Wing polymorphism in aphids II. Interaction between aphids

Summary A study has been made of the effect of crowding on both prenatal and postnatal control of wing development in aphids. In experiments on prenatal form control it was shown that brief controlled encounters of only a minute's duration between two adult aphids could cause them to switch from producing apterous to alate progeny. Aphids which had been exposed to contact with other aphids continued to produce alate progeny for several days. Evidence from a number of experiments suggests that the principle stimulus involved is tactile. In experiments on postnatal form control, it was shown that more alates developed among larvae which were reared together than among larvae reared in isolation.

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Zusammenfassung In großen Blattlauskolonien werden leichter Geflügelte gebildet als in kleinen. Dafür gibt es zwei Ursachen. Bei hoher Dichte verändern die Aphiden die Physiologie der Pflanze dergestalt, daß bei den Läusen, die an ihr saugen, die Geflügeltenbildung gefördert wird; und bei hoher Befallsdichte vermehrt sich für die Aphiden auch die Wahrscheinlichkeit, stimulatorische Wirkungen aufeinander auszuüben. In der vorliegenden Arbeit wird die Natur und Wirkungsweise der Wechselwirkung zwischen Individuen von Aphis craccivora Koch untersucht.Die Blattläuse wurden bei 20° und 10stündiger Photoperiode in völliger Isolation auf kleinen Blattscheiben gehalten und weiter vermehrt, die—aus Blättern ihrer Wirtspflanze herausgestanzt — auf einer Nährlösung schwammen. Unter diesen Bedingungen brachten sie überwiegend ungeflügelte Nachkommen hervor. Zur Erzeugung geflügelter Nachkommen konnten sie durch kurze Begegnungen mit anderen Aphiden angeregt werden. In einigen Versuchen wurden die Blattläuse für verschiedene Zeitabschnitte in kleine Käfige zusammengesperrt, in anderen wurden sie herausgefangen und veranlaßt, in rascher Folge mehrere Male übereinander hinwegzukriechen. Beide Behandlungsweisen bewirkten, daß die Aphiden von der Produktion ungeflügelter zur Produktion geflügelter Nachkommen umschalteten. Eine Anzahl von Versuchen berechtigen zu der Annahme, daß weder visuelle noch olfaktorische Reize dabei mitwirken, und es ist zu vermuten, daß in der Hauptsache Berührungsreize den Ausschlag geben. Der Versuch, einen entsprechenden künstlichen Reiz zu setzen, hatte nur zum Teil Erfolg.Die Blattläuse, welche einem Kontakt mit anderen Läusen ausgesetzt gewesen waren, wurden täglich auf frische Blattscheiben weitergesetzt und ihre Nachkommenschaft auf den Blattscheiben aufgezogen, auf denen sie geboren wurde. Auf diese Weise ließ es sich zeigen, daß kurze Begegnungen zwischen Blattläusen mehrere Tage lang einen Einfluß auf die Form der Nachkommenschaft beibehalten können, die sie danach produzieren.Die Flügelentwicklung von Aphis craccivora kann auch während der postnatalen Entwicklung beeinflußt werden und es ließ sich nachweisen, daß auch hierbei die Wechselwirkung zwischen den Larven einen wesentlichen Faktor darstellt. Isoliert auf Blattscheiben aufgezogene Larven entwickelten sich weniger häufig zu Geflügelten als solche, die in Dichten von zwei oder mehreren je Blattscheibe gehalten wurden.

     

Wing polymorphism in aphids III. The influence of the host plant

The condition of the host plant can have an important influence on the development of winged forms in Aphis craccivora Koch. Prenatal form control can be influenced both by the condition of the host on which the parent aphids have been reared and by the host on which they are currently feeding while reproducing. Postnatal form control can be influenced by the host on which larvae are reared during their first two instars. Whether or not the host influences wing development and the extent of its effect depend upon other factors such as stimulation of aphids by other individuals. Prolonged periods of starvation both of parent aphids and of young larvae were shown to favour apterous development.

