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.     

Embryogenesis and the production of males by apterous viviparae of the green peach aphid Myzus persicae in relation to photoperiod

Apterous viviparae of a holocyclic strain of Myzus persicae were given various numbers of prenatal and postnatal exposures to male-inducing, long-night scotophases (15 hr darkness per diem). The sex of the embryos in various positions within their ovarioles was determined from chromosome preparations and correlated with the sequence with which female and male larvae were deposited. An abnormal follicle was found frequently between the female and male embryos in the ovarioles of aphids raised under long nights.Apterous viviparae exposed to long nights from birth or earlier usually produced several females and then switched over to exclusive male production. The more prenatal exposures to long nights given to the aphids, the fewer females were deposited by them and the earlier they started to produce males. When apterae were continuously exposed to long nights from 2 days or more before their birth, some, and occasionally all, their ovarioles were solely androparous. In general, male-producing apterae deposited considerably fewer larvae (female as well as male) than apterae that produced females exclusively. The 1–2 day pause in larviposition that typically occurs between the deposition of the female and male larvae was attributed to a slower rate of development of the male embryos. The results are discussed in relation to the hormonal changes that may be induced by the photoperiodic treatments.     

Ovariole number and fecundity for the two emigrating generations of the bird cherry-oat aphid (Rhopalosiphum padi) in Sweden

Abstract. 1. The number of ovarioles and the potential fecundity in first (fundatrigenia) and second generation emigrants of Rhopalosiphum padi (L.) was investigated.
2. The relationships between weight and host quality on ovariole number were investigated.
3. The first generation of emigrants had a greater number of ovarioles and a higher fecundity than the second.
4. There was a positive relationship between ovariole number and weight.
5. The number of ovarioles and weight decreased with decreasing host quality.
6. The results do not agree with a recently proposed reproductive strategy of aphids where the number of ovarioles are supposed to be independent of weight and host quality.
7. In years with a high initial population on bird cherry, the migration to grasses, e.g. cereals, will be earlier, probably larger and, to a greater extent, consists of aphids which have a higher reproductive rate than in years with a small initial number on bird cherry.     

Development and reproductive potential of the cereal aphid Sitobion avenae on resistant wheat lines (Triticum monococcum)

The effect of wheat resistance in lines of Triticum monococcum L., on the reproductive performances of the cereal aphid (Sitobion avenae F.) was investigated. Aphids were reared from birth to adult moult either on resistant or susceptible wheat lines, and transferred as apterae to both host genotypes. The influence of these transfers on the subsequent adult weight, gonad status and reproductive performances was evaluated. Aphids transferred from resistant to susceptible plants proved able to compensate for their poor nymphal growth, mainly through additional embryo growth and an increase in the number of matured embryos within the first 10 days of their adult life. Most aphids transferred from susceptible to resistant plants died within the first week following the transfer. Their most advanced embryos matured and were born, but subsequent embryo growth was quickly reduced. The reproductive strategies adopted by S. avenae when facing plant resistance, and the hypothesis of a resistant mechanism based on a poor nutritional state of the resistant plants are discussed.     

Differential colonization of wheat cultivars by two biotypes of Russian wheat aphid （Homoptera： Aphididae）

