotyped sequence of behaviors when encountering a host. Once recognized, an aphid is examined and probed by the wasp with the everted ovipositor prior to oviposition. Oviposition success is influenced by aphid behavior that is related to aphid size and expressed through instarspecific escape and defense reactions. Being smaller and less able to defend themselves, first and early-second instars of pea aphid are more susceptible to successful parasitism than third and fourth instars, in that order. Observed patterns of preference by Aphelinus females for particular aphid species and instars reflect the outcome of behavioral interactions between the hosts and the parasites, rather than preference in the strict sense.     

The effects of two species of hymenopterous parasitoid on the reproductive system of the pea aphid, Acyrthosiphon pisum

Two cultures of first instar pea aphids, Acyrthosiphon pisum (Harris) (Homoptera: Aphididae), were subjected to parasitization by Ephedrus plagiator (Nees) and Aphidius ervi Haliday (Hymenoptera: Braconidae) respectively, and a third culture, of fourth instar pea aphids, was subjected to parasitization by E. plagiator. Significant parasitogenic effects on the reproductive system occurred within 24 h of exposure to parasitoids in the form of reductions in both embryo number and the size of the largest embryo, and degenerative changes in the embryos themselves. No evidence was found of direct feeding by the larvae on the host's tissues, until just before larval pupation. The largest embryos of some fourth instar aphids escaped the effects of parasitization probably because they had developed a resistant cuticle by the time of oviposition by the parasitoid. Results suggested that embryonic degeneration was an indirect effect of parasitization due to starvation, interference with the aphid's hormone levels, or both.

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Résumé Trois élevages de 100 à 200 premiers stades d'A. pisum, nés à 24 heures les uns des autres, ont été divisés chacun en un lot témoin et un lot expérimental. Deux des élevages expérimentaux ont alors été soumis au parasitisme d'Ephedrus plagiator ou de Aphidius ervi, le troisième élevage a pu se développer jusqu'au 4è stade avant d'être exposé à E. plagiator. Des échantillons de 10 pucerons ont été retirés de chaque lot après 24 heures et ensuite toutes les 48 heures pour être disséqués dans du liquide physiologique. Les nombres d'ovarioles et d'embryons et la taille du plus gros embryon ont été notés. Des cellules développées, issue de la séreuse de l'oeuf de l'hyménoptère se sont disséminées dans l'hémolymphe de l'hôte et ont servi d'aliment aux larves en développement. Seul le dernier stade larvaire des parasitoïdes s'est alimenté sur les tissus de l'hôte et ceci avait lieu juste avant la nymphose. Des effets significatifs se sont traduits après 24 heures par la réduction du nombre des embryons, par la diminution de la taille du plus grand de ceux-ci et par des altérations dans les embryons eux-mêmes.

     

Distribution in time and space of resistance to the pathogenic fungus Erynia neoaphidis in the pea aphid Acyrthosiphon pisum

Two biotypes of pea aphid, Acyrthosiphon pisum, have been recognized in Australia, by their susceptibility (S) or resistance (R) to certain isolates of the fungal pathogen Erynia neoaphidis. The responses of these two biotypes to 40 isolates of the fungus have shown that the R form is largely confined to two States, New South Wales and Victoria, but appears to have recently spread into Queensland and Tasmania. There is no evidence to suggest it occurs outside Australia. Sequential sampling of two field populations of pea aphids during 1981 and 1982 showed that the proportion of R form remained stable at 10.7±3.0 and 14.6±2.6% (mean±standard error) for the two populations. Glasshouse competition experiments run at the comparatively high temperature of 25°C resulted in the R form becoming dominant even when the initial ratio was 4:1 in favour of the S form. The ecological and genetical implications are discussed.

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Résumé Deux types biologiques d'Acyrthosiphon pisum ont été définis, en Australie, suivant leur sensibilité (S) ou leur résistance (R) à certains isolats du champignon pathogène Erynia neoaphidis. Les réponses des deux biotypes à 40 isolats du champignon ont montré que la forme R est essentiellement cantonnée à deux états: Nouvelle Galle due Sud et Victoria, mais a récemment gagné le Queensland et la Tasmanie. Aucun élément ne fait dire qu'elle existe hors d'Australie. Des échantillonnages séquentiels des deux populations de pucerons dans la nature en 1981 et 1982 ont montré que la proportion de la forme R est restée stable à 10,7 et 14,62 dans les deux populations. Des expériences de compétition en serre à la température relativement haute de 25°C ont rendu la forme R dominante, même quand le rapport initial était de 4/1 en faveur de la forme S. La discussion porte sur les conséquences écologiques et génétiques.

     

Inducing the sexual forms and hatching the eggs of pea aphids

In temperate climates, pea aphids (Acyrthosiphon pisum) produce a single sexual generation each year in response to declining photoperiod and temperature. Mating occurs in the fall and the eggs have an obligatory winter diapause. Genetic recombination during the sexual phase is thought to be an important source of genetic variability within cyclically parthenogenetic aphid populations. Methods for reliably producing sexual forms and hatching the eggs of aphids are therefore central not only to the study of evolutionary change in aphid populations, but also for a general understanding of the origin of agriculturally important variation in destructiveness within pest species.Here, sexual forms of six pea aphid clones were induced in the laboratory and eggs were successfully hatched by creating conditions that closely mimicked those found in field situations. A declining photoperiod was produced by controlling artificial lighting using a timer with variable cycle length. Using these conditions, sexual forms were successfully produced for all six clones tested, which were then mated in all combinations. Eggs were exposed to a daily cycle of freezing and thawing in an incubator under a short-day photoperiod. Egg hatch averaged 60%, but was as high as 89% for some crosses. These methods will permit testing of evolutionary hypotheses and execution of detailed genetic studies of sources of variability within pea aphid populations. They are thus important tools for both evolutionary and agricultural studies.     

The significance of gut sucrase activity for osmoregulation in the pea aphid, Acyrthosiphon pisum

The osmotic pressure of the body fluids of aphids is lower than in their diet of plant phloem sap. It is hypothesised that aphids reduce the osmotic pressure of ingested food by sucrase-mediated hydrolysis of dietary sucrose to glucose and fructose, and the polymerisation of glucose into oligosaccharides of low osmotic pressure per hexose unit. To test this hypothesis, the impact of the alpha-glucosidase inhibitor acarbose on the sugar relations and osmoregulation of aphids was explored. Acarbose inhibited sucrase activity in gut homogenates and the production of monosaccharides and oligosaccharides in the honeydew of live aphids. Acarbose caused an increase in the haemolymph osmotic pressure for aphids reared on a diet (containing 0.75 M sucrose) hyperosmotic to the haemolymph and not on the isoosmotic diet containing 0.2 M sucrose. It did not affect aphid feeding rate over 2 days, except at high concentrations on 0.75 M sucrose diet, and this may have been a secondary consequence of osmotic dysfunction. Acarbose-treated aphids died prematurely. With 5 microM dietary acarbose, mean survivorship on 0.2 M sucrose diet was 4.2 days, not significantly different from starved aphids, indicating that, although these aphids fed, they were deprived of utilisable carbon; and on 0.75 M sucrose diet, mean survivorship was just 2.8 days, probably as a consequence of osmotic failure. It is concluded that the aphid gut sucrase activity is essential for osmoregulation of aphids ingesting food hyperosmotic to their body fluids.