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Flügelpolymorphismus bei aphiden III. Der einfluss der wirtspflanze

Zusammenfassung Der Einfluß des Zustandes der Wirtspflanze auf den Flügelpolymorphismus von Aphis craccivora Koch wurde unter kontrollierten Bedingungen untersucht. Zur Untersuchung des praenatalen einflusses auf die Form wurden die Mutterläuse bei 20° und 10 Stunden Licht je Tag isoliert auf Puffbohnen-Keimlingen und auf Blattscheiben von Puffbohnen gehalten, die auf Nährlösung schwammen. Die Blattläuse wurden dann als junge Adulte auf frische Blattscheiben übertragen und während der folgenden 24 Stunden die Form der produzierten Larven festgestellt. Es ließ sich zeigen, daß die Form der Nachkommen sowohl von der Wirtspflanze beeinflußt werden kann, auf der die Mutterläuse ihre Larvalentwicklung durchgemacht haben, wie von derjenigen, auf der sie sich zur Zeit der Larvenproduktion ernährten. Es wird vermutet, daß das erstere einen Nahrungseinfluß darstellen könnte, während das letztere eine Geschmacksreaktion sein könnte. Blattläuse, die auf Blattscheiben aufgezogen wurden, brachten mehr geflügelte Nachkommen hervor als diejenigen, die sich an Keimlingen entwickelten. Unter Übervölkerungsbedingungen war der Prozentsatz der Geflügeltenproduktion vergrößert und zwar stärker bei Läusen, die auf Blattscheiben aufgezogen worden waren, als bei Läusen, die auf Keimlingen heranwuchsen.Erwachsene Läuse, die geflügelte Nachkommen erzeugten, konnten durch eine dreitägige Hungerperiode zur Produktion ungeflügelter Nachkommen veranlaßt werden. Längeres Hungern bestimmte auch larven des ersten Stadiums, die ursprünglich zur Entwicklung ovn Flügeln determiniert waren, sich zu Ungeflügelten zu entwickeln. Die Wirkung des Hungers konnte bei Larven, aber nicht bei Erwachsenen durch Übervölkerung unterbunden werden.

     

Wing dimorphism influencing resistance or toxicity tests and food uptake in Myzus persicae

Toxicity or resistance tests on the green peach aphid Myzus persicae (Sulz.) by incorporating the toxicant into artificial media are influenced by a number of factors. It was found that the morph of the aphid had a pronounced effect on results. Apteriform larvae imbibed more food and at the discriminatory concentration tested were also over five times more susceptible to phosphamidon than were alatiform larvae. No difference was observed between the two aphid morphs in the inhibition of their AChE by malaoxon.

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Résumé Le milieu artificiel d'élevage utilisé est celui de Dadd et al. (1967). Les insecticides adjoints à ce milieu sont le phosphamidon et le malaoxon. Il se révèle que des larves «aptériformes» prennent plus de nourriture que les larves «alatiformes», et sont au moins cinq fois plus sensibles que ces dernières à une concentration donnée d'insecticide. Cependant le taux d'inhibition de l'acetylcholinestérase est sensiblement le même chez ces deux formes.

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Hemiptera, Aphididae.

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This research has been financed in part by a grant made by the U.S. Department of Agriculture, under PL 480.     

Wing dimorphism in Gryllus rubens: genetic basis of morph determination and fertility differences between morphs

Summary The genetic basis of wing morph determination and fertility differences between wing morphs were studied in the wing-dimorphic cricket, Gryllus rubens. Using pair corsses, a significant effect of genotype on morph determination was documented in F₃ progeny of field-collected crickets. The effect of genotype was significantly stronger in females than in males. Results are consistent with an earlier study (Zera and Tiebel 1988) showing that wing development is more strongly buffered from environmental variation in females. Segregation patterns were consistent with a polygenic mode of inheritance and provided no evidence for the existence of genes of major effect, maternal effects, or sex linkage. Only a weak morph x sex association was observed. These results contrast those of Walker (1987) where sex-linked loci of major effect on morph determination were identified in crosses between long-wing and short-wing-selected strains of G. rubens. Short-winged female G. rubens began ovipositing earlier and oviposited significantly more eggs than long-winged females during the first 24 days after adult eclosion. The greater reproductive output of the short-winged morph was due entirely to greater oviposition during the first two weeks after adult eclosion. Preliminary results indicate that flight may further accentuate the reduced reproductive output of long-winged versus short-winged females. These data provide the foundation for investigating the endocrine basis of morph-associated fertility variation in G. rubens and its mechanistic relationship with morph determination.     