Susceptible and resistance wheat cultivars, Triticum aestivum L, were presented to two biotypes of Russian wheat aphid, Diuraphis noxia （Mordvilko）, in multiple choice tests to assay their relative acceptability as host plants. Both apterae （third and fourth instars） and alate adults were offered plants at the two-leaf stage in different cultivar combinations at 22±1℃ and 16：8 （L： D） hour photoperiod. Apterae were released from Petri dishes in the center of a circle of test plants, whereas alatae dispersed from a mature aphid colony to settle on plants arranged in rows. Both alatae and apterous nymphs of both biotypes readily colonized all cultivars tested：‘2137＇, ‘Akron＇,‘Ankor’,‘ Halt’ ,‘ Jagger’ ,‘ Prairie Red’ , ‘Stanton＇,‘TAM 107＇,‘TAM 110＇,‘Trego＇, ‘ Yuma＇, and ‘Yumar＇. Fewer biotype I apterae responded （settled and fed） in the combination containing more resistant （Dn4- and Dny-expressing） cultivars, compared to the combinations that had fewer. The reverse was true for biotype 2 apterae; more aphids responded in the combination containing the largest number of Dn4 expressing cultivars. Differential colonization of cultivars was observed in only one combination, in which biotype 2 apterae colonized Akron and Yumar in larger numbers than they did Stanton and Yuma. A separate experiment confirmed that, 48 hours after infestation, more biotype 2 apterae abandoned plants of Yuma than plants of Yumar. This differential response was likely due to genetic differences between the two ＇ near isogenic＇ lines that include the lack of Dn4 expression in Yuma. Choice tests with alatae did not result in differential rates of cultivar colonization by either biotype in any combination tested. These results suggest that young wheat plants appear to lack any meaningful antixenosis toward D. noxia, even though the aphids appear to perceive, and sometimes respond to, certain differences in cultivar suitability.     

Production of sexual morphs by apterous virginoparae of Myzus persicae (Hemiptera: Aphididae) in relation to pre- and postnatal exposure to short day conditions

The production of males and females by apterae of a holocyclic clone of Myzus persicae (Sulzer) was studied in the laboratory by transferring apterous virginoparae from long day to short day conditions at different stages of their pre- and post-natal development. Prenatal exposure was also examined in nine additional clones coming from different regions of Greece. By increasing the number of short day cycles at 17 degrees C, the aphids switched to male production earlier and produced more males. The highest mean number of male progeny was observed in apterae that were transferred to short day conditions 13-14 days prenatally. Apterae that were transferred to short day conditions after the fourth nymphal instar produced only females. Apterous virginoparae were produced by aphids that were transferred to short day conditions 0-1 days prenatally or postnatally. Aphids, given short days prenatally or in the first instar, produced alate female progeny that were all gynoparae. The later the instar aphids were transferred to short day conditions the fewer the gynoparae produced among their female progeny. A reduction in the total number of progeny and embryos per ovariole was observed as aphids were transferred to short day conditions in earlier stages of their development. The increase in male production, which follows the increase in the duration of long night exposure, was also observed in the additional eight out of nine examined clones.     

Effects of population density on alienicolae of Aphis fabae Scop.

A technique for rearing aphids in dense populations is described. Apterae were reared in high (‘C’) and low (‘U’) population densities and the proportions of alatae among their progenies were compared. Proportions of alatae were larger among progenies of ‘C’ apterae, and the proportions also increased as the density of the population in which the progenies were reared increased. Progenies of alatae never became alatae.     

A test of three hypotheses for ovariole number determination in the grasshopper Romalea microptera

Ovariole number in insects determines potential fecundity and can be influenced by genes, environmental conditions during development and parental effects. In the present study, three hypotheses are tested for ovariole number determination in the grasshopper Romalea microptera (Beauvois), which exhibits both intra‐ and interpopulation variation in ovariole number. In hypothesis 1, variation in ovariole number is a result of genetic variation. In hypothesis 2, ovariole number is influenced by nutrition during development. In hypothesis 3, ovariole number is influenced by maternal nutritional status. Females from four treatments are compared: low‐food, high‐food, daughters of low‐food, and daughters of high‐food. There is a relationship between parent and offspring ovariole number despite different environments, supporting hypothesis 1. Also, ovariole numbers are slightly, but significantly lower in individuals fed a low‐food diet compared with a high‐food diet, supporting hypothesis 2. Hypothesis 3 is not supported: starved and well‐fed females produce eggs of similar mass, as well as offspring with similar numbers of ovarioles, suggesting that the nutritional status of mothers does not influence offspring mass or offspring ovariole number. The results imply that genetic variation and developmental conditions determine ovariole number in this species but maternal environment does not. These results conflict with previous studies of ovariole determination in grasshoppers and locusts.     