Wing dimorphism and the migratory syndrome: Correlated traits for migratory tendency in wing dimorphic insects

The hypothesis that the morphological, physiological, and behavioral traits comprising the migratory syndrome in insects are genetically correlated through pleiotropic effects of genes controlling the titre of a common hormonal determinant is explored. Evidence that juvenile hormone (JH) influences the component traits of the migratory syndrome is presented, and thus JH is assumed to be the underlying, common determinant. However, readers are cautioned that this does not imply that JH is solely responsible for these traits, nor is this necessary for the arguments presented. For wing dimorphic taxa, the “correlated traits hypothesis” predicts covariance within wing morphs between JH titre and the proportion winged. Four simple genetic models for wing-morph determination are considered: single-locus with short-winged (SW) dominant; single-locus with long-winged (LW) dominant; polygenic, fixed threshold, shifting distribution; and polygenic, shifting threshold, fixed distribution. In each case, wing morphology is assumed to be a threshold trait with the liability being JH titre at some critical stage of development. All models predict covariation between %LW and the mean JH titre of at least one of the wing morphs, but the form and direction of the relationship depends critically on the genetic model used. The results suggest that we should expect the traits associated with the migratory syndrome, and hence the trade-offs associated with the evolution of wing dimorphism, to be correlated with proportion winged and, in this sense, to be frequency-dependent.     

Wing dimorphism in the migratory locust,Locusta migratoria: differentiation of wing morph and phase polyphenism

A short‐winged morph was recently discovered in the migratory locust, Locusta migratoria. It is different from the normal, long‐winged morph not only in forewing length but also in hind femur length, displaying a dimorphism. To understand the significance of this dimorphism, other morphological characters were compared between the two morphs, and the time of differentiation of wing‐pad length was investigated. Wing weights were heavier in the long‐winged morph than in the short‐winged morph. This result showed that the short‐winged morph is not formed by a failure of wing expansion. No obvious morph‐specific differences were observed in wing venation, but wing allometry studies indicated that the distal areas of the fore‐ and hindwings were disproportionally reduced in the short‐winged morph compared to the long‐winged morph. The morphological differentiation of the wing pad between the two morphs was observed at the penultimate nymphal stage. The flight muscle was well developed in the two morphs, and no sign of flight muscle histolysis was detected in either morph after adult emergence. An analysis of adult body dimensions suggested that the density‐dependent phase shifts known for the long‐winged morph of this locust were also exhibited by the short‐winged morph, demonstrating that these shifts are not specific to the migratory long‐winged morph.     

Wing shape,wing size,and sexual dimorphism in eye‐span in stalk‐eyed flies (Diopsidae)

The eyes of stalk‐eyed flies (Diopsidae) are positioned at the end of rigid peduncles (‘stalks’) protruding laterally from the head. Eye‐stalk length varies within the family and, in some species, varies between males and females. Larger eye‐stalks in males result from sexual selection for longer stalks, a trait that increases male reproductive success. In the present study, we examined whether an increase in eye‐stalk length results in an adjustment of wing size and shape to deal with the burden of bearing an exaggerated ‘ornament’. We compared wing morphology among ten species of stalk‐eyed flies that differ in eye‐span and the degree of sexual dimorphism. Mass‐specific wing length differed between males and females in seven out of the ten species. Nondimensional wing shape parameters differed between the species (P < 0.001), but mostly did not differ between males and females of the same species. Dimorphism in eye‐span closely correlated with dimorphism in wing length (r = 0.89, P < 0.001) and the correlation remained significant (r = 0.81, P = 0.006) after correcting for phylogenetic relationships. Once corrected for phylogenetic relatedness, the mass‐specific wing length of males (but not females) was weakly correlated with mass‐specific eye‐span (r = 0.66, P = 0.042). We propose that the observed proportional increase in wing length associated with increased eye‐span can facilitate aerial manoeuverability, which would otherwise be handicapped by the elevated moment of inertia imposed by the wider head. © 2009 The Linnean Society of London, Biological Journal of the Linnean Society, 2009, 98 , 860–871.     