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

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

Precocious metamorphosis of the aphid Myzus persicae induced by the precocene analogue 6-methoxy-7-ethoxy-2,2-dimethylchromene

For up to three days after being treated with the precocene analogue 6-methoxy-7-ethoxy-2,2-dimethylchromene, adult green peach aphids (Myzus persicae) gave birth to offspring (alate and apterous virginoparae, males) which underwent precocious metamorphosis. Only a few precocious gynoparae and no precocious oviparae were recorded. The precocious aphids became adultoid in the third or fourth-instar as a result of corpus allatum destruction. They were able to develop mature embryos, but could not deposit them because of incomplete differentiation of the reproductive tract. The treatment did not induce the production of alate virginoparae in the experimental clone of Myzus persicae. However, a few males were born late in the reproductive sequence of treated apterae. The corpora allata of the treated adults appeared on histological examination to be unaffected by precocene.     

Parasitoid-host relationship betweenTrioxys (Binodoxys) Indicus [Hymenoptera: Aphidiidae] andAphis Craccivora [Hemiptera: Aphididae]. VI. Impact of males on the number of progeny of the parasitoid reared on certain host plants

The present paper elucidates the impact of males on the reproductive rate ofTrioxys (Binodoxys) indicus reared on the aphids ofCajanus cajan, Dolichos lablab andSolanum melongena host plants. The F₁ offspring increases with the increase of parasitoid number. This increase is maximum inC. cajan followed byD. lablab andS. melongena reared aphids. The presence of male parasitoids caused a significant decrease in the emergence of F₁ generation.      

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.

     

Reproductive characteristics of the flower breeding Drosophila hibisci Bock (Drosophilidae) along a latitudinal gradient in eastern Australia: relation to flower and habitat features

For many insect species, egg and larval substrate characteristics are significantly correlated with interspecific differences in female reproductive allocation and egg size-number tradeoffs. We tested the hypothesis that a similar pattern occurred within the Australian drosophilid, Drosophila hibisci, that is restricted throughout its life cycle to flowers of species in the genus Hibiscus. These plants occur as small, isolated, normally monospecific stands that should facilitate differentiation of the fly populations in relation to specific oviposition and larval substrates. Data from 38 sites ranging from 20.8̀ to 34.4̀ S latitude in eastern Australia indicated no relationship between female body size, egg size, or ovariole numbers and floral size or mass among four species of Hibiscus. However, the flies did show a latitudinal cline in ovariole number that was independent of floral variation. Females averaged 15–20 ovarioles per female in the south (32–34̀ S latitude) and 10–12 ovarioles in the north (21–22̀ S latitude). The increase in ovariole number with latitude was due to divergence in the ovariole number of the largest females. In contrast, small females in the north and south had the same number of ovarioles. Reproductive allocation of female flies in the northern region was less than females in the southern region. The latitudinal divergence in ovariole number was not associated with habitat differences (density of trees, density of flies and beetles), nor with differences in floral characteristics (flower weight, petal length, yeast species present). Short term weather patterns in daily temperature and rainfall preceding collections pardy explain the variation in ovariole number. These observations in conjunction with preliminary genetic results suggest the cline is associated with genetic differences that interact with environmental determinants such as the temperature during larval development.     

Identification of immature aphids by chromatography

• 1 Immature apterae of five species of cereal aphid were distinguished by ultraviolet examination of chromatograms of their fluorescent pigments.
• 2 Non-expert examiners could identify Metopolophium dirhodum, Sitobion fragariae and Rhopalosiphumpadi with a mean reliability of 98, 83 and 71% respectively; Metopolophium festucae and Sitobion avenae could be separated from the rest but not from each other.
• 3 All species gave consistent chromatograms whether reared on oats or on wheat: some species produced atypical traces if reared on certain grasses.