Optimum body size in aphids

Abstract.

• 1 Smallness in British plant-sucking bugs is associated with feeding on the contents of individual plant cells, especially phloem.
• 2 The sizes of species of aphids living in similar habitats and the rates of water loss from those that have and lack a filter chamber do not support the contention that small size in aphids is a means of maximizing the loss of water by cuticular transpiration.
• 3 An optimum energy partitioning model, previously developed for aphids, was used to predict the interspecifie relation between adult weight and birth weight that would maximize the population growth rate, r_m. Using the observed r_m/IRGR ratio (0.8–0.9) and that the gonads are smaller than the soma at birth the predicted relation between the logarithm of the adult weight and the logarithm of the birth weight is linear and has a slope of 1.
• 4 The relation between the logarithm of the adult weight and the logarithm of the birth weight for eighteen species of aphids is linear and has a slope of 1.
• 5 Birth size differs between species of aphids and in each species is assumed to be the minimum size necessary for feeding. Accepting this constraint then the optimum adult size for maximizing r_m is approximately 15 times the birth size.

     

Ladybird-induced life-history changes in aphids

Predator-mediated plasticity in the morphology, life history and behaviour of prey organisms has been widely reported in freshwater ecosystems. Although clearly adaptive, similar responses have only recently been reported for terrestrial organisms. This is surprising as aphids are polyphenic and develop very rapidly compared with their predators and often produce very large colonies, which are attractive to predators. Therefore, one might expect terrestrial organisms like aphids to show a facultative change in their development in response to the presence of predators and other results have confirmed this. The results presented below indicate that the pea aphid responded to the tracks left by ladybird larvae by producing a greater proportion of winged offspring, which avoid the impending increased risk of predation by dispersing. Associated with this was a short-term increase in activity and reduction in fecundity. The black bean and vetch aphids, which are afforded some protection from ladybirds because they are ant attended and/or unpalatable, did not respond in this way to the presence of ladybird larvae.     

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.     

Sexual dimorphism in proconsul

One of the more important sources of variability in primate species is sexual dimorphism. Most Primates heavier than five kilos bodyweight are sexually dimorphic, both in body size and in shape of certain hard tissues. Despite these facts, most of the fossil Primates from East African Miocene deposits were originally perceived as being monomorphic, a perception which has propogated through the literature. Re-examination ofProconsul from various sites in Western Kenya results in the view that it was as dimorphic in its splanchonocranium and in bodyweight as chimpanzees and gorillas. The clearest evidence comes from Rusing Island, where adequate samples are known of two morphs, traditionally identified as two species, but more likely to represent two sexes of a single species,P. nyanzae. Co-occurrence of the two morphs is 100% at the various Rusinga sites. Less complete samples have been collected from the Tinderet sites os Koru and Songhor, yet what is available shows that similar patterns of dimorphism characterise the speciesP. africanus andP. major, and that the co-occurrence of the two morphs in each species is 100%. The identification of fossils taking into consideration the role of sexual dimorphism clarifies many of the old debates in which individual specimens frequently shifted between different species, mainly on the basis of metric rather than morphologic evidence. Consequently, the distribution of the species ofProconsul is rather different after accounting for dimorphism, than it was before.     

Pollen dimorphism in soybean

Summary Microspores of soybean plants (Glycine max (L.) Meer.) of four cultivars were cytologically analysed. The pollen grains showed a clear dimorphism when stained with propionic-carmine from binucleate stage onwards. The majority of the grains are large, deeply stained and with asymmetric division (normal type) while the remainder grains are smaller, lightly stained, uninucleate or with two similar nuclei (P-pollen). The different frequencies of P-pollen on the four cultivars suggest a genotype effect of microspore dimorphism.     