     

Photoperiodic determination of gynoparae and males of damson-hop aphid Phorodon humuli

Abstract. Sexual morph production in Phorodon humuli is controlled by daylength. Wingless aphids reared from birth in short-day conditions (LD 12 : 12 h) and transferred when adult to long-day conditions (LD 18 : 6 h) produce only gynoparae and males until death some 7 weeks later, whereas those reared in long days and transferred to short days produce 20% wingless parthenogenetic females, 50% gynoparae and 30% of males in an overlapping sequence. No winged morph capable of re-infesting hop is produced. Less mature embryos are more sensitive to short days than older embryos because 100% of the former became gynoparae after 4 days of exposure of their mothers, and 59% when their mothers were exposed for the 4 days immediately preceding the birth of their offspring. Two days of exposure to short days switches 94% of young embryos from wingless to gynopara production when mature. The response to short days is irreversible. Wingless aphids reared from birth to adult in short days produce 30% fewer offspring than those reared to the same stage in long days and are male-biased, with 76–78% of their offspring being male.     

Escaping an ecological dead-end: asexual overwintering and morph determination in the lettuce root aphid Pemphigus bursarius L.

1. Pemphigus bursarius exhibits a host-alternating life-cycle, overwintering as eggs on the primary host plant (poplar) before migrating in summer to the secondary host plant (mainly annual Compositae). A proportion of the population does not produce return migrants (sexuparae) in the autumn but remains in the soil and overwinters as asexual apterae, even after the annual plants have died in early winter. Laboratory and field experiments investigated the biology and overwintering success of these asexual apterae. 2. Both temperature and photoperiod are important in morph determination. The highest number of apterae (approximately 50% of progeny) was produced in a 20 °C, LD 18 : 6 h regime. Very few apterae were produced in an alatae-inducing environment (15 °C, LD 12 : 12 h), but the proportion increased over successive generations. 3. Apterae remaining in the soil in the autumn overwintered successfully in large numbers and were able to reinfest directly the root systems of newly planted lettuce grown in the same field in the following growing season. 4. Overwintered asexual populations also produced alates in July, which were able to colonise other lettuce plants, indicating that they were not sexuparae. Hence, P. bursarius can avoid the ecological dead-end that would occur through local patch extinction. Clones can therefore persist indefinitely as both asexual apterae and alatae without the need to return to poplar and undergo the sexual phase of the life-cycle.     

Alkaloid level in narrow-leafed lupin, Lupinus angustifolius, influences green peach aphid reproductive performance

Two viviparous parthenogenetic clones of the green peach aphid, Myzus persicae (Sulzer), one collected from Rydalmere, New South Wales (NSW), and the other from South Perth (SP), Western Australia, were reared on radish, Raphanus sativus L. cv. Scarlet Globe, under controlled conditions. The NSW clone was fed on simple artificial diets containing alkaloid extracted from narrow-leafed lupin, Lupinus angustifolius L. cv. Fest., and its reproductive performance monitored over 112 h. Forty (40) h into the experiment and thereafter, aphids on the control diet (sucrose solution) produced significantly more offspring (P<0.05) than those on diets containing alkaloid. In a separate experiment, apterae of each clone were caged on three lines (cv. Yorrel, cv. Danja and 84L:441) of narrow-leafed lupin, and allowed to reproduce. The first three offspring were retained, and all developed to 3rd or 4th instar stage. Two nymphs were removed, and the remaining nymph reared through. All three lines produced adults. The number of young produced were counted over 11 days. Fecundity of the SP clone was lower on line 84L:441, but there was no difference in the fecundity of the NSW clone. Phloem exudate and green tissue was concomitantly collected from all lines, and analysed by GC-MS for the alkaloids lupanine and 13-hydroxylupanine. Line 84L:441 contained the highest level of total alkaloids in both phloem and tissue. All experiments indicate that alkaloid level may suppress fecundity of green peach aphids.     