昆虫的雌雄二型现象

对发生雌雄二型现象的昆虫类群、生态因子及进化进行了概括总结;还特别介绍了长足虻科昆虫雌雄二型的相关方面;并简要讨论了雌雄二型与性选择的关系。     

Canine dimorphism

Canine dimorphism in many primates is exaggerated, with males possessing enormous, sharp canines that project far beyond the occlusal plane of the other teeth and females having smaller, less projecting canines. Ever since Darwin,¹ canine dimorphism generally has been attributed to sexual selection. However, recent analyses suggest that the evolution of canine dimorphism is complex and that the sexual selection hypothesis is only part of the story.     

Pelvic dimorphism in relation to body size and body size dimorphism in humans

Many mammalian species display sexual dimorphism in the pelvis, where females possess larger dimensions of the obstetric (pelvic) canal than males. This is contrary to the general pattern of body size dimorphism, where males are larger than females. Pelvic dimorphism is often attributed to selection relating to parturition, or as a developmental consequence of secondary sexual differentiation (different allometric growth trajectories of each sex). Among anthropoid primates, species with higher body size dimorphism have higher pelvic dimorphism (in converse directions), which is consistent with an explanation of differential growth trajectories for pelvic dimorphism. This study investigates whether the pattern holds intraspecifically in humans by asking: Do human populations with high body size dimorphism also display high pelvic dimorphism? Previous research demonstrated that in some small-bodied populations, relative pelvic canal size can be larger than in large-bodied populations, while others have suggested that larger-bodied human populations display greater body size dimorphism. Eleven human skeletal samples (total N: male = 229, female = 208) were utilized, representing a range of body sizes and geographical regions. Skeletal measurements of the pelvis and femur were collected and indices of sexual dimorphism for the pelvis and femur were calculated for each sample [ln(M/F)]. Linear regression was used to examine the relationships between indices of pelvic and femoral size dimorphism, and between pelvic dimorphism and female femoral size. Contrary to expectations, the results suggest that pelvic dimorphism in humans is generally not correlated with body size dimorphism or female body size. These results indicate that divergent patterns of dimorphism exist for the pelvis and body size in humans. Implications for the evaluation of the evolution of pelvic dimorphism and rotational childbirth in Homo are considered.     

Distribution of aphids in cereal crops

Cereal crops were examined weekly for aphids during 1969. Plants in twenty samples of row 0.3 m long were examined in a sheltered perimeter of a crop and along a transect 36.6 m into the crop. Aphids were usually first found within 1–4 weeks of the first alatae caught in a suction trap operating 12.2 m above ground. When first alatae caught in a suction trap operating 12.2 m above ground. When the first found from 10 to 27% of the 0.3 m lengths sampled contained aphids. Rhopalosiphum padi, first found late in May, were scarce (< 0.53/0.3 m) throughout June and July. Sitobium spp. and Metapolophium dirhodum, which appeared in mid-June, were more numerous than R. padi; most occurred during the second half of July, and populations decreased just before harvest in early August. Sitobium avenae was more abundant (max. 19.3/sample) than either S. fragariae (0.91) or M. dirhodum (2.51). More aphids occurred in oats (max. 52/0.3 m) during July than in wheat (45), and barley had fewer (6.8). S. avanae was more abundant than M. dirhodum in sheltered areas of barley and wheat, and in exposed areas of the same crop M. dirhodum was commonest. Along sheltered perimeters, the ratio of S. avenae to M. dirhodum was largest in barley (11:1), intermediate in oats (6:1) and smallest in wheat (3.7:1). Sitobium spp. were most numerous on the ears, when most M. dirhodum were on the leaves. Regression analyses of log. S² on log. m suggested that S. avenae was more evenly distributed within (36.6 m) the field (b = 1.056 + 0.109) than along the sheltered perimeter (b = 1.432 + 0.132), though it seemed similarly distributed along perimeters of barley, oats and wheat. The distributions of M. dirhodum and Sitobium spp. along sheltered perimeters of all crops were apparently similar.     

Enzyme variability in populations of aphids

Summary From 1976 to 1978 nine aphid species were investigated electrophoretically. A low degree of polymorphism was found. Only 5 of the species were polymorphic, and only at most in 3 enzyme systems out of 19.