Effect of resource gradient on age and size at maturity and their influence on early‐life fecundity in the predatory Asian lady beetle,Harmonia axyridis

Variation in food availability impacts the performance of insects in terms of their size and age to maturity and fecundity. Age at maturity determines how quickly individuals in a population can start to reproduce and how much they can reproduce. Results from studies on various insect species show that food availability influences the size and fecundity of adult females. It is predicted that under poor growth conditions, variation in size is low, but variation in age at maturity is considerable. This prediction was examined in a widely distributed lady beetle species, Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae), a predator of aphids and coccids. Using a food gradient from low to high aphid prey density, performance of females that were reared on excess food was recorded for pre‐reproductive duration, size at reproductive maturity, number of aphids consumed, and fecundity in the first 10 days of their reproductive period. Results suggested that female H. axyridis that were reared on surplus food when kept at low prey density (poor growth condition) took, on average, three times longer to attain maturity and produced, on average, 14 times fewer eggs than females that were also reared on surplus food, but kept at high prey density (good growth condition). Females performed best at a prey density of 30 aphids per female per 150 cm². Results suggested that the current food availability significantly influenced the age and size of females at maturity and their fecundity. Age and size at maturity of female lady beetles showed non‐linear responses to prey density as well as the occurrence of a minimum size of females, below which H. axyridis females fail to mature. The steep slope recorded at lower prey densities suggests relatively high variation in age at maturity but low variation in size.     

Embryogenesis and oögenesis in alate virginoparae,gynoparae, and oviparae of the aphid Myzus persicae, in relation to photoperiod

The study reports on the effects of prenatal and/or postnatal exposures to short-night or long-night conditions, and of crowding, on embryogenesis and oögenesis in alate virginoparae, gynoparae, and oviparae of a holocyclic strain of the green peach aphid, Myzus persicae, from Yakima, Washington State.In alate virginoparae raised at a density of 10–20 per radish seedling in a short-night regime (8 hr darkness per diem), 3–4 embryos occurred in each of their 10 ovarioles, when the aphids attained adulthood. More than 30 larvae were deposited by most of these alatae. However, in young adult gynoparae, raised at these densities in a long-night regime (15 hr darkness per diem), only one viable embryo (a presumptive ovipara) occurred per ovariole. The follicle containing this embryo was followed by 1–2 abnormal follicles in each ovariole, and the number of larvae deposited by a gynopara was generally less than 10. In young adult oviparae similarly raised under a long-night regime, only one egg typically occurred in each of their 10 ovarioles, and the eggs deposited by an ovipara (only after it had mated) generally numbered less than 10. Alate virginoparae and gynoparae contained an additional embryo in some of their ovarioles when these morphs were raised at a lower density (1–5 per plant).Presumptive gynoparae partially developed the reproductive features of alate virginoparae when transferred to a short-night regime at birth; the converse was true when presumptive alate virginoparae were transferred to a long-night regime early in larval life. Oviparae maintained in short nights from before birth developed the appearance of apterous virginoparae but still produced eggs rather than embryos. However, their oögenesis was enhanced and eggs (10–20) were deposited by them without prior mating. Under all regimes tested, oviparae were always deposited early in the larviposition sequence of their alate mothers, and the number of oviparae deposited never exceeded 15.The possible involvement of juvenile hormone in the regulation of these events and the ecological significance of the results are discussed.     

Predator-induced transgenerational phenotypic plasticity in the cotton aphid

Transgenerational phenotypic changes, whereby offspring have an altered trait or a distinct alternate phenotype, frequently occur in response to increased maternal predation risk. The cotton aphid, Aphis gossypii (Glover), is unique, however, as offspring consist of four distinct phenotypes (normal light green apterae, normal dark green apterae, dwarf yellow apterae, and alatae), all with divergent life history patterns and resulting population dynamics. Here, we show that increased predation risk induces transgenerational phenotypic changes in cotton aphids. When exposed to search tracks from larval or adult convergent ladybird beetles, Hippodamia convergens Guérin-Méneville, cotton aphids produced greater numbers of winged offspring. In a subsequent experiment, apterous and alate individuals on clean plants were found to have primarily normal and dwarf offspring, respectively. We suggest that elevated predation risk may cause phenotypic changes in aphids over multiple generations, resulting in a more precipitous decline in herbivore populations than could be explained solely by increased predation rates.